A new genus and species of Breyeriidae and wings of immature stages from the Upper Carboniferous,...

© Koninklijke Brill NV, Leiden, 2013 DOI 10.1163/1876312X-44032098

Insect Systematics & Evolution 44 (2013) 117–128 brill.com/ise

A new genus and species of Breyeriidae and wings of immature stages from the Upper Carboniferous, Nord-Pas-de-Calais,

France (Insecta: Palaeodictyoptera)

Jakub Prokopa,*, Zuzana Tippeltováa, Patrick Roquesb and André Nelc

aDepartment of Zoology, Faculty of Science, Charles University in Prague Viničná 7, CZ-128 44, Praha 2, Czech Republic

bAllée des Myosotis, F-93330 Neuilly sur Marne, France cCNRS UMR 7205, CP 50, Entomologie, Muséum National d’Histoire Naturelle

45 rue Buffon, F-75005 Paris, France *Corresponding author, e-mail: [email protected]

Published 10 August 2013

AbstractA new palaeodictyopterid, Aviobreyeria gracilis gen.n., sp.n., is described from an outcrop of Upper Carboniferous (Westphalian C/D) deposits at Avion in the Nord-Pas-de-Calais department (France). This new taxon is placed in the family Breyeriidae based on a comparison of the pattern of venation in its forewing venation with that of other breyeriids and possibly related genera within the Breyerioidea. In addition, wings of different immature instars of palaeodictyopterids from the same strata with a similar pattern of venation but differing in having an enlarged RP and a broader anal area (supposed hind wings) were also tentatively attributed to Breyeriidae gen. et sp. indet.. These wings have a roughly identifiable venation and exhibit the gradual development of wings through different ontogenetic stages and their position on the thorax, which is sparsely documented for Paleozoic insects. Discovery of immature and adult stages of palaeodictyopterids in the same layers also indicates they lived in similar habitats.

KeywordsLate Carboniferous; Moscovian; Palaeodictyopteroida; Breyeroidea; gen.n.; sp.n.; taxonomy; larval wing venation

Introduction

Breyeriidae is a small family of the Palaeodictyoptera with a characteristic pattern of wing venation and hind wings markedly broader than fore wings, which occur over a rather short interval from the Bashkirian to Gzhelian in Upper Carboniferous deposits in western and central Europe (Belgium, Czech Republic, England, France, Germany and The Netherlands) and USA (Tennessee) (Carpenter 1992; Brauckmann & Gröning 1996; Brauckmann & Herd 2000). Handlirsch (1906–1908) established this family

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for Breyeria Borre, 1875, Borrea Brongniart, 1893 and Megaptiloides Handlirsch, 1906. These genera are very uniform in terms of their venation, thus it is likely there are fewer valid genera as indicated by several authors (Kukalová 1959; Laurentiaux-Vieira & Laurentiaux 1964; Carpenter 1967). Kukalová (1969) revised and confirmed the synonymy of Borrea, Breyeriopsis Laurentiaux, 1949 and Pseudoborrea Handlirsch, 1919 with the type genus Breyeria. Other genera, such as Stobbsia Handlirsch, 1908, Jugobreyeria Brauckmann et al., 1985, Hasala Brauckmann, 1995; Breyeriopsis Lau-rentiaux, 1949 and Pseudoborrea Handlirsch, 1919, were designated later (Handlirsch 1919; Laurentiaux 1949a; Laurentiaux-Vieira & Laurentiaux 1964; Brauckmann et al. 1985; Brauckmann 1995).

Wings of immature stages of palaeodictyopterans are generally poorly known due to lack of suitable fossils and difficulties with systematic attribution due to the partially developed venation (e.g., Handlirsch 1906–1908; Carpenter 1948; Carpenter & Richardson 1971; Kukalová-Peck & Peck 1976; Sharov & Sinitshenkova 1977; Sinitshenkova 1979). For example, Rochdalia parkeri Woodward, 1913, first described as branchiopod crustacean (Anostraca), was re-described by Rolfe (1967) as a nymph of Palaeodictyoptera and tentatively attributed to the Breyeriidae. Wootton (1972: 666) supported its placement within Palaeodictyoptera but stated that it is impossible to confidently assign Rochdalia to any family of Palaeodictyoptera.

Materials and Methods

Fossil insects were reported in the Upper Carboniferous deposits in the Nord-Pas-de-Calais department in France by Pruvost (1912, 1914, 1919), Laurentiaux (1949a–c), Laurentiaux & Laurentiaux-Vieira (1979, 1980) and Nel et al. (2007). The recently discovered outcrop at Avion in the north of France is dated to Moscovian (Westphalian C/D or equivalent Bolsovian/Asturian). The rare fossil insects were found in ‘Terril N°7’, in rock in the slag heap of a coal mine. The members of the following insect groups were examined: Archaeorthoptera (undescribed specimen MNHN A31824), Paraneoptera (Bruayaphis oudardi Nel et al., 2012) and Thripida (Westphalothripides oudardi Nel et al., 2012) (see Nel et al. 2012a, b). Several new specimens await description.

The fossils were observed under a stereomicroscope Olympus SZX-9 in a dry state and under a film of ethyl alcohol. A line drawing of the pattern of the venation of these specimens was drawn directly with the aid of a camera lucida. Photographs were taken using a digital camera (Canon D550, with macro lenses (MP-E 65 mm and EF 50 mm)) and a Nikon D40 with a Nikon AF-S VR Micro-Nikkor 105 mm macro lens. Original photographs were processed using Adobe Photoshop CS image-editing software.

Type material (Avion No. 1, part and counterpart of a forewing) is housed in the private collection of Patrick Roques (holotype) and a plaster cast is deposited in the Muséum national d’Histoire naturelle (Paris, France) (MNHN). Specimens with imma-ture wings originally in the Patrick Rocques coll., provisionally deposited in the MNHN, but are waiting formal numbering because this collection is still to be catalogued.

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J. Prokop et al. / Insect Systematics & Evolution 44 (2013) 117–128 119

We mainly follow the wing venation nomenclature of Kukalová-Peck (1991). The venational symbols used here are specified as follows: symbols in capitals denote the longitudinal veins (ScP, subcostal posterior; RA/RP, radial anterior/posterior; MA/MP, medial anterior/posterior; CuA/CuP, cubital anterior/posterior; AA, anal anterior).

Systematic and divisions follow the concept of Palaeodictyopteroida sensu Bechly (1996, 2012), Riek (1976) and Sinitshenkova (2002).

Systematic Palaeontology

Superorder Palaeodictyopteroida Bechly, 1996Superfamily Breyerioidea Handlirsch, 1904Family Breyeriidae Handlirsch, 1904

List of genera after Carpenter (1992) supplemented by Brauckmann & Gröning (1996)

Breyeria Borre, 1875Hasala Brauckmann, 1995Jugobreyeria Brauckmann et al., 1985Megaptiloides Handlirsch, 1906Stobbsia Handlirsch, 1908.

Genus Aviobreyeria gen.n.

Type species

Aviobreyeria gracilis sp.n. by present description.

Diagnosis

Costal margin strongly curved in basal part; concave ScP weakly curved reaching RA rather distally at about 5/6 wing length; RA and RP fused basally, separating at about 1/3 wing length from base; RA simple and straight; faint simple cross-veins arranged in rows on apical part of wing; RP distally pectinate with few branches ending on pos-terior margin; stem of M and R very close for a short distance near their bases, M divided into MA and MP distad of separation of RA and RP; MA simple; MP pectinate approximately in the middle of the wing with few branches reaching poste-rior margin of wing; forks of Cu and of M rather distant; CuA simple; CuP distally forked into two main branches; anal area considerably reduced and consisting of a coarse network of cells, first anal vein AA basally close to stem of Cu with two branches ending on posterior margin of wing.

Etymology

Composite name after Avio- (derived from type locality Avion) and Breyeria (type genus); feminine in gender.

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Aviobreyeria gracilis sp.n. (Figs 1, 2)

Description

Fore wing very slender with rather thick membrane probably originally hyaline, wing covered with rather coarse network along posterior margin of wing; costal margin strongly curved in basal part; concave ScP weakly curved, running parallel to RA and reaching it 7.0 mm from the apex of the wing; area between costal margin and ScP very broad and on proximal half of wing bearing numerous cross-veins; distal part of same area not so broad; RA and RP fused basally, separating about 1/3 of the length of the wing from base; RA simple and straight, ending on anterior margin probably at wing apex; RA and RP area with small cells in the middle of the wing and numerous cross-veins on apical part; RP distally pectinate ending in four branches on posterior margin; first branch of RP a long distance (15.4 mm) from base of RP; stems of M and R very close for a short distance near their bases and slightly curved, M divided into MA and MP 1.5 mm distad of separation of RA and RP; MA simple, strongly curved distally, ending on posterior margin of wing about 6.8 mm from wing apex; base of MP about 12.6 mm from wing base; first fork of MP 5.1 mm distal to its base; MP pectinate with four branches reaching posterior margin of wing; MP area rather broad and reaching posterior margin of wing between 14.7 to 7.2 mm from wing apex; stem of Cu divided into CuA and CuP about 7.3 mm from wing base; forks of Cu and of M rather distant, 5.5 mm apart; CuA simple, slightly curved and ending on posterior margin of wing about 23.7 mm from wing base; CuP slightly curved, more or less parallel to CuA and distally forked into two branches; anal area considerably reduced, first anal vein AA basally close to stem of Cu with two branches ending on posterior margin of wing, anal area made up of a coarse network of cross-veins.

Dimensions

Holotype: length of fragment about 39.1 mm, probable total length about 41 mm, width in widest part 11.5 mm (wingspan more than 85 mm).

Fig. 1. Aviobreyeria gracilis gen.n. et sp.n., drawing of holotype Avion No. 1 (Patrick Rocques coll., cast deposited in Museum national d’Histoire naturelle, Paris, France). Scale bar = 5 mm.

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Etymology

Named after the slender shape of the wings.

Type locality

Avion, Pas de Calais, France.

Type strata

Moscovian (Westphalian C or D, equivalent Bolsovian or Asturian, in a rock fragment from the Terrile fosse N°7, Late Carboniferous (Middle Pennsylvanian), Avion, depart-ment of Nord-Pas-de-Calais, France.

Discussion

In spite of the fact that the current state of our knowledge of the higher systematics of the Palaeodictyopteroida is not satisfactory, we consider here additional support for both Riek (1976) and Sinitshenkova (2002), as well as the previously demonstrated limits (see Prokop & Nel 2004). Nevertheless there is no phylogenetic analysis of this group that we can follow.

First, following the key to the ‘Eupalaeodictyoptera’ proposed by Riek (1976: 230) Aviobreyeria gen.n. belongs in the Breyerioidea based on the following characters: ‘CuA simple’; ‘MA simple’, ‘Sc fused with R beyond middle part of wing’, ‘archeodictyon reduced, sometimes to cross veins’, ‘MP branched’ and ‘CuA widely separated from the stem of M’. According to the key of Sinitshenkova (2002: 122), Aviobreyeria can be placed also in the Breyerioidea (Dictyoneurina) with reservation for wing basally wide (not a useful character for fore wings), main veins arch basally and ScP scarcely extends beyond middle of the wing. The latter character is rather speculative, because a number of taxa have a long ScP, which reaches the distal part of RA (e.g., Breyeria bistrata Brauckmann, 1995 or Breyeria vrankeni Laurentiaux-Vieira & Laurentinaux, 1964). Nevertheless, there are similar patterns of fore wing venation in Breyeriidae and Lithomanteidae Handlirsch, 1906, especially Lithomanthis bohemica (Novak, 1880), whose forewing venation is very similar to that of Aviobreyeria in terms of RA, MA, CuA simple and RP, MP, CuP and anal vein with few branches, and the same pattern of branching of the main veins. The difference is that ScP ends at the apex of the wing in Lithomanteidae whereas it is the RA in Aviobreyeria and Breyeria. The other differ-ence between Breyeriidae and Lithomanteidae is the pattern of the cross veins, which are rather faint, numerous and often anastomose with one another in rows in the for-mer, while they are prominent, irregular and mostly un-branched in the latter family (Carpenter 1992; Kukalová-Peck, pers. comm.). Such differences are so weak that the two families may well be synonymized in the future. At this time this synonymy is also difficult to establish because the type specimen of Lithomantis carbonarius (Woodward, 1876) is a very incomplete fossil, hard to compare with other Breyeriidae and Lithomanteidae. Another difference could be in the cubito-anal area of the hind

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wing, which is distinctly broader in Breyeriidae than Lithomanteidae, at least in the taxa in which it is known. It is difficult to separate palaeopteran families based on this difference. For example, there are species with very narrow and others with very broad hind wings among Recent Odonata Libellulidae (Ris 1909–1913).

Within Breyerioidea (Breyeriidae, Stobbsiidae and Cryptoveniidae), Stobbsia Handlirsch, 1908 is based on the type species S. woodwardiana Handlirsch, 1908, which was subsequently placed in the family Stobbsiidae because the ScP terminates at the costa and branches of M and Cu do not strongly curve towards the posterior mar-gin of the wing (Lameere 1917; Laurentiaux & Laurentiaux-Vieira 1952; Kukalová 1969). It also differs from Aviobreyeria in that the numerous branches of RP are not pectinate.

Cryptovenia Bolton, 1912 (type species C. moyseyi Bolton, 1912) is a poorly known taxon based only on a wing fragment. Lameere (1917) and Handlirsch (1919) list it but Carpenter (1992) considers it as incertae sedis.

Breyeria Borre, 1875, with 13 species, i.e., B. (= Pachytylopsis) borinensis (Borre, 1875), B. barborae Kukalová, 1959, B. lachlani (Brongniart, 1893), B. boulei (Meunier, 1910), B. bistrata Brauckmann, 1995, B. harlemensis Brauckmann & Gröning, 1996, B. rappi Carpenter, 1967, B. solida Brauckmann & Herd, 2000, B. brevis Brauckmann & Herd, 2002, B. britannica Laurentiaux & Laurentiaux-Vieira, 1952, B. stopai (Laurentiaux & Laurentiaux-Vieira, 1952) and B. delruei (Laurentiaux, 1949), has the following characters: division of MA and MP nearly at the same level as that of RA and RP, RP with 5–9 branches; anal area well developed without coarse network of cells, unlike Aviobreyeria (Laurentiaux-Vieira & Laurentiaux 1963). B. brevis has five branches of RP but its ScP is not fused with RA (Brauckmann & Herd 2002). B. brauckmanni Pinto, 1992, which is based on an apical wing fragment, differs in that RP has five main branches the first branch of which is deeply bifurcated, unlike in Aviobreyeria (Pinto, 1992). Nevertheless, the attribution of B. brauckmanni to the genus Breyeria and even Breyeriidae is uncertain due to poor state of preservation. This fragment of the apex of a wing is similar to that of other families, such as, e.g., Lithomanteidae, as it has a similar pattern of venation (see matrix of characters in Prokop & Ren 2007: 270). Furthermore, the termination of ScP at RA on the apex of the wing seems to be unusual or an artifact and the detailed examination of the original figure in Pinto (1992: fig. 2) indicates there a dense pattern consisting of a network of irregular shaped small cells (archedictyon). Therefore, we propose to consider this taxon as Palaeodictyoptera family incertae sedis.

Jugobreyeria Brauckmann et al., 1985 is based on the type-species Jugobreyeria sippelorum Brauckmann, Koch & Kemper, 1985. Like Aviobreyeria it has rather elongated wings. It differs from Aviobreyeria in having a rather short ScP, RP not pectinate but with numerous secondary branches and RA & RP divided at the same level as MA and MP.

Hasala Brauckmann, 1995 is a monotypic genus based on a hind wing (Hasala inferiorsaxonica Brauckmann, 1995). Nevertheless it shares with Aviobreyeria a rather long ScP, but mainly differs from it in having a strongly curved median vein, which is nearly perpendicular to the posterior margin of the wing, and an RP with 10 branches.

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The description of Stephanian Megaptiloides (M. brodiei (Brongniart, 1893)) is based on the mid part of a hind wing. Its RP branches the same number of times as in Aviobreyeria but its ScP reaches RA at a more basal position, the first branch of RP.

Family Breyeriidae Handlirsch, 1904

Gen. et sp. indet. (Figs 3–6)

Figs 2–6. (2) Aviobreyeria gracilis gen.n. et sp.n., photograph of holotype No. 1 (Patrick Rocques coll., cast deposited in Museum national d’Histoire naturelle, Paris, France); (3–6) Breyeriidae gen. et sp. indet. (3, 4) imprint and counter imprint of a wing of an immature individual including fragments of meso- and metathoracic segments specimen No. 2 (Patrick Rocques coll., provisionally deposited in the Museum national d’Histoire naturelle, Paris, France); (5) imprint of thoracic segments and hind wing of immature specimen No. 3 (Patrick Rocques coll., provisionally deposited in the Museum national d’Histoire naturelle, Paris, France); (6) imprint of hind wing of immature specimen No. 4 (Patrick Rocques coll., provisionally deposited in the Museum national d’Histoire naturelle, Paris, France). Scale bars = 5 mm. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/conten/1876312x.

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Description

There are numerous tubercles on the surface and a median keel partly preserved on frag-ments of strongly sclerotized but incomplete proto-, meso- and meta thoracic tergites.

Specimen Nos 2, 3 (Figs 3–5): Hind wing slender rather elongated, strongly curved and probably originally with a hyaline membrane, cross-veins not well developed and creased; costal margin strongly curved in basal part; concave ScP straight, parallel to RA reaching it very close to apex of wing; costal area slightly narrows close to wing base, very broad in proximal third of wing, distal part of same area not so broad; RA and RP basally close together, separating at about 1/3 wing length from base; convex RA sim-ple, running straight to wing apex; RP distally pectinate with six discernible branches ending at posterior margin; first branch of RP a long distance (4.1 mm, No. 2) from base of RP; M divided into MA and MP distad of separation of RA and RP; convex MA simple, strongly curved distally, ending at posterior margin of wing; division of MA and MP about same level or slightly distad as RA and RP from wing base; MP pectinate with three or more faint branches reaching posterior margin of wing (Fig. 3); stem of Cu distinctly closer to wing base than division RA and RP; simple CuA slightly curved distally; concave CuP slightly curved, more or less parallel to CuA and splits into two main branches; anal area rather broad and formed by four faint branches.

Specimen No. 4 (Fig. 6): Wing of triangular shape, moderately curved with proba-bly originally a hyaline membrane, cross-veins not developed; costal margin broad and curved; wing venation partly discernible; convex RA ending at apex of wing, concave RP pectinate and ends in five branches; division of RA and RP at the same level as MA and MP; convex MA simple; concave MP ends in several branches; stem of Cu diverg-ing basally to division of CuA and CuP; convex CuA simple; concave CuP forks dis-tally and the two long branches end at the posterior margin of the wing; three or four anal veins recognizable.

Dimensions

Specimen No.2: length of hind wing 17.6 mm, width at widest part 7.0 mm; Specimen No.3: length of hind wing 30.5 mm, width at widest part 9.2 mm; Specimen No.4: length of hind wing 16.0 mm, width at widest part 6.7 mm.

Locality and layer

Avion, Pas de Calais, France; Moscovian (Westphalian C or D, equivalent Bolsovian or Asturian), in a rock fragment from the Terril of Fosse N° 7’, Late Carboniferous (Middle Pennsylvanian), Avion, department of Nord-Pas-de-Calais, France.

Discussion

Hind wings of various immature instars have been preserved separately and are attached to thoracic segments with a partially developed articulation. As the three

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immature wings preserved have very different shapes and sizes, it is likely they corre-spond to hind wings of different instars and the wide cubito-anal area indicates they are hind wings. These immature wings exhibit clear tracheation that roughly corre-sponds to the main part of the future pattern of veins on the wings of the imago described above. In addition, if they belong to the same species than they probably represent wings of different ontogenetic stages because they differ in size and angle with the thorax as follows: (1) Specimen No. 4 (Fig. 6): young larval instar with a rather sub-oval shaped wing and hardly recognizable corrugation of main veins, indis-tinguishable articulation and are positioned at an acute angle with the thorax; (2) Specimen No. 2 (Figs 3, 4): older larval instar with triangular shaped wing, broad costal area, more developed corrugation of main veins, articulation clearly visible and less acutely positioned relative to the thorax; (3) Specimen No. 3 (Fig. 5): older larva or subimaginal instar with narrow costal area, well developed corrugation of main veins, well developed articulation with discernible plates and held perpendicular to the thorax.

Our comparison with previously described wings of immature individuals of the palaeodictyopterids Rochdalia parkeri Woodward, 1913 and Idoptilus oniscoformis Wootton, 1972 revealed they have a similar pattern of main veins consisting of simple convex veins RA, MA and CuA alternating with concave branched RP, MP and CuP (Wootton 1972). However, this pattern is present in several palaeodictyopteran fami-lies, e.g., Breyeriidae, Lithomanteidae and Lycocercidae, which rules out a precise fam-ily assignment. Nevertheless this comparison supports our assumption that the wings are those of older instars because the angle of the wings with the thorax is less acute and its articulation with the thorax in specimen Nos 2, 3 is well developed (Kukalová-Peck 1983, 1991).

Conclusions

Aviobreyeria gen.n. is established for Aviobreyeria gracilis sp.n. (type species), which has the following unique combination of fore wing characters: ScP weakly curved ending in RA, stem of M divided into MA and MP distad of separation of RA and RP, MA and CuA simple, MP pectinate with few branches, anal area considerably reduced and made up of a coarse network of cells.

Three wings of immature palaeodictyopterids from the same strata are tentatively attributed to Breyeriidae gen. et sp. indet. based on the presence of a similar pattern of venation to that of Aviobreyeria but differing mainly in that the RP is enlarged and has six branches instead of four and a broader cubito-anal area. These differences are simi-lar to the well known differences between fore wing and hind wing venation in this group. However, the present wings of immature individuals have a venation that is roughly identifiable and possibly reflects the gradual development through different ontogenetic stages and their position on the thorax, which is sparsely documented for Paleozoic insects. Moreover, the discovery of immature and adult stages of palaeodic-tyopterids in the same layers indicates they possibly lived in similar habitats.

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Acknowledgements

We are grateful to Carsten Brauckmann (Clausthal University of Technology, Clausthal-Zellerfeld, Germany) for his valuable comments on the manuscript and Anthony F.G. Dixon, University of East Anglia (Norwich, UK) for improvements of the English text. The first author acknowledges the research support from the Grant Agency of the Czech Republic No. P210/10/0633.

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A new genus and species of Breyeriidae and wings of immature stages from the Upper Carboniferous,...

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