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Late Palaeozoic Paoliida is the sister group ofDictyoptera (Insecta: Neoptera)Jakub Prokopa, Wieslaw Krzeminskib, Ewa Krzeminskab, Thomas Hörnschemeyerc, Jan-Michael Ilgerd, Carsten Brauckmannd, Philippe Grandcolase & André Nelea Charles University in Prague, Faculty of Science, Department of Zoology, Czech Republic,Prahab Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków,Polandc Johann-Friedrich-Blumenbach-Institut für Zoologie & Anthropologie, Georg-August-Universität Göttingen, Göttingen, Germanyd Clausthal University of Technology, Institute of Geology and Palaeontology, Clausthal-Zellerfeld, Germanye CNRS UMR 7205, CP 50, Entomologie, Muséum National d’Histoire Naturelle, Paris, FrancePublished online: 14 Oct 2013.

To cite this article: Jakub Prokop, Wieslaw Krzeminski, Ewa Krzeminska, Thomas Hörnschemeyer, Jan-Michael Ilger, CarstenBrauckmann, Philippe Grandcolas & André Nel , Journal of Systematic Palaeontology (2013): Late Palaeozoic Paoliida is thesister group of Dictyoptera (Insecta: Neoptera), Journal of Systematic Palaeontology

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Journal of Systematic Palaeontology, 2013http://dx.doi.org/10.1080/14772019.2013.823468

Late Palaeozoic Paoliida is the sister group of Dictyoptera (Insecta: Neoptera)Jakub Prokopa∗, Wieslaw Krzeminskib, Ewa Krzeminskab, Thomas Hornschemeyerc, Jan-Michael Ilgerd,

Carsten Brauckmannd, Philippe Grandcolase and Andre Nele

aCharles University in Prague, Faculty of Science, Department of Zoology, Praha, Czech Republic; bInstitute of Systematics andEvolution of Animals, Polish Academy of Sciences, Krakow, Poland; cJohann-Friedrich-Blumenbach-Institut fur Zoologie &

Anthropologie, Georg-August-Universitat Gottingen, Gottingen, Germany; dClausthal University of Technology, Institute of Geologyand Palaeontology, Clausthal-Zellerfeld, Germany; eCNRS UMR 7205, CP 50, Entomologie, Museum National d’Histoire Naturelle,

Paris, France

(Received 12 September 2012; accepted 24 March 2013)

Paoliida is an insect group of highly controversial composition and equally controversial affinities. Based on comprehensivereinvestigations we propose a new delimitation of the insect order Paoliida sensu nov. associating the families Paoliidae andBlattinopsidae on the basis of the following main wing venation characters: veins CuA convex and CuP concave separatingfrom a rather long basal stem Cu; a short, more or less distinct, but generally convex arculus brace (crossvein) between Mand CuA (more distinct in forewing than in hind wing), and a broad area containing veinlets, between CuP and CuA, witha general course of CuA making a double curve (autapomorphy); CuP straight or sigmoidal. The Paoliida is consideredas a neopteran clade and potential sister group of the Dictyoptera on the basis of the presence of well-defined anteriorbranches of CuA, with the same convexity as the median vein and more concave than the posterior branches of the samevein (synapomorphy). The polarities of the other characters shared by these clades are discussed. New diagnoses of the orderPaoliida and the family Paoliidae are provided after re-examination of the type material. We attribute Herbstiala herbsti toPaoliidae and consider Herbstialidae as a junior synonym of this family. The grylloblattid family Ideliidae is consideredas a junior synonym of Paoliidae, transferring genera Stenaropodites, Aenigmidelia, Archidelia, Sojanidelia, Micaidelia,Acropermula and Mongoloidelia to this family. Kochopteron hoffmannorum and Protoblattina bouvieri are newly includedin Paoliidae. Protoblattinopsis stubblefieldi is reinterpreted as a hind wing having highly specialized cubito-anal structuresfunctionally analogous to the anal loop structure of the hind wing of the Mesozoic Isophlebioidea (Odonatoptera). Weattribute Protoblattinopsis to Paoliida and consider Protoblattinidae as a junior synonym of Paoliidae. Furthermore, a newPaoliidae, Silesiapteron jarmilae gen. et sp. nov. is described from Upper Carboniferous sphaerosiderite concretion of Poland.

http://zoobank.org/urn:lsid:zoobank.org:pub:296A17BD-2A84-4C1F-B19E-737DDFE7E17C

Keywords: Insecta; Dictyoptera; Paoliidae; Blattinopsidae; systematics; Late Carboniferous

Introduction

The Palaeozoic insect group Paoliida Handlirsch, 1906a(= Protoptera Sharov, 1966) is of crucial interest forthe phylogeny of Pterygota given that it is currentlyconsidered: as the most inclusive group of Pterygota(Sharov 1966; Rasnitsyn 1980, 2002a); as the earliestneopteran insects (Grimaldi & Engel 2005); as a subgroupof the ‘Scarabaeones Laicharting, 1781’ (sensu Rasnitsyn1980); as a group comprising all winged insects except thePolyneoptera Martynov, 1938 (= Gryllones Laicharting,1781); or, together with the Blattinopsidae (Kukalova-Peck& Brauckmann 1992), as an ‘ancestor’ group of a part ofthe Hemiptera (‘paoliid line’). They were also consideredas palaeodictyopteran insects (Handlirsch 1906a, b, 1919,1937), as Protoblattoidea Handlirsch, 1906a (at least for theBlattinopsidae, i.e. “insects with wings intermediate in type

∗Corresponding author. Email: jprokop@natur.cuni.cz

between those of the Palaeodictyoptera and Blattoidea”;Bolton 1925, p. 18), or as a ‘blattoid-like’ protorthopterangroup (Hennig 1981; Carpenter 1992).

The phylogenetic relationships of the Paoliida remaincontroversial due in great part to divergent opin-ions about the main subdivisions of the Pterygota(Palaeoptera–Neoptera versus Scarabaeones–Gryllones).The diagnoses and apomorphies of the Paoliida and theincluded families greatly vary by author. For example,Kukalova-Peck & Brauckmann (1992) considered thisgroup as comprising the Paoliidae, Eucaenidae, Strepho-cladidae, Blattinopsidae, Synomaloptilidae and Cymbop-sidae, whilst Rasnitsyn (2002a) restricted them solely toPaoliidae.

The historical background of research and systematicsof Paoliidae was summarized by Prokop et al. (2012).After several recent studies of new material and revision

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of previously described taxa that are considered to bemore or less related to Paoliidae (Brauckmann 1984, 2007;Brauckmann et al. 1985, 2003; Bethoux & Nel 2002;Prokop & Nel 2007; Quetscher & Ilger 2007; Ilger &Brauckmann 2008), it is now possible to propose a redefini-tion of Paoliida, of Paoliidae, and of their possible affinities,based on a reinterpretation of the body structures and wingvenation. This study has important consequences for theunderstanding of the evolution of Pterygota and especiallypolyneopteran insects, by clarifying the affinities of severalfamilies currently considered as ‘Grylloblattodea’. This lastgroup artificially combines many fossils with recent taxa(Jarvis & Whiting 2006). It is suspected to be para- or poly-phyletic, and it is not supported by any known apomorphy(Storozhenko 2002, p. 279; Grimaldi & Engel 2005).

Material and methods

The fossil specimens were observed using OlympusSZX-9 and Nikon SMZ 645 stereomicroscopes in a drystate and also under a film layer of ethyl alcohol. Thevenation patterns were drawn using a stereomicroscopewith a camera lucida and finally overlain on the photo-graph. Photographs were taken with digital cameras NikonD80 equipped with macro lens Nikon AF-S VR Micro-Nikkor 105 mm, and Canon D550 equipped with macrolens MP-E 65mm or EF-S 60 mm. Original photographswere processed using the image-editing software AdobePhotoshop 8.0.

The wing venation nomenclature generally followsthe conception of Kukalova-Peck (1991). Paoliid taxon-omy follows Carpenter (1992), partially Kukalova-Peck &Brauckmann (1992), and the recent update by Prokop &Nel (2007).

Institutional abbreviationsSpecimens were examined from the following institu-tional collections: NHML: The Natural History Museum,London, UK; BU: Lapworth Museum of Zoology, Univer-sity of Birmingham, UK; FM: The Field Museum,Chicago, USA; GD NRW: Geologischer Dienst NRW,Krefeld, Germany; IRSNB: Institut Royal des SciencesNaturelles de Belgique, Brussels, Belgium; ISEZ PAN:Natural History Museum of the Institute of Systemat-ics and Evolution of Animals PAS, Krakow, Poland;MMO: Municipal Museum of Ostrava, Ostrava, CzechRepublic; MNHN: Palaeontology Department, Museumnational d’Histoire naturelle, Paris, France; NNMN:Nationaal Natuurhistorisch Museum Naturalis, Leiden, TheNetherlands; PIN: Paleontological Institute of RussianAcademy of Sciences, Moscow, Russia; PrF UK – Paleo:collection of Institute of Geology and Palaeontology,Charles University, Praha, Czech Republic; WMf.N: LWL-

Museum fur Naturkunde, Westfalisches Landesmuseummit Planetarium, Munster/Westfalen, Germany.

Anatomical abbreviationsWing venation abbreviations: ScP, subcosta poste-rior; RA/RP, radius anterior/posterior; R, radius; M,median vein; CuA/CuP, cubitus anterior/posterior; A1/A2,first/second anal vein (see Fig. 1A, B). Reticulate vena-tion or a dense pattern of crossveins, the so-called ‘arche-dictyon’ is considered sensu Wootton & Kukalova-Peck(2000).

Systematic palaeontology

Paoliida Handlirsch, 1906a (= Protoptera Sharov, 1966)

Included families. Paoliidae Handlirsch, 1906a (IdeliidaeZalessky, 1928 junior synonym, Herbstialidae Schmidt,1953 junior synonym), Blattinopsidae Bolton, 1925.

Remarks. Rohdendorf (1977) erected this order butattributed it to Handlirsch (1906a) who defined the familyPaoliidae. If we followed Dubois (2006) who recommendedusing different rules for taxa of ‘class series’ and ‘familyseries’, it would be necessary to consider that the authorof Paoliida is Rohdendorf. Thus, the anteriority rule wouldfavour the name Protoptera Sharov, 1966. Nevertheless, asthe commonly used name is Paoliida, we prefer to extendthe rules of the ‘family series’ and consider that the authorof Paoliida is the same as that of the family Paoliidae, thusHandlirsch.

Diagnosis. For discussion, see below (Figs 1A, B, 2C–G).Fore and hind wings nearly homonomous, hind wingbasally broader than forewing and subtriangular in shape;crossveins simple or reticulate often forming a net of irreg-ular cells (archedictyon); concave ScP ending in C or RAin distal half of wing; strongly convex RA simple or withweak, short but distinct anterior branches, unusually promi-nent steep elevation from ScP to RA forming a charac-teristic wing profile; costal area between ScP and costalmargin with series of rather irregular, simple or branch-ing, sigmoidal or oblique veinlets; RP rather concave withnumerous branches; M rather concave; convex CuA andconcave CuP separating from a rather long basal stemCu; CuA is not in contact with the stem R + M; a short,more or less distinct, but generally relatively convex arcu-lus brace (crossvein) between M and CuA (more distinct inforewing than in hind wing) (Fig. 1A, C); area between CuPand CuA broad compared to the median and radial areas(autapomorphy), a general course of CuA making a doublecurve (autapomorphy); CuA has convex posterior branches,more or less developed (depending on the species), plussome weaker concave anterior branches (synapomorphywith Dictyoptera, see below); CuP simple or with a short

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Systematic position of Paoliida (Insecta: Neoptera) 3

Figure 1. Wing venation of Paoliidae and Blattinopsidae showing crossvein m-cua, and separation of convex CuA and concave CuP fromneutral common stem Cu (A–C) and roof-like position of wings at rest (D–E). A, Zdenekia silesiensis Prokop et al., 2012 (Paoliidae),paratype ISEZ PAN No. MP ISEA I-F/MP/1540/25/09 (negative imprint). B, Blattinopsis ornata Bolton, 1925 (Blattinopsidae) withenlarged distal part of CuA, holotype NHML No. P7291. C, Paoliidae gen. et sp. indet., ISEZ PAN No. MP ISEA I-F/MP/1492/342/09.D, Kemperala hagenensis Brauckmann, 1984 (Paoliidae), specimen WMf.N No. P.20500. E, Blattinopsis fimbriata Bolton, 1925 (Blat-tinopsidae), holotype NHML No. P7284. For abbreviations see text. Arrows indicate position of arculus, scale bars represent 10 mm.

terminal twig, straight or more or less sigmoidal, notposteriorly curved; anal fan strongly reduced on both wingpairs, archedictyon usually present; wings at rest in a roof-like position (Fig. 1D, E).

Family Paoliidae Handlirsch, 1906a

Type genus. Paolia Smith, 1871.

Included taxa. Paolia vetusta Smith, 1871, type species(USA, Indiana, The Netherlands, South Limbourg); othertaxa: Darekia sanguinea Prokop et al., 2012 (Poland, UpperSilesian Coal Basin); Holasicia vetula Kukalova, 1958a(Czech Republic, Upper Silesian Coal Basin); Holasiciarasnitsyni Brauckmann, 1984 (Germany, Hagen-Vorhalle);Kemperala hagenensis Brauckmann, 1984 (Germany,

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Figure 2. Paoliid prothoracic paranotal extensions (A–B) and pattern of fore- and hind wing venation (C–G). A, Kochopteron hoffman-norum Brauckmann, 1984, specimen WMf.N No. P.26093 (arrow indicates possible limit of the prothoracic extensions). B, Protoblattinabouvieri Meunier, 1909, holotype MNHN No. DP R52936 (Arrows indicate position of arculus). C, Paolia gurleyi (Scudder, 1885),holotype FM No. PE 6393. D, Zdenekia grandis Kukalova, 1958a, holotype MMO No. B1006. E, Zdenekia silesiensis Prokop et al., 2012,holotype ISEZ PAN No. MP ISEA I-F/MP/1488/2ab/08. F, Zdenekia cf. grandis Kukalova, 1958b specimen MMO No. B1009. G, Paoliacf. vetusta Smith, 1871, specimen NNMN No. B1002, scale bars represent 5 mm.

Hagen-Vorhalle); Kochopteron hoffmannorum Brauck-mann, 1984 (Germany, Hagen-Vorhalle); Mertovia sustai(Kukalova, 1958a) (Czech Republic, Upper Silesian CoalBasin); Olinka modica Kukalova, 1958a (Czech Repub-lic, Upper Silesian Coal Basin); Paoliola gurleyi (Scudder,1885) (USA, Indiana); Protoblattina bouvieri Meunier,1909 (see below; France, Commentry); Pseudofouquea

sp. (England, Bickershaw; Anderson et al. 1997); Pseud-ofouquea cambrensis (Allen, 1901) (Wales, LlanbradachColliery); Silesiapteron jarmilae gen. et sp. nov. (Poland,Upper Silesian Coal Basin); Sustaia impar Kukalova, 1958a(Czech Republic, Upper Silesian Coal Basin); Zdenekiagrandis Kukalova, 1958a (Czech Republic, Upper SilesianCoal Basin); Zdenekia occidentalis Laurentiaux-Vieira &

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Laurentiaux, 1986 (Belgium, Charbonnages de Ressaix);Zdenekia silesiensis Prokop et al., 2012 (Poland, UpperSilesian Coal Basin).

The ‘grylloblattid’ family Ideliidae Zalessky, 1928is herein considered as a junior synonym of Paoli-idae, with the following genera transferred to Paoli-idae: Acropermula Aristov & Storozhenko, 2011 (CzechRepublic, Obora, Permian), Aenigmidelia Sharov, 1961(Russia, Kaltan, Permian), Archidelia Sharov, 1961(Russia, Kaltan, Permian), Micaidelia Aristov, 2004(Russia, Chekarda, Sylva River, Permian), MongoloideliaStorozhenko, 1992 (Mongolia, Bor-Tologoi, Permian),Sojanidelia Storozhenko, 1992 (Russia, Chekarda, SylvaRiver, Permian), Stenaropodites Martynov, 1928 (Russia,Tikhie Gory, Permian), Sylvidelia Martynov, 1940 (Russia,Chekarda, Sylva River, Permian). Herbstialidae Schmidt,1953 (Herbstiala herbsti Schmidt, 1953, Langsettian,Germany) is also considered as a junior synonym ofPaoliidae.

Previous diagnoses and paoliid venation. Carpenter(1992) proposed a diagnosis based on wing venation alonewhile some body characters are known. Rasnitsyn (2002a,p. 83) indicated that Paoliidae have no known apomorphiesand supplemented the diagnosis as follows: “Large insectswith long but otherwise apparently unmodified legs andlong (longer than body), filiform antennae. Pronotum lack-ing wide paranota, other body structures unknown. Wingresting position low roof-like. Wings moderately wide, withfore margin weakly convex or nearly straight, and withapex often rather narrow. Costal space moderately narrow,with series of rather irregular, simple or branching, obliqueveinlets. Main veins are weakly concave except for convexR [herein RA], M5 [herein brace between M and CuA](+ CuA) and A in forewing, only R in hind wing, anddistinctly concave SC [herein ScP] and, in forewing, CuP.SC meeting R near apical quarter of wing. SC, R and oftenalso CuP and anal veins lacking long branches. RS, Mand CuA irregularly branching, with M and/or CuA havingprevalence over RS [herein RP]; forewing CuA often withseries of rather uniform hind branches in its middle part. RSstarting near basal quarter, sometimes third of wing length.M5 forming short, oblique, strong crossvein between Mand CuA (not very distinct in Pseudofouquea Handlirsch).Crossveins simple or, mainly, abundantly branching andoften forming coarse archedictyon. Hind wing subtriangu-lar, with anal area more or less widened, not tucking downin folded wing.”

We discuss below several points of this diagnosis plussome supplementary characters of interest for Paoliidae:

1. Wings at rest position low roof-like (Fig. 1D, E).Such positions of the wings are identical to whatoccurs in Neoptera. The large body size of Paoliidaecan have the consequence of the wings being disposedmore or less in lateral positions relative to the body. It

is not possible to infer a more precise condition exceptfor a support of an attribution to Neoptera (if ‘foldedwings backward at rest’ is considered as more derivedcondition) apomorphy (Martynov 1925; Grimaldi &Engel 2005).

2. Filiform antennae (Fig. 1D).In fact, the paoliids have filiform, linear and slenderantennae with long antennomeres. This character iswidespread, especially in Polyneoptera. So it is notvery useful for a determination of the affinities ofPaoliidae.

3. Pronotum lacking wide paranota (Fig. 2A, B).This character is present in many palaeopteran orneopteran clades. Furthermore, the homology andpolarity of this structure among the different ptery-gote groups remains unstudied. Paoliidae certainlydid not have wide paranota, and it seems that theyhad small ones, after the herein proposed attribu-tion of Kochopteron Brauckmann, 1984 to this family(see below). Ilger & Brauckmann (2008) indicatedthat this taxon has small wing-like elongations, so-called ‘winglets’. After our re-examination of speci-men WMf.N P26093/4 we consider that these struc-tures could probably be derived from the pronotum assmall prothoracic elongations (Fig. 2A). The state ofpreservation of this fossil does not allow us to see moredetails or exact delimitation. Other paoliid specimens,either are isolated wings or body structures, seem tobe too poorly preserved to determine if they had thesestructures.

4. Termination of ScP on radial vein (Fig. 2C, D).Maples (1989) already noted that in Paolia vetustaSmith, 1871, ScP ends on the costal margin and noton R (Fig. 5C). The same occurs for Mertovia sustai(Kukalova, 1958a), while it ends on R in Zdenekiagrandis Kukalova, 1958 (Kukalova 1958a). Thus, thesituation is variable within the family and the character‘ScP meeting RA’ cannot be used in the family diag-nosis. Nevertheless, ScP ends near the apical quarterof the wing in Paoliidae.

5. Presence of a branch ‘M5’ versus ‘arculus’ brace(Figs 1A–C, 2E).Rasnitsyn (2002a) did not follow the hypothesis ofKukalova-Peck & Brauckmann (1992) regarding the‘arculus’. Instead of considering it as a special-ized crossvein, he assumed that it is a branch ofthe Media, namely ‘M5’. Rasnitsyn’s hypothesis isunlikely because: (1) this vein is convex (in some taxaas convex as CuA itself, see Olinka modica Kukalova,1958a), while the median vein is concave in Paoliidae(at least more concave than CuA); in the general insectwing pattern, the convex branch of a main vein is theanterior one, not in a position posterior to a concavebranch; and (2) in some Paoliidae (Paolia cf. vetustaSmith, 1871; see Fig. 2G), this arculus is stronglyreduced, suggesting that it is not a posterior branch

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of the Media but simply a more or less strengthenedcrossvein. Similar reinforced crossveins, looking likeposterior branches of median vein, occur in some otherinsect groups, one of the most typical being the poste-rior side of the arculus in Odonata (Nel et al. 1993).

6. Median vein (Fig. 1A).In Paoliida the median vein is more concave than CuA.Nevertheless, it forks into an anterior branch and aposterior one rather near to its base. But as the anteriorbranch is not distinctly more convex than the posteriorone, there is no clear evidence of the presence of areally convex MA that would be independent froma (relatively) concave MP or that would appear as aposterior ‘branch’ of RP. Therefore we choose to namethe median vein ‘M’.

7. Convex versus concave branches of CuA in Paoliidae(Figs 1A, 2E).The CuA has convex posterior branches that are moreor less developed (depending on the species), plussome (generally weaker) concave anterior branches(more precisely, branches with the same convexityas the branches of M). The same situation occursin Dictyoptera, as figured by Rehn (1951, pls 1–3)for several modern Blattodea (i.e. Polyphaga aegyp-tiaca (L., 1758), Periplaneta americana, Calolam-pra irrorala (Fabricius, 1775), etc.) (pers. obs.), andnoted by Bethoux & Wieland (2009) and Bethouxet al. (2009, 2010, p. 750, fig. 4) who hypothe-sized that “provided that the concave oblique veinis CuA, the convex anterior branches diverging fromit likely belong to M”. These veins could corre-spond to branches of M that are translocated onCuA (for definition of translocation see Bethoux2007, 2009; Bethoux & Wieland 2009). Bethouxet al. (2009, fig. 5) proposed two models of homol-ogization of the short ‘archimylacridaean arculus’(= Rasnitsyn’s ‘M5’ vein) between M and CuA,i.e. a crossvein versus a posterior branch of M thatfuses with CuA and re-emerges distally into theconcave anterior branches of CuA. The argumentsabove rather support the crossvein hypothesis, butdo not explain the presence of concave branchesof CuA. Nevertheless, the presence of this convex‘arculus’ brace plus the presence of concave ante-rior branches of CuA in Paoliidae and Dictyoptera(but also in Blattinopsidae, see Fig. 1B), supportsthe hypothesis that these structures are homologousamong these groups. Bethoux et al. (2009, p. 149)already discussed this point but did not reach a defini-tive conclusion on the exact origin of these veins.Bethoux et al. (2011) confirmed the presence of thecapture of a branch of concave M by convex CuAin the Palaeozoic blattid Phyloblatta gaudryi (Agnus,1903), through a ‘reticulated fusion’ between theseveins.

8. Cubital veins and cubital area.The taxa that are currently included in the ‘orderEoblattida’ (Eoblattidae Handlirsch, 1906b, typegenus Eoblatta Handlirsch, 1906b, but also the generaStenoneura Brongniart, 1893, Ctenoptilus Lameere,1917, Nectoptilus Bethoux, 2005, Lobeatta Bethoux,2005, Sinopteron Prokop & Ren, 2007, etc.) aresupposed to have a pattern of cubital veins superfi-cially similar to Paoliidae in the presence of a ratherbroad area containing veinlets, between main cubitalveins, with a general course of CuA making a doublecurve (Rasnitsyn & Aristov 2010). But the structuresconcerned are not homologous in the two groups.While the ‘Eoblattida’ have a CuP forking into twobranches, the anterior one reaches the convex CuAthat emerges from a common stem with (R +) M(typical pattern of archaeorthopteran wing venationsensu Bethoux & Nel 2002; see also Bethoux & Nel2005; Bethoux 2005; Prokop & Ren 2007). On thecontrary the Paoliidae clearly have a convex CuAand a concave CuP separating from a rather longcommon stem Cu. CuA is not in contact with thetwo strongly approximate veins R and M (see Prokopet al. 2012). As the concerned veins are different (viz.CuA + CuPa/CuPb, versus CuA/CuP, it is not possi-ble to homologize the paoliid broad area between CuAand CuP with the ‘eoblattid’ broad area between thebranches of CuP (in basal part), and CuA and CuP inthe distal part. The difference between paoliids and‘eoblattids’ can be detected only in fossils having apreserved convexity/concavity of the concerned veins.This similarity between the two groups is probably dueto similar requirements with mechanical constraintslinked to flight.

9. Shape of hind wing anal area (Fig. 2C, F, G).Paoliidae are supposed to have fore and hind wingsnearly homonomous, even if the hind wing is basallybroader than the forewing, and subtriangular in shape.However, the corresponding structure is poorly docu-mented in these insects. The known hind wings haveat most the first branches of AA (1A) preserved(Kukalova 1958a, figs 2, 8, 10, 16, 20; Prokop & Nel2007, figs 4–7; Prokop et al. 2012). Some of themshow a hind wing distinctly broader in its basal thirdthan the forewing (Zdenekia Kukalova, 1958a, SustaiaKukalova, 1958a), while others (Paolia Smith, 1871)had a rather narrow hind wing (Fig. 5A–C). The analarea seems to have been variable in shape and sizeamong Paoliidae. The hind wing anal area is also verypoorly known in the Blattinopsidae (Hornschemeyer& Stapf 2001). The majority of the taxa previ-ously included in the ‘grylloblattid’ family IdeliidaeZalessky, 1928 have a forewing venation of paoliidtype, and thus we transfer them to Paoliidae, seebelow. The hind wing is generally unknown in these

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Systematic position of Paoliida (Insecta: Neoptera) 7

‘Ideliidae’. Apart from Archidelia elongata Sharov,1961 (Sharov 1961), the best-preserved describedfossil hind wing currently attributed to an ‘ideliid’ isan incomplete one described by Storozhenko (1992)as Stenaropodites sojanensis Storozhenko, 1992 (theholotype of this species being a forewing). This hindwing shows a partially preserved anal area with longsimple vein parallel to the cubito-anal fold and an analvein posteriorly pectinate, reminiscent of the dicty-opteran type sensu Haas & Kukalova-Peck (2001,fig. 21).

It follows that the hind wing anal area of Paoliida asdelimited herein has a great variability, ranging from(apparently) a broad anal fan to the highly specializedanal area of Protoblattinopsis stubblefieldi Laurenti-aux, 1953 (see below). The same situation occurs inmodern Blattodea, ranging from an anal area coveringmore than 50% of the entire wing to very small ones(Fig. 3A, B). These variations in size and shape of thehind wing anal areas do not contradict our proposal ofattributions of all the taxa listed above in the Paoliidae.

Emended diagnosis. The diagnostic characters of Paoli-idae are as follows.

Forewing: CuA and CuP separating from a rather longbasal stem Cu; a short, more or less distinct, but gener-ally relatively convex arculus brace (crossvein) between Mand CuA (more distinct in the forewing than in the hindwing); area between CuA and CuP extremely broad, withdistinct veinlets (autapomorphy), with a general course ofCuA making a double curve; anal veins with few branches;CuP simple or with a short terminal twig, straight or slightlysigmoidal; CuA with convex posterior branches, more orless developed, plus some concave anterior branches; Mrather concave against CuA; areas of M and CuA broaderthan that of RP; RA simple or with weak, short but distinctanterior branches; costal area between ScP and costalmargin with series of rather irregular, simple or branching,oblique veinlets; RP rather concave, with fewer branchesthan in Blattinopsidae; RA convex; crossveins simple or,mainly, abundantly branching and often forming a coarsenet of irregular cells (so called archedictyon); hind wingsubtriangular, with vannus strongly reduced; large insectswith long but otherwise apparently unmodified and ratherlong legs (longer than the body); filiform antennae; wingsin a roof-like position at rest.

Phylogenetic relationships of Paoliidaeand Blattinopsidae

Previous hypothesesSeveral hypotheses have been considered concerning thephylogenetic relationships of Paoliidae and Blattinopsi-dae:

Figure 3. Recent blattodean hind wing venation showing diver-sity in size of the anal areas. A, Pelmatosilpha guianae Hebard,1926 (Blattidae) MNHN coll. B, Polyphaga aegyptiaca (L., 1758)(Polyphagidae) MNHN coll. Scale bars represent 5 mm.

1. Sharov (1966, p. 115) hypothesized that Paoliidaebelong to the infraclass ‘Archaeoptera’, together withEopteridae Rohdendorf, 1961, which were later recog-nized as Eumalacostraca (Rohdendorf 1972; Schram1980), and regarded them as a basal stem group ofPterygota. He noticed that the wings in adults ofPaoliidae are directed backwards at a certain angleto the body at rest, a position similar to many Palaeo-zoic larvae and, according to him, distinguishing thisgroup from Neoptera. The value of this character isvery difficult to estimate, because of deformations ofthe fossils due to compression and diagenesis. Such awing position in Paoliidae could also be related to the

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modification of neopteran insects with wings cover-ing large bodies (see Brauckmann et al. 2003, text-figs 15–17). Prokop et al. (2012) demonstrated thatPaoliidae had the typical neopteran wing articulation.Sharov (1966, pp. 114–117, fig. 50A–C) reviewedother characters in comparison to Neoptera, such asputative absence of any folds along the cubital and analveins. This assumption, based on the study of speci-mens in which the wing bases are missing, is clearlyerroneous, as it can be seen on very well-preservedspecimens known from sphaerosiderite concretions inPoland (Prokop et al. 2012). A new order Protopterawas created by Sharov (1966, p. 115, fig. 50A–C)for Paoliidae on the basis of fore and hind wingsbeing homonomous, the absence of an anal fan (twocharacters that could be specialized), and a specificposition of the wings at rest (discussed above). Hebelieved that the group might have given rise bothto Palaeoptera and Neoptera. Furthermore, Sharov(1966) also noticed the presence of an archedictyon inthe Paoliidae, widely considered as a plesiomorphiccharacter. The presence of an archedictyon is sharedby many palaeopteran and neopteran groups (Palaeo-dictyoptera, Dictyoptera, etc.); thus, this characteralone is not sufficient to support a basal position for thePaoliidae, contra Sharov, independently of its polarity.

2. Schneider (1984, pl. 2) proposed that Paoliidae couldbe a link between a hypothetical neopteran protowingand the orthopteroid lineage (represented by Oedis-chia Brongniart, 1885), the other line of evolutionbeing the dictyopteran lineage. This hypothesis iscontradicted by the very different organization ofthe cubital veins in Paoliidae and Archaeorthoptera(Bethoux & Nel 2002; see below).

3. Carpenter (1992) considered Paoliidae Handlirsch,1906a as a protorthopteran family, but it has beenextensively demonstrated that this last large group ispara- or polyphyletic (Bethoux & Nel 2002). Carpen-ter’s hypothesis does not solve the problem of paoliidaffinities.

4. Kukalova-Peck & Brauckmann (1992), followed byHaas & Kukalova-Peck (2001), supposed that the‘paoliid’ group comprised several families. However,many of them have been assigned to different groupsby various authors, in particular:– Eucaenidae Handlirsch, 1906a, family based on

a single species Eucaenus ovalis Carpenter &Richardson, 1976, considered by Kukalova-Peck &Brauckmann (1992) as representative of the ‘paoliidline’, thus to have a convex CuA and concave CuPseparating from a common basal stem. Rasnitsyn(2002b) put it in Eoblattida Handlirsch, 1906b,but, as indicated above, Eoblatta has a very differ-ent, archaeorthopteran, type of venation. Accord-ing to Carpenter & Richardson (1976), CuA and

CuP emerge from a common stem, which seemsalso to correspond to the reconstruction proposedin Rasnitsyn (2002b, fig. 363). If this interpreta-tion is correct, Eucaenidae would not be an archae-orthopteran ‘eoblattid’. It remains that the presenceof a common basal stem Cu is not sufficient alone tosupport a relationship between Eucaenus and Paoli-ida as redefined above. At this stage we considerEucaenus as Polyneoptera of uncertain affinity;

– Strephocladidae Martynov, 1938, a family currentlyconsidered as a junior synonym of the Anthracop-tilidae Handlirsch, 1922 in the order Hypoperl-ida Martynov, 1928 (Rasnitsyn & Aristov 2004),but more recently as composed of stem-Mantodeaaccording to Bethoux & Wieland (2009) andBethoux et al. (2010);

– Blattinopsidae Bolton, 1925, a family currentlyconsidered as unique representative of the orderBlattinopseida Bolton, 1925 by Rasnitsyn (2002d),in the superorder Caloneuridea Handlirsch, 1906b(see below);

– Synomaloptilidae Martynov, 1938, a familycurrently considered as a Hypoperlida, but Bethoux& Nel (2002) indicated that a revision of Synoma-loptila Martynov, 1938 with an examination of theconvexity of the cubital veins is necessary;

– Cymbopsidae Kukalova, 1965, a family based onthe single genus and species Cymbopsis excelsaKukalova, 1965. Rasnitsyn (2002d, p. 106) consid-ered that it “could be an aberrant Blattinopseida[sic] with M stock hardly discernible and RSbranching disorganized”. The type lacks nearly allthe cubital area, which renders its attribution uncer-tain (Kukalova 1965).

Kukalova-Peck & Brauckmann (1992) supposed thatthis ‘paoliid’ group was an ‘ancestor’ line for apart of the Hemiptera. Affinities of Paoliidae withParaneoptera are unlikely because the former havea crossvein m-cua in the forewing (arculus sensuKukalova-Peck & Brauckmann 1992), plus a convexCuA and a concave CuP separating from a ratherneutral common stem Cu (see Fig. 1C), which isvery different from the pattern of venation observed inParaneoptera (a concave Cu(P) with a crossvein cua-cup anteriorly ‘branching’ on it, proximally concaveand distally convex, and a CuA fused at the wing basewith stem R + M; Nel et al. 2012). Of course theParaneoptera have not the main paoliid apomorphy,viz. ‘area between CuP and CuA broad compared tothe median and radial areas’.

5. Rasnitsyn (2002a) restricted the Paoliida Handlirsch,1906a to the unique family Paoliidae, and consideredit as the most inclusive group of Pterygota, follow-ing the opinion of Sharov (1966). Rasnitsyn (2002a)followed the division of Pterygota nec Paoliida

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Systematic position of Paoliida (Insecta: Neoptera) 9

into Scarabaeones (a group comprising the Parane-optera, Holometabola plus the Palaeodictyoptera andOdonatoptera, etc.), and Gryllones (a group compris-ing the Dictyoptera, Plecoptera, and the orthopteroidlineage). All recent morphological and molecularphylogenetic analyses contradict this hypothesis ofphylogeny, and support the more ‘traditional’ divisionof Pterygota into Palaeoptera and Neoptera (Hennig1981; Haas & Kukalova 2001; Hovmoller et al. 2002;Kjer et al. 2006), or Palaeoptera as a paraphyleticgrade to Neoptera (e.g. Boudreaux 1979; Kristensen1981; Wheeler 1989; Ogden & Whiting 2003; Kjer2004; Whitfield & Kjer 2008). Rasnitsyn (2002e,p. 81) based his hypothesis on the relative extensionsof the main vein areas in the wings, viz. in this char-acter, some paoliids are close to Gryllones in havinglarge Cu, but no one can be related to Scarabaeonesbecause of the enlarged RS. At the same time, paoliids,like plesiomorphic Scarabaeones, keep their wingsroof-like at rest (see Rasnitsyn 2002a, figs 74–76) andhave no foldable anal lobe in the hind wing. ‘Charac-ters’ like the respective sizes of the cubital, medianor radial areas do not convincingly support suchlarge groups that are supposed to comprise severalorders. For instance, among many other cases, the‘scarabaeone’ Meganeuridae have very large radialsector while many other Odonatoptera have a narrowerone. The other issue is that, if pterygote insects aredivided into two main groups, Scarabaeones and Gryl-lones, as proposed by Rasnitsyn, it is simply impossi-ble to determine which one shows the plesiomorphicwing venation on the basis of an outgroup compari-son with non-pterygote insects, also because Rasnit-syn did not propose clear phylogenetic relationshipsbetween the Paoliida and the Scarabaeones and Gryl-lones, establishing which one of these two groupsis the sister group of Paoliida. The same problemoccurs with the hypothesis that the ‘wings roof-like atrest’ would be plesiomorphic (or apomorphic). Suchassumptions cannot be supported by any outgroupreasoning simply because the potential sister groupsof Pterygota are apterous. Rasnitsyn (2002a, p. 83, figs74–76) also indicated that Paoliida have no synapo-morphies, which makes the characterization of thisorder particularly difficult.

6. Grimaldi & Engel (2005, p. 188) considered the Paoli-idae as a potential “stem group to all other Neoptera”,“characterized by numerous primitive features”, andlacking “any derived traits (at least none observablein preserved specimens)”.

A new hypothesis of monophyly for PaoliidaePaoliidae and Blattinopsidae have the roof-like position ofwings at rest, typical of Neoptera and probably also thecorresponding wing basal joints (for Blattinopsidae, see

Brauckmann 2007, fig. 15; also see Fig. 1E). But the polar-ity of these characters remains uncertain due to the absenceof information in the apterous sister group of the Ptery-gota and the current uncertainty affecting the phylogeneticrelationships between the most basal pterygote lineages.

Paoliidae share very similar patterns of wing venationswith Blattinopsidae and Dictyoptera. In particular they havethe same pattern of median and cubital veins, viz. CuA andCuP separating from a long basal common stem + medianvein rather close or connected with radius at wing baseand separating again distally. This pattern is also sharedby many neopterous orders, i.e. Plecoptera (in which Mand R are basally separated) and Holometabola, but notArchaeorthoptera and Paraneoptera (Bethoux & Nel 2002;Nel et al. 2012). Therefore, the putative affinities of theBlattinopsidae with the Caloneuridae (type family of theCaloneuridea sensu Rasnitsyn 2002d) are unlikely, becausethe latter have a wing venation of archaeorthopteran type asdemonstrated by Bethoux et al. (2004). The three groupsPaoliidae, Blattinopsidae, and Dictyoptera also haveprothoracic paranota, rather narrow in Paoliidae and Blat-tinopsidae (see Fig. 2A, B; Brauckmann 2007, p. 189, fig.15), more or less broad in Dictyoptera. Nevertheless thischaracter is probably a plesiomorphy in Pterygota.

Paoliidae also share with Blattinopsidae and Dictyopterathe presence of a convex and short ‘arculus’ betweenconcave M and convex CuA. This character can be consid-ered as present in the groundplans of these three groupscontra Kukalova-Peck & Lawrence (2004, p. 109) whoconsidered that this ‘arculus’ or ‘brace mp-cua is “almostcertainly not part of Blattoneoptera groundplan”, becauseit would be “absent in ancestral Blattoneoptera. . .”. In thecase of Dictyoptera, this convex ‘arculus’ is present inseveral Palaeozoic taxa (e.g. Archimylacris Scudder, 1868,Manoblatta Pruvost, 1919, Mesoptilus Lameere, 1917,Miroblattites Ozdikmen, 2009, Phyloblatta Handlirsch,1906a; see Figs 4A, B, 5I; Laurentiaux-Vieira & Lauren-tiaux 1980, 1987; Schneider 1983, pers. comm.; Bethoux& Wieland 2009; Ozdikmen 2009), especially in the taxathat are currently considered as the most basal represen-tatives of Dictyoptera (Schneider 1984). Nevertheless this‘arculus’ is absent in many modern Dictyoptera (many Blat-todea including the Isoptera, even if it is still present inthe hind wing of the blattodean Periplaneta americana(Linnaeus, 1758), in the forewing of some specimens ofPolyphaga aegyptiaca, or the mantodean Chaetessa valida(Perty, 1833)) (A.N. pers. obs.). The strength of the ‘arcu-lus’ also greatly varies among the different Paoliidae, froma strongly convex vein to a very weak one. A convex ‘arcu-lus’ between concave M and convex CuA is also present inmany neopterous orders, i.e. Plecoptera and Holometabola,but not in Archaeorthoptera and Paraneoptera (Bethoux &Nel 2002; Nel et al. 2012). Its presence seems to be relatedto the pattern of median and cubital veins characteristicof these orders. It could belong to a potential common

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Figure 4. Palaeozoic blattodean forewings showing a convex ‘arculus’ (A, B) and other taxa attributed to Paoliida (C–H). A, Miroblattitescostalis (Laurentiaux-Vieira & Laurentiaux, 1987) with detail of arculus, holotype IRSNB No. 15.576. B, Phylloblatta brongniartiHandlirsch, 1906b with detail of arculus, holotype NHML No. I.7282. C, Archidelia ovata Sharov, 1961 (Ideliidae), holotype PIN No.1197-287b. D, Protoblattina bouvieri Meunier, 1909, holotype MNHN No. DP R52936, detail photograph of forewing anterobasal part.E, Acropermula acra (Kukalova, 1964), holotype PrF UK – Paleo No. 69/1963. F, Herbstiala herbsti Schmidt, 1953, holotype GD NRWNo. Kar 46. G, Protoblattinopsis stubblefieldi Laurentiaux, 1953, holotype BU No. 687. H, Kochopteron hoffmannorum Brauckmann,1984, holotype WMf.N No. P22596. For abbreviations see text. Arrows indicate position of arculus; scale bars represent 5 mm.

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Systematic position of Paoliida (Insecta: Neoptera) 11

ground plan of the Neoptera, and thus would not constitutea synapomorphy of a clade (Paoliida + Dictyoptera).

The presence of anterior concave branches of CuA(possibly posterior branches of M translocated onto CuA,after Bethoux et al. 2009, 2011) is much more likely to be asynapomorphy of a clade (Blattinopsidae, Paoliidae, Dicty-optera) (see discussion above for the presence of these veinsin the fossil and recent Dictyoptera). Other wing venationcharacters of both Paoliida and Dictyoptera correspond toplesiomorphies of Neoptera, thus this clade remains ratherweakly supported.

Blattinopsidae and Paoliidae share the presence of abroad area between CuA and CuP, plus the absence of ananal fan in the hind wing, which we propose as potentialsynapomorphies (for Blattinopsidae, see Hornschemeyer &Stapf 2001). All the other polyneopteran orders have a largeanal fan (if not, their wings are reduced as in Zoraptera orEmbioptera). Thus we propose to include both of them inthe clade Paoliida sensu nov. The potential synapomorphyof Paoliidae is the very broad area between CuA and CuP.The Blattinopsidae have a broad area between CuA andCuP but less than in Paoliidae (note that some of themhave a very broad area, viz. Blattinopsis sp. (Laurentiaux& Teixeira 1948); they have a pectinate RP with numer-ous parallel branches in the forewing (Fig. 5D), which isa potential synapomorphy of Blattinopsidae relatively toPaoliidae (list of taxa of this family and revision of Blat-tinopsidae in Hornschemeyer & Stapf 2001). A few recentBlattodea also have a broad area between CuA and CuP (viz.Polyphaga aegyptiaca (L., 1758), pers. obs.), but anothersignificant difference between Dictyoptera and Paoliida isthe shape of CuP, posteriorly curved in the former while itis straight or slightly sigmoidal in the latter.

It is hardly possible to define wing venation structuresas synapomorphies of the Dictyoptera. Grimaldi & Engel(2005, p. 228, table 7.3) proposed “forewing tegminous;with arched, groove-like claval suture”. If this is moreor less exact for modern representatives, it is not so formany Palaeozoic taxa (see Fig. 4A). Rasnitsyn (2002c,pp. 260–261) characterized his superorder BlattideaLatreille, 1810 by the characters “forewing with R andRS weakly individualized”, and the “anal area (clavus)wide, lanceolate, with veins gently curved according itsfore margin, and weakly branching, if at all”. The firstcharacter concerning the radius is not very clear, but RAwith long anterior and posterior branches could indeedcharacterize Dictyoptera: Schneider (1984, pl. 2) hypoth-esized that a branched RA could be plesiomorphic for adictyopteran–orthopteran lineage. This is contradicted byKukalova-Peck (1991) who considered that in the pterygote‘groundplan’ the convex anterior vein RA is simple (or withonly weak anterior branches). The situation in Palaeodicty-optera, Ephemeroptera, Odonatoptera, and basal Neopteracorresponds to the hypothesis of Kukalova-Peck. Thus it isprobable that a simple RA or with short anterior branches

is a plesiomorphy present in Paoliida, and the stronglypectinate RA could then be a synapomorphy of Dictyoptera.

The second character proposed by Rasnitsyn is presentin Paoliidae and Dictyoptera, but also in some Archae-orthoptera (e.g. Stenoneura fayoli Brongniart, 1893)(Bethoux & Nel 2002). Thus it cannot constitute a potentialsynapomorphy of a clade (Paoliida + Dictyoptera).

A further putative synapomorphy of the Dictyoptera isthe presence of a deeply concave CuP. This character isonly present in this group, clearly visible in Blattodea andMantodea, and even in the Mastotermitidae, most basalfamily of the Isoptera.

Note that Kukalova (1959, p. 4) previously supposedthe relationship of the family Blattinopsidae with Paoliidaeand Cacurgidae on the basis of the following characters:MA absent [in fact, a character very difficult to assess,the median vein being generally more concave than CuA,which does not accurately demonstrate that MA is reallyabsent], M basally close to stem of R, CuA and M coalescedat wing base [in fact, CuA approaches the median andradial veins but is not coalescent with them in paoliids], Mdiverges from CuA in first fourth of wing length. Bethoux(2006) revised Cacurgus Handlirsch, 1911 and demon-strated that it has a different venation pattern of basal orga-nization typical for Archaeorthoptera sensu Bethoux & Nel(2002).

Other taxa potentially attributableto PaoliidaeThe list of fossil taxa examined below and candidate taxato be included in the paoliid clade is not exhaustive. Animportant difficulty to define what to add to the Paoliidais the fact that the order ‘Grylloblattodea’, based on extantsecondarily apterous insects, currently comprises numerousextinct taxa put together on the basis of similarities in theirwing venations, even if they can have very different body orleg structure (Huang & Nel 2007). Furthermore, the bodycharacters shared by the modern and the fossil ‘grylloblat-todeans’ are plesiomorphies (Huang et al. 2007). Thus thisorder could be para- or polyphyletic. Some ‘grylloblattids’(but not all of them) have a common stem of CuA and CuP, arelatively broad cubital area, broader than the area betweenCuA and M, a convex ‘arculus’ between M and CuA, anda double curve to the anterior branch of Cu. These taxa areproper candidates to be related to Paoliida. As an exam-ple, Camptoneurites reticulatus (Martynov, 1928) (Camp-toneuritidae) has clearly all these characters, plus anteriorconcave branches of CuA, after the original photograph ofits holotype (Martynov 1928, pl. 8, fig. 3). Demopterumgracile Carpenter, 1950, currently included in the samefamily (Aristov et al. 2010), could also have the same struc-tures (see Carpenter 1950, pl. 2, fig. 2), but a direct exam-ination of the type material would be necessary before aformal transfer of these insects from the ‘Grylloblattida’

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Figure 5. Line drawings of wing venation showing the main pattern of longitudinal veins omitting dense crossveins (archedictyon).A, Zdenekia grandis Kukalova, 1958a (Paoliidae), forewing. B, Paolia vetusta Smith, 1871 (Paoliidae), hind wing. C, Zdenekia cf.grandis Kukalova, 1958a (Paoliidae), hind wing. D, Blattinopsis ornata Bolton, 1925 (Blattinopsidae). E, Archidelia ovata Sharov, 1961(Ideliidae). F, Acropermula acra (Kukalova, 1964). G, Protoblattina bouvieri Meunier, 1909, forewing. H, Kochopteron hoffmanno-rum Brauckmann, 1984, forewing. I, Miroblattites costalis (Laurentiaux-Vieira & Laurentiaux, 1987), forewing. J, Protoblattinopsisstubblefieldi Laurentiaux, 1953, hind wing. For abbreviations see text.

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Systematic position of Paoliida (Insecta: Neoptera) 13

into (Paoliida + Dictyoptera). Camptoneurites could alsobe more closely related to Paoliidae than to Blattinopsidae,because of the presence of a very broad cubital area.

Protoblattina bouvieri Meunier, 1909 (Figs 2B, 4D, 5G)is another potential Paoliida. Kukalova-Peck (1983, captionof fig. 17) indicated: “In paoliid venation, concave sectionof MP- bracing towards CuA is currently misinterpreted byRussian entomologists as ‘M5’, a supposedly nonsymmet-rical fifth branch of M. In fact, MP– fuses with CuA+ andseparates from it again as MP±, while plesiomorphic MA±is richly branched”. Her interpretation is based on the studyof Protoblattina bouvieri Meunier, 1909. This taxon wasrestudied by Bethoux et al. (2005) who restored the familyProtoblattinidae Meunier, 1909 and included it in the orderGrylloblattida, but following the opinion of Storozhenko(1998, pp. 86–87), without formal rediscussion on its posi-tion. A re-examination of the holotype (MNHN R52936)shows that the concave CuP and the convex CuA sepa-rate from the stem of Cu independently from M or R; Mis basally relatively convex and closely parallel to R, theproblematic ‘M5’ or ‘MP–’ is in fact a very weak butconvex brace between M and CuA, exactly correspond-ing to the paoliid ‘arculus’. The area between CuA andCuP is broad but with relatively weak and distinctly convexposterior branches of CuA; CuA has distinctly concavedistal anterior branches; all the branches of M are concave(i.e. as concave as CuP and more concave than RP andof course CuA). Kukalova-Peck (1983, caption of fig. 17)also indicated that Protoblattina had typically neopterouswing folding, and not wings at rest “supposedly orientedobliquely backwards and supposedly gave rise both to later-ally oriented wings of Palaeoptera, and to posteriorly foldedwings of Neoptera”. Direct examination of the holotypeconfirms Kukalova-Peck’s opinion on this point. Protoblat-tina bouvieri shows all the characters typical of Paoliidae,as listed above in the revised diagnosis. It also has smallprothoracic paranota (Fig. 2B). Thus we propose to considerthat this species is in fact a paoliid, and we consider thefamily Protoblattinidae Meunier, 1909 as a junior synonymof Paoliidae.

The Lower Permian Acropermula acra (Kukalova, 1964)(originally in the genus Permula Handlirsch, 1919, latertransferred to the genus Alicula von Schlechtendal, 1913by Carpenter [1992], and recently to the monospecificgenus Acropermula by Aristov & Storozhenko [2011])has all the characters typical of the Paoliidae, viz. inforewing, CuA and CuP emerging from a common stem,a convex ‘arculus’, a very broad cubital area, presence ofconcave anterior branches of CuA, a median vein moreconcave than CuA, very similar patterns of M, R, and ScP(see Figs 4E, 5F). Von Schlechtendal (1913, pl. 2, fig.10a, b) only provided photographs of ‘Alicula lebachen-sis’. Aristov & Storozhenko (2011) revised the grylloblat-todean family Aliculidae Storozhenko, 1997a (type genusAlicula von Schlechtendal, 1913, and type species Alic-

ula lebachensis von Schlechtendal, 1913) and changedthe family name to Permulidae Aristov & Storozhenkobecause the genus name Alicula was preoccupied andthus replaced by its junior synonym Permula Handlirsch,1919. Aristov & Storozhenko (2011) excluded Acroper-mula from the Permulidae and considered it as belong-ing to the grylloblattodean family Ideliidae Zalessky, 1928because of the shape of the forewing CuA. The generalpattern of forewing venation of the taxa currently consid-ered as permulids is also very similar to the paoliid type(even in the presence of concave anterior branches ofCuA), except for the area between the convex CuA andthe concave CuP that is comparatively very narrow. Thus,Permuliidae are certainly not Paoliidae, but the affinitiesof this family remain uncertain, viz. they could belong tothe paoliid–dictyopteran lineage, rather than to an uncer-tain group Grylloblattodea that is currently not supportedby any apomorphy. Storozhenko (1997a, p. 11) revisedthe Ideliidae. He characterized this group on the basisof the presence of “a few proximal branches of CuA,which terminating on CuP or disappearing in CuA-CuParea in forewing”, which is a character also present inPaoliidae. He included in this family numerous generathat have rather different wing venations. Some of themhave a forewing venation identical to those of Paoli-idae, in the shape of all veins, including the median veinmore concave than CuA (see Sharov 1961, pl. 11, fig.59; Fig. 4C), viz. Stenaropodites Martynov, 1928, Aenig-midelia Sharov, 1961, Archidelia Sharov, 1961 (Fig. 5E),Sojanidelia Storozhenko, 1992, Micaidelia Aristov, 2004,Acropermula Aristov & Storozhenko, 2011, MongoloideliaStorozhenko, 1992, and Sylvidelia Martynov, 1940 (notethat this last taxon has prothoracic paranota identical tothose of Protoblattina, see Aristov & Novokshonov 2000).But some others seem to have a distinctly narrower areabetween CuA and CuP (Ideliopsina Storozhenko, 1997b,even if some taphonomic deformation of the wings remainspossible for these fossils) (Storozhenko 1997b, 1998; Aris-tov 2004). Thus, it is possible that this family is in facta polyphyletic grouping. We therefore propose to trans-fer the taxa listed above into Paoliidae on the basis of thepresence of all forewing venation characters and apomor-phies. As we consider Stenaropodites (type genus of Ideli-idae) as belonging to Paoliidae, we have also to considerthe former family as a junior synonym of the latter.Furthermore, some taxa currently considered as Ideliidaehave to be excluded from Paoliidae, at least provision-ally: Rachimentomon Zalessky, 1939 (based on a ratherpoorly preserved fossil); Kolvidelia Zalessky, 1956 (witha reduced wing venation, which renders its attributionto the Paoliidae more dubious than for the other taxalisted here); Ideliopsina Storozhenko, 1997b; AnaideliaStorozhenko, 1997b; Madygenidelia Storozhenko, 1997b;Pseudoshurabia Storozhenko, 1997b (area between CuAand CuP narrower than in Paoliidae).

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14 J. Prokop et al.

The Permian Paolekia perditae Riek, 1976 from SouthAfrica is based on a single wing apex and originally placedin Paoliidae (Riek, 1976). Carpenter (1992, p. 133) consid-ered it as a “Protorthoptera of family uncertain”. It lacksinformation on nearly all the most important wing struc-tures but it shares with Paoliidae a long ScP, a RA with fewshort apical branches, RP and M both concave and a generalorganization of the wing apex very similar to those of theother paoliids. The absence of a convex median vein alsosupports an attribution to this family. But its exact affinitiesremain very uncertain.

Prokop et al. (2012, p. 13, appendix 1) listed Ampeliptera,Stygne and Katerinka as “unassigned taxa probably closelyrelated to Paoliida or inclusive”. Bethoux & Nel (2002,p. 17) revised Ampeliptera limburgica Pruvost, 1927 andindicated that it belongs to the Archaeorthoptera. Stygneroemeri Handlirsch, 1906b could be related to the paoli-ids. Its CuA is clearly convex and apparently begins froma common stem with CuP, but it seems that it has no clearconcave distal anterior branches of CuA; the basal part ofthe area between CuA and M and vein CuP are rather poorlypreserved. It has a long, convex and distally forked vein inthe area between main CuA and CuP. This vein is prob-ably a basal branch of CuA (Schwarzbach 1939, fig. 8).Katerinka candida Prokop & Nel, 2007 (type species ofthe Katerinkidae Prokop & Nel 2007) has the characters ofthe paoliid line (convex CuA and concave CuP emergingfrom a common stem, a convex ‘arculus’, anterior concavebranches of CuA, a broad area between CuA and CuP).It differs from Paoliidae in the short ScP ending on RA,but some Camptoneuritidae have also a short ScP (JabloniaKukalova, 1964, some Camptoneurites Martynov, 1931)(Aristov et al. 2010). Thus Katerinka could as well bea Camptoneuritidae, but a revision of the taxa currentlyattributed to the Camptoneuritidae is needed before draw-ing any conclusion on this point.

Herbstiala herbsti Schmidt, 1953 (in the monospecificfamily Herbstialidae Schmidt, 1953) has all the diagnos-tic characters of a paoliid forewing (Fig. 4F). It was firstattributed to the group ‘Protocicada’ (Schmidt, 1953).Carpenter (1992, p. 119) considered it as “related toCacurgidae (probably synonymous)”. The main differencewith this orthopteroid family is the clearly convex CuAand the concave CuP both separating from the stem Cu.Rasnitsyn (2002a, p. 83) considered it as a “Scarabaeonaincertae sedis (winged insects of obscure taxonomic posi-tion)”. We attribute Herbstiala herbsti to the Paoliidaeand consider the Herbstialidae as a junior synonym ofthis family. Kochopteron hoffmannorum Brauckmann, 1984is another enigmatic taxon that has a forewing venationlooking like that of a paoliid (Figs 4H, 5H). It was firstattributed to Cacurgidae Handlirsch, 1911 (Brauckmann1984; Brauckmann et al. 1985), a group considered asPanorthoptera by Bethoux & Nel (2002), challenged byBethoux (2006) who considered the type genus as a more

basal Archaeorthoptera. Later, Brauckmann et al. (2003)and Ilger & Brauckmann (2008, p. 281) put in doubtthis attribution; these later authors avoided any precisesystematic assignment for Kochopteron, but indicated thatit “resembles Holasicia rasnitsyni Brauckmann, 1984”, ataxon that is currently included in Paoliidae (Prokop et al.2012). Kochopteron hoffmannorum is of great interest forthe presence of prothoracic paranota in an additional spec-imen (WMf.N P.26093, pers. obs.). Thus its potential attri-bution to Paoliidae is of importance. After our present re-examination, Kochopteron has a convex CuA and a concaveCuP separating from a basal common stem (paoliid type).A comparison with the archaeorthopteran Heterologopsisruhrensis Brauckmann & Koch, 1982 is decisive. After thevery clear photographs of Brauckmann et al. (1985, pl. 19,fig. 1b and pl. 21, fig. 1) and the direct re-examination ofKochopteron, it appears that Heterologopsis has a stronglyconvex CuA separating from a common stem with R + Mand a more concave vein between CuA and concave CuP,while Kochopteron has a clearly convex CuA emergingfrom a common stem with CuP, but also a distinctly convex‘arculus’ between CuA and M. It also has a very broad areabetween CuA and CuP, anterior branches of CuA of thesame elevation as the distal branches of M. This pattern ofvenation is typically of a paoliid type. Thus we propose toconsider Kochopteron as a Paoliidae.

Rasnitsyn (2002a, p. 81) indicated that the paoliids(i.e. Zdenekia Kukalova, 1958a) closely resemble some“plesiomorphic Scarabaeones (Ampeliptera Pruvost, 1927,Limburgina Laurentiaux, 1950, Propachytylopsis Lauren-tiaux & Laurentiaux-Vieira, 1981 [sic], HeterologopsisBrauckmann & Koch, 1982, etc.)”. Ampeliptera limburgicaPruvost, 1927 is an Archaeorthoptera (see above), whileHeterologopsis and Protopachytylopsis are Panorthoptera(Bethoux & Nel, 2002). Limburgina antiqua Laurentiaux,1950 is based on an incomplete forewing, with basal thirdmissing. Thus, even if it seems that the CuA is emergingfrom a common stem with CuP after the general direc-tions of these veins (Laurentiaux, 1950), there is no clearevidence of this crucial point, “making its assignment tothe ‘paolid line’ or to the Archaeorthoptera impossible”,as was already indicated by Bethoux & Nel (2002, p. 22).Thus, the set of arguments based on the convexity of thebasal parts of the main longitudinal veins and patterns oforganization of the median and cubital veins, as proposedby Bethoux & Nel (2002), Nel et al. (2012), etc., allowus to place these taxa in lineages that strongly differ fromPaoliida.

The enigmatic Carboniferous Westphaloblattinopsisedwardsae Bethoux & Jarzembowski, 2010 could be relatedto Paoliida if we admit the preferred interpretation of thewing venation of Bethoux & Jarzembowski (2010, p. 91),i.e. CuA emerging from a common stem with CuP, plusthe fact that it seems to have a concave anterior branch ofconvex CuA plus a broad area between CuA and CuP. But

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Systematic position of Paoliida (Insecta: Neoptera) 15

these characters are not conclusive because of the poor stateof preservation of this fossil.

Protoblattinopsis stubblefieldi Laurentiaux,1966Protoblattinopsis stubblefieldi (Figs 4G, 5J) is based onthe basal two-thirds of a wing, with the extreme base alsomissing. Laurentiaux (1953, p. 447, fig. 42) figured thisfossil and named it ‘Protoblattinopsis stubblefieldi Lauren-tiaux, 1952’ with the figure caption “Protoblattinopsisstubblefieldi Laurentiaux, Westphalien inferieur de Deep-field (South Staffordshire). Base d’une aile mesothoraciquegauche, gr. nat. Genotype (d’apres Laurentiaux)”, but nopublication by this author in 1952 is concerned with thistaxon. In 1953, he did not give any diagnosis or descrip-tion. So it was a nomen nudum. Later Laurentiaux (1966,p. 828) indicated that it was described by him in 1953:“1953. Protoblattinopsis stubblefieldi Laurentiaux [p. 447,fig. 42]”, and cited by Kukalova (1958b, p. 130, 1959, p.7) under the name “Protoblattinopsis stubblefieldi Lauren-tiaux, 1952”. The first description of this taxon is actuallyin Laurentiaux (1966).

Laurentiaux (1953, 1966) and Bethoux et al. (2009)considered that it is a forewing, but without clear argument.An implicit one could be that it is supposed to belong to thedictyopteran lineage in which the hind wing has an anal fan.But this fossil has a very curious pattern of the CuP and analveins: the concave CuP, the convex first anal vein A1, andthe concave second anal vein A2 are strongly curved andnearly perpendicular to the main direction of the wing, withone row of cells between them. Also the anal area is trian-gular, with its posterior margin following a fissure in therock, except at its very base where it is complete. A similarsituation occurs in Odonatoptera: Isophlebioidea in whichthe convex CuA and the concave AA are closely parallel,make a curve and become perpendicular to the main direc-tion of the wing; also basal to AA there is a rather small analarea, rounded in female and triangular in male, nearly iden-tical to the anal area of Protoblattinopsis (Nel et al. 1993).Similar structures are also present in the Odonatoptera:Libelluloidea, viz. the anal loop is a structure perpendicu-lar to the general direction of the wing, limited by convexCuA, a concave Cuspl and a convex branch of AA. To acertain point, the Odonatoptera: Aeschnidiidae also have intheir four wings a structure with a similar function, i.e. theelongate transverse discoidal triangle. In these groups ofOdonatoptera such transverse structures support and rein-force proximally the anal area and distally the cubital area.They only occur in the hind wings because they are broaderthan the forewings, except in the Aeschnidiidae that havevery broad forewings and hind wings (Fleck & Nel 2003).

This morpho-functional similarity suggests that the typespecimen of Protoblattinopsis is a hind wing. This hypoth-esis would imply that this taxon belongs to a group with-

out an anal fan in hind wing. The presence of relativelyconcave anterior branches of CuA is the synapomorphythat we proposed above for the (Paoliida + Dictyoptera).An attribution of Protoblattinopsis to Dictyoptera does notfit well with the absence of an anal fan in the hind wing.Dictyoptera have an anal fan in the hind wing, even ifit may be less developed than in the archaeorthopteranlineage. Paoliidae could be a better candidate as closestrelative of Protoblattinopsis because they have a ratherreduced anal fan in the four wings. Interestingly the onlydifferences in the forewing venation of a paoliid and thedictyopteran Archimylacris are as follows: presence of avery broad cubital area in fore and hind wings (synapo-morphy of the Paoliidae); RA simple or posteriorly weaklybranched in Paoliidae whereas it is anteriorly pectinate orbranched in Archimylacris and other Dictyoptera. Otherveins have a very similar organization in both groups, evenin the presence of a brace mp-cua ‘arculus’ (Schneider1984; Bethoux et al. 2009). These characters are clearlypresent in Protoblattinopsis. The interpretation of the typespecimen of Protoblattinopsis stubblefieldi as a hind wingis also congruent with the absence of an ‘arculus’. Nearlyall Paoliidae have an ‘arculus’ in their forewing whereasthis structure is weaker or absent in their hind wing.

Bethoux et al. (2009) considered that the base of RP inProtoblattinopsis is far distally from that of M, which isvery different from the situation in Paoliidae. This couldconstitute a counter-argument against an attribution of thistaxon to this family. Nevertheless, Protoblattinopsis has an‘anterior branch’ emerging from M and running parallel toR, exactly in the same position as the ‘normal’ paoliid RP.This concave vein emits a posterior branch just before theputative base of RP reaches it nearly at right angle. Theputative distal base of RP is also becoming weaker near itsfusion with the putative ‘anterior branch of M’. This situa-tion can easily be interpreted as a capture of the true base ofRP by M, from which RP re-emerges very quickly, and afterhas a ‘normal’ course, except for the presence of a second‘distal base’ of RP (probably in fact a modified crossvein,similar to the subnodus in Odonatoptera). A similar struc-ture also occurs in the recent Neuroptera: Hemerobiidae inwhich there can be from one to three different ‘bases’ andbranches of RP emerging more or less independently fromR (Oswald 1993). It remains that Protoblattinopsis has ahighly specialized hind wing.

Description of a new paoliid genusand species

Genus Silesiapteron gen. nov.

Type species. Silesiapteron jarmilae sp. nov.

Etymology. Composite name after Silesia, a historicalregion in Central Europe (in the Czech Republic and

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Figure 6. Silesiapteron jarmilae Prokop et al., gen. et sp. nov., holotype ISEZ PAN No. MP ISEA I-F/MP/1488/1a-b /08. A, photographof forewing; B, drawing of forewing venation. For abbreviations see text. Scale bar represents 10 mm.

Poland) where the outcrop is located, and pteron (Greekfor wing); neuter in gender.

Diagnosis. Forewing characters only. Absence of archedic-tyon, area between CuA and CuP very broad but with onlysix rows of large cells, CuA very close to M in their basalparts, ‘arculus’ mp-cua absent or in a very basal position,anal area narrow, RA with posteroapical branches, M withtwo main branches, secondarily forked.

Silesiapteron jarmilae Prokop sp. nov.(Fig. 6A, B)

Etymology. The epithet honours Dr Jarmila Kukalova-Peck (Carleton University, Ottawa, Canada), world-famouspalaeoentomologist.

Diagnosis. As for the genus.

Material. Holotype specimen No. I-F/MP/1488/1a /08imprint and No. I-F/MP/1488/1b /08 counter-imprint ofthe medial four-fifths of well preserved forewing insphaerosiderite concretion, housed in the Muzeum Przy-rodnicze Institutu Systematyki Ewoluciji Zwierz

↪at PAN

(Krakow, Poland).

Occurrence. Upper Carboniferous, WestphalianA (Langsettian), Mudstone series (Zał

↪e_ze beds),

Sosnowiec–Klimontow, originally Por↪abka–Klimontow

Mine, Upper Silesian Coal Basin, Poland.

Description. Based on nearly complete forewing, withapex and very basal mid part missing; membrane of wingdark with no spot or other coloration pattern, rather coarse

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Systematic position of Paoliida (Insecta: Neoptera) 17

network of crossveins; wing length about 71 mm, maximumwidth 25 mm in first and second third of wing; an indenta-tion of posterior wing margin at the end of CuP, defining theclaval area; anterior wing margin basally strongly curved,ScP basally curved reaching RA in distal fifths about 17 mmfrom wing apex, numerous oblique simple crossveins inbroad area between ScP and costal margin; stem of convexR probably basally connected to M (not preserved but thetwo veins are strongly approximate); division of RA andRP 21.8 mm from wing base, convex RA running paral-lel to costal margin and ending with four terminal poste-rior branches, all reaching wing apex, first 17 mm fromwing apex secondarily bifurcated; neutral to convex RPdeeply bifurcated 28.3 mm from wing apex, both branchesof RP terminally twigged ending on posterior wing margin;concave M diverging from R about 16.7 mm from wingbase, just basal to level of base of RP, first bifurcation of Mclose to mid wing, both branches secondarily branched, andanterior one three-time twigged, all reaching posterior wingmargin with five branches; CuA and M strongly approxi-mated near their bases; area between CuA and CuP verybroad, with five-six rows of large pentagonal or hexagonalcells in widest part; basal part of both veins not preserved,but their general direction suggests that they separated ina rather open angle, convex CuA with three anterior andthree posterior branches ending on posterior wing margin,strongly concave CuP simple and curved, well delimitingthe anal area; anal area strongly reduced with five main moreor less parallel veins, with one row of simple crossveinsbetween them.

Discussion. This fossil can be considered as a forewingbecause of the reduced anal area and its general shape.It bears a combination of diagnostic characters of severalfamilies within ‘Protorthoptera’ and ‘Protoblattoidea’.However, both groups have been considered as polyphyletic(Carpenter 1966) for a long time. Because of the convexCuA not captured by M and absence of a common stemR + M, this fossil is not related to the archaeorthopteranclade (Bethoux & Nel 2002).

Silesiapteron jarmilae gen. et sp. nov. can be placed closeto Paoliidae in bearing an oval shape of the wing, a consider-ably broad costal area with numerous veinlets, ScP endingon RA in distal part, division of RA & RP proximal ofmidwing, M concave and well developed, division of CuA& CuP not close to wing base and probably separating inan open angle, with CuA probably basally curved, CuAwith numerous branches, the anterior ones being relativelyconcave, area between CuA and CuP very broad with largecells and secondary branches of CuA, anal area weaklydelimited by a marginal indentation at the end of CuP andstrongly reduced. On the other hand, S. jarmilae bears asimple coarse network of crossveins instead of an archeo-dictyon, typical for most paoliids. The occurrence of anarculus cannot be ascertained, due to poor preservation in

Figure 7. Hypothetical tree of relationships of Paoliida sensu nov.and Paoliidae with dictyopteran orders based on wing venationcharacters.

the corresponding area between veins M and CuA in S.jarmilae.

Silesiapteron jarmilae can be easily separated from allspecies currently included in Paoliidae by the followingcharacters: absence of archedictyon, area between CuA andCuP very broad but with only six rows of large cells, CuAvery close to M in their basal part, plus maybe ‘arculus’mp-cua absent or in a very basal position.

Conclusions

Since Handlirsch’s time (e.g. Handlirsch 1906b) the posi-tion of the family Paoliidae among winged insects has beenrather uncertain and greatly varied. Our present analysisprovides new evidence for a close relationship betweenthe families Paoliidae and Blattinopsidae, resulting in aredefinition of the order Paoliida sensu nov. on the basisof new, putative synapomorphies in their wing venations(Fig. 7). After the re-examination of the type material wealso propose a new diagnosis for the family Paoliidae.Herein we consider the Paoliida as an undisputed neopteranclade, currently ranging from the early Late Carboniferousthrough the Late Permian. We also consider it as the sister

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group of Dictyoptera on the basis of the presence of ante-rior concave branches of CuA in the forewing (synapomor-phy) (see Fig. 7). Our re-examination provides evidence forconsidering the family Herbstialidae and the ‘grylloblattid’family Ideliidae as junior synonyms of the Paoliidae, and fora subsequent reassignment of the genera of these families,but also of the genera Kochopteron Brauckmann, 1984 andProtoblattina Meunier, 1909 into Paoliidae. Furthermore,the present study provides a new interpretation of the wingvenation of Protoblattinopsis stubblefieldi Laurentiaux,1953, suggesting a highly specialized cubito-anal area func-tionally analogous (but not homologous) to the anal loop ofthe hind wing of the Isophlebioidea (Odonatoptera). Afterevaluation of the characters, we propose to include Proto-blattinopsis within the Paoliida. Our discovery of Silesi-apteron jarmilae gen. et sp. nov. from a Late Carboniferous(Langsettiian) sphaerosiderite concretion in the Upper Sile-sian Coal Basin (Poland) is of great interest, as this taxonbears a strongly reduced anal area and a coarse network ofcrossveins instead of fine meshwork of crossveins typicalfor most paoliids.

The family Paoliidae is among the oldest known wingedinsects, which could have been an argument favouring aninclusive position in Pterygota. Nevertheless, it is contem-poraneous with the oldest known Odonatoptera, Archae-orthoptera, Dictyoptera, etc. It is currently known from arather short interval during the early Late Carboniferous(Late Namurian to Langsettian) of Euroamerican deposits(Kukalova 1958a; Brauckmann et al. 2003; Prokop & Nel2007; Prokop et al. 2012), but the present study extendsits range into the Late Permian by the addition of severaltaxa previously included in other families (see below).The group shows relatively high abundance with surpris-ingly low morphological diversity in comparison with theother groups of neopteran insects that are diverse in theDuckmantian/Bolsovian (Carboniferous: Pennsylvanian).Prokop et al. (2012) hypothesized a correlation between asudden extinction of the paoliids and a decrease of humiditycalled the first ‘drier interval’ at the Langsettian/Bolsovianboundary (Carboniferous: Pennsylvanian). This event iswell documented in the plant fossil record and consideredas a major event for Euroamerican coalswamp vegetation(Phillips & Peppers 1984; Galtier 1997).

After our transfer of several Late Carboniferous andPermian taxa into Paoliidae, the hypothesis of the suddenextinction of Paoliida at the Langsettian/Duckmantianboundary no longer stands. Nevertheless, it seems that thislineage was flourishing during the early Late Carbonifer-ous, had an important decrease in diversity during the latestCarboniferous, and rediversified during the Early Permian.In parallel, the cockroaches were rare during the first phaseof diversification of Paoliidae, and they became much morediverse when the paoliids had their first decrease. Blattinop-sidae are known during the same period of time as the Paoli-idae but they were frequent and diverse during the Stepha-

nian while Paoliidae had their first decrease. As nothing isknown about the biology of these insects, it is not possi-ble to establish accurate links between their diversity andpalaeoclimatic changes.

Several other Palaeozoic to early Mesozoic taxa,currently placed in Grylloblattodea, could also be related tothe paoliid and dictyopteran clades. Revising these fossilswill help to empty this taxonomic wastebasket. Lastly,the new position of the Paoliida as a neopteran cladeclosely related to Dictyoptera contradicts their previoussituation as the most basal Pterygota and also the hypothe-sis of the subdivision of this last clade into Gryllones andScarabaeones (sensu Rasnitsyn 2002a).

Acknowledgements

We thank Dr Annelise Folie for access to the collec-tions of the Royal Belgian Institute of Natural Sciences(Brussels, Belgium), Mgr Eva Mertova for access to thecollection of the Ostrava Museum (Ostrava, Czech Repub-lic), Dr Paul Mayer (The Field Museum, USA), and DrsAlfred Hendricks and Lothar Schollmann for access to thecollection of the LWL-Museum fur Naturkunde (Munster,Germany). We are grateful to Dr Jirina Daskova (CzechAcademy of Sciences) for loan of a specimen from theLapworth Museum, University of Birmingham (UK) and toJason Hilton for previous access to the Birmingham collec-tion. The authors are grateful to Prof. Alexander Rasnit-syn and Dr Daniil Aristov (Russian Academy of Sciences,Moscow, Russia) for a photograph of Archidelia ovataSharov, 1961 in the Palaeontological Institute collections.The first author (JP) expresses thanks for his visit to theInstitut royal des Sciences naturelles de Belgique (Brus-sels) supported by the SYNTHESYS Project financed bythe European Community Research Infrastructure Actionunder the FP7 Integrating Activities Programme (BE-TAF-1004) and acknowledges research support from the GrantAgency of the Czech Republic No. P210/10/0633. Thesecond and third authors (WK and EK) are grateful forresearch support from the Polish Committee for ScientificResearch (KBN), grant No. N N303 345535. The Clausthalauthors (JMI and CB) gratefully acknowledge the financialsupport of the Deutsche Forschungsgemeinschaft (DFG)project BR 1253/4-1 and 4-2. TH was supported by DFGgrants HO 2306/6-1 and 7-1.

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