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Zoological Journal of the Linnean Society, 2009, 156, 79–113. With 23 figures OLIVIER BÉTHOUX1,2* and FRANK WIELAND3 1 Freiberg University of Mining and Technology, Institute of Geology, Department of Palaeontology, Bernhard-von-Cotta Str. 2, D-09596 Freiberg, Germany 2 State Natural History Collections of Dresden, Museum of Zoology, Königsbrücker Landstraße 159, 01109 Dresden, Germany 3 Georg-August-Universität, Johann-Friedrich-Blumenbach-Institut für Zoologie und Anthropologie und Zoologisches Museum, Abt. Morphologie, Systematik und Evolutionsbiologie, Berliner Str. 28, 37073 Göttingen, Germany Received 26 February 2008; accepted for publication 30 May 2008 Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species †Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re-interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem-Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total-group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem-representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock-crawlers), suggesting a survivorship of several main Pterygota lineages at the end-Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113. ADDITIONAL KEYWORDS: Chaeteessa – evolution – Mantoida – Mesoptilus – Metallyticus – Palaeozoic – phylogeny – Protorthoptera – Pterygota – wing venation. INTRODUCTION As a major component of continental biodiversity in terms of ecological interactions, species richness, and biodisparity, winged insects (Pterygota) constitute a pertinent model for studying biotic macroevolutionary patterns, and determining the nature of the main processes that framed the observed extant diversity. In that respect, the identification of fossil taxa as stem groups of extant major clades is essential. However, this identification is difficult to achieve *Corresponding author. E-mail: [email protected] for taxa that evolved during (pseudo)radiation events such as Pterygota, because appropriate outgroups are unknown or are not readily identified. Attempts to improve our knowledge of the timing of early events in Pterygota evolution are impeded by the lack of well-documented insect faunas older than the Late Carboniferous, when the group was already represented by taxa as phylogenetically distant as Odonatoptera (damselflies, dragonflies, and stem-relatives) and Archaeorthoptera (grasshoppers, crickets, and stem-relatives) (Wootton, 1981; Belayeva et al., 2002; Grimaldi & Engel, 2005). This lack of data concurs with the difficulties in assigning © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 79 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology 80 O. BÉTHOUX and F. WIELAND MATERIAL AND METHODS We use the insect wing venation nomenclature proposed by Lameere (1922, 1923), modified by Kukalová-Peck (1991), and discussed by Béthoux (2005c). Following the latter contribution, we consider the assumption that main vein sectors were distinct in the pterygotan groundplan (Kukalová-Peck, 1991; among other contributions) as based on insufficient evidence. The M5 pattern favoured by Rasnitsyn (2007; among other contributions) is discussed and discarded by Béthoux (in press). Unless specified, vein and vein sector will be referred to as ‘vein’. Wing venation abbreviations relevant for this contribution are repeated for convenience: ScP, posterior subcosta; RA, anterior radius; RP, posterior radius; M, media; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; AA1, first anterior analis; AA2, second anterior analis. The labels ‘CuA1’ and ‘CuA2’ account for the anterior and posterior branches of CuA, respectively. The two last labels do not imply a hypothesis of homology with the veins CuA1 and CuA2 as usually referred to in the Grylloblattida sensu Storozhenko (2002). We introduce the abbreviation RP*, accounting for the anterior branch of RP. We follow the vein system colourcoding proposed by Kukalová-Peck & Lawrence (2004): blue for the radius system (R); red for the media system (M); green for the cubitus system (Cu). The term ‘arculus’ refers to a strongly sclerotized structure, cross-vein like, free of trachea, and located between two main veins, such as the one occurring between M and CuA in Plecoptera (see Haas & Kukalová-Peck, 2001; Béthoux, 2005c). The term ‘translocation’ refers to the fusion of a vein with another from the base of the latter vein (see Béthoux, 2007c for evidence of this type of transformation in the fossil record; and below). Veins involved in a translocation exhibit no evident independent origin. Specimen preparation follows Béthoux (2005c) except in the following points. Wings were removed from specimens by cutting the body cuticle surrounding the wing base using Vanna’s scissors. Unless specified, right and left wings were mounted on their ventral and dorsal side, respectively. The generic acronym IWC OB accounts for Insect Wing Collection Olivier Béthoux. The acronyms NHM IWC OB and MNCN IWC OB account for insect wing collections prepared by one of us and housed at the Entomology Department, Natural History Museum (London) and the Entomology Department, Museo Nacional de Ciencias Naturales (Madrid), respectively. Specimens referred to as IWC OB only (i.e. without institutional acronym) belong to the personal collection of one of us (OB), temporarily housed in the Museum of Zoology, State Natural History Collections (Dresden). Some specimens involved in this contribution are housed in the collection of the Museum of Zoology, State Natural History Collections (Dresden), in the Oxford University Museum of Natural History (Oxford), and of the Entomology Department, Muséum National d’Histoire Naturelle (Paris). These institutions are referred to as MTD, OUMNH, and MNHN, respectively. For all institutional collections, slides are housed next to the bodies from which wings were removed, or information is provided allowing the rest of the body to be identified. Regarding IWC OB, bodies are housed either in OB’s or FW’s collection. Specimens belonging to institutional collections and mentioned in this contribution are not all illustrated, but our observations can be checked thanks to specimens numbers. Photographs of these specimens can be provided on request. Photographs were taken using a Canon EOS 400D digital camera coupled with a 50 mm macro lens and an extension tube, and driven by corresponding Canon software. High magnification photographs were taken with a Zeiss AxioCam MRc 5 installed on a microscope Nikon Eclipse 600, driven by the soft- © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 known Late Carboniferous insect species to natural groups. This issue resulted in the long-lasting existence of ‘wastebasket groups’, the most important of which is unarguably the ‘Protorthoptera’ as lately defined by Carpenter (1992). The ‘Protorthoptera’ ranges from the Late Carboniferous to the Permian, with fewer representatives in the Mesozoic. It has recently been demonstrated that a number of ‘Protorthoptera’ are genuine Archaeorthoptera (Béthoux & Nel, 2002, 2004, 2005; Béthoux, 2005b, 2007a, b; among others). Béthoux (2005a) suggested that the Archaeorthoptera experienced an unexpected extinction of moderate importance at the end of the Carboniferous (but see Béthoux, 2007a). Therefore, determining the affinities of the remaining ‘Protorthoptera’ might significantly modify our understanding of the early steps of Pterygota evolution, as currently recorded. Hennig (1981) and Carpenter (1992) predicted that the stem-group of the distinctive praying mantids (Mantodea) was to be found within the problematic Palaeozoic ‘Protorthoptera’ group. However, virtually all recent authors have hypothesized that the origin of Mantodea was not earlier than the Early Cretaceous or Late Jurassic (with the oldest material being 135 Myr old; Zherikhin, 2002; Grimaldi, 2003; Lo, 2003). We suggest that the former guess is correct, based on an investigation of the forewing morphology of extant praying mantids, allowing putative ‘protorthopteran’ relatives of Mantodea to be identified. Character states relevant for the inner phylogeny of Mantodea are discussed throughout the paper. CARBONIFEROUS MANTODEA IDENTIFIED MORPHOLOGICAL COMPARATIVE ANALYSIS WING VENATION OF EXTANT MANTODEA There are two main points of the wing venation of extant Mantodea to be investigated. The first point relates to the nature of the vein interpreted as M by Smart (1956), MP by Ragge (1955), and MA by Sharov (1962) (hereafter referred to as the composite stem), and the second being the nature of the vein interpreted as a posterior branch of R (Smart, 1956) or RP (Rs in Ragge, 1955). Further on we will investigate a peculiar case relevant for a few genera within Mantodea, as well as other relevant aspects of the forewing venation of Mantodea. Nature of the composite stem Typically, the first fork of the composite stem [i.e. the stem located between the anterior radius sector (RA) and the most anterior branch of the anterior cubitus sector (CuA)] is located distally, as observed in Metallyticus spp. (Figs 2, 6A, 20A; Wieland, 2008: fig. 14) and other mantids (Ragge, 1955; Smart, 1956; Ramsay, 1990). Regarding the homologization of this composite stem, one of the most illuminating forewings we investigated belongs to Metallyticus violaceus (Burmeister, 1838; Fig. 1). In this atypical wing, the composite stem forks at the first quarter of the wing. The resulting anterior branch, which is significantly stronger than the posterior one, is directed towards RA, runs parallel to it, and departs from it a few millimetres further (Fig. 1B). The location of the point of divergence of this vein from RA is similar to that of the point of divergence of RP from RA, as observed in polyneopteran insects such as Orthoptera (grasshoppers & crickets; Ragge, 1955; Sharov, 1971; Béthoux & Nel, 2002), Plecoptera (stoneflies; Needham & Claassen, 1925; Béthoux, 2005c), Grylloblattida (i.e. putative winged fossil relatives of rock-crawlers; Storozhenko, 1998, 2002), and stem-Dictyoptera (Bolton, 1922; Laurentiaux, 1958; Laurentiaux & Laurentiaux-Vieira, 1980), among others. This similarity of position suggests that the anterior branch of the composite stem, as observed in the atypical forewing of Met. violaceus, is RP. The posterior branch of the composite stem is unarguably provided with a trachea, as indicated by annulation (Fig. 1B, C). This trachea runs within a moderately sclerotized structure, from which it slightly diverges, and runs towards the anterior branch of CuA. According to our observations it ends before reaching CuA. Another narrow trachea diverges from RP whereas this vein runs side by side with RA (Fig. 1B, D). Distal to the apparent end of this narrow trachea, another trachea ‘begins’ (Fig. 1D). The latter runs for a long distance in a weakly sclerotized structure. It forks after some distance (Fig. 1E), with its anterior branch directed towards RP (diverging from RA), but the trachea vanishes before reaching it. The posterior stem of this trachea runs for some distance in a weakly sclerotized structure and vanishes in the area between RP and the anterior branch of CuA. The apparent end of the second trachea diverging from RP (Fig. 1B, D) is likely to be an artefact because there is no other putative origin for the trachea whose beginning is located nearby. We hypothesize that the trachea, for some reason, was damaged at this point. Although large insect species can have very narrow tracheae or lacunae occurring in cross-vein-like structures (e.g. in some phasmids and mantids, O. Béthoux, pers. observ.), this is not the case in Metallyticus spp. In a single case we observed a narrow annulated trachea inside a cross-vein located between the posterior branch of CuA, and CuP (Fig. 1F). We still cannot rule out that the trachea is a vestigial posterior branch of CuA. In the left forewing of the same specimen we observed no lacunae or tracheae inside cross-veins located in the area under scrutiny. In the atypical forewing of Met. violaceus, distal to the divergence of RP from RA, no trachea is present inside the cross-veins occurring between RP and CuA. This absence stands despite their strength and their orientation similar to that of actual RP branches. Therefore, despite the narrowness of the posterior branches of the composite stem, there is no reason to consider that they belong to a tracheal network of secondary importance. The alternative interpretation is that this set of tracheae belongs to a vein system distinct from RP, with which RP could have been fused at the wing base. Such a system located between RP and CuA could only be M (CuA is consistently identified by all previous researchers). If so, M emits a posterior branch when RP diverges from its regular course © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 ware AxioVision 4.4. Original photographs were processed using Adobe Photoshop 7.0. When possible, broken fragments of wings were re-adjusted. Dust and remains not belonging to the wings were manually erased from photographs using the clone stamp tool. We offer to provide original files on request (provided that a CD is sent to the corresponding author). Lastly, we follow Donoghue’s (2005) recommendation for naming fossil groups with respect to extant taxa (i.e. stem-, crown-, and total-groups; see references therein). The Linnaean taxonomic system will be followed for convenience, because no alternative system has ever been applied to the Mantodea. By convention, we use the term Blattodea for the group including cockroaches and termites. 81 82 O. BÉTHOUX and F. WIELAND (Fig. 1B, C), while an anterior branch keeps fused with RP. This anterior branch diverges from RP after some distance (Fig. 1B) and forks (Fig. 1B, E). Under this interpretation M is three-branched, with a simple posterior stem and a forked anterior stem. The first posterior stem is directed towards CuA, the posterior branch of the anterior stem vanishes between RP and CuA, and the anterior-most branch is directed towards RP. Another specimen belonging to Met. violaceus exhibits a different atypical morphology consistent with our interpretation (Fig. 2). In the first fifth of the wing, a supernumerary trachea occurs between RA and M (or composite stem; Fig. 2B). It diverges from RA, deviates towards M, and becomes indiscernible from M. At the second fifth of the wing a similar trachea diverges from M, runs parallel to it, and re-fuses with it (Fig. 2C). Supernumerary narrower © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 1. Forewing venation of Metallyticus violaceus (Burmeister, 1838), atypical morphology (MNHN, Paris, 씸; right forewing, dorsal view; abbreviations: ScP, posterior subcosta; RA, anterior radius; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis). A, photograph. B, detail of the wing venation as located on A. C–F, detail of the wing venation as located on B; arrows indicate various tracheae (see text for explanations). CARBONIFEROUS MANTODEA IDENTIFIED 83 tracheae occur between RA and M at the wing base (Fig. 2B). A plausible interpretation is that the supernumerary main trachea is RP, or a part of it. Anyhow, the occurrence of a supernumerary trachea suggests that the vein occurring between RA and the anterior branch of CuA is composite, as seen in the usual morphology exhibited by Metallyticus spp. and other mantids. The most plausible interpretation is that it is composed of RP and of M. We collected additional evidence of the fusion of (a part of) RP with M in Mantodea. The actual fusion of (a part of) RP with M, occurring at the wing base, was observed in two specimens belonging to Creobroter sp. In the specimen IWC OB 126 (Fig. 3A, B), observation of the ventral side of the right forewing revealed that the fusion of RP with M is not achieved at the wing base, but distal to the divergence of CuA and CuP. The course of RP is distinct from the sclerotized structure in which M runs (Fig. 3A), allowing RP and M tracheae to be distinguished and identified after their annulated structure (Fig. 3B). This atypical organization was also observed on the left forewing of the same individual. In the left forewing of the specimen IWC OB 144 (Creobroter sp.), RP and M run parallel for a short distance before they fuse (Fig. 3C, D). Although less obvious, this was also observed on the right forewing of the same individual. In both cases M is visible more distinctly from the ventral side. A more complicated uncommon organization was observed in the left forewing of the specimen IWC OB 249, belonging to Hymenopus coronatus (Olivier, 1792) (Fig. 3E, F). Viewed from the ventral side, the median trachea is located below another trachea emerging from R (i.e. RP, or a part of it). These tracheae intersect without fusing. As a result, the posterior radial trachea (or a part of it) takes a position posterior with respect to that of the median trachea. After a short distance, both tracheae fuse into the composite stem. The typical organization of this area in praying mantids involves a posterior bulge of the sclerotized © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 2. Forewing venation of Metallyticus violaceus (Burmeister, 1838), atypical morphology (NHM IWC OB 22, London; left forewing, ventral view; abbreviations: ScP, posterior subcosta; RA, anterior radius; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; M, media). A, photograph. B–C, detail of the wing venation as located on A; arrows without labels indicate a supernumerary tracheae. 84 O. BÉTHOUX and F. WIELAND structure in which the radial trachea runs, with M (and its trachea) running underneath this bulge. We hypothesize that the fusion of RP (or a part of it) with M occurs in this area. In Hy. coronatus and Deroplatys desiccata Westwood, 1839 the posterior bulge of the sclerotized structure in which R runs cannot be clearly distinguished from the sclerotized structure in which the composite stem runs (Fig. 3G, H, respectively). Our interpretation that RP would be fused with M from the wing base in all recent Mantodea implies a translocation. This sort of transformation of the wing venation pattern has been evidenced in the fossil group Pantcholmanvissiida (Béthoux, 2007c). We collected © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 3. Evidence for a fusion of posterior radius (RP) with media (M) at the wing base in Mantodea (abbreviations: ScP, posterior subcosta; RA, anterior radius; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis). A, Creobroter sp., detail of forewing base (IWC OB 126, 씹; right forewing, ventral view, reversed); arrows without label indicate the course of the median trachea. B, detail of forewing base, as located on A; arrows indicate the course of the median trachea. C, Creobroter sp., detail of forewing base (IWC OB 144, 씸; left forewing, ventral view). D, detail of forewing base, as located on C. E, Hymenopus coronatus (Olivier, 1792), detail of forewing base (IWC OB 249, 씸; left forewing, ventral view). F, detail of forewing, base as located on E. G, Hymenopus coronatus (Olivier, 1792), detail of forewing base (IWC OB 238, 씹; right forewing, dorsal view). H, Deroplatys desiccata Westwood, 1839, detail of forewing base (IWC OB 155, 씹; right forewing, ventral view, reversed). CARBONIFEROUS MANTODEA IDENTIFIED 85 additional evidence that such a transformation is plausible. In the undetermined Mantinae specimen MNCN IWC OB 27 (Fig. 4), the left forewing exhibits a typical morphology with RP + M distinct from RA (Fig. 4A), whereas R and M form a single stem in the right forewing (Fig. 4B, C). Near the wing base, a narrow trachea that might be the median one fuses with R (Fig. 4C). This atypical case illustrates that complete fusion of a vein with another one, from the wing base (i.e. ‘translocation’ as defined above), occurs in the wild. The atypical morphology exhibited by forewings of the specimen NHM IWC OB 40 (Miomantis paykullii Stål, 1871; right forewing illustrated on Fig. 5E; typical morphology of the species illustrated on Fig. 5A, C) might be considered as an ‘intermediate’ stage, in which RP + M is fused for a long distance with RA (and see below). This case suggests that the apparent suddenness of translocations might be the result of incomplete documentation. Nature of the posterior stem diverging from RA The nature of the vein interpreted as a posterior branch of R (Smart, 1956), or as RP (Rs in Ragge, 1955), generally occurring in Mantodea (indicated by * on Figs 5A, C–E, 7–17), must then be discussed. Importantly, it is absent in Metallyticus spp. (Figs 1, 2, 6A, 20A), Chaeteessa spp. (see Smart, 1956; Figs 6C, 20B), Mantoida spp. (see Smart, 1956: fig. 3, contra the corresponding figure caption, inverted with fig. 4; Figs 6E, 20C), †Lithophotina floccosa Cockerell, 1908 (see Sharov, 1962), and †Arvernineura insignis Nel & Roy, 1996 (for additional fossil taxa, see Nel & Roy, 1996; Grimaldi, 2003), all considered as plesiotypic taxa within Mantodea. Whether this branch belongs to RA or is an anterior branch of RP + M translocated onto RA is to be determined. This vein will be referred to as vein* in the discussion, later on as RP*. In the first instance this vein is emitted posteriorly (i.e. the anterior stem is aligned to the parent stem whereas the posterior stem is oblique), which would be unusual for a branch belonging to RA. In Metallyticus spp. (Figs 1, 2, 6A, 20A), Chaeteessa spp. (Figs 6C, 20B), †L. floccosa and †A. insignis, branches emitted by RA are directed anteriorly, i.e. the posterior branch resulting from a fork is aligned with © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 4. Wing venation of an undetermined Mantinae (MNCN IWC OB 27, 씹; abbreviations: ScP, posterior subcosta; RA, anterior radius; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RP, posterior radius; M, media). A, left forewing (ventral view), typical morphology. B, right forewing (ventral view, reversed), atypical morphology. C, detail of the wing base, as located on B; arrow indicates a narrow trachea directed towards radius (R), interpreted as the median trachea. 86 O. BÉTHOUX and F. WIELAND the parent-stem. This is the typical organization of RA in polyneopteran orders. This peculiar orientation of the vein* probably prompted Ragge (1955) to consider that it belongs to RP. We follow this view but argue that this branch is an anterior stem of RP secondarily translocated from RP + M onto RA. There are two main lines of evidence supporting our view. First, we documented ‘intermediate’ steps between the states ‘anterior branch of RP + M distinct from RA’ (as observed in Metallyticus spp. and Chaeteessa spp.), and ‘anterior branch of RP + M fused with RA from the wing base and diverging near wing apex’ (as observed in most extant praying mantids). In addition, individuals exhibiting atypical morphologies © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 5. Wing venation of Miomantis paykullii Stål, 1871 (씹; abbreviations: ScP, posterior subcosta; RA, anterior radius; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RP, posterior radius; M, media). A, B, specimen NHM IWC OB 38. A, left forewing (ventral view), typical morphology. B, right forewing (dorsal view), atypical morphology. C, D, specimen NHM IWC OB 39. C, left forewing (ventral view), typical morphology. D, right forewing (dorsal view), atypical morphology. E, specimen NHM IWC OB 40, right forewing (dorsal view), atypical morphology. CARBONIFEROUS MANTODEA IDENTIFIED 87 © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 6. Wing venation of praying mantids exhibiting no vein*. A, Metallyticus splendidus Westwood, 1835 (IWC OB 192, 씸; left forewing, ventral view; abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis). B, detail of the area between anterior radius (RA) and posterior radius + media (RP + M), as located on A; arrows without labels indicate short sclerotized structures located between RA and an irregularly sclerotized structure, itself located between RA and RP + M. C, Chaeteessa sp. (MTD, Dresden; right forewing, dorsal view). D, detail of the area between RA and RP + M, as located on C. E, Mantoida maya Saussure & Zehntner, 1894 (specimen IWC OB 102, 씹; left forewing, dorsal view, reversed). F, detail of the area between RA and RP + M, as located on E; arrow indicates an isolated cross-vein. 88 O. BÉTHOUX and F. WIELAND suggesting that the vein* is homologous to a branch of RP + M recur. In forewings of Chaeteessa sp. (Figs 6C, 20B) and of Mantoida maya Saussure & Zehntner, 1894 (Figs 6E, 20C) the course of the anterior branch of RP + M is marked by an inflexion point located opposite the origin of the vein* as in praying mantids considered as more derived. The inflexion point and its location constitute the first evidence of homology between the two structures (similar course and location). Notably, the anterior branch of RP + M is simple in these species. In a specimen belonging to Amorphoscelis sp. (IWC OB 243; Fig. 7), the anterior branch of RP + M is forked opposite of its point of inflexion. In the left forewing (Fig. 7A), both branches remerge into a single one, whereas in the right forewing the anterior branch vanishes in the wing membrane (Fig. 7B). A similar fork of the anterior branch of RP + M was observed in the specimen IWC OB 190, belonging to Amorphoscelis sp. (Fig. 8). However, unlike in the specimen IWC OB 243, comparison of the left and right forewings reveals that the anterior branch of RP + M, or the anterior branch of the latter, fuses with RA in the specimen IWC OB 190. In the left forewing (Fig. 8A, B) the anterior branch of RP + M forks, its anterior trachea fuses with RA, diverges from it after some distance, and re-fuses with the posterior branch of the anterior branch of RP + M (indicated by ° on Fig. 8B). Although we could not observe the course of the trachea along RA, we hypothesize that the trachea fusing with RA and the one emerging from RA after some distance are of the same origin. Areas surrounding the free parts of the trachea fusing with RA are weakly sclerotized. This is unlike the posterior branch of the anterior branch of RP + M, around which sclerotization is continuous. In the right forewing of the same specimen (Fig. 8C–F), the sclerotization surrounding the anterior branch of RP + M stops at some point, and the corresponding trachea fuses with RA without forking (Fig. 8E). The course of the posterior branch of the anterior branch of RP + M as in the left forewing is only materialized by a line of darker setae and is free of trachea (Fig. 8D, E). After some distance a posterior trachea is emitted from RA (Fig. 8D, F). Interestingly, this trachea is distinct from a weakly sclerotized structure whose origin is proximal to that of the trachea, but later joins it. A strongly sclerotized area surrounds this trachea when it reaches the location of the posterior branch of the anterior branch of RP + M as demonstrated in the left forewing (indicated by ° on Fig. 8D). Another trachea later diverges from RA and joins the sclerotized structure just mentioned (Fig. 8D, G), implying that the anterior branch of RP + M is forked while being fused with RA. In Amorphoscelis pulchra Bolivar, 1908 the actual course of the anterior branch of RP + M, whether diverging from RP + M, partly fused with RA, or seemingly diverging from RA, can only be determined after the course of the relevant tracheae. In the right forewing of the specimen NHM IWC OB 23 (Fig. 9A, C), the pattern of sclerotized structures suggests that vein* diverges from RP + M (Fig. 9A). This is contra- © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 7. Wing venation of Amorphoscelis sp. (IWC OB 243, 씹; abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis). A, left forewing (ventral view); left arrow indicates the fork of the anterior branch of posterior radius + media (RP + M), right arrow indicates the branching of RP + M and the fusion of the resulting branches. B, right forewing (dorsal view); arrow indicates the first fork of the anterior branch of RP + M. CARBONIFEROUS MANTODEA IDENTIFIED 89 dicted by the tracheal pattern, which is provided with a broad trachea diverging from RA, whereas a narrower trachea diverges from RP + M and directs towards vein*. In the left forewing of the same individual, the origin of vein* from RA is more evident based on macroscopic observation (Fig. 9B, D), although a trachea of secondary importance diverges from RP + M and fuses with vein*. In the left forewing of the specimen NHM IWC OB 24, the origin of vein* is ambiguous based on macroscopic and micro- © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 8. Wing venation of Amorphoscelis sp. (IWC OB 190, 씹; abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis). A, left forewing (ventral view). B, detail of the area between anterior radius (RA) and posterior radius + media (RP + M), as located on A; left arrow indicates the anterior branch of the anterior branch of RP + M; right arrow indicates the anterior branch of the anterior branch of RP + M emerging from RA; ° indicates the posterior branch of the anterior branch of RP + M. C, left forewing (dorsal view). D, detail of the area between RA and RP + M, as located on C; ° indicates the posterior branch of the anterior branch of RP + M. E, detail of the fusion of the anterior branch of RP + M with RA, as located on D; arrow indicates the RP + M trachea. F, detail of the divergence of the posterior branch of the anterior branch of RP + M from RA, as located on D; arrow indicates the trachea of the posterior branch of RP + M. G, detail of the divergence of the anterior branch of the anterior branch of RP + M from RA, as located on D; arrow indicates the trachea of the anterior branch of RP + M. 90 O. BÉTHOUX and F. WIELAND scopic observations (Fig. 9E, G): a network of narrow tracheae occurs between RA and RP + M. In the right forewing of the same specimen the origin of vein* is ambiguous based on macroscopic observations (Fig. 9F), whereas it apparently diverges from RP + M based on the tracheal pattern (Fig. 9H). In summary, a fork of the anterior branch of RP + M occurs in several Amorphoscelidae [the mono- © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 9. Wing venation of Amorphoscelis pulchra Bolivar, 1908; in C, D, G, H, thickness of arrows indicates strength of indicated tracheae (abbreviations: ScP, posterior subcosta; CuP, posterior cubitus; AA, anterior analis). A–D, specimen NHM IWC OB 23. A, left forewing (ventral view). B, right forewing (dorsal view). C, detail of the connection of the anterior branch of posterior radius + media (RP + M) with anterior radius (RA), as located on A. D, detail of the connection of the anterior branch of RP + M with RA, as located on B. E–H, specimen NHM IWC OB 24. E–F, arrows without label indicate the origin of a branch of anterior cubitus (CuA). E, left forewing (ventral view). F, right forewing (dorsal view). G, detail of the connection of the anterior branch of RP + M with RA, as located on E. H, detail of the connection of the anterior branch of RP + M with RA, as located on F. CARBONIFEROUS MANTODEA IDENTIFIED branch of CuA diverges from RP + M, or from CuA, in the left and right forewing, respectively (Fig. 10A and B, respectively; origin of this branch indicated by an arrow). In the specimen MNCN IWC OB 31, assigned to Oxypilinae, both forewings exhibit a vein* diverging from RA (Fig. 11A, B). In the right forewing, RP + M is provided with an anterior stem fusing with RA (Fig. 11A, C), whereas RP + M is simple in this area in the left forewing (Fig. 11B, D). It suggests that the fusion of vein* with RA is incomplete in the right forewing and complete in the left forewing. It must be noted that the anterior-most branch of CuA diverges from CuA in the right forewing, and from M in the left forewing. In the specimen IWC OB 141, belonging to Creobroter sp., vein* originates from RA in the left forewing (Fig. 12A, B), whereas it originates from RP + M in the right forewing (homology is ascertained by a similar location and a similar number of branches; Fig. 12C, D). This pattern is reminiscent of that observed in Metallyticus spp. (Figs 1, 2, 6A, 20A). Some cases are less evident and do not allow the homology statement we propose to be positively ascertained. It is, however, still worth mentioning them. In the specimen NHM IWC OB 38, belonging to Mi. paykullii, the left forewing exhibits a vein* diverging from RA and a three-branched RP + M (Fig. 5A). The Figure 10. Wing venation of an undetermined Mantinae, atypical morphology (MNCN IWC OB 28; abbreviations: ScP, posterior subcosta; CuP, posterior cubitus; AA, anterior analis). A, left forewing (dorsal view); arrow without label indicates the origin of a branch of anterior cubitus (CuA) from posterior radius + media (RP + M). B, right forewing (ventral view); arrow without label indicates the origin of the same branch of CuA as in A. C, detail of the area between the anterior branch of RP + M and anterior radius (RA) as located on A; arrows indicate inflexion points of the anterior branch of the supernumerary branch of RP + M (see text). D, detail of the area between the anterior branch of RP + M and RA as located on B; arrows indicate the points of fusion and divergence of the supernumerary branch of RP + M with RA. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 phyly of which is uncertain (e.g. Handlirsch, 1925; Chopard, 1949a; Wieland, 2003), therefore they will be referred to as amorphoscelidaeans hereafter]. The anterior branch resulting from this fork approximates RA, is connected to it via a network of tracheae, or is almost fully fused with it. It can then be reasonably hypothesized that a gradual reduction of the tracheal network could have resulted in a definitive and complete fusion of the anterior branch of RP + M with RA, from the wing base. Atypical morphologies exhibited by derived taxa provide additional support to this hypothesis. In the specimen MNCN IWC OB 28, belonging to an undetermined Mantinae, none of the forewings exhibit a vein diverging from RA (Fig. 10). However, the composite stem RP + M is provided with a supernumerary simple anterior stem absent in related taxa (in Mantinae RP + M is typically posteriorly pectinate; see Fig. 4A; see also Fig. 18, relevant for Angelinae). The absence of co-occurrence of the supernumerary stem and of vein* suggests that these structures are homologous. In addition, the supernumerary branch runs parallel to RA for some distance in the left forewing (Fig. 10A, C), whereas it is fused with RA for some distance in the right forewing (Fig. 10B, D). This is clearly reminiscent of the morphology observed in the genus Amorphoscelis Stål, 1871. It must be noted that the anterior-most 91 92 O. BÉTHOUX and F. WIELAND right forewing of the same specimen exhibits no vein diverging from RA, and a two-branched RP + M (Fig. 5B). It cannot be excluded that vein* is fully fused with RA in the right forewing of this specimen. In the specimen IWC OB 237, belonging to Deroplatys lobata (Guérin-Méneville, 1838), the left forewing is provided with a vein* diverging from RA, and RP + M is two-branched (typical morphology; Fig. 12E). This is different from the right forewing, which has a simple RA and a three-branched RP + M (Fig. 12F). This case would imply that vein* is homologous to the anterior branch of the anterior branch of RP + M, rather than to the anterior branch of RP + M (as above). However, it could be hypothesized that, in the right forewing of the specimen IWC OB 237, vein* is fully fused with RA, and that RP + M produces a supernumerary branch in response to this atypical transformation. Such ‘compensation’ is well known in cockroach forewings, in which a reduced number of branches from one vein can be compensated by a higher number of branches of a neighbouring vein. As a result, the ‘density’ of main vein branches is preserved (see Schneider, 1977; Laurentiaux-Vieira & Laurentiaux, 1980; among others; also compare the number of branches of RP + M and of CuA on Fig. 18A and B, respectively). The specimen NHM IWC OB 36, belonging to Statilia maculata (Thunberg, 1784), exhibits an atypical morphology occurring in both forewings, in which no stem diverges from RA, and RP + M is forked (see Fig. 13B, right forewing). In the typical morphology of the species, a vein* diverges from RA, whereas RP + M is simple (Fig. 13A). A compensation cannot be completely ruled out in the specimen NHM IWC OB 36. The similar location of the divergence of vein* from RA in the typical morphology, and of the fork of RP + M in the atypical morphology, however, suggest that the typical vein* is homologous to the atypical supernumerary anterior branch of RP + M. In the specimen NHM IWC OB 33, belonging to Amantis reticulata (De Haan, 1842), we observed that the left forewing has a branch diverging posteriorly from RA and a forked RP + M. The right forewing has a three-branched RP + M. However, the respective location of these branches does not allow a homology statement to be positively ascertained. In summary, the recurrence of atypical patterns suggesting that vein* as exhibited by most mantids is homologous to the anterior branch of RP + M as exhibited by Metallyticus spp. (Figs 1, 2, 6A, 20A) and Chaeteessa spp. (Smart, 1956; Figs 6C, 20B), among others, strongly supports the conclusion drawn from the investigation of the amorphoscelidaean wing venation. Although a full fusion of vein* with RA, resulting in a supernumerary compensatory branch of RP + M, could explain some cases of the atypical © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 11. Wing venation of an undetermined Oxypilinae sp. (MNCN IWC OB 31, 씹; abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis). A, right forewing (dorsal view). B, left forewing (ventral view). C, detail of the connection of the anterior branch of posterior radius + media (RP + M) with anterior radius (RA), as located on A. D, detail of the connection of the anterior branch of RP + M with RA, as located on B. CARBONIFEROUS MANTODEA IDENTIFIED 93 morphologies we document, the most parsimonious explanation, with respect to the transition sequence documented in amorphoscelidaeans, is to consider these atypical morphologies as representing rare occurrences of a plesiomorphic condition. It is unlikely that the median vein takes part in the vein*, therefore we hereafter refer to it as RP*. A further step in the homologization of RP* concerns its number of branches. We observed a number of specimens exhibiting an atypical morphology suggesting that an anterior branch of RP* runs fused with RA. These specimens exhibit a fork of RP* near its point of divergence from RA (Fig. 14). The resulting anterior branch fuses with RA in the left forewing of the specimen IWC OB 242 (Parasphendale affinis Giglio-Tos, 1915; Fig. 14C, D; the right forewing, exhibiting a typical morphology, is illustrated on Fig. 14A, B). In the left forewing of the specimen IWC OB 76 [Ameles decolor (Charpentier, 1825); Fig. 14F], it runs parallel to RA (the right forewing of the specimen IWC OB 76, exhibiting the typical morphology, is illustrated on Fig. 14E). We interpret the atypical morphology of the right forewing of the specimen IWC OB 71 (A. decolor; Fig. 14G) in a similar way: © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 12. Intra-individual variations in the origin of the anterior branch of posterior radius (RP*); on B, D, arrows indicate the origin of RP* (abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; M, media; RA, anterior radius). A–D, forewings of Creobroter sp. (IWC OB 141, 씸). A, left forewing (ventral view). B, detail of left forewing apex. C, right forewing, atypical morphology (dorsal view). D, detail of right forewing apex. E–F, forewings of Deroplatys lobata (Guérin-Méneville, 1838) (IWC OB 237, 씹). E, left forewing (ventral view). F, right forewing, atypical morphology (dorsal view). 94 O. BÉTHOUX and F. WIELAND RP* apparently emits two successive branches (see also Fig. 5C, D), although an additional translocation of a branch of RP + M onto RA cannot be completely ruled out in this case. A fork of RP* was observed in Hierodula sp. (right forewing; MNCN IWC 30), in Omomantis zebrata (Charpentier, 1843) (observed on a single specimen housed at the NHM), and Polyspilota sp. (single specimen housed at the MNCN). Therefore, we hypothesize that RP* is usually forked, and that its anterior branch is usually fused with RA. The compelling evidence we gathered strongly supports the view that in all praying mantids but Metallyticus spp., Chaeteessa spp., Mantoida spp., some taxa currently assigned to the Amorphoscelidae, and some fossil taxa, the anterior branch of RP + M runs fused with RA from the wing base, and diverges distally from RA (or its posterior stem). We propose not to differentiate the stem of RP* that keeps fused with RA, and not to differentiate the stem of RP that keeps fused with M. Additional evidence of the RP + M fusion Now that the fusion of RP* with RA is assessed as a trait of most praying mantids, and because this fusion is absent in Metallyticus spp., Chaeteessa spp., and Mantoida spp., it is worth examining the organization of the area located between RA and RP + M in the latter. As mentioned above, the divergence of RA and RP, from the stem of R, is usually located in the distal part of the wing in polyneopteran insects. Therefore, a proximal area between RA and RP is a trait unknown in these insects. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 13. Wing venation of Statilia maculata (Thunberg, 1784) (씹; abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A, specimen NHM IWC OB 37, left forewing (ventral view), typical morphology. B, specimen NHM IWC OB 36, right forewing (dorsal view), atypical morphology. C, specimen NHM IWC OB 35, left forewing (dorsal view), atypical morphology. D, detail of the connection of the anterior branch of posterior radius (RP*) with posterior radius + media (RP + M), as located on C. CARBONIFEROUS MANTODEA IDENTIFIED 95 In Metallyticus spp. (Fig. 6A, B), short sclerotized structures connect RA to an irregularly sclerotized structure. This structure is free of trachea, parallel to both RA and RP + M, and we interpret it as the ‘remnant’ of the sclerotized structure in which the RP trachea ran before becoming completely fused with M. Along its course, the composite vein RP + M actually switches from this structure to the ‘regular’ one in the aberrant specimen illustrated on Figure 2 (also in the specimen IWC OB 126, Fig. 3A). In our opinion, the occurrence of this ‘remnant’ constitutes by itself additional evidence of the fusion of RP with M in © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 14. Evidence of the branching of the anterior branch of posterior radius (RP*) (abbreviations: CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A–D, forewings of specimen IWC OB 242, belonging to Parasphendale affinis Giglio-Tos, 1915 (씸). A, right forewing (dorsal view). B, detail of right forewing apex, as located on A. C, left forewing (dorsal view, reversed). D, detail of left forewing apex, as located on B; arrows indicate the two branches of RP*. E–G, forewings of specimens belonging to Ameles decolor (Charpentier, 1825). E, right forewing of the specimen IWC OB 76 (dorsal view; 씹). F, left forewing of the specimen IWC OB 76 (ventral view; 씹). G, right forewing of the specimen IWC OB 71 (dorsal view; 씹). 96 O. BÉTHOUX and F. WIELAND Metallyticus spp. In this taxon regular cross-veins occur between RA and RP + M distal to a distinctive point of inflexion of RP + M (Figs 1A, 2A, 6A, 20A). We interpret this point of inflexion as homologous to the point of divergence of RP from R as in other polyneopteran insects (see above). In the specimens of Chaeteessa spp. and Mantoida spp. that we could examine (see Fig. 6C, D and Fig. 6E, F, respectively; and O. Béthoux, pers. observ.), very few cross-veins occur in this area, if any. As in Metallyticus spp., regular cross-veins occur distal to a point of inflexion of RP + M or distal to the first fork of RP + M. We interpret the absence of cross-veins in the basal part of the area between RA and RP + M as evidence of RP being fully fused in the composite stem. As a result of this fusion, an area between RA and RP, which does not occur in putative sister-groups of Mantodea, is present in Chaeteessa spp. and Mantoida spp., with the absence of crossveins being indicative of this novelty. Our opinion is supported by the fact that the absence of cross-veins is more or less correlated with the absence of fusion of RP* (or the anterior branch of RP + M) with RA. In amorphoscelidaeans (lacking this fusion or not), very few or no cross-veins occur between RA and RP + M basal to the first fork of RP + M (based on specimens NHM IWC OB 23–26 and IWC OB 190, 243). In Mantodea exhibiting the fusion of RP* with RA, cross-veins occur in the area that is posterior to RA + RP* [Fig. 13, Fig. 18 and O. Béthoux, pers. observ. on Mantis religiosa (Linnaeus, 1758), among other taxa]. In these taxa, however, this area separates RA + RP* and RP + M, and hence is an area between branches of RP, not between RA and RP. It is then consistent that this area has cross-veins, as is usual in areas separating RP branches in polyneopteran insects (although usually located in a more distal position). Secondary fusion of RP* with RP + M The most derived case in mantodean forewing morphology is probably that exhibited by the species belonging to Schizocephala Serville, 1831, Parathespis Saussure, 1869, Hoplocorypha Stål, 1871, and Danuria Stål, 1856 (among those genera investigated in our survey). In these, we suggest that RP*, diverging from RA + RP*, is secondarily fused with RP + M. This proposition is supported by several observations. In the left forewing of the specimen MNCN IWC OB 24, belonging to Schizocephala bicornis (Linnaeus, 1758), RP* is connected to the single branch of RP + M via two arculuses (i.e. two strongly sclerotized structures free of trachea), the respective tracheae having no direct contact (Fig. 15A, C). In the right forewing of the same specimen, the RP* trachea and RP + M trachea share the same sclerotized structure for some distance and run side by side (Fig. 15B, D). The degree of connection of RP* with RP + M is variable in Hoplocorypha nigerica Beier, 1930: RP* can be distinct from RP + M (Fig. 16A, C), connected to RP + M via a strong cross-vein (Fig. 16B, D), briefly © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 15. Wing venation of Schizocephala bicornis (Linnaeus, 1758) (MNCN IWC 24, 씹; abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A, left forewing (ventral view). B, right forewing (dorsal view). C, detail of the connection of the anterior branch of posterior radius (RP*) with posterior radius + media (RP + M), as located on A. D, detail of the connection of RP* with RP + M, as located on B. CARBONIFEROUS MANTODEA IDENTIFIED 97 connected to RP + M (Fig. 16E, G), or exhibit no clear origin (Fig. 16F, H). In both forewings of the specimen IWC OB 239 (Fig. 17), assigned to Hoplocorypha sp., a short oblique, and moderately sclerotized structure occurs. It possesses a trachea, diverges from RA, and fuses with RP + M. After some distance, the composite stem resulting from the fusion of this ‘vein’ with RP + M forks. Based on the observations mentioned above, we interpret this organization as RP* fusing with RP + M for some distance and then diverging. This is arguably a derived state within Mantodea. Interestingly, this organization occurs as a rare atypical case in S. maculata (Fig. 13C, D). Branching pattern of CuA Another distinctive feature of forewings of most mantids is the branching pattern of the posterior © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 16. Wing venation of Hoplocorypha nigerica Beier, 1930 (씹; abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A, C, specimen NHM IWC OB 29, right forewing (dorsal view). B, D, specimen NHM IWC OB 31, right forewing (dorsal view). C, detail of the origin of the anterior branch of posterior radius (RP*), as located on A. D, detail of the origin of RP*, as located on B. E–H, specimen NHM IWC OB 30. E, left forewing (ventral view). F, right forewing (dorsal view). G, detail of the origin of RP*, as located on E. H, detail of the origin of RP*, as located on F. 98 O. BÉTHOUX and F. WIELAND Figure 18. Wing venation of Euchomenella heteroptera (De Haan, 1842) (NHM IWC OB 28; 씹; abbreviations: CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; M, media; RP, posterior radius); arrows without label indicate the alternative origin of a branch of anterior cubitus (CuA), presumably homologous in both forewings (apparent fusion of posterior subcosta (ScP) with anterior radius (RA) is a result of artefacts of preparation). A, left forewing (ventral view). B, right forewing (dorsal view). branch of CuA (referred to as CuA2), which is generally anteriorly pectinate (i.e. successive anterior branches emitted by CuA2 are simple; see Figs 1A, 2A, 4A, B, 6A, 10B, 13A–C, 14A, C, E, F, 15A, B, 16B, E, F, 18A, 20; but see below). The anterior branch of CuA (CuA1), when branched, is generally posteriorly pectinate (Figs 1A, 2A, 4A, B, 6A, 13A–C, 18A). This branching pattern was noticed by Ragge (1955: 135) and is referred to as internally pectinate. Whether this pattern is diagnostic of Mantodea as a whole must be discussed, as several mantids do not exhibit it, and it can be variable at the levels of individuals and species. In Metallyticus spp., anterior branches of CuA2 can be branched distally, but typically distal to the fork of the corresponding posterior ‘sister-branch’ (i.e. anterior branches of CuA/CuA2 are branched distal to posterior branches of CuA2; Figs 1A, 2A, 6A, 20A; see scheme in Fig. 19A). This is a constant feature of Metallyticus spp. forewing venation pattern (see figures; eight forewings in IWC OB; and 18 observed forewings of OUMNH, of which 16 exhibit an anteriorly pectinate CuA2, 2 whereas two exhibit an ambiguous pattern). In the specimen NHM IWC OB 28 belonging to Euchomenella heteroptera (De Haan, 1842), the first anterior branch of CuA2 in the left forewing (Fig. 18A) is arguably homologous to the first posterior branch of CuA1 as in the right forewing (Fig. 18B; compare also Fig. 16A, B). Similar ‘switches’ occur relatively commonly in mantids. Four out of 13 forewings of Ameles decolor (Charpentier, 1825) present in IWC OB and relevant for comparison exhibit an anteriorly pectinate CuA2 (and CuA) as schematized in Figure 19B (see Fig. 14E, F). Conversely, the remaining nine forewings exhibit a dichotomously branched CuA as schematized in Figure 19C (see Fig. 14G; a single individual can exhibit both morphologies). The point of origin of the branch indicated by ‘c’ in Figure 19B is located opposite the point of origin of the branch indicated by ‘c’ in Figure 19C. It suggests that these branches are homologous, and that a dichotomously branched CuA could derive from a posteriorly pectinate CuA. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 17. Wing venation of specimen IWC OB 239, belonging to Hoplocorypha sp. (씹; abbreviations: CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A, left forewing, ventral view. B, detail of the origin of the anterior branch of posterior radius (RP*), as located on A; arrow without label indicates the origin of (the posterior branch of) RP*. CARBONIFEROUS MANTODEA IDENTIFIED A C a b c d D a b c d E a b c d a b c d Figure 19. Possible transition from an anteriorly pectinate branch of the posterior branch of anterior cubitus (CuA2) to a posteriorly pectinate anterior cubitus (CuA). A, branching pattern of CuA as in Metallyticus spp. B, one of the branching patterns of CuA exhibited by Ameles decolor (Charpentier, 1825). C, most frequent branching pattern of CuA as in A. decolor (Charpentier, 1825). D, branching pattern of CuA as in Amorphoscelidae. E, branching pattern of CuA as in Chaeteessa spp. In amorphoscelidaeans, CuA is typically dichotomously branched, with a very basal first fork of the posterior branch, as schematized in Figure 19D (see Figs 7, 8, 9A, B, E). This morphology could have equally derived (1) from the pattern presented in Figure 19A, as a consequence of the reduction of the number of CuA2 branches; or (2) from the pattern presented in Figure 19C, as a consequence of a relocation of the point of origin of the ‘a’ and ‘b’ branches, and the point of origin of the ‘c’ and ‘d’ branches. Interestingly, in the right forewing of the specimen NHM IWC OB 24 (Fig. 9F), belonging to A. pulchra, the branch indicated by ‘b’ in Figure 19D and E diverges from the stem that gives rise to branches ‘c’ and ‘d’, resulting in the pattern presented in Figure 19E (whereas the typical morphology of the species is schematized in Fig. 19D). This pattern is identical to that exhibited by Chaeteessa spp. (Figs 6C, 20B; see also Nel & Roy, 1996: fig. 3), in which CuA as a whole is posteriorly pectinate. From the available data, in Chaeteessa spp., the second fork of CuA is consistently closer to the first fork than to the third fork. This suggests that the branch indicated by ‘b’ in Figure 19E is homologous to the branch indicated by ‘b’ in Figure 19D, and that the branching pattern of CuA exhibited by Chaeteessa spp. is likely to be derived from a dichotomously branched CuA. In summary, all branching patterns of CuA as observed in Mantodea could have derived from an anteriorly pectinate pattern. However, the polarity of character state transformation is not evident. Ascertaining the position of Metallyticus Westwood, 1835 with respect to other mantids is essential in this situation. At this step, it can only be considered as plausible that CuA is anteriorly pectinate in the groundplan of Mantodea forewing venation. A similar pattern is present in Cnemidolestodea (see Béthoux & Nel, 2004, 2005; Béthoux, 2005a, 2007a), but this condition is considered as derived within Archaeorthoptera, as plesiotypic representatives of the latter taxon exhibit a posteriorly pectinate CuA (as it is, fused with CuPa) (Béthoux, 2003, 2006; see also Béthoux, 2005b; Prokop & Ren, 2007). Regarding Plecoptera, the families Eustheniidae, Diamphipnoidae, and Austroperlidae have an anteriorly pectinate CuA (see Zwick, 1979; McLellan, 1996; Béthoux, 2005c). According to the phylogenetic scheme of Zwick (2000), this is likely to represent the ancestral state of Antarctoperlaria (in Gripopterygidae CuA is forked or simple, therefore the character describing the branching pattern is not applicable). In Euholognatha the character is not applicable as CuA is forked or simple (Béthoux, 2005c). The ancestral condition in Systellognatha cannot be ascertained: whereas Pteronarcyoidea exhibit an anteriorly pectinate CuA, Perloidea exhibit a posteriorly pectinate CuA (when applicable; see Needham & Claassen, 1925; Béthoux, 2005c). Thus, although an ancestral plecopteran condition ‘CuA anteriorly pectinate in forewings’ can be drawn from the available information (and based on the phylogenetic scheme in Zwick, 2000), the robustness of this inference is low. It is equally plausible that the plecopteran groundplan involves a simple or forked CuA, hence the character describing the branching pattern of CuA would be inapplicable. Regarding putative stem-Dictyoptera (Bolton, 1922; Laurentiaux, 1958; Laurentiaux & Laurentiaux-Vieira, 1980) and stem-Blattodea (see Rehn, 1951; Schneider, 1977, 1978, 1980a, 1980b, 1982, 1983), virtually all of them exhibit a posteriorly pectinate CuA. Within Blattodea, some taxa such as the Polyphagidae exhibit an anteriorly pectinate CuA2 (see Rehn, 1951), but the wing venation of the members of this family is extremely variable. The available data on extant taxa (morphology and phy- © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 B 99 100 O. BÉTHOUX and F. WIELAND logeny) do not allow the ancestral Blattodean condition to be ascertained. Anyhow, a posteriorly pectinate CuA2 could be diagnostic of Mantodea, or of (some) Dictyoptera. Summary A synthesis of our comparative morphological analysis is presented in Figure 20, with a sample of various mantodean forewings. The left forewing of the specimen IWC OB 193 (Met. splendidus; Fig. 20A) is atypical in that RP + M has two main stems (RP + M is typically posteriorly pectinate in Metallyticus spp., see Figs 1A, 2A, 6A). Branches resulting from the fork of RP + M might be RP and M diverging distally. If so, M would be three-branched. This atypical morphology is presented as a matter of record. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 20. Homologization of forewing venation in Mantodea (abbreviations: CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; M, media; RA, anterior radius; RP, posterior radius). A, Metallyticus splendidus Westwood, 1835 (based on specimen IWC OB 193, left forewing, 씸). B, Chaeteessa filata Burmeister, 1838 (based on specimen described by Smart, 1956, housed at OUMNH). C, Mantoida maya Saussure & Zehntner, 1894, first interpretation (based on specimen IWC OB 97, left forewing, 씹). D, Tarachodula pantherina (Gerstaecker, 1869) (based on specimen IWC OB 96, left forewing, 씹). CARBONIFEROUS MANTODEA IDENTIFIED FOSSIL MATERIAL This section is based on literature data (see below) and new investigations. We reinvestigated material assigned to †Mesoptilus dolloi Lameere, 1917, a Late Carboniferous species yielded by the famous French deposit of Commentry (Lameere, 1917; Fig. 21; c. 310 Mya). The holotype specimen MNHN DP R51159 (Fig. 21) and the specimen MNHN A27122 (Fig. 22) were observed. Although the wings are poorly preserved on the specimen MNHN A27122, it is assigned to the same species as MNHN DP R51159 based on: the similar size; identical wing coloration; similarly long, oblique, and numerous branches of ScP; similar (fore-)leg morphology. A new restoration of the wing venation of †Mes. dolloi is provided on Figure 21C. The area where the origin of RP (diverging from R) is likely to have occurred is poorly preserved on the holotype, and is not visible on the specimen MNHN A27122. The location of the origin of RP is inferred from related taxa (see below). We also consider the leg morphology of the specimen MCZ 5875, paratype of †Homocladus grandis Carpenter, 1966. Forewing morphology †Mes. dolloi is affiliated with a number of Palaeozoic genera such as †Strephoneura Martynov, 1940, †Spargoptilon Kukalová, 1965, †Homocladus Carpenter, 1966, †Paracladus Carpenter, 1966, and †Graticladus Novokshonov & Aristov, 2004. Most of these genera have been assigned to the family †Strephocladidae by Carpenter (1992). As the monophyly of this family is uncertain (see below), the corresponding species will be referred to hereafter as strephocladidaeans. All of them exhibit an anteriorly pectinate CuA or CuA2, suggesting that they have dictyopteran or mantodean affinities (see above). Carpenter (1966) noted the difficulty in identifying the median system in these taxa. In most of them, at least a branch of M is connected to RP, and it cannot be ruled out that some anterior branches of CuA fuse with M for some distance. However, in his interpretation of the wing venation of †Ho. grandis, Carpenter (1966) consistently identified M as a system constituted by a forked anterior branch and a simple posterior branch. In †Graticladus severus Novokshonov & Aristov, 2004, in which there is no connection of M with RP, M is also composed of a forked anterior branch and a simple posterior branch, just as in the atypical forewing of Met. violaceus (Fig. 1). We hypothesize that this is also the case in †Mes. dolloi. If so, in this taxon, the most anterior stem of M is fused with RP, the second stem of M (the first posterior branch of the anterior branch) runs free, and the posterior stem of M is fused for some distance with the anterior branch of CuA (such a fusion of the anterior branch of CuA with M has been observed in extant Mantodea, see Fig. 10A). Whether the anterior branch of CuA is fused with M from the wing base or fuses with it later, via an arculus (as indicated on Fig. 21C), cannot be determined with certainty. Although the location of the origin of RP and the exact number of branches belonging to M in †Mes. dolloi could be debated, the anterior-most branch of M unarguably fuses for some distance with RP. This is evidenced by the long distance separating the point of divergence of the anterior branch of M from more distal branches (as it is, of RP). It is noteworthy that the location of actual RP branches is similar to that observed in forewings of Metallyticus spp. This is also the case in †Ho. grandis and †Homocladus ornatus Carpenter, 1966 (see Carpenter, 1966; figs 4, 5, respectively). An incomplete fusion of M with RP in forewings was also acquired within Archaeorthoptera (= totalOrthoptera) (see Béthoux & Nel, 2003, 2004, 2005), but the most likely plesiotypic members of this taxon lack this fusion (Béthoux, 2003, 2006). It is absent in forewings of Plecoptera (Needham & Claassen, 1925; Béthoux, 2005c), occurs sporadically in Grylloblattida in which it is likely to be a derived character state (see Storozhenko, 2002: fig. 393, nodes 30, 64; atypical morphologies exhibiting a fusion of an anterior branch of M with R occur sporadically, see Tillyard, 1928a: fig. 14; Tillyard, 1928b: fig. 5), and is absent in stem-Dictyoptera (Bolton, 1922; Laurentiaux, 1958; Laurentiaux & Laurentiaux-Vieira, 1980), among others. Therefore, coupled with the branching pattern of CuA reminiscent of that observed in Metallyticus spp., the character state ‘anterior branch of M fused with RP for a long distance’ is strong evidence for considering †Mes. dolloi and its relatives as stemMantodea. Leg morphology Several legs are preserved on the holotype of †Mes. dolloi, but none can positively be identified as a foreleg from their respective position, because of the © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 In conclusion, crown-Mantodea are characterized by the following forewing character state: RP and M fused from the wing base (as opposed to: M and RP distinct). A putative diagnostic character state is: CuA internally pectinate (hence CuA2 anteriorly pectinate; as opposed to: posterior branch CuA simple, or without consistent branching pattern). Within Mantodea, plausible plesiomorphic character states of the forewing morphology are as follows (mainly based upon the morphology exhibited by Metallyticus spp., typical and atypical, Chaeteessa spp., and fossil taxa mentioned above): RA with anterior branches; and M branched. 101 102 O. BÉTHOUX and F. WIELAND © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 21. Mesoptilus dolloi Lameere, 1917 (holotype specimen MNHN DP R51159; abbreviations: CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; M, media; R, radius; RA, anterior radius; RP, posterior radius). A, habitus (under ethanol, polarized light). B, detail of a leg, as located on A (dry-ethanol composite photograph); arrows indicate remains of spines. C, drawing of the right forewing. D, right forewing (negative imprint, reversed; dry-ethanol composite photograph, polarized light). CARBONIFEROUS MANTODEA IDENTIFIED 103 disarticulation of the individual (Fig. 21A). However, one leg preserves evidence of stout spines located on the femur (Fig. 21B). They are incompletely preserved as they are broken off along the line of splitting (the opposite imprint of the specimen is unfortunately not available). Body parts are also partly disconnected in the specimen MNHN A27122 (Fig. 22A). In this specimen, however, stout spines with a bent acron and a broad base are well preserved on the femur of one leg, and two spines are preserved on the tibia (Fig. 22B). The leg is overall poorly preserved (the opposite imprint of the specimen is not available). The shape of the spines located on the femur as observed on the specimen MNHN A27122 are indicative of a raptorial leg, most probably a foreleg. Connection with the morphology exhibited by extant praying mantids is made difficult by the poor preservation of the fossil material, but it is likely that †Mes. dolloi had raptorial forelegs. Regarding †Ho. grandis, Carpenter (1966: 62) mentioned that an incomplete leg, with two rows of heavy spines on the tibia and smaller spines on the femur, is preserved next to the wing of the paratype. This leg is likely to belong to the same individual (Fig. 23A). Examination of photographs of the specimen revealed that two parallel rows of spines occur on the ventral side of the tibia, without additional spines between these rows (Fig. 23B). The spines are sturdy, pointed, and variable in length. About 20 spines are discernible on each row. They stand close together, point slightly apicad, and are relatively long with respect to the diameter of the tibia. These characters (number, closeness, and length of the spines) can be taken into account for a comparison with other polyneopteran taxa. In extant Mantodea the fore tibia similarly carries two parallel rows of long spines. Some other groups of Polyneoptera display spines along the ventral side of their fore tibiae, arranged in rows, often in correlation with a carnivorous lifestyle. Among them, for instance, are the Mantophasmatodea (see Klass et al., 2003: fig. 3; Klass, Zompro & Adis, 2003: fig. 10.1A, B; Zompro et al., 2003: fig. 4), and some Ensifera, e.g. Listroscelidinae (Beier, 1955: fig. 45; Groll & Günther, 2003), Saginae (Kaltenbach, 1967: fig. 127, tab. 7, Kaltenbach, 1990: figs 2–5), and Gryllacrididae (Karny, 1937: pl. 1: figs 5, 9, pl. 3: figs 5, 9; Chopard, © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 22. †Mesoptilus dolloi Lameere, 1917 (specimen MNHN A27122). A, habitus (under ethanol, polarized light). B, detail of a leg, as located on A (dry-ethanol composite photograph). C–D, details of a leg, as located on B (dry-ethanol composite photographs); arrows indicate spines. 104 O. BÉTHOUX and F. WIELAND 1949b: fig. 297). However, we observed no such arrangement of spines in the Blattodea we investigated (Blattidae, Blaberidae, Blattellidae, Lamproblattidae, Cryptocercidae, Polyphagidae). Instead, in cockroaches the spines are arranged around the tibia in an unordered manner (see Roth, 1991: figs 19.1, 19.2B, 19.8). The number of sturdy spines in a single row on the tibia of †Ho. grandis was probably about 20. Exhibiting more than 15 spines is a state shown by some Mantodea, e.g. some Hymenopodinae, Mantidae, Toxoderidae, and Empusidae (Roy, 1999; F. Wieland, pers. observ.). These taxa, however, are generally considered to be derived within Mantodea (Beier, 1968; Grimaldi, 2003; Svenson & Whiting, 2004; Grimaldi & Engel, 2005). Most extant species exhibit fewer than 15 spines per row. This is inclusive of all putative basal Mantodea, in addition to the fossil species identified so far (see Grimaldi, 2003). The same is true for other Polyneoptera. In Mantophasmatodea the number of tibial spines is lower (see Klass et al., 2003: fig. 3D; Engel & Grimaldi, 2004: fig. 3; Arillo & Engel, 2006: fig. 5). In Ensifera, the number of tibial spines is often much lower than 15, with an exception being Saga Charpentier, 1825, in which they may reach a number of up to 15 spines per row (Kaltenbach, 1967: tab. 7). Some Ensifera exhibit strongly elongated spines, although in smaller numbers (e.g. Listroscelidinae and Gryllacrididae; see Karny, 1937: pl. 1: figs 5, 9, pl. 3: figs 5, 9; Chopard, 1949b: fig. 297; Beier, 1955: rig. 45; Hale & Rentz, 2001: fig. 6.2A; Groll & Günther, 2003). Otherwise, the spines are smaller than those of †Ho. grandis. As far as we are aware, among Polyneoptera, the high number and closeness of spines present on the tibia of †Ho. grandis is displayed only by Mantodea. A number of important features could not be observed on the specimen. For example, foreleg spines in extant Mantophasmatodea are mere cuticular out- growths (F. Wieland, pers. observ. in Karoophasma biedouwensis Klass et al., 2003; and K.-D. Klass, pers. comm.), whereas in Ensifera, Mantodea, and Blattodea we observed distinct sutures between the spines and the tibia, indicating that the spines are of setal origin. Unfortunately, the occurrence of sutures separating the spines from their bases cannot be assessed in †Ho. grandis. In addition, an apical spur on the fore tibia, formed by the enlarged distal anteroventral spine, is present in all extant (and most fossil) Mantodea except Chaeteessa spp. (e.g. Beier, 1968; Roy, 1999; Grimaldi, 2003; Klass & Meier, 2006). Whether this trait is plesiomorphic (Beier, 1968) or represents a secondary loss in Chaeteessa spp. (Klass & Meier, 2006; also implicated by the phylogenetic trees by Grimaldi, 2003 and Grimaldi & Engel, 2005), is still debated. Unfortunately, the fore-tibial apex of †Ho. grandis is not preserved, therefore neither the insertion of the tarsus on the tibia, nor the occurrence of an apical spur, can be assessed. In conclusion, the arrangement of spines on the fore tibia excludes close relationships with Blattodea, and the number and closeness of spines suggest that the known preserved leg of †Ho. grandis is a raptorial foreleg mostly similar to that of some extant Mantodea. It must be acknowledged that important traits of the tibial morphology that would help to diagnose †Ho. grandis as a Mantodea are unknown (presence or absence of tibial spur, lateral or terminal tarsal insertion, setal origin of tibial spines). If a close relationship with Mantodea is assumed, the high number of tibial spines would be homoplasic, as it is shared by derived extant Mantodea, the wing venation of which is far more derived than that of †Ho. grandis. Summary As a result of this comparative analysis, †Mes. dolloi and its Palaeozoic relatives can be considered as © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Figure 23. †Homocladus grandis Carpenter, 1966 (paratype specimen MCZ 5875, courtesy of the Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA). A, left forewing and associated leg remain. B, detail of the leg, as located on A (dry-ethanol composite photograph). CARBONIFEROUS MANTODEA IDENTIFIED DISCUSSION ATAVISM, POLYMORPHISM, AND SAMPLE SIZE Our investigation relies in part on the occurrence of atypical features exhibited by few individuals. In the case of Met. violaceus, we observed two forewings on which two tracheal systems occur between RA and the anterior branch of CuA. We interpret these tracheal systems as RP and M. A unique trachea was observed in other specimens we investigated (eight forewings in IWC OB). Whether the unusual morphology exhibited by these specimens is an atavism (i.e. resurgence of an ancestral character state), or if these specimens are extreme cases of a regular polymorphism, is unknown. In any case, the occurrence of these atypical features has to be explained. We interpret the anterior trachea as RP because we see no other explanation consistent with the apparent lack of RP in this species and in Chaeteessa spp. and Mantoida spp. This explanation is also consistent with the morphology exhibited by putative fossil relatives. In species belonging to Amorphoscelis it is clear that the connection of the anterior branch of RP + M (i.e. RP*) with RA is a matter of intra-individual to intraspecific polymorphism. The apparent absence of a connection between RP* with RA (as in Fig. 8) was observed in Amorphoscelis singaporana Giglio-Tos, 1915, Amorphoscelis siebersi Werner, 1933, and Amorphoscelis annulicornis Stål, 1871 (three, one, and four specimens, respectively; observed under a binocular microscope, without preparation allowing course of trachea to be observed; NHM, London). The formation of an eye as illustrated on Figure 9 was observed in Amorphoscelis orientalis Giglio-Tos, 1913 (with few cases of complete fusion over 17 observed specimens; NHM, London), Amorphoscelis tigrina Giglio-Tos, 1913, Amorphoscelis austrogermanica Werner, 1923 (eight and two specimens observed, respectively), and Amorphoscelis pulchra (about 20 specimens observed). The state of fusion of RP* with RA ranges from absent to complete in Amorphoscelis nigriventer Beier, 1930. The fusion is consistently complete in Amorphoscelis grisea Bolivar, 1908 and Amorphoscelis pulchella Giglio-Tos, 1913 (four and five specimens observed, respectively). Considering the sample size, a typical morphology can hardly be outlined for this group of species. This variability could be explained by the fact that these species might be successive ‘intermediates’ between the more plesiotypic taxa in which RP* diverges from RP + M and the more derived taxa in which RP* diverges from RA + RP* (like some genera currently assigned to the Amorphoscelidae, such as Perlamantis Guérin-Méneville, 1843; Paramorphoscelis Werner, 1907), or by poor sampling. We recurrently observed extant specimens exhibiting an RP* presumably diverging from RP + M, although the typical morphology of their respective species is a divergence of RP* from RA (Figs 5B, 10, 11A, 12C, D, F, 13B). Similar to the case of Met. violaceus, it is difficult to determine whether such morphology is an atavism or an extreme case fitting within the range of a regular polymorphism. In any case, the fact that these atypical morphologies recurrently occur indicates that they are the result of a consistent phenomenon. Considering the variability of this character state as exhibited by Amorphoscelis spp., the most plausible explanation is to consider the character state ‘RP* diverging from RP + M’ as a plesiomorphic condition. Regarding the secondary fusion of RP* (diverging from RA + RP*) with RP + M as observed in some species currently assigned to the Thespidae, all ‘intermediate’ conditions could be observed, relevant at the level of intraspecific to intra-individual variability. Again, in most cases, a typical morphology cannot be properly outlined without a significant sample. In summary, regarding the fusion of RP with M, only rare occurrences of atypical morphologies are documented (as it is, in Met. violaceus). Taxa in which this putative fusion occurred are extinct, and a good sample of relevant taxa is not available from the fossil record. In the case of the translocation of RP* onto RA, species lacking the translocation, polymorphic species (including relevant variation at the level of individuals), and the rare occurrence of individuals lacking the translocation but belonging to species in which the presence of this transformation is typical, are all properly documented. Regarding species in which the secondary fusion of RP* (diverging from RA + RP*) with RP + M is the typical morphology, the rare occurrence of specimens exhibiting a lack of this fusion is not properly documented. We assume that the sample we investigated was not significant enough for sufficient documentation of this case. It must be pointed out that the atypical morphologies documented on Figures 5B, D, E, 12C, D, and 13B, D, were picked out of a sample of about 40 specimens (NHM, London), 20 specimens (IWC OB), and about 40 specimens (NHM, London), respectively. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 stem-Mantodea on the basis of two forewing character states: anterior branch of M fused with RP for a long distance (completely fused in crown-Mantodea; as opposed to M and RP distinct); and CuA2 anteriorly pectinate (as opposed to posteriorly pectinate, or no consistent branching pattern). This proposition is supported by evidence suggesting that strephocladidaeans exhibited raptorial forelegs, the morphology of which is still, however, poorly known. 105 106 O. BÉTHOUX and F. WIELAND MODE OF VEIN FUSION The fusion of RP* (emerging from RP + M) with RA is probably the best documented vein fusion assessed in winged insects. Based on our observation, successive steps were hypothesized. The first step is a branching of the anterior branch of RP + M (i.e. RP*), with the possible re-fusion of branches resulting from this fork (Fig. 7). The next step is a brief fusion of the anterior branch of RP* with RA (Fig. 8A). The following step is a fusion of the whole RP* with RA for a short distance, with distinct origins of its posterior and anterior branches from RA, both branches re-merging into a single RP* (Fig. 8B). From this point, various degrees of fusion via a network of tracheae can be observed, (Fig. 9), and, thereafter the full fusion of RP* with RA for a short distance (Figs 10B, D, 11A), followed by a complete translocation of RP* with RA from the wing base (Figs 11A, 12A, B, E, 13A, 14A, E, 15), are hypothesized. The essential part of the scenario is the formation of an ‘eye’ delimited by the anterior and posterior branch of the vein undergoing fusion as a preliminary step towards a complete fusion. Future investigations in insect wing venation might provide evidence for a recurrent pattern. We demonstrate additionally that, at least in case of a fusion of veins, the tracheal network pattern could be distinct from the sclerotization pattern (i.e. the ‘classical venation pattern’; notably Fig. 8F). This fact is to be taken into account in future investigations, in particular for homologization of wing venation of fossil taxa, for which only information on the sclerotization pattern is accessible (at least for those fossils preserved as compression). UNRESOLVED ISSUES The number of branches of the composite stem RP + M to be assigned either to RP or M is difficult to assess. In one of the atypical specimens of Met. violaceus, the free part of M is forked, with its anterior branch forked after some distance (Fig. 1). Based on our survey, in Mantodea as a whole, RP + M can either be simple or branched. Some specimens belonging to species generally exhibiting a simple RP + M may have it branched (as found in Ma. religiosa). Mapping the states of this character onto a phylogeny of Mantodea might help to resolve the point. Homologization of the forewing anal area is not easy. In Metallyticus spp. the first convex vein posterior to CuP (so-called AA1) can either be simple or forked, and always reaches the posterior wing margin. This is unlike in Chaeteessa spp., Mantoida spp., and †A. insignis, in which AA1 is always simple and vanishes in the wing membrane between CuP and AA2. In contrast, several amorphoscelidaeans exhibit an organization similar to that observed in Metallyticus spp. (see Figs 7–9), although these taxa are considered as derived with respect to Chaeteessa spp. and Mantoida spp. This leaves some doubt as to the homologization of this vein in more derived mantids, in which it can vanish in the wing membrane, as in Chaeteessa spp. and Mantoida spp. (although it can also reach CuP, or fuse with another anal vein). It cannot be ruled out that the anterior branch of AA1 as exhibited in Metallyticus spp. and some amorphoscelidaeans actually fuses with CuP in more derived mantids. In Chaeteessa spp. and Mantoida spp., AA1 could be primarily simple. Alternative homologizations should be tested by congruence with other characters. Finally, homologization of the hind wing anal area is out of the scope of this contribution. A survey of hind wing anal area of the Dictyoptera as a whole, including fossil material, would be necessary. The interpretation provided by Smart (1956) regarding the morphology of Chaeteessa spp. hind wings must be reviewed. FOREWING VENATION AND PHYLOGENY The question of the homologization of mantodean wing venation with respect to that of other winged insects was unresolved because of multiple and inconsistent propositions (see Ragge, 1955; Smart, 1956; Sharov, 1962; Balderson, 1991; Deitz, Nalepa & Klass, 2003). This situation is a common case for polyneopteran insects (see a summary of various interpretations regarding Orthoptera in Béthoux & Nel, 2002: tabs. 1, 2; for Plecoptera see Béthoux, 2005c: tab. 1; for Blattodea, see Rehn, 1951: 14–15), and precluded a correct identification of fossil taxa. Resulting from our comparative analysis, all crownMantodea exhibit a fusion of the main stem of RP with M, occurring at the forewing base. It must be regarded as an apomorphic trait of the order, as it is absent in Blattodea (Rehn, 1951; Schneider, 1977, © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 Interestingly, we recurrently observed a rare atypical morphology in which RP + M, or one of its anterior branches, fuses for a more or less long distance with RA + RP* [Figs 4B, 5E, and O. Béthoux, pers. observ. on specimens belonging to Ma. religiosa, Blepharopsis mendica (Fabricius, 1775) (both in IWC OB), Creobroter sp. and D. lobata (both in FW’s collection), specimen MNCN IWC OB 35, belonging to Parastagmatoptera flavoguttata (Serville, 1839), and specimens belonging to Hierodula sp. housed at the MNCN]. The fixation of this trait, which can be seen as part of a trend involving the fusion of RP* with RA, might be the next step of forewing venation evolution in Mantodea. CARBONIFEROUS MANTODEA IDENTIFIED SISTER-GROUP AND ORIGINATION DATE OF MANTODEA Our work suggests that taxa more closely related to the modern Mantodea than to any other modern group, i.e. belonging to stem-group Mantodea, existed as early as the Late Carboniferous (c. 310 Mya). This contrasts with the views of several recent authors, (Zherikhin, 2002; Grimaldi, 2003; Lo, 2003; among others), who stated that the origin of the order Mantodea was not to be assumed to be earlier than the Early Cretaceous or Late Jurassic (with the oldest material being 135 Myr old). However, the question of a Mesozoic sister-group of Mantodea is controversial. Vršanský, Vishniakova & Rasnitsyn (2002) indicated that the families †Blattulidae, Polyphagidae, †Umenocoleidae, and the order Mantodea are related on the basis of ‘venation simplified, with hind wing M fewer than five-branched, CuP simple’. The fact that this statement could allow the order Plecoptera to be part of this assemblage speaks for the vagueness of this argument. Notably, a ‘simple CuP’ is common to all Neoptera but Archaeorthoptera. Moreover, this proposal obliterates the plethora of character states that differentiates the Blattodea from the Mantodea, the most important of which is the loss of a differen- tiation of R into RA and RP. This differentiation is a strict apomorphy of the former group (see above), whereas Mantodea exhibits the antonymic plesiomorphic state (i.e. RA and RP differentiated). Besides the absence of clearly defined synapomorphies, Vršanský et al.’s (2002) scenario implies a reversion of this character. The confusion increases once the conclusions drawn by Vršanský et al. (2002) are compared to those of Vršanský (2002): in the former, the family †Raphidiomimidae is unrelated to the Mantodea, whereas in the latter, it is suggested that they are sister-group related on the basis of ‘predacious way of life, indicated RS [RP] in the forewing, wings less rigid’ (Vršanský, 2002: fig. 27, node 24). However, strict homology between the predacious organs involved in the respective group has never been demonstrated. Even Vršanský (2002: 14) states that the legs of the †Raphidiomimidae and of the Mantodea differ in the length of the tibia. Once again, the vagueness of Vršanský’s (2002) ‘diagnosis’ leaves enough room for the inclusion of the orthopteran †Mesotitanini Tillyard, 1925 (as defined in Béthoux, 2007c). Importantly, we were unable to identify RA from RP in the representatives of the †Raphidiomimidae. In the same paper Vršanský (2002) suggests that the family †Liberiblattinidae also possesses a differentiated RP in forewings, although this family is supposedly more basal than the †Raphidiomimidae and Mantodea (after fig. 27 therein). Our observations of the available illustration of the holotype of the type-species of †Liberiblattina Vršanský, 2002 (Vršanský, 2002: fig. 8), the type-genus of the family, led us to believe that there is no differentiated RA and RP in this taxon. Other characters provided by Vršanský (2002: 10) as indicative of a close relationship of the †Liberiblattinidae with the Mantodea are inconsistent [notably, the character state ‘forewing long, straight’ is shared by the group Blattoidea + Isoptera + Polyphagoidea + (†?)Umenocoleoidea (fig. 27, node 11)], not shared by basal Mantodea, not present in †Liberiblattinidae, and/or vague. Lastly, a branched ScP, the only putative apomorphy of the †Liberiblattinidae + Mantodea, is also present in the Carboniferous taxa we consider as stem-Mantodea, and might be a plesiomorphy at the level of Neoptera. In summary, there is no strong positive evidence for a sister-group of Mantodea occurring in the Late Jurassic and Early Cretaceous. In contrast, our proposition, which traces the origin of mantids further 175 Myr back, offers plenty of Carboniferous putative stem-Dictyoptera (Bolton, 1922; Laurentiaux, 1958; Laurentiaux-Vieira & Laurentiaux, 1980), whose relationships are poorly known, as potential sister-groups to the Mantodea. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 1978, 1980a, 1980b, 1982, 1983), stem-Dictyoptera (Bolton, 1922; Laurentiaux, 1958; Laurentiaux-Vieira & Laurentiaux, 1980), Orthoptera (Ragge, 1955; Sharov, 1971; Béthoux & Nel, 2002), Plecoptera (Needham & Claassen, 1925; Béthoux, 2005c), and Grylloblattida (Storozhenko, 1998, 2002). The main stem of the vein sector RP diverges from R at the wing base and passes by the sclerotized structure usually ‘occupied’ by the median trachea. All mantids but Metallyticus spp., Chaeteessa spp., Mantoida spp., and †L. floccosa, and some other fossil taxa, have an RA + RP* composite stem, which is an apomorphic trait for a monophyletic group within Mantodea. Some taxa currently assigned to the Amorphoscelidae exhibit various states of this fusion, which supports the view of earlier workers stating that the traditional Amorphoscelidae are paraphyletic (e.g. Handlirsch, 1925; Chopard, 1949a; Wieland, 2003). As a result of our investigation, RA is differentiated from RP in Mantodea, unlike in Blattodea in which RA is fully fused with RP. The latter state is apomorphic within Neoptera, as evidenced by the morphology exhibited by Orthoptera (Ragge, 1955; Sharov, 1971; Béthoux & Nel, 2002), Plecoptera (Needham & Claassen, 1925; Béthoux, 2005c), Grylloblattida (Storozhenko, 1998, 2002), and stem-Dictyoptera (Bolton, 1922; Laurentiaux, 1958; Laurentiaux-Vieira & Laurentiaux, 1980). 107 108 O. BÉTHOUX and F. WIELAND ORIGIN OF THE OOTHECA IMPLICATIONS FOR INSECT PALAEOBIODIVERSITY This contribution supports the view that the ‘Protorthoptera’ must no longer be considered wholly as an extinct group. A preliminary review suggests that about 6% of ‘protorthopteran’ genera assigned to families in the latest compilation (Carpenter, 1992; hereafter referred to ‘protorthopteran’ genera) are to be considered as relatives of modern mantids. Other recent taxonomic reviews (Béthoux & Nel, 2002, 2003, 2004, 2005; Béthoux, 2003, 2005a, 2006, 2007a, b; and unpubl. data) reveal that about 16% of protorthopteran genera are stem relatives of Orthoptera (or even genuine Orthoptera, see Béthoux & Ross, 2005). Additionally the fossil Grylloblattida, encompassing about 53% of protorthopteran genera, are representatives of an extant order (Storozhenko, 2002). Finally, with 75% of protorthopteran genera to be viewed as stem-groups of total-groups having modern representatives, it can be argued that the estimation of a low end-Permian diversity and subsequent extinction of the ‘Protorthoptera’ at the beginning of the Triassic (Labandeira & Sepkoski, 1993); see also (Ross & Jarzembowski, 1993) was based on a taxonomic artefact. It could yet be conjectured that a number of these stem taxa constitute extinct monophyletic clades, within their new respective total-groups. For example, as defined by Carpenter (1992), the taxon †Strephocladidae is monophyletic, occurred in the Permian, and has no Triassic representatives. Then, we would observe an equally high level of family extinction at the end-Permian. However, the validity of such lower-rank groups belonging to stem-groups is uncertain. Several taxonomic revisions have already demonstrated that a significant number of Carpenter’s (1992) protorthopteran families are para- or polyphyletic (Béthoux & Nel, 2004, 2005; Béthoux, 2007c). In light of the current analysis, it is uncertain whether the taxon †Strephocladidae is monophyletic or paraphyletic. This taxon could indeed be composed of successively closer relatives of crown-Mantodea. If so, considering extinction rates makes sense at the genus or species level only. In fact, our knowledge of phylogenetic relationships among stem-Orthoptera, stem-Mantodea, and stemGrylloblattida taxa is minimal. As exemplified by the present contribution, the main reason is that major issues in insect taxonomy rest at the super-ordinal to ordinal taxonomic levels [see Inward, Beccaloni & Eggleton (2007) and subsequent comments (Lo et al., 2007; Eggleton, Beccaloni & Inward, 2007, and Béthoux (2007c) for recent updates of insect taxonomy at the ordinal level]. Attaining a more accurate view is still out of reach without further taxonomic revisions, and descriptive studies. In summary, the total-group Mantodea was distinct from the total-group Blattodea from the Carboniferous, implying that both groups survived the Permian– Triassic biocrisis. Accordingly, the importance of the end-Permian faunal turnover will have to be re-evaluated in the future. This contribution suggests that our current view of the early steps of Pterygota evolution could yet be significantly modified by outcomes of taxonomic revisions. Rather than families, genera and species used as a proxy are more likely to provide accurate results on Pterygota evolution. However, attention will have to be paid to the sensitivity of the data to the quality of the fossil record at the level of deposits, as well as to the relevance of the associated taxonomic literature. CONCLUSION Our comparative analysis demonstrates that †Mes. dolloi is a Carboniferous relative of modern mantids. A number of ‘protorthopteran’ taxa will have to be re-examined and re-interpreted in light of the present discovery. It implies that the divergence between the © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79–113 Downloaded from https://academic.oup.com/zoolinnean/article/156/1/79/2596002 by National Science & Technology Library user on 04 July 2024 It is generally admitted that laying eggs within an ootheca is an apomorphy of the Dictyoptera (as represented by modern taxa; Hennig, 1981; Grimaldi & Engel, 2005; among others). The assumption that stem-Mantodea existed in the Carboniferous implies that this sort of oviposition existed at the time. Although controversial (Brown, 1957; Hennig, 1981), it cannot be completely ruled out that fossils of Carboniferous oothecae are actually recorded (Pruvost, 1919, 1930; Laurentiaux, 1960). Laurentiaux (1960) suggested that a part of the Late Carboniferous stemBlattodea produced oothecae. The corresponding evolutionary steps highlighted by Grimaldi (1997) could well have occurred during the Late Carboniferous, or even earlier. Our discovery does not imply that the ootheca was acquired twice, as stem-lineages of crown-Dictyoptera could well have diverged since the Late Carboniferous. It must be noted that scenarios regarding the origin of the ootheca and of the related morphological traits in Carboniferous stem-Dictyoptera are highly speculative, considering that body remains (other than wings) that are sufficiently well preserved are extremely rare for insect species of this period, and that the phylogeny of stem-Dictyoptera is virtually unknown. The morphology of modern taxa alone suggests that oothecae and related morphological traits were acquired only once (Klass, 2003). This is not in contradiction with a Carboniferous origin of the Mantodea. CARBONIFEROUS MANTODEA IDENTIFIED Chaeteessa sp., belonging to the collection of the MTD, to be dissected by one of us (OB). The authors also thank B. Archibald (MCZ, Harvard University, Cambridge), W. Farrum (MCZ, Harvard University, Cambridge), L. Clunie (Landcare Research, Auckland), M. Mostovski (Natal Museum, University of KwaZuluNatal, Pietermaritzburg) who provided photographs of fossil specimens, or material. We also thank the President and Fellows of Harvard College for permission to use MCZ copyrighted material (photographs in Fig. 23). The authors are grateful to J.-M. Pacaud (MNHN, Paris) who allowed the holotype of Mesoptilus dolloi and the specimen MNHN A27122 to be investigated. The authors are also grateful to Z. Simmons (Oxford University Museum of Natural History, Oxford) for allowing a loan of the specimens belonging to the genera Chaeteessa and Metallyticus (including the material described by Smart, 1956) housed at the OUMNH. The authors are grateful to R. J. Rinas (USA) for improvement of the English grammar. The authors thank D. Bauer (Würzburg) and C. Schwarz (University of Würzburg) for their help during a collecting mission in Southern France (August 2006). FW is thankful to S. Materna (Erlangen), J. Mehl (University of Erlangen), and K. Schütte (University of Hamburg) for their help and support during a collecting mission in Malaysia (March 2007), and to M. Deyrup (Archbold Biological Station, Lake Placid, Florida) for providing specimens. Finally, we are grateful to the members of the Interessengemeinschaft Mantodea (IGM) who provided several of the specimens examined in this study. ACKNOWLEDGEMENTS The first author is a postdoctoral research fellow of the Alexander von Humboldt Foundation. This research project received support from the SYNTHESYS Project (http://www.synthesys.info/), which is financed by European Community Research Infrastructure Action under the FP6 ‘Structuring the European Research Area’ Program (visits to the MNCN, Madrid, July 2007, and to the NHM, London, August 2007), attributed to one of us (OB). This study was partly funded by the German Research Foundation (DFG) as part of the research project DFG Wi599/12 (participation of FW in collecting missions to France and Malaysia). One of us (OB) is grateful to I. Izquierdo Moya (MNCN) and G. Beccaloni (NHM) for their help and support during the visit to the collections they respectively are in charge of, and for allowing dissection of relevant material. The authors are grateful to R. Roy (MNHN, Paris) for allowing specimens of Metallyticus violaceus, belonging to the collection of the MNHN, to be dissected by one of us (OB). Similarly, K.-D. 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