Guyiling Jianboni Gen. Et Sp.N., an Antlion-Like Lacewing, Illuminating

Systematic Entomology (2012), 37, 617–631

Guyiling jianboni gen. et sp.n., an antlion-like lacewing, illuminating homologies and transformations in Neuroptera wing venation CHAOFAN SHI1, OLIVIER BETHOUX´ 1,2, CHUNGKUN SHIH1 and D O N G R E N1

1College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100048, China and 240 rue d’Aveillans, 38770 La Motte d’Aveillans, Isere,` France

Abstract. A new genus and species of antlion-like fossil lacewing, Guyiling jianboni gen. et sp.n. (Insecta: Neuroptera) are described based on a single specimen from the Yixian Formation (Liaoning Province, China; Early Cretaceous). The new taxon exhibits derived traits such as distally dilated antennae and well-developed anterior Banksian line (known in Myrmeleontidae), but also genuine plesiomorphies (at the level of Myrmeleontiformia), such as the divergence of a distinct CuA1 stem from MP2 + CuA1 (forewing), and a basal origin of MA (diverging from RP + MA; both wing pairs). This combination is unique among the ‘Palaeoleontidae’, a group better considered as a paraphyletic assemblage of various stem-Myrmeleontiformia. The wing morphology of the new species is considered in the light of a survey of wing venation topological homologies (and implied transformations) among several Neuroptera families. The survey includes a revision of the holotype of Leptolingia jurassica Ren (Grammolingiidae; Jiulongshan Formation, Daohugou locality, Inner Mongolia Autonomous Region, China; middle Jurassic). The forewing morphology of Guyiling jianboni gen. et sp.n. demonstrates that the fusion of MP2 with CuA, and the differentiation of CuA into two distinct main stems (namely CuA1 and CuA2) are traits shared with Myrmeleontidae and Ascalaphidae (and possibly Nemopteridae). However, the survey also demonstrates that a fusion of MP2 with CuA occurred repeatedly among Neuropterida, although by means of various modalities (namely translocation vs regular fusion). The ‘pectinate fusion’ of CuA(1) with MP2 [i.e. CuA1 has no distinct stem and emits successive branches from MP2 + CuA(1) partim] is a further step in this fusion, and occurred repeatedly as well (at least in the hind wings of Osmylidae and Nymphidae, and both fore- and hindwings of a sub-group of Myrmeleontiformia including Myrmeleontidae and Ascalaphidae, and possibly Nemopteridae). It is anticipated that the current contribution will constitute useful background information for further studies, focusing on particular transformation cases, ideally including a documentation of intraspeciﬁc variation.

Introduction New, 1991); the patterns are commonly used for identiﬁcation at various taxonomic levels and for identiﬁcation of fossil taxa The amazing variety of wing venation patterns of Neuropterida (e.g. Yang et al., 2009; Wang et al., 2009; Shi et al., 2011). has attracted the attention of many researchers (e.g. Brongniart, Yet a rapid survey demonstrates that contradictory conjec- 1893; Tillyard, 1915; Carpenter, 1940; Aspock¨ et al., 1980; tures of topological homology, at the level of Neuroptera, can be found in recent literature data. For example, the forewing Correspondence: Dong Ren, College of Life Sciences, Capital ‘oblique vein’ indicated as ‘O’ by Tillyard (1915) and vari- Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing ous other authors, and commonly occurring in forewings of 100048, China. . E-mail: [email protected] Myrmeleontidae, has been interpreted as a posterior branch of

MP fusing with a Cu branch (Tillyard, 1915; Aspock¨ et al., Cu, Cubitus; CuA, anterior Cubitus (royal blue); CuA1, ante- 1980; MP itself according to Menon & Makarkin, 2008), but rior branch of CuA (royal blue); CuA2, posterior branch of is neglected by New (1985, 2003), who does not recognize CuA (light blue); CuP, posterior Cubitus (brown); AA1, first the composite nature of the resulting vein (labelled ‘CuA’). anterior Analis. Notice that RP is often referred to as ‘RS’ or In addition, the occurrence of this element is uncertain in ‘Rs’ in the previous literature. Following New (1985, 1989, Nemopteridae, for example (see section Systematic palaeon- 2003; and see Tillyard, 1915), we refer to the cross-veins tology). As is often the case with wing venation, this situation aligned along the longitudinal axis of wings, and connected likely is the consequence of a ‘morphological saturation’ of with main veins whose course is shortly reorientated along the various crown-groups making up Neuropterida, coupled the longitudinal axis, as ‘Anterior Banksian line’. Presectorial with a lack of broad comparative analyses incorporating recent cross-veins are located in the area between R (+ MA) and MP, and fossil species (Bethoux,´ 2009). The discovery of a well- basal to the origin of RP (+ MA). It is very common that main preserved, intricate and singular fossil specimen from the Yix- stems are forked near the posterior wing margin in Neuroptera. ian Formation (Liaoning Province, China; Early Cretaceous) In such cases the actual number of branches of a vein is difficult prompted us to scrutinize wing venation patterns documented to determine, because an actual vein can hardly be differen- in the Myrmeleontiformia (i.e. Myrmeleontidae, Ascalaphidae, tiated from a cross-vein. Where appropriate we indicate the Nemopteridae, Nymphidae, Psychopsidae), one of the most number of ‘main branches’, those occurring before distal twigs. consistently monophyletic clades in the order (Aspock,¨ 2002; Some particular types of transformation are (re-)introduced Grimaldi & Engel, 2005; Winterton et al., 2010; and references here. Vein ‘translocation’ is defined as the fusion of a vein therein). Aiming at determining a plausible ground plan for the with another from the base of the latter. In other words the group, we found ourselves embarked in a wider survey. Our ‘translocated’ vein is no longer provided with a basal dis- results are presented in the following. tinct stem. Such transformation was first documented in Per- mian and Triassic titanopteridan Orthoptera (Bethoux,´ 2007), and in extant Mantodea (Bethoux´ & Wieland, 2009). Several Material and methods instances of ‘pectinate fusion’ are presented herein, but it has to be introduced prior to the description of the new species. This The specimen newly described herein is housed at the Key transformation involves the successive pectinate emergence of Lab of Insect Evolution and Environmental Changes, College branches of a vein fused with another, the former lacking a dis- of Life Sciences, Capital Normal University, Beijing, China tinct main stem. This organization has already been assumed (CNU; Ren Dong, Curator). Extant specimens were prepared for RA and RP in various Hemerobiidae species, in particular for the comparative analysis. They all belong to the personal those assigned to the genus Wesmaelius Kruger¨ [the common collection of one of us (OB), referred to as IWC OB. Wings stem RA + RP (partim) is indicated as ‘R + Rs’ in Aspock¨ were cut off and mounted in white Euparal medium (Asco et al. (1980: fig. 113); and see, among others, Oswald (1993)]. Laboratories, Manchester, UK). The venation patterns and vein widths of the specimen Systematic palaeontology CNU-NEU-LB2011014 and of selected extant specimens were drawn by OB with a SteREO Discovery V8 stereomicroscope Order Neuroptera Linnaeus equipped with a pair of W-PL 10×/23× eye pieces, a PlanApo S 1.0× FWD objective, and a camera lucida. Wings of the Suborder Myrmeleontiformia holotype of Leptolingia jurassica Ren were drawn by OB using Family incertae sedis a Leica MZ12.5 dissecting microscope equipped with a camera lucida. Final drawings were inked manually and ‘polished’ Genus Guyiling gen.n. using Adobe Photoshop. Fossil material was observed dry Type and only species. Guyiling jianboni gen. et sp.n. and under ethanol. Photographs were taken using a Canon EOS 450D/550D digital camera coupled with a Canon 50 mm macro lens, or a Canon MP-E 65 mm macro lens, and were Diagnosis. By monotypy, that of the type species. processed using Adobe Photoshop. Photographs referred to as ‘composite’ are a combination of photographs of a specimen Etymology. Pinyin transliteration of ‘Gu’, ‘Yi’ and ‘Ling’, both dry and immersed in ethanol. meaning ‘ancient’, ‘ant’ and ‘lacewing’ in Chinese, respec- The traditional nomenclatural procedure is followed herein. tively; name according to the ‘myrmeleontoid’ affinities of this This does not imply support for this approach on the part fossil taxon. of OB. We follow the serial insect wing venation ground- plan Lameere (1922, 1923). The corresponding wing venation Comments. Although the new genus exhibits several charac- nomenclature is repeated for convenience, with indication of ters diagnostic of the Myrmeleontidae (namely distally dilated colour coding: ScP, posterior Subcosta; RA, anterior Radius; antennae; well-developed anterior Banksian line), the occur- RP, posterior Radius; M, Media; MA, anterior Media (red in rence of traits presumably plesiomorphic, such as the diver- the corresponding figures); MP, posterior Media; MP1, anterior gence of a distinct CuA1 stem from MP2 + CuA1, and a basal branch of MP (orange); MP2, posterior branch of MP (green); origin of MA, prevented us from assigning it to this taxon. It

© 2012 The Authors Systematic Entomology © 2012 The Royal Entomological Society, Systematic Entomology, 37, 617–631 Guyiling jianboni gen. et sp.n. 619 is better compared with members of the ‘family’ Palaeoleon- (10) (reticulate cubital triangle) is documented in Myrmeleon- tidae Martins-Neto, 1992. The proposed diagnostic characters tidae (Fig. 8C, E; Aspock¨ et al., 1980), Ascalaphidae (Fig. 9A; of this assemblage are to be discussed first. Aspock¨ et al., 1980), and Nemopteridae (Tjeder, 1967). These The diagnosis of the Palaeoleontidae was revised by Heads characters (7, 8, 10) are absent in more ‘basal’ Neuroptera et al. (2005) and Menon & Makarkin (2008), with the families, and therefore represent synapomorphies of several following list of characters: (1) antennae long, filiform, neither families, in addition to the new genus and members of the dilated nor clavate distally; (2) rich wing venation; (3) wings ‘Palaeoleontidae’. longer than wide; (4) RP originating from R close to wing The relevance of the character (9) (namely proximal fork of base; (5) presectorial cross-veins absent in both wings; (6) MA MP into MP1 and MP2) proposed by Heads et al. (2005) was branching from RP proximally of the most basal branch of RP; debated by Menon & Makarkin (2008). According to the lat- (7) ‘oblique vein’ present in forewing; (8) anterior Banksian ter, ‘MP1’ in the material described by Heads et al. (2005) is line well developed; (9) in forewing MP two-branched, with actually the proximal part of a longitudinal vein-like fold. Also, MP1 profusely branched distally and MP2 simple; (10) CuA which vein is to be understood as ‘MP1’ (with many branches) forked, forming characteristic myrmeleontoid reticulate cubital and ‘MP2’ (simple) according to these authors (and the litera- triangle; and (11) CuP and AA1 separate. ture data) is not evident. In any case, according to our compar- Unlike other characters, character (1) would exclude the ative analysis, the MP1/MP2 fork is proximal in several Neu- new genus from the Palaeoleontidae, because it possesses roptera (sub-)families, including some Osmylidae (Fig. 6D, E), distally dilated antennae (rather than filiform). A ‘dilated’ and Grammolingiidae (Fig. 7A, B). It possibly is a plesiomor- condition similar to that observed in the new genus occurs phy with respect to Myrmeleontidae and Ascalaphidae. in Myrmeleontidae (Aspock¨ et al., 1980) and Araripeneuridae Finally, due to the preservation of the basal wing part in (Martins-Neto & Rodrigues, 2010). Notice that it can be argued our specimen, the condition of CuP and AA1 (11) cannot be that the ‘clavate’ condition observed in Ascalaphidae (Aspock¨ observed. In any case, for a similar reason, this character is et al., 1980) is a particular condition of the ‘dilated’ one. not documented in many ‘Palaeoleontidae’. The characters (2) and (3) are commonly found among most At this stage, we propose that the ‘family Palaeoleontidae’ families within Myrmeleontiformia (Grimaldi & Engel, 2005: could be a paraphyletic assemblage, because there is no fig. 9.14), but also in some members of more distantly related character that definitively indicates its monophyly; instead it families, such as Osmylidae (New, 1989: figs 25–27) and is characterized by a mix of plesiomorphic and apomorphic Ithonidae (Winterton & Makarkin, 2010; New, 1989: fig. 21; conditions shared by other families. This assemblage includes among others). various stem relatives of a sub-group of Myrmeleontiformia, The origin of RP located near wing base (4), and the absence including Myrmeleontidae and Ascalaphidae at least, possibly of presectorial cross-veins (5) are characters present in most Nemopteridae. Therefore, we consider the family affinities of neuropteran families, such as Dilaridae, Osmylidae, Nymphi- the new genus as uncertain. The erection of a new genus is dae and Grammolingiidae. These characters are absent in discussed below. Myrmeleontidae, Araripeneuridae, Ascalaphidae, Nemopteri- The new species mostly resembles those assigned to the dae and Babinskaiidae (Tjeder, 1967; Martins-Neto, 1997; palaeoleontidan genera, including Baisopardus Ponomarenko, New, 2003; Martins-Neto & Rodrigues, 2010). Given the cur- Cretoleon Ponomarenko, Metahemerobius Makarkin, Neuras- rent phylogenetic scheme for Neuropterida (Winterton et al., tenyx Martins-Neto, Palaeoleon Rice, Paraneurastenyx 2010), it is clear that characters (4) and (5) are plesiomorphies, Martins-Neto, Parapalaeoleon Menon & Makarkin and Sam- and are likely correlated; character (6) occurs in a large num- sonileon Ponomarenko. It must be emphasized here that the ber of Neuroptera families. Therefore, none of these characters interpretation of Baisopardus cryptohymen by Heads et al. ascertains a placement of the new taxon within the ‘family’ (2005) is problematic (Menon & Makarkin, 2008). Based on Palaeoleontidae. original data and new photographs (provided by K. Wolf- The forewing ‘oblique vein’ (or ‘O’; below interpreted as Schwenninger, personal communication, 2011), we assume MP2) (7) is not unique to the members of the Palaeoleonti- that this species has a multibranched MA, a simple MP1 dae, as this character also occurs in Myrmeleontidae (Fig. 8C, and a distinct stem of CuA1 (diverging from MP2 + CuA1; E; Aspock¨ et al., 1980; among many others), Ascalaphidae i.e. without pectinate fusion of CuA1 with MP2). These (Fig. 9A; ‘Mp2’ on fig. 252 in Aspock¨ et al., 1980), and traits, in particular the distinct stem of CuA1, tend to indi- possibly in Nemopteridae [‘m-cu’? on fig. 1931 in Tjeder cate that this species is not closely related to Baisopardus (1967); strong oblique ‘cross-vein’ represented on fig. 1 in banksianus Ponomarenko (known from hindwings only), but Mansell (1983a), and figs 7–11 in Mansell (1983b)]. As for more closely related to Neurastenyx spp. Indeed, based on the developed anterior Banksian lines (8), many Myrmeleon- photographs of the material of B. banksianus (provided by tidae possess this line [Fig. 8C, E; Aspock¨ et al., 1980; A. Khramov, personal communication, 2012), we assume that among many others; it must be noticed here that the devel- no fusion of MP2 with CuA occurs in this species, which opment of anterior Banksian lines is of somewhat subjec- otherwise has important similarities with the new taxon, such tive interpretation, as many Myrmeleontidae species exhibit as a simple MP1 [MP sensu Ponomarenko (1992)], and a shade of various conditions, ranging from absent to well posteriorly pectinate MP2 and CuA [‘cubital veins’ sensu developed, probably due to secondary loss(es)]. The character Ponomarenko (1992)], supporting our interpretation. The lack

© 2012 The Authors Systematic Entomology © 2012 The Royal Entomological Society, Systematic Entomology, 37, 617–631 620 C. Shi et al. of MP2 + CuA(1) fusion in B. banksianus makes the new simple; abdomen elongate, about 35 mm long; wings: both material readily identifiable as belonging to a distinct taxon. wing pairs subequal, elongate and slender; cross-veins widely Only B. cryptohymen will be considered in the following. spaced; RP + MA arising close to wing base; MA and MP1 The new genus is different from Parapalaeoleon in its simple (in preserved area) and straight; RP with more than ten distally dilated antennae (a putative apomorphy; as opposed main branches, no presectorial cross-veins observed; anterior to ‘filiform antennae’). The new genus can be distin- Banksian line present; forewing: at least 44 mm long, 8.5 mm guished from Metahemerobius, Cretoleon, Parapalaeoleon, wide; forewing costal area narrow, cross-veins in area between Samsonileon and Paraneurastenyx by the lack of pectinate anterior wing margin and ScP simple; area between ScP and fusion of CuA1 with MP2 (i.e. by the presence of a distinct RA without visible cross-veins; close to its point of origin, main stem of CuA1 diverging from MP2 + CuA1; a putative MP2 (* on Fig. 2C, D) fused with CuA for 1.0 mm/1.1 mm plesiomorphy). The new genus differs from Metahemerobius, (forewing 1/2, respectively); CuA2 diverging from MP + CuA Neurastenyx, Parapaleoleon and B. cryptohymen in its simple 0.67 mm/0.77 mm distal to fusion of MP2 with CuA; MP2 and MA (or forked near margin, at best). The new genus differs CuA1 diverging as distinct stems, both posteriorly pectinate; from Metahemerobius and Palaeoleon in the presence of a MP2 with about seven main branches, CuA1 with five main well-developed anterior Banksian line. The new genus dif- branches; CuA2 with three branches (without distal twig); CuP fers from Paraneurastenyx in its more basal origin of MA long, parallel to posterior wing margin, posteriorly pectinate, (a putative plesiomorphy). The new genus differs from Meta- with about seven branches, distally connected to CuA2 by hemerobius, Palaeoleon, Parapaleoleon and Samsonileon in a short cross-vein; hind wing: venation similar to that of its elongate wing shape. Owing to the very incomplete docu- forewing, except for MP1 and MP2 separating close to wing mentation on Palaeoleon (known based on the distal half of base, MP2 distinct from CuA, and CuA not clearly forked into a single wing), comparison with the new material is not easy. two main stems. The new genus differs from Palaeoleon in the configuration of Type locality and horizon. Huangbanjigou, Beipiao City, MP2, which is posteriorly pectinate in the new genus, whereas Liaoning Province, China; early Cretaceous, Yixian Formation. it (probably) is internally pectinate in the latter. In summary, the unique combination of plesiomorphic Etymology. The species name is dedicated to Mr. Jian Bon (divergence of a distinct CuA1 stem from MP2 + CuA1; Shih (son of CKS), for demonstrating initiatives, creativity and basal origin of MA) and derived conditions (distally dilated innovation in his study and work, and for providing inspiration antennae; well developed anterior Banksian line) exhibited by and support to CKS. the new species prevents its assignment to any of the previously existing genera. Erection of a new genus is therefore necessary. Specimen examined. Holotype CNU-NEU-LB2011014, composed of positive and negative imprints (which polarity Guyiling jianboni gen. et sp.n. is hardly determinable; all photographs except Fig. 1B represent the same side, which has incomplete antennae), with an (Figs 1–2) almost complete body and four wings overlapping.

Diagnosis. Antennae shorter than half of forewing length, Comments. The available specimen is composed of a single gradually enlarged distally; wings: RP + MA arising close individual with the four wings overlapping. Yet the wing to wing base; anterior Banksian line present; elongate shape; venation can be traced if observed with sufficient magnification forewing: no cross-veins in the area between ScP and RA; (Fig. 2G, H). Due to a fault located near wing apices, wing MP2 (‘oblique cross-vein’ of previous authors) fused with CuA venation in this area could not be associated with either one of for a short distance; a single main stem of CuA1 diverging the wings. Therefore, apices are not represented on Fig. 2C–F. from MP2 + CuA1 (as opposed to pectinate fusion of CuA1 Although fore- and hindwing can be identified, as well as with MP2); CuA divided into two main stems; occurrence of both wings belonging to the same single side, it cannot be a short cup-cua2 cross-vein; hind wings: MP2 distinct from determined to which side they belong. Therefore, wings of the CuA/CuA1; CuA not obviously divided into two main stems. same side are indicated by the same number in figure captions. One of the most important characters exhibited by the new species is the anterior Banksian line occurring in the Description. Head, legs, abdomen: head about 3 mm long RP area. It is weakly imprinted in the specimen. Coupled (excluding antennae); compound eye large, occupying most with the overlap of the four wings, it is difficult to observe. space of head; no ocellus observed; antenna short, about Typically, anterior Banksian lines are composed of short cross- 11 mm long, less than half of forewing length, with about veins connecting two main veins, whose course is shortly 30 flagellar articles; scape larger than pedicel and flagellar reorientated parallel to the wing longitudinal axis (e.g. Fig. 8). articles, with scattered, fine setae; flagellum gradually dilated, There is one particularly clear indication of the occurrence most apical flagellar article much longer and wider than basal of such typical short cross-veins in the specimen CNU-NEU- ones; mandible falciform, with blunt apex; legs long and LB2011014, on the hindwing 1 (Fig. 2E). In detail, it connects slender, with dense setae; tibia with a pair of apical spurs; fifth MA and the first posterior branch of RP (Fig. 2G, H; cross-vein ◦ tarsomere longer than remaining four tarsomeres; tarsal claw indicated by , first posterior branch of RP by arrows).

Fig. 1. Guyiling jianboni gen. et sp.n., holotype specimen CNU-NEU-LB2011014. (A) Photograph (composite, ﬂipped horizontally) and drawing of habitus (apart from apical area, venation that could not be assigned with certainly to any of the wings was omitted). (B) Detail of antennae, as located on A (composite).

Comparative analysis analysis carried out below (it can be referred to as ‘red vein’ only), a preliminary discussion on this topic appears necessary. Preliminary discussion Most authors (e.g. Carpenter, 1940, 1947; Aspock¨ et al., 1980; New, 2003) assumed it to be MA (hence fused with R/RP at It must be mentioned here that the nature of the ﬁrst vein some point). In contrast Makarkin et al. (2009: 967) suggested diverging from the genuine RP stem, in Kalligrammatidae, and that this stem is merely a branch of RP, and that ‘in Neuroptera Neuropterida in general, has been recently debated (Makarkin the radius and media are not fused basally’. However, there is et al., 2009; and see Oswald, 1993: 172–173). Although the a body of evidence suggesting that MA actually fuses with RP actual nature of this stem does not infringe on the comparative in various Neuropterida.

Fig. 2. Guyiling jianboni gen. et sp.n., holotype specimen CNU-NEU-LB2011014 (all photographs ﬂipped horizontally). (A) Photograph of thorax, head and legs, as located on Fig. 1A (under ethanol). (B) Photograph of tarsi, as located on Fig. 1A (under ethanol). (C) Forewing 1. (D) Forewing 2. (E) Hindwing 1. (F) Hindwing 2. (G) Detail of hindwing 1 (as located on Fig. 1A and E), photograph (arrows indicate the course of the ﬁrst RP ◦ branch; under ethanol), and drawing ( indicates a cross-vein of the anterior Banksian line). (H) Detail of hindwing 1, as located on G (indications as in G; under ethanol).

First, several taxa show a free portion of MA actually Le. jurassica (based on new observations), Phymatosmylus fusing with RP, such as in the hindwing of Dilar saldubensis caprorum Adams (based on literature data; Adams, 1969: Navas´ [Aspock¨ et al., 1980: fig. 423, and Fig. 5B; and ‘b’ fig. 1A), Spilosmylus obliquus New (based on literature on various figures by Carpenter (1947)]. Second, in the data; New, 1986: figs 29, 30), Nesydrion nigrinerve Esben- forewing of D. saldubensis as documented by Aspock¨ et al. Petersen (based on literature data; New, 1981: figs 85, 86), (1980: fig. 423, and Fig. 5A), there is a stem diverging Myrmecaelurus trigrammus (Pallas) (based on observation of from R prior to the origin of the genuine RP. This stem actual material), and Libelloides coccajus Denis & Bitsch can only be MA – hence this species (specimen?) possesses (based on observation of actual material). The comparative aR+ MA stem (a similar condition was observed in an analysis focuses on the course of the veins MP1, MP2, CuA unusual Nemopteridae to be described elsewhere; OB, personal and CuP, which are those undergoing the most important observation). Even if this particular case is based on an transformations. Vein MA will be used as a landmark. unusual morphology for the family, the genus, or the species, We propose to begin our comparative analysis with the provided that other aspects of the wing venation pattern Dilaridae, because the family occupies a ‘basal’ position are unchanged as compared to other Dilaridae, it remains within Neuroptera, at least with respect to Myrmeleontiformia demonstrative of the occurrence of R + MA and RP + MA (Aspock¨ & Aspock,¨ 2008; Winterton et al., 2010). Our survey common stems in the family. Among many other contributions, of literature data (Brongniart, 1893; Carpenter, 1940, 1947; New (1983a) describes a ‘b’ stem in both wing pairs of Aspock¨ et al., 1980; New, 2003; Winterton et al., 2010; among Osmylidae representatives, which clearly is MA. The same others) suggests that characters which are the focus of our com- stem was observed in the holotype of Le. jurassica Ren parative analysis exhibit a plesiomorphic condition in members (Fig. 4B). Therefore, there is good evidence of a fusion of of this family (but just as in Hemerobiidae and Sisyridae, MA with R/RP in Neuroptera. among others, except for the organization of Cu in hindwing). Investigating correspondence in venation patterns among Other species were selected based on the availability of mate- fore- and hindwings will prove to be an essential step in rial and of reliable literature data, and on their ‘explanatory this analysis. However, as rightfully stated by Makarkin et al. power’ (i.e. because they are putative key taxa). For example (2009), vein elevation in hindwings of Neuroptera (-ida?) P. caprorum was selected because the (presumed) transloca- has been altered significantly, especially for CuA and CuP. tion of MP2 onto CuA in the hindwing is more evident in this This is evident in the material of Le. jurassica (Figs 3–4; species that in most other Osmylidae (in which MP2 and CuA Grammolingiidae; Jiulongshan Formation, Daohugou locality, are more tightly integrated in a single stem). Inner Mongolia Autonomous Region, China; middle Jurassic). Identification of main veins in forewings is straightforward Dilar saldubensis Navas´ in Laguna (Dilaridae) (Fig. 3A, B), especially once the claval fold is located (arrows (Fig. 5) on Fig. 4C). In hindwings (Fig. 3C, D), MP2, CuA1 and CuA2 are rather easy to identify, based on their location and Conjectures of topological homology we propose for this respective number of branches, very similar to the condition species are described as follows: of the corresponding veins in forewings (Fig. 3A, B) (in particular MP2; compare Fig. 3A and C). However, MP2, • Forewing (Fig. 5A). MA coalesced with R stem at wing base CuA and CuP are concave, convex and concave (respectively) and diverged before RA and RP separated; MP1/MP2 fork in forewings (Fig. 3A), and exhibit exactly the opposite located distal to the divergence of MA from R + MA; CuA elevation in hindwings (Figs 3D, 4D). However, it must be and CuP separated slightly proximal to the divergence of emphasized that numerous elevation shifts were observed MA from R + MA; CuA forked distally (three branches); in forewings of Le. jurassica (Figs 3A, 4E), suggesting that CuP posteriorly pectinate (five branches). opposed elevation might be the result of similar shifts affecting • Hindwing (Fig. 5B). MA fused for some distance with hindwings. Therefore, vein elevation should be given cautious RP near the origin of the later, then diverged from RP; consideration when conjecturing on topological homology in MP1/MP2 fork located just distal to the origin of RP; CuP hindwings of Neuroptera. Indeed the ‘opposed elevation’ was without distinct main stem. ignored in our interpretation of the hindwing venation of Le. jurassica as represented in Fig. 7B. Our interpretation basically follows that proposed by Finally, regarding elevation shifts in forewings of Carpenter (1947; at least for forewings) and Aspock¨ et al. Le. jurassica, we observed that four of these shifts are more or (1980). Identification of MA, MP1 and MP2 in hindwings, less aligned along the longitudinal axis of the wing, in a loca- according to their location in forewings, is straightforward. tion similar to that of the anterior Banksian line (grey line on Provided that the number of MA, MP1 and MP2 branches are Fig. 3A). This organization might be a ‘precursor’ of the gen- very similar in fore- and hindwings, it is assumed that it is uine anterior Banksian line observed in various ‘Palaeoleonti- also the case for CuA and CuP. Therefore, in hindwings, it is dae’ genera (including Guyiling gen.n.) and Myrmeleontidae. assumed that a secondary fusion of CuP with CuA, with no For our comparative analysis, in addition to the new species free stem of CuP and instead successive emergence of CuP described above, we selected Dilar saldubensis Navas´ in branches from a CuA + CuP partim stem (Fig. 5C), occurs. Laguna (based on literature data; Aspock¨ et al., 1980: fig. 423), Such a pattern is referred to as pectinate fusion herein.

Fig. 3. Letpolingia jurassica Ren, holotype specimen NN99007, wing venation with indication of elevation of selected veins. (A) Left forewing (grey line, presumed ‘pre-’ Banksian line; grey frame, location of area reproduced on Fig. 4E). (B) Right forewing. (C) Left hindwing. (D). Right hindwing.

Fig. 4. Letpolingia jurassica Ren, holotype specimen NN99007 (all photographs light-mirrored). (A) Photograph of habitus (composite). (B) Photograph of right forewing base, as located on A (composite; arrow indicates the presumed free part of MA basal to its fusion with RP). (C) Photograph of left forewing base, as located on A (dry; arrows indicate the claval fold). (D) Photograph of basal part of the left hindwing, as located on A (dry). (E) Photographs of left forewing middle area, as located on A (and frame on Fig. 3A; dry).

A A

Fig. 5. Conjectures of wing venation topological homologies in Dilar saldubensis Navas´ (Dilaridae; modiﬁed from Aspock¨ et al., 1980: D ﬁg. 423). (A) Forewing. (B) Hindwing. (C) Scheme representing the CuA – CuP pectinate fusion.

Phymatosmylus caprorum Adams (Osmylidae) (Fig. 6A, B) E Conjectures of topological homology we propose for this species are described as follows:

• Forewing (Fig. 6A). MA fused with R at wing base, diverging from RP near the origin of the latter; MA simple with terminal twig; MP1/MP2 fork located near wing mid-length; MP1 forked near wing margin; MP2 branched shortly after separation of MP1 and MP2, with about 1.5 Fig. 6. Conjectures of wing venation topological homologies in Phy- more branches than MP1; CuA and CuP diverging close to matosmylus caprorum Adams (A–C), and Spilosmylus obliquus New wing base; CuA branched distally, with similar number of (D, E) (both Osmylidae; modified from original descriptions). (A, B) branches as MP2; CuP posteriorly pectinate, with numerous Favoured conjectures. (A) Forewing. (B) Hindwing. (C) Conjectures proposed by Adams (1969: fig. 1A) for hindwing. (D) Forewing. branches. (E) Hindwing. • Hindwing (Fig. 6B). MA course and organization as in forewing; MP1 individualized basally, branched distally, with few branches near wing margin; MP2 and CuA fused branches of these veins as observed in the forewing. Under our for a long distance, diverging near wing mid-length, oppo- interpretation, MP1, MP2, CuA and CuP have similar locations site the MP1/MP2 fork in forewing; both MP2 and CuA of their first branching point in fore- and hindwings. In the lat- branched at about the same level as MP1, with respective ter, if traced backwards, ‘our’ ‘MP2’ and ‘CuA’ diverge from a numbers of branches similar to those of the correspond- common stem. This organization can be explained by a translo- ing veins in forewing; CuP as in forewing, with fewer cation of MP2 onto CuA. The position of the divergence point branches. of ‘MP2’ along the wing longitudinal axis, similar in both wing pairs under our interpretations (but either from MP, or from Compared to D. saldubensis, this species has a more dis- CuA), provides a strong support to it. Overall the degree of cor- tal MP1/MP2 fork in the forewing, but identification of these respondence between the interpretations presented in Fig. 6A, veins appears straightforward. Regarding the hindwing, our B is higher than that between the interpretations presented interpretation differs from that in the original description in Fig. 6A, C, and allows us to favour that represented in (Fig. 6C), in particular in the identification of MP2, CuA and Fig. 6B. In the current case a MP2 + CuA stem occurs in CuP. We identified these veins primarily after the number of hindwings.

The hindwing venation of several Osmylidae taxa can easily A be interpreted in the same way as for P. caprorum. Based on New (1983b: figs 1–4) and Kimmins (1940: pls 3–5), members of the subfamilies Porisminae and Eidoporisminae, and genera such as Isostenosmylus Kruger¨ and Australysmus Kim- mins, clearly possess a MP2 (basal stem of New’s ‘MP’ in forewing) translocated onto CuA, with a distinct main stem B of CuA, in hindwings. This is also very likely to be the case of representatives of the subfamily Kempyninae, whose hindwing venation was probably improperly interpreted by New (1983a: fig. 2): his forked ‘MP2’ is probably composed of MP2 and CuA. However, many Osmylidae exhibit a more elaborated condition, as exemplified by S. obliquus, treated below. C

Spilosmylus obliquus New (Osmylidae) (Fig. 6D, E)

Conjectures of topological homology we propose for this D species are described as follows:

• Forewing (Fig. 6D). MA fused with R at wing base, diverging from RP near the origin of the latter; MP1/MP2 fork located just distal to the origin of MA (from RP + MA); MP1, MP2, CuA and CuP posteriorly pectinate; MP1 and MP2 parallel, close, simple for a long distance, Fig. 7. Conjectures of wing venation topological homologies in Lep- with similar numbers of main branches (four, and five, tolingia jurassica Ren (Grammolingiidae; modified from original respectively); CuA and CuP diverging close to wing base; description and according to new observations) (A, B) and Nesy- CuA branched distally, with similar number of branches drion nigrinerve Esben-Petersen (Nymphidae; modified from original description) (C, D). (A, C) Forewing. (B, D) Hindwing. as MP1 and MP2; CuP with numerous branches; ultimate MP1, MP2, CuA, and CuP branches connected to MA, MP1, MP2 and CuA, respectively, by a short cross-vein, Leptolingia jurassica Ren (Grammolingiidae) basal to the first fork of MA, MP1, MP2 and CuA, (Fig. 7A, B; and Figs 3, 4) respectively. • Hindwing (Fig. 6E). Mostly as in forewing, except for MA Conjectures of topological homology we propose for this with a slightly more distal origin, and CuA and MP2 forming species are described as follows: a common stem, from the wing base. • Forewing (Fig. 7A). MA with a distinct stem, fused with Unlike in P. caprorum, but as in D. saldubensis,the RP at the origin of the latter; MP1/MP2 fork opposite the MP1/MP2 fork is located basally in the forewing of emergence of MA (from RP + MA); both MA and MP1 S. obliquus. More importantly, in the hindwing, we assume forked near wing margin, with similar number of branches; a fusion of MP2 with CuA, from the wing base. As for MP2, CuA and CuP more or less dichotomously branched; P. caprorum, this assumption is supported by the respective first MP2 fork located near wing mid-length; MP2 with numbers of terminal branches of MP1 and CuP, and by the as many branches as MA and MP1 altogether; CuA and sum of branches of MP2 and of CuA, similar in fore- and CuP diverging close to wing base; first fork of CuA located hindwings under our interpretation. The fact that MP2 and opposite the emergence of the first RP branch, with a basal CuA form a common stem in the hindwing of P. caprorum,a veinlet and two main stems (CuA1, CuA2; the latter in light related species, is also supportive of our interpretation. How- blue on Fig. 7); CuP branched proximal to fork of CuA, ever, there is an important difference between P. caprorum and with fewer branches than CuA. S. obliquus in the organization of this fusion: in the former • Hindwing (Fig. 7B). Mostly as in forewing, except for CuA CuA retains a distinct stem, while in the latter CuA branches without basal veinlet, and CuP without evident fork. are successively emitted from a MP2 + CuA partim stem. The latter qualifies for a pectinate fusion of CuA with MP2. There- As opposed to the Osmylidae investigated above, but as fore, the actual number of MP2 and CuA branches cannot in D. saldubensis, this species retains distinct MP2 and CuA be determined with certainty. Our interpretation relies on the stems in hindwings. Unlike in previous species, two main forewing venation of the species. stems of CuA can be identified.

Nesydrion nigrinerve Esben-Petersen (Nymphidae) A (Fig. 7C, D)

Conjectures of topological homology we propose for this species are described as follows:

• Forewing (Fig. 7C). MA fused with R at wing base, B diverging from RP near the first third of wing length; MP1/MP2 fork located basal to the emergence of MA (from RP); MP1 with about five main branches; MP2 posteriorly pectinate, with six main branches; CuA and CuP diverging close to wing base; first fork of CuA located basal to the emergence of MA (from RP); CuA more or less dichotomously branched, with three to four main branches; C CuP posteriorly pectinate, with eight branches. • Hindwing (Fig. 7D). Mostly as in forewing, except for MP2 fused with CuA from wing base, CuA without main stem, no evident CuA1/CuA2 fork.

As above, comparison of venation patterns of fore- and hindwing of N. nigrinerve indicate that a common stem D MP2 + CuA occurs on the latter. Indeed, if this interpretation is followed, the respective numbers of branches of MP1, MP2 + CuA and CuP in fore- and hindwings match. As in the S. obliquus hindwing (Fig. 6E), it is assumed that a pectinate fusion of CuA with MP2 occurs. The occurrence of a posterior stem of CuA (namely CuA2) is suggested. E Guyiling jianboni gen. et sp.n. (Fig. 8A, B)

Conjectures of topological homology we propose for this species are described as follows (and see formal description above): F

• Forewing (Fig. 8A). MA diverging from RP + MA, simple for most of its length; MP1/MP2 fork basal to the emergence of MA (from RP); MP1 simple (with distal twig?); MP2 short and oblique at its origin, fused for some distance with CuA; distal free part of MP2 posteriorly pectinate, with five branches, first fork located opposite the emergence of the Fig. 8. Conjectures of wing venation topological homologies in first RP branch; CuA forked into two main stems, CuA1 Guyiling jianboni gen. et sp.n. (A, B) restored based on holotype and CuA2, between the fusion and divergence MP2; CuA1 material, and Myrmecaelurus trigrammus (Pallas) (Myrmeleontidae; IWC OB 99, right wings) (C–F). (A) Forewing. (B) Hindwing. (C, D) and CuA2 with five and three main branches, respectively; Favoured conjectures. (C) Forewing. (D) Hindwing. (E, F) Conjectures CuP with several branches; proposed by New (2003: fig. 177). (E) Forewing. (F) Hindwing. • Hindwing (Fig. 8B). MA diverging from RP + MA, simple for most of its length; MP1/MP2 fork located at wing base; MP1 simple (with distal twig?); MP2 distinct from hindwings. If traced backwards (from their distal branches), CuA, posteriorly pectinate, with 5 branches, first fork MP2 and CuA necessarily form a common stem. The origin of located opposite the emergence of the first RP branch; this fusion is evident, as the basal free portion of MP2 is very CuA posteriorly pectinate, with 11 branches; CuP with few strongly imprinted in the available specimen (Fig. 1). The iden- branches, short. tification of CuA1 and CuA2 stems in forewings is prompted by the observation of such organization in the forewing of In this particular case, the hindwing venation pattern is illu- Le. jurassica (Figs 3, 7A), and possibly in that of N. nigrinerve minating regarding the forewing one. The location of MP2 (Fig. 7C). This identification of CuA1 and CuA2 stems in and CuA in the forewings is conjectured based on the num- forewings will be corroborated based on the morphology of ber of branches of each vein (and located on first fork) in the M. trigrammus (see below). The occurrence of distinctive

CuA1 and CuA2 stems in hindwings is not evident (but this fore- (Fig. 8E) and hindwing (Fig. 8F) (among other authors). could be the consequence of a pectinate fusion of CuA2 with The higher degree of correspondence among our conjectures CuA1). above, at different levels (between fore- and hindwings in M. trigrammus, and between M. trigrammus and G. jianboni Myrmecaelurus trigrammus (Pallas) (Myrmeleontidae) gen. et sp.n.), indicates that our conjectures are more plausible. (Fig. 8C, D) Libelloides coccajus Denis & Bitsch (Ascalaphidae) Conjectures of topological homology we propose for this (Fig. 9A, B) species are described as follows: The wing venation patterns of Li. coccajus are not fun- • + Forewing (Fig. 8C). RP MA diverging from R distally; damentally different from those observed in M. trigrammus. + MA diverging from RP MA, simple for most of its length; Apart from MA branched more basally, and the CuP simple, + MP1/MP2 fork basal to the divergence of RP MA; MP1 major features are the same, namely: simple (with distal twig); CuA ﬁrst fork just basal to the fusion with MP2; MP2 + CuA1 posteriorly pectinate, with • In forewing (Fig. 9A). MP2 short before its fusion with about ten main branches (possibly, MP2 and CuA1 with CuA1; pectinate fusion of CuA1 with MP2. ﬁve branches each); CuA2 with about four main stems; • In hindwing (Fig. 9B). MP2 fused with CuA/CuA1 from CuP fused with AA1 at wing base; CuP + AA1 posteriorly wing base; pectinate fusion of CuA1 with MP2. pectinate (actual number of CuP branches uncertain). • Hindwing (Fig. 8D). Mostly as in forewing, except for MP2 fused with CuA/CuA1 from wing base; and CuP distinct from AA1. Discussion

One of the most important points of our conjectures on Conjectures of topological homology proposed herein exhibit Myrmeleontidae and Ascalaphidae wing venation patterns a high degree of correspondence, at several levels. At the regards the identification of CuA1 and CuA2. This identifica- individual level, our favoured conjectures between fore- and tion is related to the course of MP2. As conjectured, based hindwing venation patterns are more consistent than some on the G. jianboni gen. et sp.n. forewing venation pattern previously proposed (compare Fig. 6A, B to Fig. 6A, C and (Fig. 8A), the presence of a comparatively strong oblique stem Fig. 8C–F). Conjectures within families (e.g. see Fig. 6A, B, between MP and CuA (actually CuA1) is indicative of the D, E) and across families (Figs 8A–D, 9) are also highly course of MP2: it is composed of a short basal free portion and consistent. Although a wider sampling would be necessary to immediately fuses with CuA (actually CuA1). However, unlike test the relevance of the favoured conjectures at the level of in G. jianboni gen. et sp.n., the stem (in royal blue in Fig. 8C) the whole Neuroptera, a survey of the literature data provided with which MP2 fuses is forked just basal to the fusion. The no conflicting evidence. first posterior branch (in light blue in Fig. 8C) can hardly be interpreted as CuP, as it would imply a significant alteration of the organization of this vein with respect to species investigated A above. It is more plausible to assume that both the anterior and posterior branches belong to CuA. If so, CuA2, as identified in Le. jurassica (Figs 3, 4, 7A, B) and G. jianboni gen. et sp.n. (and possibly N. nigrinerve; Fig. 7C), is readily identified in M. trigrammus. In addition, MP2 fuses with CuA1 – not CuA. The interpretation demonstrates that CuA is provided with two distinct stems in the species, and confirms the identification of B these branches in G. jianboni gen. et sp.n. (see above). Postulating the occurrence of a fusion of MP2 with CuA/CuA1 from the wing base in hindwings, and of a pectinate fusion of CuA1 with MP2 in both wing pairs, is not extraordinary, given that similar patterns have been conjectured in hindwings of S. obliquus (Fig. 6E) and N. nigrinerve (Fig. 7D). Due to the pectinate fusion of CuA1 with MP2, the actual number of branches of each vein cannot be determined with certainty. Our conjecture for M. trigrammus primarily relies on the fact that MP2 and CuA1 share a similar number of distal branches in G. jianboni gen. et sp.n. Fig. 9. Conjectures of wing venation topological homologies in Our interpretation of M. trigrammus wing venation signif- Libelloides coccajus Denis & Bitsch (Ascalaphidae; IWC OB 846, icantly differs from that by New (2003) for Myrmeleontidae right wings) (A) Forewing. (B) Hindwing.

The wing morphology G. jianboni gen. et sp.n. proved Conclusion to be essential for the clarification of wing venation conjectures in Myrmeleontidae and Ascalaphidae, mostly because it Our comparative analysis encompasses fore- and hindwing shows a fusion of MP2 with CuA, but with distinct MP2 and venation homologies of several major lineages in Neuroptera. CuA1 distal main stems. The situation in Myrmeleontidae and We demonstrate that a common stem MP2 + CuA was Ascalaphidae is more intricate, as the pectinate fusion of CuA1 acquired several times independently, albeit by means of with MP2 makes distal portions of both veins undistinguish- various modalities (namely translocation vs regular fusion). able. As a matter of fact, the MP2 + CuA1 stem in forewings We also newly formalize a transformation type – pectinate has been considered by some as composed of a single stem fusion – involving the successive pectinate emergence of (Fig. 8E). Our interpretation concurs with that of Aspock¨ branches of one vein fused with another, the former lacking et al. (1980: figs 211, 252), who assume a MP2 + CuA1 a distinct main stem. This formulation adequately describes fusion in the Myrmeleontidae and Ascalaphidae. Also, tak- the fusion of CuA with MP2 in hindwing of some Osmylidae ing into account several cases within Neuroptera allowed us and Nymphidae, the fusion of CuA1 with MP2 in wings of to demonstrate the plausibility of some particular transforma- Myrmeleontidae and Ascalaphidae, and the possible fusion of tion which affected the wing venation of Myrmeleontidae and CuP with CuA in hindwings of Dilaridae. Ascalaphidae, such as the fusion of MP2 with CuA at the Several wing venation characters useful for phylogenetic wing base in hindwings, and the pectinate fusion of CuA1 investigations can be extracted from our contribution. How- with MP2. ever, although it constitutes a large attempt to unravel wing Our survey coupled with phylogenetic data (Winterton venation homologies across Neuroptera, our main focus was to et al., 2010) demonstrates that several similar conditions were highlight a number of enlightening instances of transformation acquired repeatedly among Neuroptera. The fusion of MP2 (i.e. point out putative key taxa). It is anticipated that the cur- with CuA/CuA1 is one of them. It occurs in hindwings of rent contribution will constitute a useful ground for further Osmylidae (Fig. 6B, E), Nymphidae (Fig. 7D), Myrmeleon- studies, focusing on particular transformation cases, ideally tidae (Fig. 8D) and Ascalaphidae (Fig. 9B). In the case of including a documentation of intraspecific variations. The more the two latter families, obviously this acquisition was inde- comprehensive picture of wing venation in Neuroptera that pendent from that in the two former ones, as one of their could be derived from such additional works will allow a more putative stem relatives, namely G. jianboni gen. et sp.n.,lacks reliable assignment of fossil taxa, and improve our knowledge this trait. And according to the phylogeny of Neuroptera pro- of the timing of the group evolution. posed by Winterton et al. (2010), Osmylidae and Nymphidae are only remotely related, and are separated by successive taxa lacking this trait (e.g. Dilaridae, Ithonidae). Interestingly Acknowledgements the fusion of MP2 with CuA/CuA1 in various hindwings is probably the consequence of a translocation, while a ‘sim- We thank S. L. Winterton and another, anonymous, reviewer ilar’ fusion occurred by the mean of a regular fusion in for useful comments. We are very grateful to V. N. Makarkin forewings of G. jianboni gen. et sp.n. (Fig. 8A), Myrmeleon- for comments on primary data on the newly described species. tidae (Fig. 8C), and Ascalaphidae (Fig. 9A). Also, it must We thank Y.-Y. Cui, K. Wolf-Schwenninger and A. Khramov, be noticed that Carpenter (1947) reported a similar fusion to for assistance with photographs of fossil material housed at occur in the forewings of a few Dilaridae species, namely Nal- various institutions. We thank U. and H. Aspock¨ for assistance lachius americanus McLachlan, 1866, and Neodilar hermosa with the determination of recent specimens, and J. Lapeyrie for Banks, 1913. assistance with collecting some of these. We thank the editorial Repeated acquisitions of a pectinate fusion of CuA/CuA1 board of Systematic Entomology, and in particular F. de with MP2 are evident as well. In hindwings, some Osmyli- Guzman for her dedication. This research was supported by dae lack it (Fig. 6B; presumably plesiotypic condition), while the National Basic Research Program of China (973 Program) others possess it (Fig. 6E; presumably derived condition). Its (2012CB821906), the National Natural Science Foundation of occurrence in Nymphidae hindwings (Fig. 7D) can only be China (No. 31071964, 31172143), Scientific Research Key accounted for by a convergence. Finally, it is clear that the Program (KZ200910028005), the China Geological Survey same fusion was acquired in both wing pairs of Myrmeleonti- (1212011120116) and PHR Project of Beijing Municipal dae (Fig. 8C, D) and Ascalaphidae (Fig. 9) independently from Commission of Education (20090509, 201107120). other lineages, as G. jianboni gen. et sp.n. lacksthistrait. 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