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 © 2012 The Authors Systematic Entomology © 2012 The Royal Entomological Society 617 618 C. Shi et al. 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