[Papers in Palaeontology, 2019, pp. 1–12] STEM-GROUP FOSSILS OF SYMPHRASINAE SHED LIGHT ON EARLY EVOLUTION OF MANTISPIDAE (INSECTA, NEUROPTERA) by CHAOFAN SHI1,2,* , QIANG YANG2,3,4, SHAUN L. WINTERTON5, HONG PANG4 and DONG REN2,* 1School of Earth Sciences and Engineering, Sun Yat-sen University, Guangzhou, 510275, China 2College of Life Sciences, Capital Normal University, Beijing, 100048, China; [email protected] 3School of Life Sciences, Guangzhou University, Guangzhou, 510006, China 4State Key Laboratory of Biocontrol, Key Laboratory of Biodiversity Dynamics and Conservation of Guangdong Higher Education Institute, Ecology and Evolution, School of Life Sciences, Sun Yat-sen University, Guangzhou, 510275, China; 5California State Arthropod Collection, California Department of Food and Agriculture, Sacramento, CA 95832, USA; *Corresponding authors Typescript received 17 September 2018; accepted in revised form 13 December 2018 Abstract: Two new mantid lacewing genera and species, illuminate the evolutionary history of Symphrasinae, as well Archaeosymphrasis pennyi gen. et sp. nov. and Habrosym- as providing insights into the group’s historical biogeogra- phrasis xiai gen. et sp. nov., are described from mid-Cretac- phy. They also provide new evidence for divergence time eous Myanmar amber. Phylogenetic analysis recovered the estimates of extant mantispid subfamilies. new genera as stem-group Symphrasinae, sister to the rest of the subfamily. These represent the oldest fossil record of the Key words: Symphrasinae, Mantispidae, Neuroptera, fossil, subfamily Symphrasinae (Neuroptera, Mantispidae) and evolution. M ANTISPIDAE are a distinctive family of Neuroptera, venational similarities to Drepanicinae (Panfilov 1980; notable for their raptorial forelegs in the adult and Ansorge & Schluter€ 1990; Makarkin 1990; Makarkin & brood-parasite biology of larvae (Lambkin 1986). The Menon 2005; Wedmann & Makarkin 2007), as well as family comprises over 400 species distributed in all major Doratomantispa burmanica (Late Cretaceous of Myanmar) biogeographical regions (Ohl 2004, 2005). They are on both venational and foreleg characters (Poinar & divided into four extant subfamilies, that is Symphrasinae, Buckley 2011). However, the monophyly of the subfamily Drepanicinae, Calomantispinae and Mantispinae, and one is supported by characters of female genitalia (Lambkin extinct subfamily Mesomantispinae, except for a few 1986; Liu et al. 2015), which could not be observed in the extinct genera that are difficult to assign to any existing fossil specimens of the four species mentioned above. Ceno- subfamily (Lambkin 1986; Wedmann & Makarkin 2007; zoic mantispids are typically better preserved and have all Liu et al. 2015; Jepson et al. 2018a, b). Fossil mantispids been assigned to extant subfamilies, namely Mantispinae are known as far back as the Early Jurassic (Ansorge & and Symphrasinae, except for four unnamed larvae (Cock- Schluter€ 1990; Wedmann & Makarkin 2007); the Meso- erell 1921; Jarzembowski 1980; Nel 1989; Poinar 2006; Engel zoic Mantispidae were all restricted to Eurasia, where the & Grimaldi 2007; Wedmann & Makarkin 2007; Ohl 2011; centre of origin is assumed to be (Wedmann & Makarkin Wunderlich 2012). From the Miocene, fossil mantispids 2007; Jepson 2015). Most fossils ranging from Middle have been found beyond Eurasia, that is southern North Jurassic to Early Cretaceous have been assigned to Meso- America (Poinar 2006; Engel & Grimaldi 2007). mantispinae (Makarkin 1996; Jepson et al. 2013, 2018a, b; Symphrasinae are a small subfamily of Mantispidae Khramov 2013; Jepson 2015), although the assignment of (Neuroptera), which are considered to be the most ple- Mesozoic mantipids to extant subfamilies has always been siomorphic clade among extant mantispids, and placed as controversial due to their relatively poor preservation and sister to the remaining three extant subfamilies Drepanici- limited characters visible to diagnose them. For example, nae, Calomantispinae and Mantispinae (Lambkin 1986; Liassochrysa stigmatica (Early Jurassic of Germany), Pro- Liu et al. 2015). The subfamily consists of three extant mantispa similis (Late Jurassic of Kazakhstan) and Ger- genera, Anchieta, Plega and Trichoscelia, entirely restricted staeckerella asiatica (Late Cretaceous of Kazakhstan) show to South America and southern North America (Lambkin © The Palaeontological Association doi: 10.1002/spp2.1265 1 2 PAPERS IN PALAEONTOLOGY 1986; Ohl 2004; Oswald 2018; Appendix 1). In addition, of Insect Evolution & Environmental Changes, College of one extinct genus, Symphrasites, represented by one com- Life Sciences, Capital Normal University, Beijing, China pressed fossil specimen from the Middle Eocene of Ger- (CNUB; Dong Ren, Curator). The holotype of Habrosym- many, was assigned to Symphrasinae (Wedmann & phrasis xiai sp. nov. (LPAM BA180001) is housed in the Makarkin 2007) and considered as the earliest fossil Lingpoge Amber Museum, Shanghai, China. The specimens record of the subfamily heretofore. Symphrasines possess were examined using a Nikon SMZ 25 microscope, illus- unique characters of forelegs and wing venation that dis- trated with the aid of a drawing tube, and photographed tinguish them from the other mantispids, namely foretar- with a Nikon DS-Ri 2 digital camera system. The final sus subdivided into four tarsomeres, apical elongation of drawings and photographs were prepared with the aid of the basitarsus, the forewing with two r1-rs crossveins and Adobe Illustrator CS6 and Adobe Photoshop CS6. the hind wing with one r1-rs crossvein (Lambkin 1986; Wing venation and terminalia terminology follows Aspock€ & Mansell 1994). However, the specimen of Sym- Lambkin (1986). Abbreviations for wing venation: 1A–3A, phrasites was only preserved with one incomplete forew- first to third anal veins; CuA, cubitus anterior; CuP, cubi- ing. Consequently, it is difficult to infer whether the tus posterior; hv, humeral vein; MA, media anterior; MP, venational and foreleg characters of the subfamily were media posterior; R1, radial first branch; RS, radial sector; derived simultaneously or gradually. This further resulted Sc, subcosta. in the question regarding whether the extant symphrasi- nes acquired and developed their significant foreleg mor- Phylogenetic analysis phology before, or after, their dispersal from Eurasia. Furthermore, during their evolutionary history of over A phylogenetic analysis was conducted to investigate the 180 MA, at which period did Symphrasinae diverge from phylogenetic placement of the new amber taxa with regard the other lineages? to Symphrasinae and the other mantispid subfamilies. Herein we describe two new genera and two new species (Archaeosymphrasis pennyi gen. et sp. nov. and Habrosym- Morphological examination. It has been demonstrated that phrasis xiai gen. et sp. nov.) from mid-Cretaceous Myan- the reconstruction of the phylogenetic affinity of fossil taxa mar amber, dated at c. 99 Ma. The new taxa demonstrate is extremely sensitive to the strategy used to interpret non- characters of the significant foreleg and venation concor- preserved characters. Whether treated as missing or absent, dant with Symphrasinae, yet a few venational characters taphonomic loss of present characters causes taxa to slide indicating more plesiomorphic states. Phylogenetic analysis down trees, toward the root (Sansom 2015). Analyses of liv- recovered them as stem-group members of the Symphrasi- ing and fossil taxa based on morphological characters that nae, sister to the Cenozoic and extant symphrasines. The are observable and significant for the lineage mostly result discovery of these well-preserved, intricate amber speci- in the position of fossil taxa remaining unresolved, presum- mens prompted us to scrutinize the early evolution of ably due to large amounts of missing data (unpublished Symphrasinae, which also sheds light on subfamilial diver- data). Based solely on observable characters we included gence estimates of Mantispidae during the Late Mesozoic. nineteen morphological characters from prothorax, foreleg, forewing and hind wing venation, which were coded for the outgroup and ten ingroup taxa. Morphological characters MATERIAL AND METHOD used in the phylogenetic analysis are listed in Appendix 2. We coded 17 characters as binary and 2 as multistate. Taxonomy Unknown characters were coded as ‘?’. The specimens described here were collected from the Exemplar selection. The ingroup included the two new Hukawng Valley of the northern state of Kachin in Myan- described genera, Archaeosymphrasis and Habrosymphrasis; mar (26°2000N, 96°3600E). The site within the Hukawng the other fossil genus (Symphrasites), all three extant genera Basin is comprised of sedimentary (volcanic) lithic clasts, of Symphrasinae (Anchieta, Plega and Trichoscelia), along with minor fragments of quartz and feldspar (Cruick- with representatives from the four other subfamilies of shank & Ko 2003; Shi et al. 2012). Based upon U–Pb dat- Mantispidae, (i.e. Drepanicinae, Calomantispinae, Man- ing of zircons, the Myanmar amber was probably formed tispinae and Mesomantispinae). The three extant genera at (but not earlier than) c. 98.79 Æ 0.62 Ma, in the earli- and four subfamilies were respectively represented as single est Cenomanian (Grimaldi et al. 2002; Cruickshank & Ko composite taxa in the analysis. Especially for Mesoman- 2003; Poinar & Buckley 2011; Shi et al. 2012, 2015; Chen tispinae, an almost complete set of characters were coded et al. 2019;