[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 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 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 (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 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. 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 0.62 Ma, in the earli- and four subfamilies were respectively represented as single est (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; Zhang et al. 2018). from different species of the subfamily, instead of many The holotype of Archaeosymphrasis pennyi sp. nov. exemplars each with large amounts of missing data. Poly- (CNU-NEU-MA2018065) is housed in the Key Laboratory morphisms were scored as multiple states in these taxa. SHI ET AL.: AMBER SYMPHRASINAE FROM MID-CRETACEOUS MYANMAR 3

Phylogenetic analysis. Parsimony analyses were conducted produced into a large spinose process, with the second in TNT (Goloboff et al. 2008) using an heuristic search tarsomere arising from the centre of its medial face. Pre- that included 500 random addition sequence replicates, tarsus with a pair of simple claws and an arolium. Tibia holding 10 trees per replication after tree bisection and and tarsus together shorter than femur. Forewing humeral reconnection (TBR) for branch swapping and 90 itera- vein recurrent. Costal crossveins mostly forked. Sc fused tions of ratchet (Nixon 1999). In all cases, branch support with R1 distally. One sc-r1 and three r1-rs present. Rs was assessed by Jackknife calculated from 3000 pseu- with less than five branches. One gradate series present. doreplicates of re-sampled data sets. All characters were Hind wing with two r1-rs. Hind wing CuA longer than treated as unordered and with equal weight. Parsimony half of wing length. analyses were also conducted using WinClada v. 1.00.08 (Nixon 2002) and NONA v. 2.0 (Goloboff 1998; Goloboff Remarks. Archaeosymphrasis is assigned to Symphrasinae et al. 2008), using a branch and bound search protocol to based on foreleg and wing venational characters such as find all trees (options: set to hold 10 000 trees, 1000 foretarsus with four tarsomeres; basitarsus enlarged and replications, 100 starting tree replication, multiple spinously produced; forewing CuP very closely approxi- TBR + TBR search strategy). Bootstrap support values mating 1A proximally; pterostigma situated between were determined using PAUP*4.0b10 (Swofford 1998) cal- costal margin and Sc+R1 (see Discussion below). The culated from 1000 heuristic search (TBR) pseudoreplicates genus can be distinguished from the extant symphrasine of re-sampled data sets, each with 30 random additions genera (Anchieta, Plega and Trichoscelia) by forewing long (constant characters excluded). Parsimony analyses were recurrent humeral vein present with branches; costal conducted using the morphological matrix containing 19 crossveins mostly forked; three r1-rs crossveins; MP sepa- characters and 11 taxa (Shi et al. 2019), as well as the same rated from R proximally; hind wing with two r1-rs cross- matrix but excluded Symphrasites, due to large amounts of veins; hind wing Sc fused with R1 distally, costal space missing data in this genus. Character states were mapped almost as long as wing length. Archaeosymphrasis differs on a most parsimonious tree (MPT) using WinClada from Symphrasites on forewing three r1-rs (two r1-rs in v. 1.0 (Nixon 2002), showing only unambiguous changes. Symphrasites), fewer than five Rs branches (Rs with 11 branches in Symphrasites), MP2 forked distally (MP2 (in- ferred as MP in Wedmann & Makarkin 2007) forked SYSTEMATIC PALAEONTOLOGY proximally in Symphrasites). by Chaofan Shi, Qiang Yang and Dong Ren Archaeosymphrasis pennyi sp. nov. Figure 1 Class INSECTA Linnaeus, 1758 Order NEUROPTERA Linnaeus, 1758 Family MANTISPIDAE Leach, 1815 LSID. urn:lsid:zoobank.org:act:55922730-471D-41D4-B9DF- A4FB77B97179 Subfamily SYMPHRASINAE Navas, 1909 Genus ARCHAEOSYMPHRASIS nov. Derivation of name. The species is dedicated to Dr Nor- LSID. urn:lsid:zoobank.org:act:8459A249-AF9F-4E0E-81CA- man D. Penny, who contributed greatly to the taxonomy 8E46853090B2 of various lacewing families, especially to Symphrasinae, and helped with the extant specimens in this project. Derivation of name. Archaeosymphrasis is derived from the Greek archaeo-, meaning ancient, primitive, combined Material. Holotype: CNU-NEU-MA2018065. with symphrasis, a combination of the Greek sym- and phrasis, derived from the subfamily name Symphrasinae, Diagnosis. As for the genus. referring to primitive symphrasines. Gender feminine. Description. Body length c. 9.1 mm. Forewing length 8.9 mm, Type species. Archaeosymphrasis pennyi sp. nov. width 3.7 mm; hind wing length 8.0 mm, width 3.0 mm. Forecoxa length 1.9 mm, width 0.26 mm; forefemur length 2.2 mm, width 0.55 mm/0.65 mm (left/right femur, respectively); foretibia length Diagnosis. Forefemur wide, bearing closely spaced spine- 1.9 mm, first to fourth tarsomere length 0.53, 0.16, 0.07, 0.20 mm. like setae along lateroventral ridge. Foretibia arched, bear- Body. Head wider than high. Vertex and frons slightly domed, ing a row of closely spaced, short, thick, prostrate setae with tiny tubercles on the surface. Compound eyes large, covered with apices pointing distally along the ventral ridge. Fore- nearly half of the head surface. Antenna moniliform, with 25 tarsus with four tarsomeres. The basitarsus enlarged and antennomeres. Scape wider than and almost twice as long as 4 PAPERS IN PALAEONTOLOGY

A B

C

D

E

F

FIG. 1. Archaeosymphrasis pennyi sp. nov. A, habitus in dorsal view. B, camera lucida drawing in dorsal view with minor modifica- tions from photo images. C, abdomen in ventral view. D, forewing venation. E, hind wing venation. F, foreleg in inner lateral view showing foretrochanter, forefemur, foretibia and foretarsus. Scale bars represent: 1 mm (A–E); 0.5 mm (F). Colour online. SHI ET AL.: AMBER SYMPHRASINAE FROM MID-CRETACEOUS MYANMAR 5 pedicel and flagellomere. Prothorax moderately elongate. Prono- from the subfamily name Symphrasinae, referring to the tum with distinct hump (Fig. 1A, B). Foreleg raptorial. Forecoxa delicate feature and finely preservation of the holotype. located at the anterior of the prothorax. Femur wide, not laterally Gender feminine. compressed along the ventral margin. One short cuticular spine with equally long apical setae present on the ventral surface of Type species. Habrosymphrasis xiai sp. nov. femur, located inside the midway of basitarsus. Two rows of spine-like setae along lateroventral ridge of femur, arranged inside Diagnosis. Forefemur stout, bearing closely spaced spine- and outside tibia. Tibia arched. Tarsus with four tarsomeres. The like setae along lateroventral ridge. Foretibia arched, bear- basitarsus enlarged with spinose process, with the second tar- somere arising from the centre of its medial face. Tibia and the ing a row of closely spaced, short, thick, prostrate setae basitarsus with a row of closely spaced, short, thick, prostrate setae with apices pointing distally along the ventral ridge. Fore- with apices pointing distally along the ventral ridge. The second tarsus with four tarsomeres. The basitarsus enlarged and and fourth tarsomeres distinctly longer than the third. Pretarsus spinately produced, with the second tarsomere arising with a pair of simple claws and an arolium (Fig. 1F). from the centre of its medial face. Pretarsus with a pair Wings oval, with trichosors along entire wing margin except of simple claws and an arolium. Tibia and tarsus together for the very base. One trichosor present between adjacent veins. shorter than femur. Forewing humeral vein recurrent. Forewing humeral vein recurrent. Costal area broad, narrowed Costal crossveins mostly simple. More than one trichosors distally. Costal crossveins mostly forked. Sc fused with R1 near present between adjacent veins along wing margin. Sc distal one-third of wing length. Pterostigma formed by costal convex and approaching costal margin before sharply margin, partial Sc and Sc+R1, with the incorporated crossveins recurving and fusing with R1. One sc-r1 and three r1-rs faint. One sc-r1 crossvein present, located in the midway of sub- costal area. Rs separated from R1 near basal one-third of wing present. Rs with less than five branches. One gradate ser- length. Three r1-rs crossveins present. Rs with four branches. ies present. Hind wing with two r1-rs. One gradate series present. MA separated from Rs near the mid- dle of wing length. CuP concave and approaching 1A near base. Remarks. Habrosymphrasis is assigned to Symphrasinae One short cup-1a connected the concave point of CuP and 1A based on same foreleg and wing venational characters as (Fig. 1D). Hind wing narrower and slightly more pointed at the Archaeosymphrasis. Habrosymphrasis can be distinguished apex than forewing. Costal area narrow. Costal crossveins simple from the extant symphrasine genera (Anchieta, Plega and before the fusion of Sc and R1. Two r1-rs present. Rs separated Trichoscelia) by forewing having long recurrent humeral from R near basal one-fourth of wing length. Rs with four vein present with branches; three r1-rs crossveins; hind branches. MA and MP separated near basal one-eighth of wing wing with two r1-rs crossveins; hind wing Sc fused with R1 length. The basal MA sinuous, fused with Rs for a short distance distally, costal space almost as long as wing length. around the basal one-third of wing length. CuA and CuP sepa- Habrosymphrasis differs from Symphrasites on forewing rated proximal to the separation of MA and MP. CuA long and having three r1-rs, fewer than five Rs branches, MP2 parallel with posterior margin, terminated in the distal half of wing length. CuP long and distally creeping. CuP touching 1A forked distally. Habrosymphrasis mostly resembles near base. The distal half of CuP parallel and close to posterior Archaeosymphrasis, but can be differentiated from the latter margin, with marginal twiggings. One long and sinuous cua-cup by forewing having costal crossveins mostly simple (mostly and one short cup-1a present closely before the CuP marginal forked in Archaeosymphrasis), multiple trichosors present twiggings. 1A and 2A long. All anal veins with marginal twig- between adjacent veins along proximal costal margin (one gings. 1a–2a present (Fig. 1E). trichosor between adjacent veins in Archaeosymphrasis), Sc Abdomen with nine tergites and ectoproct visible. The sev- convex and approaching costal margin before fusing with enth sternite with lateral edges protruded. Genitalia not pre- R1 (Sc not approaching costal margin in Archaeosymphra- served or unobservable (Fig. 1C). sis), hind wing CuA not distinctly longer than half of wing length (hind wing CuA distinctly longer than half of wing Type locality and horizon. Hukawng Valley, Kachin State, length, with more branches in Archaeosymphrasis). Myanmar; earliest Cenomanian, of Late Cretaceous, dated at 98.79 0.62 Ma based on a 206Pb/238U concordia Habrosymphrasis xiai age (Shi et al. 2012). sp. nov. Figures 2, 3

Genus HABROSYMPHRASIS nov.

LSID. urn:lsid:zoobank.org:act:EEF296B0-A076-44E0-9D83- LSID. urn:lsid:zoobank.org:act:E7C8082E-4018-42BB-9EBE- AFDCEFB378D3 5A21CB06772A

Derivation of name. Habrosymphrasis is derived from the Derivation of name. The specific name is dedicated to Mr Greek habr-, meaning delicate, combined with symphrasis, Fangyuan Xia, Director of the Lingpoge Amber Museum, a combination of the Greek sym- and phrasis, derived for his contribution to the study of . 6 PAPERS IN PALAEONTOLOGY

A

B

C

FIG. 2. Habrosymphrasis xiai sp. nov. A, habitus in dorsolateral view. B, head and thoraces in ventrolateral view. C, camera lucida drawing in dorsolateral view, with minor modifications from photo images. Scale bar represents 1 mm. Colour online.

Material. Holotype: LPAM BA180001.

Diagnosis. As for the genus.

Description. Body length 6.0 mm. Forewing length c. 6.0 mm, width 1.9 mm; hind wing length c. 4.7 mm, width c. 1.3 mm. Forecoxa length 1.4 mm, width 0.30 mm; forefemur length 1.4 mm, width 0.38 mm; foretibia length 1.1 mm, basitarsus length 0.25 mm. Body. Head wider than high. Vertex relatively flat. Compound eyes large. Antenna moniliform, with 29 antennomeres. Scape longer and wider than pedicel and flagellomeres. Prothorax moderately elongate. The posterior part of pronotum ventrally extended and merged in the midline of ventral. Forecoxa elon- gate and located at the anterior of prothorax. Femur slightly wider than coxa, not laterally compressed along the ventral mar- FIG. 3. Habrosymphrasis xiai sp. nov. camera lucida drawing gin. One basal femoral spine formed of partial cuticular and api- with minor modifications from photo images: A, forewing vena- cal spine-like setae present inside the midway of basitarsus, tion; B, hind wing venation. Scale bar represents 1 mm. SHI ET AL.: AMBER SYMPHRASINAE FROM MID-CRETACEOUS MYANMAR 7 slightly longer than others. A row of spine-like setae arranged length, with four branches. One gradate series present. along the lateroventral ridge of femur, outside the tibia, includ- MA and MP separated around basal one-sixth of wing ing one long half cuticular spine with apical setae near the tibia length. MA basal part sinuous, fused with Rs for a short apical. Femoral spine or setae inside the tibia mostly unobserv- distance around the basal one-third of wing length. Two able. Tibia arched. The basitarsus enlarged and spinously pro- mp-cua present. The basal one close to the separation of duced, with the following tarsomere arising from the centre of MA and MP. CuA slightly longer than half of wing its medial face. Pretarsus with a pair of simple claws (Fig. 2). length. CuP long and distally creeping. CuP distal part Wings elongate oval. Forewing with trichosors along costal margin except for the basal area and apical half of posterior parallel and close to posterior margin, with marginal margin. More than one trichosors present between adjacent twiggings. One long and sinuous cua-cup present closely veins on costal margin. Humeral vein recurrent. Costal area before the CuP marginal twiggings. 1A long and distal broad near base, narrowed distally. Sc convex and approaching, forked. cup-1a moderately long (Fig. 3B). nearly touching costal margin for a short distance before fused Abdomen with nine tergites and ectoproct visible. One elon- with R1. Sc fused with R1 near distal one-third of wing length. gate and curved structure extended from the end of abdomen, Pterostigma distinct, formed by costal margin, partial Sc and ventral to ectoproct (Fig. 2A, C). Sc+R1, with the incorporated crossveins faint. One sc-r1 present, located in the midway of subcostal area. Rs sepa- Type locality and horizon. Hukawng Valley, Kachin State, rated from R1 near basal one-third of wing length. Three Myanmar; earliest Cenomanian, Late Cretaceous, dated at r1-rs present. Rs with four branches. One gradate series 98.79 0.62 Ma based on a 206Pb/238U concordia age present. MP separated from R around basal one-sixth of (Shi et al. 2012). wing length. One cua-cup present. 1A long. 1A and 2A distally forked (Fig. 3A). Hind wing with trichosors along PHYLOGENETIC RESULTS posterior margin and apical area. Costal area narrow. Sc fused with R1 near distal one-third of wing length. Two Parsimony analysis of the morphological matrix contain- r1-rs present. Rs branched from midway along wing ing 19 characters and 11 taxa recovered a single MPT

A

E

B

F

C

G

D

FIG. 4. Phylogenetic relationship among Mantispidae. Single tree recovered from parsimony analysis. Tree topology shows the analy- ses result of the intact matrix of 11 taxa. Character state changes are plotted on each node, as well as subtended by statistical support values (Bootstrap: Bremer support). Nodes support values are the result from the restricted matrix excluded Symphrasites. Arrows on the geological time scale indicate fossil ages of: Mesomantispinae; Archaeosymphrasis and Habrosymphrasis; Symphrasites. Foreleg diver- sity of Mantispidae and wing venation evolution of Symphrasinae: A, Longipronotum benmaddoxi (Mesomantispinae: Late Jurassic), modified from Jepson et al. (2018a, fig. 1B); B, Trichoscelia varia (Symphrasinae: Recent), modified from Tjeder (1959, fig. 247); C, Drepanicus gayi (Drepanicinae: Recent); D, Dicromantispa electromexicana (Mantispinae: Miocene), modified from Engel & Grimaldi (2007, fig. 6); E, Archaeosymphrasis pennyi sp. nov. (Symphrasinae: Late Cretaceous); F, Habrosymphrasis xiai sp. nov. (Symphrasinae: Late Cretaceous); G, Trichoscelia remipes (Symphrasinae: Recent), modified from Hoffman (2002, fig. 542). Colour online. 8 PAPERS IN PALAEONTOLOGY

(length = 27; consistency index (CI) = 0.77; retention wing margin; (viii) multiple trichosors between adjacent index (RI) = 0.87) (Fig. 4). The MPTs recovered by TNT marginal veins. Characters (i) to (iv) were considered to and NONA were congruent in topology, except for minor be synapomorphies of Symphrasinae by Lambkin (1986), differences among Drepanicinae, Calomantispinae and while (v) to (vii) were considered to be the plesiomorphic Mantispinae. Parsimony analysis of the restricted matrix conditions distinguished from Drepanicinae, Caloman- recovered a single MPT of identical topology of the ten tispinae and Mantispinae. Character (viii) was considered taxa to the tree based on the intact matrix, but with to be a synapomorphy of Mantispidae by Aspock€ & Man- increase in the consistency index and retention index sell (1994), although this is only present in Symphrasinae (length = 26; CI = 0.80; RI = 0.88) and notable increase and some Drepanicinae (Wedmann & Makarkin 2007). in support values on every node. Support values for boot- But this character is difficult to identify in compression strap (BS) and Bremer support (B) of the analysis based fossils, so we decide to exclude this character from the on restricted matrix are plotted on individual nodes in matrix in the phylogenetic analysis. Figure 4. Archaeosymphrasis and Habrosymphrasis differ from the Mantispidae were recovered as monophyletic. Within crown group of Symphrasinae, that is Anchieta, Plega and the family, there was a distinctive basal dichotomy sepa- Trichoscelia by characters: (ix) forewing with three r1-rs rating two clades. One comprises Drepanicinae, Caloman- crossveins (forewing two r1-rs in the three extant genera); tispinae and Mantispinae. The monophyly of this clade (x) hind wing with two r1-rs crossveins (hind wing one was well supported (BS: 88; B: 3), although the relation- r1-rs in extant genera); (xi) forewing MP separated from ships among the three subfamilies differ between different R proximally, basal to 1 m-cu (MP fused with R for a analysis. The other clade comprises Mesomantispinae and considerable distance distal to 1 m-cu in extant genera); Symphrasinae. But the monophyly of this clade was (xii) hind wing Sc fused with R1 distally, costal space weakly supported (BS: 56; B: 1), possibly due to the large almost as long as wing length (hind wing Sc ended in amounts of missing data for Mesomantispinae. This sug- costal margin, resulting in costal space terminating in gests that the phylogenetic status of Mesomantispinae basal half of wing length in extant genera). The above needs further study. four characters possessed by Anchieta, Plega and Tri- Symphrasinae containing extant and extinct genera choscelia were considered as synapomorphies of Sym- were recovered as monophyletic (BS: 81; B: 3) and were phrasinae (Lambkin, 1986). Archaeosymphrasis and supported in this analysis by multiple apomorphic char- Habrosymphrasis are also distinguished by (xiii) forewing acters, including forewing CuP and 1A approximating long recurrent humeral vein present with branches. This proximally (character 6: state 1), foretarsus four subdivi- character is considered plesiomorphic, as it is also present sion (18:1), and basitarsus elongated and spinously pro- in Mesomantispinae and occasionally in Plega and Tri- duced distally (19:1). The extant symphrasines were choscelia. Archaeosymphrasis is distinguished from other recovered as strongly monophyletic (BS: 97; B: 4), sister Symphrasinae by (xiv) forewing costal crossveins mostly to Symphrasites. Habrosymphrasis was recovered as sister- forked, which is considered to be plesiomorphic in Man- group of the clade including Symphrasites and extant tispidae; this character is also found in Mesomantispinae symphrasines; Archaeosymphrasis was recovered sister to and some Drepanicus (Fig. 4E–G). all the other symphrasines. One other fossil symphrasine from the Eocene, Symphr- asites, was represented by a specimen preserving one forewing. The forewing venation of Symphrasites possess DISCUSSION two r1-rs crossveins (related to ix) and probable pteros- tigma situated between costal margin and R1, formed by Phylogeny of Symphrasinae Sc recurving sharply and fusing with R1 (iv). It is worthy of note that in the original description of Symphrasites, Archaeosymphrasis and Habrosymphrasis are assigned to the sharply recurving Sc distal was interpreted as 3sc-r Symphrasinae based on: (i) foretarsus with four tar- (Wedmann & Makarkin 2007). Moreover, the forewing someres; (ii) basitarsus enlarged and spinously produced trichosors present only in apical portion of wing margin with the second tarsomere arising from the centre of the (related to vii), which is considered as autapomorphy of medial face; (iii) forewing CuP very closely approximating Symphrasites. 1A proximally; (iv) pterostigma situated between costal We consider Archaeosymphrasis and Habrosymphrasis as margin and R1, formed by Sc recurving sharply and fus- stem-group Symphrasinae based on these data. In this ing with R1; (v) forefemoral spines composed of long sense, the synapomorphies of the subfamily Symphrasinae black setae and cuticular spines with black setae distally, are characters (i) to (iv): foretarsus four subdivision, with (vi) hind wing with the stem MA sinuous and longitudi- basitarsus enlarged and spinately produced with the sec- nal; (vii) wing with trichosors along almost the entire ond tarsomere arising from the centre of its media face; SHI ET AL.: AMBER SYMPHRASINAE FROM MID-CRETACEOUS MYANMAR 9 forewing CuP very closely approximating 1A proximally, dispersal-local extinction/ousted relict scenario applies to pterostigma situated between costal margin and R1, Mantispidae (Wedmann & Makarkin 2007). With regards formed by Sc recurving sharply and fusing with R1. It to the distribution of extant symphrasines being confined reveals that these foreleg and forewing venation characters to the tropics and subtropics, the dispersal of symphrasi- were already present in mantispids as early as the mid- nes is hypothesized to have taken place along a southern Cretaceous. Thulean route from southern Europe to North America Forewing with two r1-rs (related to ix) should be via Iceland and Great Britain (Tiffney 1985), where the synapomorphy of the lineage including Symphrasites, climate was inferred to be mostly subtropical during the Anchieta, Plega and Trichoscelia, which form the Cenozoic Paleocene/Eocene (Sluijs et al. 2006). But as this connec- symphrasines. It indicates this character might have tion was severed during the early Eocene (Tiffney 1985), evolved in Symphrasinae since around the Paleocene. the dispersal to North America should have happened Other characters that were inferred to be synapomorphies prior to this. Subsequently, the symphrasines dispersed to of Symphrasinae are: (xv) hind wing with a small South America after closure of the Central American Sea- emargination on proximal posterior margin; (xvi) female way in Middle Miocene (Montes et al. 2015). Alterna- ninth gonocoxites strongly elongated and upcurved to tively, the Burmese amber has also been suggested to have form a long ovipositor; and (xvii) female ectoprocts fused a Gondwanan origin (Rangin 2017; Poinar 2018), which with ninth tergite (Lambkin, 1986). Holotypes of both would give rise to an alternative hypothesis that Mantispi- A. pennyi sp. nov. and H. xiai sp. nov. have separated dae initially had a wider historical distribution, followed ninth tergite and ectoproct. H. xiai sp. nov. possessed an by rampant extinction leading to the present disparate elongate and curved structure similar to an ovipositor. distribution pattern. A more complete fossil history, more But genitalia are mostly unobservable on both specimens, information for the palaeogeography of the West Burma making it difficult to determine their gender. Therefore, Block and future historical biogeographic analysis are characters xv to xvii are still difficult to identify confi- needed to investigate the hypotheses, and the extinction dently in the fossil taxa. Herein, we tentatively consider of the stem-group symphrasines. the synapomorphies of crown group symphrasines, that is Anchieta, Plega and Trichoscelia, as forewing MP fused with R for a considerable distance distal to 1m-cu; hind Early evolution of Mantispidae wing Sc ended in costal margin, resulting in costal space terminating in basal half of wing length, with one r1-rs, Mantispidae have a geological history extending to Early and a small emargination on proximal posterior margin; Jurassic (Ansorge & Schluter€ 1990; Wedmann & Makar- female ninth tergite fused with ectoprocts, ninth gonocox- kin 2007), and exhibit diverse morphologies, especially in ites strongly elongated and upcurved to form a long the foreleg (Fig. 4A–D). Most Mesozoic mantispids have ovipositor, related to (x, xi, xii, xv, xvi, xvii). been assigned to Mesomantispinae (Jepson et al. 2013, 2018a–b; Khramov 2013; Jepson 2015), whereas a few others are still controversial regarding their subfamilial Historical biogeography placement. Among the extant subfamilies, Caloman- tispinae and Mantispinae possess distinct synapomor- The newly documented stem-group Symphrasinae from phies, such as fore, mid and hind tarsal claws bifid the Late Cretaceous of Myanmar represent members from (Calomantispinae), forecoxa with a transverse sulcus, the early divergence of the Mantispidae. The Burmese foretarsus with one simple claw, without arolium, fore- amber mines are located on the West Burma Block, wing MP stem forming a small triangular with R (Man- which was inferred to have become part of Southeast Asia tispinae). These make it easy to identify fossil taxa of during Cretaceous by Metcalfe (2013, 2017) and Zahi- these subfamilies and points to them being relatively rovic et al. (2014). This agrees with the hypothesis of a derived groups within the Mantispidae (Lambkin 1986; Eurasian origin of Mantispidae and their marked diversi- Liu et al. 2015). Fossil Mantispinae have been found in fication during late Mesozoic. Furthermore, the occur- Eocene aged deposits (Cockerell 1921; Jarzembowski rence of Symphrasites in the Eocene of Germany implies 1980), but no fossil calomantispines have yet been their appearance in Europe during the early Cenozoic, reported (Jepson 2015). Drepanicinae were considered along with the diversification of Symphrasinae resulting monophyletic, weakly supported by female genitalia char- in both foreleg and wing venation differentiation (charac- acters (Lambkin 1986; Liu et al. 2015), which makes it ters i–iv, ix, this analysis) (Wedmann & Makarkin 2007). difficult to assign fossil taxa to this subfamily confidently, However, all extant symphrasines are restricted to although some Mesozoic taxa (Liassochrysa, Promantispa, Neotropical and southern Nearctic regions (Ohl 2004; Doratomantispa) were considered to be drepanicines Oswald 2018). It has therefore been proposed that a based on wing venation and foreleg similarities (Poinar & 10 PAPERS IN PALAEONTOLOGY

Buckley 2011; Liu et al. 2015). Archaeosymphrasis and ENGEL, M. S. and GRIMALDI, D. A. 2007. The neuropterid Habrosymphrasis are the earliest fossil taxa which can be fauna of Dominican and Mexican Amber (Neuropterida: Mega- confidently assigned to the extant subfamily. They pro- loptera, Neuroptera). American Museum Novitates, 3587,1–58. vide important fossil evidence for the origin of Sym- GOLOBOFF, P. A. 1998. NONA, Version 2.0. Published by phrasinae and divergence time of the extant subfamilies. the author, Tucuman, Argentina. -FARRIS, J. S. and NIXON, K. C. 2008. TNT, a free Regarding the uncertain phylogenetic status of Mesoman- program for phylogenetic analysis. Cladistics, 24, 774–786. tispinae, it indicates that Symphrasinae diverged from the GRIMALDI, D., ENGEL, M. S. and NASCIMBENE, P. other extant subfamilies during the Early Cretaceous, 2002. Fossiliferous Cretaceous amber from Myanmar (Burma): which agrees with the estimated divergence time based on its rediscovery, biotic diversity, and paleontological signifi- molecular sequence data (Winterton et al. 2010). cance. American Museum Novitates, 3361,1–72. HOFFMAN, K. 2002. Family Mantispidae. 251–275 (text), Acknowledgements. We are grateful to Dr Ivonne J. Garzon- 419–432 (figures). In PENNY, N. D. (ed.) A guide to the Orduna~ for advices on parsimony phylogenetic analysis, and Dr lacewings (Neuroptera) of Costa Rica. Proceedings of the Cali- Michael Ohl for providing of extant species images for this fornia Academy of Sciences, Series 4, 53 (12), 161–457. study. Our thanks go to Sally Thomas and two anonymous ref- JARZEMBOWSKI, E. A. 1980. Fossil from the Bem- erees for helpful suggestions on earlier drafts of this paper. This bridge Marls, Paleogene of the Isle of Wight, southern Eng- study was supported by National Natural Science Foundation of land. Bulletin of the British Museum of Natural History China (grant nos. 31501881, 31711530649, 41602014, 31730087), (Geology), 33, 237–293. Basal Research Fund of Sun Yat-sen University (grant no. 17l- JEPSON, J. E. 2015. A review of the current state of knowledge gzd04), Program for Changjiang Scholars and Innovative of fossil Mantispidae (Insecta: Neuroptera). Zootaxa, 3964, Research Team in University (grant no. IRT-17R75), Support 419–432. Project of High-level Teachers in Beijing Municipal Universities -HEADS, S. W., MAKARKIN, V. N. and REN, D. (IDHT20180518), and The Basic Work Special Project of the 2013. New fossil mantidfies (Insecta: Neuroptera: Mantispi- National Ministry of Science and Technology of China dae) from the Mesozoic of north-eastern China. Palaeontology, (2013FY111500). 56, 603–613. -KHRAMOV, A. V. and OHL, M. 2018a. New Meso- mantispinae (Insecta: Neuroptera: Mantispidae) from the DATA ARCHIVING STATEMENT Jurassic of Karatau, Kazakhstan. 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APPENDIX 1 CHECKLIST OF EXTINCT AND EXTANT SYMPHRASINAE GENERA

Anchieta Navas, 1909 Distribution: Neotropic †Archaeosymphrasis gen. nov. Distribution: Myanmar; Age: Late Cretaceous †Habrosymphrasis gen. nov. Distribution: Myanmar; Age: Late Cretaceous Plega Navas, 1928 Distribution: Nearctic and Neotropic †Symphrasites Wedmann & Makarkin, 2007 Distribution: Germany; Age: Eocene Trichoscelia Westwood, 1852 Distribution: Neotropic

APPENDIX 2 MORPHOLOGICAL CHARACTER DESCRIPTIONS

1. Forewing humeral vein 0 recurrent; 1 simple 2. Forewing costal crossveins 0 mostly forked; 1 mostly simple 3. Forewing Sc termination 0 fused with R1; 1 fused with C 4. Forewing r1-rs crossveins number 0 three or more; 1 two 5. Forewing MP stem separation from R 0 proximal; 1 distal to 1m-cu 6. Forewing CuP and 1A proximally 0 detached; 1 approximating or touching 7. Hind wing costal space 0 almost as long as wing length; 1 shorter than half wing length 8. Hind wing r1-rs crossveins number 0 two or more; 1 one 9. Hind wing MA stem 0 longitudinal; 1 not longitudinal 10. Hind wing CuP 0 entirely present; 1 entirely or basal absent 11. Wing trichosors range 0 almost entire wing margin; 1 wing apex or absent 12. Wing pterostigma 0 inconspicuous; 1 distinct 13. Prothorax 0 not elongated posterior to forecoxae; 1 elongated posterior to forecoxae 14. Pronotum maculae 0 absent; 1 present 15. Foreleg 0 cursorial; 1 raptorial 16. Forefemoral basal spine 0 spine absent; 1 same as other spines; 2 distinct elongate; 17. Forefemoral spines 0 short thin setae; 1 mainly long, thick setae; 2 cuticular spines; 18. Foretarsus 0 five tarsomeres; 1 four tarsomeres 19. Basitarsus 0 not elongated; 1 elongated apically