Cladistics

Cladistics (2015) 1–36 10.1111/cla.12104

Phylogeny of split-footed lacewings (Neuroptera, Nymphidae), with descriptions of new Cretaceous fossil species from China

Chaofan Shia,b, Shaun L. Wintertonc and Dong Renb,*

aSchool of Earth Science and Geological Engineering, Sun Yat-sen University, Guangzhou 510275, China; bCollege of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100048, China; cCalifornia State Arthropod Collection, California Department of Food and Agriculture, Sacramento, CA 95832, USA Accepted 27 October 2014

Abstract

A phylogeny of the lacewing family Nymphidae based on morphology and DNA sequences is presented including representa- tives of all living genera and selected fossil genera. Widely distributed Jurassic and Cretaceous genera gave rise to recent taxa now restricted to Australasia. Two previously defined clades (i.e. Nymphinae and Myiodactylinae) were recovered and reflect the diverging adult and larval morphology of members of these two subfamilies. From Chinese Cretaceous deposits, a new genus (Spilonymphes gen. nov.) is described with one new species, as well as new species described in the genera Baissoleon Makarkin and Sialium Westwood. © 2015 The Authors Cladistics published by John Wiley & Sons Ltd on behalf of Willi Hennig Society.

Nymphidae are a small family of distinctive Diagnostic features of nymphid imagos include neuropterans characterized by a medially divided aro- elongated filiform antennae, ocelli absent, legs with a lium, leading to their common name of split-footed bifid arolium, wings with trichosors present, nygmata lacewings. The family is considered one of the more absent and thyridiate (incomplete) crossveins present plesiomorphic clades of Myrmeleontiformia, sister to a of varying length along the subcostal space. Nymphi- group comprising at least Myrmeleontidae (antlions) dae are one of the few families of lacewings that lay and Ascalaphidae (owlflies) (Aspock€ et al., 2001, 2012; eggs on silken stalks (New, 1982, 1983a). Arguably, Aspock,€ 2002; Engel and Grimaldi, 2008) or also the most specialized arrangement of eggs on silken including Nemopteridae (spoon-winged lacewings) stalks found in Neuroptera is produced by the rela- (New, 1984; Oswald, 1998; Winterton et al., 2010). tively derived genus Nymphes Leach. Eggs are laid Extant Nymphidae are restricted entirely to the Aus- with every second egg produced with a silken stalk, tralasian region, the greatest diversity being found in and intermediate eggs produced perpendicular to the eastern mainland Australia with a few species found previous one and used to connect the stalked eggs such in western Australia and New Guinea (New, 1987). A that the egg mass is arranged in a large “U”-shaped single species has been described from the Philippines pattern. Immature stages are distinctive and are (i.e. Myiodactylus chrysopoides Navas) but the type known for the genera Osmylops Banks, Nymphes and has been lost and no specimens are known to confirm Norfolius Navas (New, 1982, 1983a, 1989a; New and this anomalous record (New, 1981). Eight extant gen- Lambkin, 1989). Immatures typically have a large era containing 33 species are described with an armoured head with six stemmata and broadly sepa- additional 14 extinct genera containing 23 species, rated jaws that often articulate beyond 180° (Mac- from both Mesozoic and Palaeogene deposits. Leod, 1964). The jaw has a large single tooth midway along its length, separating it from most other neurop- *Corresponding author: teran larvae. Antennae are shorter than half the length E-mail address: [email protected] of the mandible with the flagellum being distinctly

© 2015 The Authors Cladistics published by John Wiley & Sons Ltd on behalf of Willi Hennig Society This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made. 2 C. Shi et al. / Cladistics 0 (2015) 1–36 thinner than the scape and pedicel. Labial palpi have in the family in a broad sense, and reflected in the sub- three segments and scoli are present along the lateral familial rankings followed herein (i.e. Nymphinae and edges of the mesothorax, metathorax, and abdomen. Myiodactylinae). New (1984) discussed the intrafamilial relationships Nymphinae adults typically have elongated wings, and biogeography of Nymphidae. Two major types of tibial spurs present, brown to orange body coloration, nymphids have been defined previously, “nymphoid” and male genitalia with gonarcus arms widely sepa- and “myiodactyloid,” which have been elevated to fam- rated medially. This group includes the genera Nesyd- ily-level status by some authors (Handlirsch, 1906– rion Gerstaecker, Austronymphes Esben-Petersen and 1908; Tillyard, 1926). By contrast, some authors consid- Nymphes Leach (Fig. 1a, b). Nymphinae larvae are ered this a simple issue of rank, with the clades treated typically brownish, with a relatively elongated body, as sister groups. Handlirsch (1906–1908), for example, short thoracic and abdominal scoli, and usually carry considered Myiodactylidae to be more distantly related a large trash packet; these larvae inhabit leaf litter and to Myrmeleontiformia, most typically in a clade with soil. In contrast, adults of Myiodactylinae are typified Osmylidae. As reviewed by New (1981), Myiodactylidae by broader, rounded wings, lack of tibial spurs, fre- was proposed by Handlirsch (1906–1908), and followed quent green body coloration and the male gonarcus by various authors (e.g. Esben-Petersen, 1914; Com- ends very close or touching medially. Genera in this stock, 1918; Navas, 1921; Withycombe, 1925; Tillyard, subfamily include the remaining extant genera, Osmy- 1926). Although Handlirsch (1906–1908) did not pro- lops Banks, Myiodactylus Brauer, Norfolius Navas, vide a diagnosis or list of included genera for the fam- Umbranymphes New and Nymphydrion Banks (Fig. 1c, ily, the family name Myiodactylidae was used in the d) (New, 1984, 1987). Myiodactylinae larvae also have discussion of neuropteran familial relationships; no ear- a typically greenish body colour, with a flattened dis- lier references have been found mentioning Myiodac- coid body shape with elongated scoli and little to no tylidae. Myiodactylidae is considered synonymous with trash packet; these larvae are arboreal (New, 1982, Nymphidae here, although the two names reflect the 1983a, 1989a), although New and Lambkin (1989) notable dichotomy of adult and larval body form found described the larva of Norfolius as a litter dweller.

(a) (b)

(c) (d)

Fig. 1. Living Nymphidae. (a) Nymphes nigrescens New, Western Australia (photo: Fred and Jean Hort); (b) Nymphes myrmeleonoides Leach, Queensland (photo: Shaun L. Winterton); (c) Norfolius howensis (Tillyard), Queensland (photo: Shaun L. Winterton); (d) Myiodactylus osmylo- ides Brauer, Queensland (photo: Shaun L. Winterton). C. Shi et al. / Cladistics 0 (2015) 1–36 3

The earliest fossil nymphids are recorded from Jurassic deposits that correspond very well with the (a) proposed origin of the family based on DNA sequence divergence estimates by Winterton et al. (2010). Four genera with six species have been described from local- ities in China, Kazakhstan and Germany (Makarkin et al., 2013b; Shi et al., 2013). Nymphidae were apparently more diverse and widely distributed during the Cretaceous. To date, ten genera with 14 species have been described from the Cretaceous of England, Russia, Myanmar, Brazil and China, including the new genus and species described herein. They were from the Jehol Biota in China, which was a diverse fauna in Early Cretaceous inclusive of abundant and remarkable insects (Ren, 1998; Ren et al., 2010; Gao et al., 2013; Yao et al., 2014). Besides, three species of Nymphidae from the Eocene have been described. One species from the Early Eocene of USA belongs to the extant genus Nymphes, which is the earliest representa- tive of any extant genus (Archibald et al., 2009). The other two species are described from Baltic amber, belonging to the genus Pronymphes Kruger€ (Hagen, (b) 1854, 1856; Kruger,€ 1923; Archibald et al., 2009). A phylogeny of Nymphidae is presented herein based on morphological scoring of all extant species plus a selection of relatively complete fossil exemplars, and combined with DNA sequence data for a subset of extant species. The long held concept of a basal dichotomy of living nymphids is tested, confirming two subfamilies Nymphinae and Myiodactylinae, and the placement of various fossil genera (including new genera described herein) is investigated relative to the total evidence phylogeny.

Materials and methods Fig. 2. Wing venation terminology. Forewings and hind wings of: (a) Norfolius howensis (Tillyard); (b) Austronymphes insularis Esben- Petersen. Major wing veins are highlighted in colour: radial sector Terminology (Rs): green; medial anterior (MA): blue; medial posterior (MP): pink; anterior branch of MP (MP1): yellow; posterior branch of MP Wing venation terminology used here generally (MP2): orange; cubital anterior (CuA): magenta; cubital posterior follows New (1981) except for indication of MA, MP1 (CuP): purple; analis (1A, 2A, 3A): brown. Inset shows detail of thy- ridiate crossveins in Sc space. and MP2 on the hind wing (Fig. 2). The correspond- ing venation is indicated with colour coding in Fig. 2: 1A–3A, analis (brown); CuA, cubital anterior (1997) treated MP1 (yellow) as MA and MP2 (orange) (magenta); CuP, cubital posterior (purple); MA, med- as MP. The obvious sigmoid vein in the hind wings of ial anterior (blue); MP, medial posterior (pink); MP1, many Neuroptera indicates that MA fused with Rs/R anterior branch of MP (yellow); MP2, posterior basally, as identified by various authors in studies on branch of MP (orange); R1, radial anterior; Rs, radial Plega signata (Hagen) (Mantispidae; Ferris, 1940: fig. sector (green); rv, recurrent humeral veinlet; Sc, subco- 15B), Wesmaelius concinnus (Stephens) (Hemerobiidae; sta. Wing venation terminology in Nymphidae has Oswald, 1993b: fig. 51) and Dilar saldubensis Navas been treated in two ways in previous work by New (Dilaridae; Aspock€ et al., 1980: fig. 423; Shi et al., (1981) and Oswald (1997). The main difference is the 2012: fig. 5), and exemplified by the oblique vein in homology statement and delineation of MA in the the forewing (and hind wing) of Norfolius howensis forewing, essentially whether or not MA is assumed as (Tillyard) (Nymphidae; Figs 2b and 3e), Dilar saldub- fused with Rs/R basally. New (1981) identified fore- ensis Navas (Dilaridae; Aspock€ et al., 1980: fig. 423; wing MA as shown in Fig. 2 (blue vein), while Oswald Shi et al., 2012: fig. 5) and Osmylidae (New, 1983b: 4 C. Shi et al. / Cladistics 0 (2015) 1–36

(a) (b)

(c) (d)

(e) (f)

Fig. 3. Nymphidae wings. (a) Austronymphes insularis Esben-Petersen (forewing length (FWL) = 20.5 mm); (b) Nesydrion fuscum Gerstaecker (FWL = 27.4 mm); (c) Myiodactylus osmyloides Brauer (FWL = 21.3 mm); (d) Osmylops sejunctus (Walker) (FWL = 23.4 mm); (e) Norfolius howensis (Tillyard) (FWL = 23.2 mm); (f) Nymphes myrmeleonoides Leach (FWL = 38.8 mm).

figs 1 and 2). They are typically present on forewings and CuP, which suggests MP2 and CuA in the hind and hind wings in many families of Neuroptera, which wing are not fused. provides sufficient evidence to suggest that MA basally Genitalic terminology generally follows New (1981) fused with Rs/R on both forewing and hind wings and and Oswald (1998), as shown in Fig. 21a. The mediun- is the plesiomorphic condition within the order. The cus defined here (green) was inferred as the arcessus hind wing venational terminology used in this study is by New (1981), and 11th gonostyli by Aspock€ and also different from that used by Shi et al. (2012: fig. Aspock€ (2008). The gonarcus (magenta) here is equiva- 7D). Shi et al. (2012) suggested MP2 (orange vein in lent to the 11th gonocoxite of Aspock€ and Aspock€ Fig. 2) as MP2 + CuA, and CuA (magenta) as CuP. (2008). The paramere (dark blue) is equivalent to the Examination of hind wing base of Nymphidae displays 9th gonocoxite of Oswald (1998) and Aspock€ and coalescence of MP1 and MP2, and coalescence of CuA Aspock€ (2008). The gonapsis (light blue) here is C. Shi et al. / Cladistics 0 (2015) 1–36 5 equivalent to the 10th gonostyli of Aspock€ and the analysis). These included Liminympha makarkini Aspock€ (2008). Genitalia were macerated in 10% Ren et Engel (Middle Jurassic, China), Baissoleon simi- KOH to remove soft tissue, then rinsed in distilled lis sp. nov. (Early Cretaceous, China), Baissoleon water and dilute glacial acetic acid, and dissected in cretaceous Makarkin (Early Cretaceous, Russia), 80% ethanol. The genitalia preparation was placed in Nymphites bimaculatus Shi, Makarkin et Ren (Middle glycerine in a genitalia vial mounted on the pin Jurassic, China), Sialium sinicus sp. nov. (Early beneath the specimen. Cretaceous, China), Sialium minor sp. nov. (Early Cre- taceous, China) and Spilonymphes major gen. et sp. Exemplar selection nov. (Early Cretaceous, China). Outgroups were included from lacewing families Ithonidae (Ithone fulva Nymphidae have an extensive fossil record (23 spe- Tillyard) (EU734865, EU839734, EU815247, EU860118) cies in 14 genera) comparable to the number of extant and closely related families also in Myrmeleontiformia, taxa (33 species in eight genera) (Appendix 1). While i.e. Psychopsidae (Psychopsis barnardi Tillyard) all living genera are restricted to the Australasian (EU734897, EU839764, EU815280, EU860149) and Ne- region, extinct representatives of the family have been mopteridae (Nemoptera coa (Linnaeus) (EU734878, found far more widely, being recorded from deposits EU839745, EU815259, EU860131) and Chasmoptera in China, Russia, Kazakhstan, Germany, England, hutti (Westwood) (EU734851, EU839723, EU815232, Myanmar, USA and Brazil (Makarkin et al., 2013b). EU860106)). Outgroups were selected based on the No fossil Nymphidae are recorded from anywhere in most recent phylogenetic analyses of lacewing higher- the extant distribution of the family in Australia level phylogeny by Winterton et al. (2010), and chosen northwards into New Guinea. Nymphidae are rela- as a mix of the two closest related familes of Myrme- tively rare lacewings and are uncommon in collections, leontiformia (Psychopsidae and Nemopteridae) with a although one species of Nymphes (N. myrmeleonoides more distantly related family Ithonidae, a group still Leach) can be locally abundant in dry sclerophyll for- closely related to Myrmeleontiformia. These families ests with sufficiently dense understorey, and is also were preferentially selected not only due to their close known to enter houses frequently. All extant genera phylogenetic relationship to Nymphidae, but also were included in the morphological analysis by at least because they exhibit numerous putative plesiomorphic one species. These included the genera Osmylops (four character conditions in their wing venation and termi- species) (Fig. 3d), Myiodactylus (two species) (Fig. 3c), nalia. Such features are highly modified or highly Norfolius (one species) (Fig. 3e), Nesydrion (three spe- reduced in other potential candidate outgroups such as cies) (Fig. 3b), Austronymphes (one species) (Fig. 3a), Ascalaphidae, Myrmeleontidae and Chrysopidae, Nymphes (three species) (Fig. 3f), Umbranymphes (one thereby obscuring likely homology and weakening any species) and Nymphydrion (one species). Umbranym- homology statements proposed here. phes is known from only the type specimen and was scored from the literature. DNA nucleotide sequences Morphological examination are known for four species of Nymphidae, having been included previously in analyses of Neuropterida phy- Sixty-eight adult and larval characters were numeri- logeny by Winterton et al. (2010). Sequences of two cally coded for the four outgroup and 23 ingroup taxa. ribosomal genes (16S rRNA and 18S rRNA) and two Morphological characters used in the phylogenetic protein-encoding genes [cytochrome oxidase I (COI) analysis are listed in Appendix 2. Thirty-nine charac- and the CPSase region of carbamoyl-phosphate syn- ters were coded as binary and 29 as multistate. Inap- thetase-aspartate transcarbamoylase-dihydroorotase plicable and unknown characters were respectively (CAD)] were retrieved from GenBank for Nymphes coded as “-” and “?”. The data matrix is given in Sup- myrmeleonoides Leach (EU734884, EU839751, EU815268, porting Information Tables S1 and S2. EU860137), Osmylops armatus (McLachlan) (EU734886, EU839754, EU815270, EU860140), Myiodactylus osm- Sequence alignment and phylogenetic analysis yloides Brauer (EU734873, EU839741, EU815255, EU860126) and Norfolius howensis (Tillyard) (EU734882, Sequences were aligned manually, with CAD and EU839749, EU815264, EU860134), and were all COI aligned with reference to translated amino acid included in the combined analysis along with sequences (standard eukaryote and invertebrate mito- sequences for all outgroup taxa. Additional extinct chondrial sequences, respectively) using Mesquite (ver. genera were also included in the analysis and selected 2.75) (Maddison and Maddison, 2011). Ambiguously according to relative completeness of the specimen and aligned regions of 16S, where positional homology the resultant proportion of missing characters in the could not be inferred with a reasonable level of confi- morphological character matrix (i.e. those with more dence, were excluded prior to analysis. Parsimony observable characters were preferentially included in analyses were conducted using WinClada ver. 1.00.08 6 C. Shi et al. / Cladistics 0 (2015) 1–36

(Nixon, 2002) and NONA ver. 2.0 (Goloboff, 1993; generation. The first three million trees were discarded Goloboff et al., 2003, 2008), using a branch and as burn-in. A majority rule consensus tree was com- bound search protocol to find all trees. All characters puted with posterior probabilities (PP) for each node. were treated as unordered and with equal weight. Bootstrap support values were determined using PAUP*4.0b10 (Swofford, 1999) calculated from 1000 Results heuristic search (TBR) pseudoreplicates of re-sampled data sets, each with 30 random additions (constant Phylogeny characters excluded). Character states were mapped on a most parsimonious tree (MPT) using WinClada ver. Parsimony analysis of the morphological matrix 1.0 (Nixon, 2002), showing only unambiguous containing 68 characters across all extant Nymphidae changes. Bayesian analyses were performed using recovered a single MPT [length = 213; consistency MrBayes 3.1.2 (Ronquist and Huelsenbeck, 2003). The index (CI) = 0.52; retention index (RI) = 0.65] (Fig. 4). data were partitioned by data type (DNA sequence, When DNA sequence data for a select few ingroup morphology), by locus, and by the remaining two and all outgroup taxa were combined with the mor- codon positions for each protein-coding locus. A sepa- phological scoring, it returned a single MPT of identi- rate GTR + c nucleotide substitution model was cal topology to the tree based on morphology alone applied to each DNA partition. The mk1 model but with notable increase in the consistency index (Lewis, 2001), with coding set to variable, was applied (length = 1180; CI = 0.73; RI = 0.63). The tree recov- to the morphology partition. Each analysis consisted ered from the Bayesian analysis was overall congruent of four Monte Carlo Markov chains run simulta- in topology, but failed to recover some terminal nodes neously for ten million generations, until stationarity in Nymphidae, particularly in Myiodactylinae. Support was reached. Trees were sampled every 1000th values for bootstrap (BS), Bayesian PP (for congruent

Fig. 4. Phylogeny of living Nymphidae. Single tree recovered from parsimony analyses of morphological characters alone (length = 213; CI = 0.52; RI = 0.65), and from combined analysis with DNA sequence data (length = 1180; CI = 0.73; RI = 0.63). Character state changes are plotted on each node, as well as subtended by statistical support values (Bootstrap: Bayesian posterior probability: Bremer support). Bayesian posterior probability values are included only on congruent nodes, as the Bayes tree lacked some resolution in some terminal nodes. Nymphinae and Myiodactylinae are clearly recovered. Images of representative adults and larvae are included. Image credits: larval drawings: CSIRO; photo- graphs: Nymphes nigriscens (Fred and Jean Hort), Osmylops sp. (Anne Marie McKinnon). C. Shi et al. / Cladistics 0 (2015) 1–36 7 branches) and Bremer support (B) are plotted on indi- nificantly weakened the statistical support for most vidual nodes in Fig. 4. nodes, with resultant loss of resolution across the phy- Nymphidae were recovered as strongly monophyletic logeny. Still, a very similar topology was recovered for (BS: 98; PP: 98; B: 6) and were supported in this analysis the phylogeny when fossil taxa were included (Fig. 5). by multiple apomorphic characters, including a bifid Of note was the placement of the Jurassic genus arolium (character 5: state 1) and wing subcostal thyri- Liminympha as sister to all other nymphids, and a sub- diate crossveins present (15:1–3) in the adult, and larval sequent basal dichotomy of Nymphinae and Myiodac- characters such as stemmata number (62:2), antennal tylinae. Myiodactylinae was recovered as monophyletic flagellum shape (65:1), number of segments of the labial with Spilonymphes in a sister group relationship with palp (66:1), a single large tooth on the jaw (67:1), and Umbranymphes as the sister to the remaining genera in presence of thoracic and abdominal scoli (68:1). One or the clade. All other fossil taxa included in the analysis more of the larval apomorphies defined here are found were placed in Nymphinae, with Baissoleon recovered in some species of Ascalaphidae or Myrmeleontidae (e. as an intermediate group between Nesydrion and more g. abdominal and thoracic scoli), but in combination derived genera. Sialium formed a clade with Nymphites, they are clearly unique to Nymphidae. but was rendered paraphyletic by the latter. Within Nymphidae there was a distinctive basal dichotomy separating the two defined subfamilies, i.e. Taxonomy Nymphinae and Myiodactylinae. Nymphinae com- prises the extant genera Nesydrion, Austronymphes and Nymphes and was recovered with moderate statistical Family Nymphidae Rambur, 1842 support (BS: 73; PP: 84; B: 2) with Nesydrion sister to the clade Austronymphes + Nymphes. Characters sup- Type species: Nymphes myrmeleonoides Leach, 1814 porting the monophyly of this lineage include the pres- Nymphides Rambur 1842; type genus: Nymphes ence of tibial spurs (3:1), narrow costal space (10:2), Leach, 1814. many rows of crossveins between forewing CuA Nymphitidae Handlirsch 1906–1908; type genus: branches (24:0), 1A short (30:1), hind wing MP Nymphites Haase, 1890. branches with multiple rows of linking crossveins Nymphesidae Handlirsch 1906–1908; type genus: (35:1) and, in the male genitalia, lateral lobes on the Nymphes Leach, 1814. mediuncus (43:1) and the gonarcus arms separated Myiodactylidae Handlirsch 1906–1908; type genus: medially (49:1). Austronymphes is a monotypic genus, Myiodactylus Brauer, 1866. while the monophyly of both Nymphes and Nesydrion was relatively well supported. Diagnosis. Medium to large lacewings; antennal Myiodactylinae was recovered with moderate statisti- flagellum elongate, filiform; ocelli absent; prothorax cal support (PP: 70; B: 2) and consists of the extant gen- slender, usually longer than wide; arolium bifid; era Umbranymphes, Osmylops, Norfolius, Nymphydrion trichosors present; nygmata absent; ScA bulge present; and Myiodactylus. Characters supporting the mono- Sc and R1 fused before termination; subcostal area phyly of this lineage include the absence of tibial spurs with thyridiate (incomplete) crossveins present and of (3:0), broad costal space (10:0–1), no crossveins present varying length, true crossveins rarely present (if so, between forewing CuA branches (24:2), 1A long (30:0), then only basally); larva with myrmeleontoid body hind wing MP branches with a few linking crossveins form, with narrow filiform antenna; single tooth on (35:0), absence of mediuncus lateral lobes (43:0) and go- jaw; scoli present; six stemmata; egg produced on narcus dorsal apical touching or very close (49:0). Um- silken stalk with oviruptor present. branymphes is a monotypic genus and was recovered as sister-group of the other genera. Osmylops was recov- Comments. A character apparently unique among ered as paraphyletic. This genus was previously divided Neuroptera, and a putative synapomorphy for the into two species groups: the sejunctus species group and family, is the presence of thyridiate crossveins in the armatus species group (Oswald, 1997). The monophyly wing subcostal cell (Fig. 6; sensu Oswald, 1998). In of the sejunctus species group was well supported, while our examination, they are present in all extant the armatus species group was recovered as paraphylet- nymphids and always originate on Sc regardless of ic. Norfolius and Nymphydrion are monotypic and were length and number. The basal one or two subcostal recovered as sister groups. The monophyly of Myio- crossvein(s) are complete or elongated thyridiate (thus dactylus was relatively well supported. appearing complete). The remaining are thyridiate of While there is strong support for many of the nodes varying length, ranging from nearly complete with on the single tree recovered from the analyses of radial end adjacent to R1 (Fig. 6a), to short barely extant species, inclusion of fossil exemplars and the noticeable remnants arising from Sc (Fig. 6c). The associated increase in missing data in the analysis sig- length of thyridiate crossveins is variable even within 8 C. Shi et al. / Cladistics 0 (2015) 1–36

Fig. 5. Phylogeny of living and extinct Nymphidae. Single tree recovered from parsimony analyses of combined analysis with morphological characters and DNA sequence data. Character state changes are plotted on each node with Bootstrap support values (> 60%). genera. This structure was mentioned by New (1981, barely evident (e.g. Myiodactylinae: Norfolius, 1984) and drawn by Oswald (1997, 1998). The most Nymphydrion; Nymphinae: Nymphes). In Nymphes and common condition for thyridiate crossveins Nesydrion the length and number of thyridiate throughout the family is medium length, although in crossveins can be highly variable among species, from both Nymphinae and Myiodactylinae the sister taxon very few short crossveins to numerous very long to the remaining taxa in each clade has medium crossveins. length. In intermediate and derived genera in each A vein-like structure was observed in the basalmost subfamily there is a progressive shortening of the portion of the costal area on both forewing and hind length of the thyridiate crossveins so that they may be wing of all extant nymphids and Ithone Newman C. Shi et al. / Cladistics 0 (2015) 1–36 9

(a)

(b)

(c)

Fig. 6. Forewing subcostal area, thyridiate crossvein arrangement in Nymphidae. (a) Nymphes myrmeleonoides Leach; (b) Osmylops armatus (McLachlan); (c) Osmylops sejunctus (Walker). Scale bar = 1 mm.

(Ithonidae) (Fig. 7). There are three possible phes Makarkin, Yang, Shi et Ren, 2013, Nymphites terminologies for this structure: ScA, indicating it as a Haase, 1890, Mesonymphes Carpenter, 1929, Sialium branch of the longitudinal vein Sc, as used in Westwood, 1854, Cretonymphes Ponomarenko, 1992, Makarkin et al. (2013b); recurrent humeral veinlet, Baissoleon Makarkin, 1990a, Olindanymphes Martins- indicating it as a crossvein in costal area, e.g. in Psyc- Neto, 2005, Santananymphes Martins-Neto, 2005, hopsidae and Ithonidae (e.g. Winterton and Araripenymphes Menon, Martins-Neto et Martill, 2005, Makarkin, 2010); and ScA bulge (Kukalova-Peck and Spilonymphes gen. nov., Elenchonymphes Engel et Lawrence, 2004), indicating membranous thickenings. Grimaldi, 2008, Dactylomyius Makarkin, 1990b, In this study, we prefer to use the terminology ScA Pronymphes Kruger,€ 1923, Austronymphes Esben- bulge, although the structure actually lacks the bulge- Petersen, 1914, Myiodactylus Brauer, 1866, Nesydrion like, oval shape and broad size as drawn in figs 5 and Gerstaecker, 1885, Norfolius Navas, 1922, Nymphes 6 in Kukalova-Peck and Lawrence (2004). In our Leach, 1814, Nymphydrion Banks, 1913, Osmylops examination, it is as thin as or slightly wider than Banks, 1913, Umbranymphes New, 1987. actual crossvein/longitudinal veins, but fainter, without setae, and sometimes the same colour as the costal Key to genera of extant Nymphidae (modified after crossveins. On the hind wing, it is sometimes weakly New, 1981, 1987) connected with Sc. Both the ScA bulge and the recur- rent humeral veinlet are present in Ithone (Fig. 7), 1 Short tibial spurs present on at least hind tibia; while the recurrent humeral veinlets were not clearly forewing costal margin narrower; body colour usu- indicated in figs 5 and 6 by Kukalova-Peck and Law- ally brown to orange in live specimen ...... rence (2004), and in Daonymphes (Makarkin et al., ...... 2. 2013b: fig. 6A) it is not a recurrent veinlet. Due to the – Tibial spurs absent; forewing typically with broad weak connection between ScA bulge and Sc in the costal margin; body colour frequently green in live hind wing, we prefer not to use ScA for this structure. specimen ...... 4. New (1981, 1984) mentioned Myiodactylidae as being 2 Forewing with veins 2A and 3A forming a large first proposed by Handlirsch (1906–1908), although closed loop (Figs 2b and 3a)...... Handlirsch only mentioned Myiodactylidae twice, once ...... Austronymphes Esben-Petersen. in a discussion of relationships among several families, – Forewing anal veins not forming closed loop- and then in a checklist. Various subsequent authors (e.g...... 3. Esben-Petersen, 1914; Comstock, 1918; Navas, 1921; 3 Hind wing with two rows of cells behind CuA Withycombe, 1925; Tillyard, 1926) have used Myiodac- (Fig. 3f)...... Nymphes Leach. tylidae, and none attributes the concept to any author – Hind wing with a single row of cells behind CuA earlier than Handlirsch (1906–1908). (Fig. 3b)...... Nesydrion Gerstaecker. 4 Forewing costal cells with veins linking costal Included genera (see Appendix 1 for a checklist of crossveins at least in basal half of wing, forming one species). Liminympha Ren et Engel, 2007, Daonym- or more rows of cells (Figs 2a and 3e)...... 10 C. Shi et al. / Cladistics 0 (2015) 1–36

(a) (Fig. 3d)...... Osmylops Banks. – Forewing CuP space broad, typically twice width of intracubital space (except M. osmyloides); forewing MP forked in distal part of wing (level with outer gradate crossveins) appearing simple (Myiodactylus), or forked at base of wing (Nymphydrion); forewing costal space very broad especially at base of wing (Fig. 3c)...... 7.

7 Forewing MP forked at outer gradate series (appear- ing simple)...... Myiodactylus Brauer. – Forewing MP forked at base of wing ...... Nymphydrion Banks.

Subfamily Myiodactylinae Handlirsch, 1906–1908, stat. nov

Myiodactylidae Handlirsch 1906–1908 [as synonym (b) of Nymphidae Rambur: see New 1984] Myiodactylini Navas 1922; type genus: Myiodactylus Brauer, 1866 Nymphydrini Navas 1922; type genus: Nymphydrion Banks, 1913

Type genus. Myiodactylus Brauer, 1866.

Diagnosis. Medium to small nymphids; tibial spurs absent; wing costal area broad (often with multiple forked costal crossveins); no crossveins between forewing CuA branches; 1A long; hind wing MP branches with a few linking crossveins; dorsal ends of gonarcus arms close to touching.

Fig. 7. Wings of Nymphidae and Ithonidae. (a) Osmylops sejunctus (Walker); (b) Ithone fulva Tillyard. Insets show detail of ScA bulge. Included genera. Umbranymphes New, 1987, Spilonymphes gen. nov., Osmylops Banks, 1913, ...... 5. Norfolius Navas, 1922, Nymphydrion Banks, 1913, Myiodactylus Brauer, 1866. – Forewing costal cells in a single row, usually with pectinate forking (e.g. Fig. 3c)...... 6. 5 Forewing subcostal space with a single elongated crossvein basally, vein MP forks in basal half of Spilonymphes major gen. et sp. nov. wing; male genitalia with gonapsis present Figs 8 and 9 (Fig. 3e)...... Norfolius Navas. – Forewing subcostal space with multiple elongated Type species. Spilonymphes major gen. et sp. nov. crossveins along length, vein MP forks in apical half of wing; male genitalia with gonapsis absent...... Etymology. From Greek spilos [rpιkος, m], spot, ...... Umbranymphes New. and Nymphes. Species epithet is from the Latin, 6 Forewing CuP space narrow, typically less than 1.5 magnus, -a, -um (comparative major), great (greater). times width of intracubital space; forewing MP forked at least in basal one-third to basal half of Diagnosis. Forewing basal rs-m present, proximal to wing; forewing costal space typically narrow the oblique vein; MA separated from Rs close to the C. Shi et al. / Cladistics 0 (2015) 1–36 11

(a) (b)

Fig. 8. (a) Photograph and (b) line drawing of Spilonymphes major gen. et sp. nov. Holotype CNU-NEU-LB2014001. origin of Rs; CuP space broad, nearly three times as terminating. 1A long and pectinate forked. 2A and wide as intracubital space; hind wing CuA space 3A forked. Hind wing shorter than forewing with broad. similar venation pattern, except MP forked from very near wing base, MP2 pectinately branched around Description. Forewing preserved length 22.5 mm, midway along wing length with more branches, up to preserved width 6.6 mm. Hind wing preserved length ten. CuA parallel to posterior margin, with broad 21.1 mm, preserved width 6.2 mm. Prothorax longer space between them. CuA pectinately branched with than wide. Pterothorax well developed. Mesothoracic seven branches terminating on posterior margin. CuP prescutum divided by median notal suture. Wings long and pectinately branched. CuA and CuP elongated, with apex pointed. Irregular wing branches continuously forked before terminating. markings present medially, at pterostigma and in distal part. Forewing costal area dilated distal to the Type material. Holotype CNU-NEU-LB2014001. pterostigma. Costal crossveins seldom forked and more numerous towards apex. Sc and R1 Locality and horizon. Huangbanjigou Village, approximating along length, then fused at Chaomidian County, Beipiao City, Liaoning Province, approximately distal one-quarter of wing length. Rs China. Yixian Formation, Early Cretaceous. separated from R1 near wing base. Seven crossveins present between R1 and Rs. MA separated from Rs Comments. The new genus has a forewing costal stem around the basal one-quarter of wing length. Rs space slightly broader than typical Nymphinae genera, with 17 branches. Outer gradate crossveins and and is more similar to recent myiodactyline genera. proximal irregular crossveins present among Rs Our analyses place this genus in Myiodactylinae as branches. MP forked before the separation of MA sister to Umbranymphes. A broad forewing costal from Rs. Basal rs-m present, proximal to the oblique space is not common in fossil nymphids and is only vein. MP1 forked until near wing margin, with few found in four monotypic genera (although being branches. MP2 straight and strong, pectinately somewhat continuous in character it may be difficult branched from the midway of its length, with to delineate): Spilonymphes gen. nov., Daonymphes numerous branches terminating on outer margin. Makarkin, Yang, Shi et Ren, Dactylomyius Makarkin, CuA parallel to posterior margin with broad space Elenchonymphes Engel and Grimaldi. The new between them. CuA pectinately branched with four genus has a forewing MP forked proximal to the or five branches midway along wing length. CuP separation of MA, which is found in all fossil space broad, about three times as broad as nymphids, and considered as a plesiomorphic intracubital space. CuP pectinately branched with character in Nymphidae. Forewing MP is forked distal about six branches terminating on posterior margin. to the separation of MA in the recent myiodactyline CuA and CuP branches continuously forked before genera. 12 C. Shi et al. / Cladistics 0 (2015) 1–36

(a)

(b)

(c)

(d)

Fig. 9. Wing venation of Spilonymphes major gen. et sp. nov. Holotype CNU-NEU-LB2014001. (a) Left forewing; (b) right forewing; (c) left hind wing; (d) right hind wing. C. Shi et al. / Cladistics 0 (2015) 1–36 13

Spilonymphes gen. nov. differs from Elenchonymphes and B. similis sp. nov. from the Yixian Formation and Dactylomyius by the denser spaced Rs branches, (China). and MP2 forked in the basal half of wing length on both forewing and hind wing. Elenchonymphes has Remarks. Baissoleon shares characters with typical fewer Rs branches and MP2 forked in distal half of Nymphinae genera, e.g. tibial spurs, elongated wings, wing length in both wings. The hind wing of Dactyl- and narrow costal area. Yet it differs from these omyius has fewer Rs branches, and MP2 forked near genera by the narrow costal distal area, Sc + R1 wing margin. Moreover, the hind wing of Dactylomyi- entering margin before wing apex and a narrow CuA us is distinctly dilated in distal half, while the hind space. Baissoleon is similar to Nesydrion based on wing of Spilonymphes gen. nov. is elongated and MP forked proximal to the separation of MA. smoothly dilated in the distal half. Baissoleon also resembles Nymphes and The new genus resembles Daonymphes most closely, Austronymphes based on a distinct pterostigma in but can be easily distinguished from the latter by fore- both wings. wing MP2 forked in basal half, CuP space nearly three Compared with fossil nymphids, Baissoleon and Olin- times wide as intracubital space, instead of forewing danymphes (from the late Aptian Crato Formation in MP2 forked in the distal half, and the CuP space Brazil) are closely related genera. Besides general simi- about twice as wide as intracubital space in Daonym- larity of their venation, they share one significant con- phes. dition in contrast to all other genera, i.e. forewing CuP is non-pectinate. Although in all known Jurassic Subfamily Nymphinae Rambur, 1842 nymphids CuP is strongly pectinate in the forewing, a non-pectinate condition found in these Cretaceous gen- Nymphini Navas 1922; type genus: Nymphes Leach, era is probably a plesiomorphic condition. A similar 1814 configuration of CuA and CuP is characteristic of the Late Jurassic genus Chrysoleonites Martynov, which is Type genus. Nymphes Leach, 1814 thought to be closely related to ancestors of the myr- meleontoid + chrysopoid clades (Yang et al., 2012). Diagnosis. Large to medium nymphids; tibial spurs present at least on hind legs; costal area narrow; Baissoleon similis sp. nov. crossveins present between branches of CuA; 1A short; Figs 10 and 11 hind wing MP branches with multiple rows of linking crossveins; male genitalia with arms of gonarcus far Etymology. From Latin similis,-e, similar, in separated medially, mediuncus with large lateral lobes. reference to the close resemblance between this species and B. cretaceous. Included genera. Nesydrion Gerstaecker, 1885, Baissoleon Makarkin, 1990a, Sialium Westwood, 1854, Diagnosis. Very similar to B. cretaceous, but easily Nymphites Haase, 1890, Austronymphes Esben- distinguished by the longer CuP and 1A (CuP Petersen, 1914, Nymphes Leach, 1814. terminating distal to the fork of MP, and 1A terminating nearly at level of fork of MP in B. similis Baissoleon Makarkin, 1990 sp. nov.; CuP terminating nearly at level of fork of MP, and 1A terminating much more proximally to the Baissoleon Makarkin, 1990a: 125 (Nymphitidae) fork of MP in B. cretaceous). Makarkin et al., 2012: 57, 65 (Nymphidae) Description. Body length (from apex head to Type species. Baissoleon cretaceous Makarkin, abdomen) 15.2 mm. Head elongated. Mandible 1990a, by original designation. protruding with apex pointed. Compound eye medium- sized. Antenna appear short (probably incomplete). Revised diagnosis. Prothorax longer than wide; tibial Prothorax elongated, slightly narrower than head. spurs present at least on hind legs; wings slender; Mesonotum and metanotum distinctly broader than pterostigma distinct; Sc + R1 entering margin before prothorax. Legs slender, with femora and tibiae quite wing apex; costal area not dilated after the fusion of long. Hind tibia with apical spurs. Abdomen slender, Sc and R1; single row of crossveins present between with nine tergites, apical segments dilated. First tergite forewing CuA branches; forewing CuP dichotomously poorly preserved, short. Second to seventh tergites forked. rather long; eighth to ninth tergites short, transverse. Forewing length 18.2 mm, width 5.2 mm. Costal Species included. Two Early Cretaceous species, space narrow. Costal crossveins simple before B. cretaceous from Baissa in Transbaikalia (Russia) pterostigma, very shallowly forked distally. Pterostig- 14 C. Shi et al. / Cladistics 0 (2015) 1–36

(a) proximal to this series between Rs branches (except between three distal-most). MP forked at midway between the origin of Rs and the separation of MA. One oblique vein between Rs stem and M. MP1 slightly arched, pectinately branched distally. MP2 strongly zigzagged; pectinately branched with three branches terminating on outer margin; all branches simple except proximal-most branch in left wing which is rather deeply forked. CuA long, zigzagged, running rather close to posterior margin; pectinately branched with five simple branches; four proximal branches connected by one row of crossveins. CuP short, deeply forked. Four (left wing) to five (right wing) intracubital crossveins. Basal CuP and 1A clo- (b) sely approach or touch. Between CuP and 1A one distal crossvein detected (connecting CuP and ante- rior branch of 1A fork). 1A long, deeply forked. 2A short, probably simple. 3A short, shallowly forked. Between 1A and 2A, 2A and 3A one crossvein pres- ent respectively. Hind wing length 15.3 mm, width ca. 4.5 mm (when restored); smaller than forewings. Costal space some- what narrower than in forewing. All costal crossveins simple before pterostigma, some forked after pterostig- ma. Pterostigma distinct. MA separated from Rs stem far from wing base, before midway of wing length. Rs (c) with seven branches, all but one forked (except dichot- omous distal-most branches). Outer gradate series of crossveins complete; several crossveins irregularly arranged proximal to this series between Rs branches (except between three distal-most). MP forked at wing base. MP1 long, almost straight, pectinately branched very distally, with two to three simple short branches. MP2 pectinately branched with four to five branches, simple to deeply forked once. Three proximal MP2 branches connected by crossveins. CuA short, pecti- nately branched, with three branches. CuP short, prob- ably simple. 1A and 2A short, simple.

Fig. 10. (a, b) Photograph and (c) line drawing of Baissoleon similis Type material. Holotype CNU-NEU-NN2014001PC, sp. nov. Holotype CNU-NEU-NN2014001PC. a nearly complete well-preserved specimen, with hind wings strongly crumpled. ma well developed. Subcostal space rather narrow, slightly dilated distally. Sc and R1 fused below Type locality and horizon. Liutiaogou Village, pterostigma; Sc + R1 entering margin before wing Ningcheng County, Chifeng City, Inner Mongolia apex. Rs separated at acute angle from R1 near Autonomous Region, China. Yixian Formation, Early wing base, slightly zigzagged in middle part. R1 Cretaceous. space narrower basally and distally, dilated most at middle part; 12 crossveins sparsely distributed before Sialium Westwood, 1854 fusion of Sc and R1; hypostigmal cell long. Rs with seven branches; MA separated from Rs stem very Sialium Westwood, 1854: 390, 396 (Neuroptera, far from its origin, nearly at midway of wing length. allied to Sialis) Most branches of Rs rather shallowly forked, except Geinitz, 1884: 571 (?Orthoptera) distal branches having additionally minute forks Scudder, 1886: 472 (Blattoptera) (dichotomous). Outer gradate series of crossveins Geinitz, 1887: 200 (Blattoptera) complete; several crossveins irregularly arranged Scudder, 1891: 130 (Blattoptera) C. Shi et al. / Cladistics 0 (2015) 1–36 15

(a)

(b)

(c)

(d)

Fig. 11. Wing venation of Baissoleon similis sp. nov. Holotype CNU-NEU-NN2014001PC. (a) Left forewing; (b) right forewing; (c) left hind wing; (d) right hind wing. 16 C. Shi et al. / Cladistics 0 (2015) 1–36

Handlirsch, 1906–1908: 609 (Nymphitidae) Description. Head large, wider than long. Handlirsch, 1939: 159 (Nymphitidae) Compound eye medium sized. Antenna filiform. Martynova, 1949: 167 (Nymphitidae) Prothorax almost as large as head, but longer than Lambkin, 1988: 451, 454 (Nymphidae) wide. Pterothorax larger than prothorax. Legs slender. Whalley, 1988: 46 (Nymphitidae) Hind tibia with a pair of apical spurs. Jarzembowski, 1993: 178 (Neuroptera) Forewing preserved length 41 mm, preserved width Jepson et al., 2012: 44 (Nymphidae) 11.8 mm. Hind wing preserved length 37.5 mm, pre- served width 10.5 mm. Wings subequal, elongated. Posterior and outer mar- Type species. Sialium sipylus Westwood, 1854. gins waved. Forewings bearing discontinuous markings in subcostal area, around pterostigma area, in the basal Revised diagnosis. Tibial spurs present at least on posterior part and apical area. Hind wing bearing dis- hind legs; wings proximally narrow; forewing humeral continuous markings in subcostal area and in distal veinlet recurrent; intracubital space dilated before part, one large spot near distal one-third of wing length. CuA forked; two or more rows of crossveins present Forewing costal crossveins seldom forked before between CuA branches; CuP space broad, nearly pterostigma area, and progressively denser with more twice as wide as basally intracubital space; hind wing forks from pterostigma area to termination. Humeral MP pectinately branched with more than eight veinlet recurrent. Sc and R1 closely parallel. Rs sepa- branches; CuP long, pectinately branched with rich rated from R1 at basal one-tenth of wing length. MA branches. separated from Rs stem at basal one-fifth of wing length. MP forked between the origin of Rs and the Species included. Sialium sipylus Westwood, 1854; separation of MA. One oblique vein present closely Sialium sinicus sp. nov.; Sialium minor sp. nov. before the fork of MP. MP1 straight and unbranched. MP2 pectinately branched with numerous branches Sialium sinicus sp. nov. terminating on outer margin. One series of crossveins Figs 12 and 13. present among them. MP2 branches continuously forked before termination. Basal mp-cua oblique. CuA Etymology. Latinized name of China. and CuP separated below the oblique crossvein. CuA pectinately branched with four branches terminating on Diagnosis. Wing markings present in subcostal area the midway of wing margin. Crossveins present between and whole apical area in both wings, markings them. CuA branches continuously forked before termi- sparsely distributed on posterior part of forewing, one nation. The space between CuA and CuP dilated before large spot present near distal one-third of wing length CuA forked. CuP parallel with posterior margin, with on hind wing. space between them broad. CuP pectinately branched

(a) (b)

Fig. 12. (a) Photograph and (b) line drawing of Sialium sinicus sp. nov. Holotype CNU-NEU-NN2014002. C. Shi et al. / Cladistics 0 (2015) 1–36 17

(a)

(b)

(c)

(d)

Fig. 13. Wing venation of Sialium sinicus sp. nov. Holotype CNU-NEU-NN2014002. (a) Left forewing; (b) right forewing; (c) left hind wing; (d) right hind wing. 18 C. Shi et al. / Cladistics 0 (2015) 1–36 with numerous branches continuously forked. 1A elon- Type material. Holotype CNU-NEU-NN2014002. gated and forked. 2A forked. 3A short. Hind wing costal crossveins forked more than in Locality and horizon. Liutiaogou Village, Ningcheng forewing. Rs separated from R1 before basal one-tenth County, Chifeng City, Inner Mongolia Autonomous of wing length. MA separated from Rs stem before Region, China. Yixian Formation, Early Cretaceous. basal one-quarter of wing length. MP forked before the origin of Rs. MP1 straight, shallowly forked near Sialium minor sp. nov. termination. MP2 long, pectinately branched, with 11 Figs 14–17. closely spaced branches forked before termination. CuA parallel to posterior margin with a broad space Etymology. From Latin parvus, -a, -um (comparative between the two. CuA pectinately branched with eight minor), small (smaller). branches continuously forked before terminating on posterior margin. One series of crossveins discontinu- Diagnosis. Wing markings present midway between ously interlinking CuA branches. CuP long, pectinately Sc and R1, MP1 and MP2, along outer gradate crossvein branched with six branches, without crossveins series area on forewing, and the area surrounding the between them. 1A and 2A short and forked. fusion of Sc and R1 on both forewing and hind wing.

(a)

(b)

Fig. 14. (a) Photograph and (b) line drawing of Sialium minor sp. nov. Holotype CNU-NEU-LB2014002. C. Shi et al. / Cladistics 0 (2015) 1–36 19

(a)

(b)

(c)

Fig. 15. Wing venation of Sialium minor sp. nov. Holotype CNU-NEU-LB2014002. (a) Left forewing; (b) left hind wing; (c) right hind wing.

Description. Holotype CNU-NEU-LB2014002. Fore- Rs stem near basal one-third of wing length. MA wing preserved length 14.3 mm, preserved width dichotomously forked proximally, almost at the same 4.1 mm. Hind wing preserved length 13.6 mm, level as the origin of the first Rs branch. The first Rs preserved width 3.8 mm. Wings elongated. Forewing branch separated from Rs stem distally, almost midway bearing small markings in the basal posterior part, along wing length. Rs with 12 branches. MP forked midway between Sc and R1, MP1 and MP2, along outer between the origin of Rs and the separation of MA. gradate crossvein series area on forewing, and the area One oblique vein present between Rs stem and M stem. surrounding the fusion of Sc and R1. Hind wing bearing MP1 forked closely before termination. MP2 pectinate- small markings dispersed in the distal part. ly branched with at least seven branches. CuA long, Forewing costal crossveins seldom forked before with few branches. CuP pectinately branched with pterostigma area, and progressively more numerous numerous branches. 1A long and forked. from pterostigma to the termination of Sc + R1. Rs Hind wing venation similar to that on forewing, but separated from R1 near wing base. MA separated from MP forked before the origin of Rs. MP2 pectinately 20 C. Shi et al. / Cladistics 0 (2015) 1–36

(a) (b)

Fig. 16. (a) Photograph and (b) line drawing of Sialium minor sp. nov. Paratype CNU-NEU-LB2014003. branched with 10–11 branches, forming a triangular from Rs remote from wing base. MP1 forked near area midway along wing. CuA pectinately branched termination. MP2 pectinately branched with about with fewer branches than MP2. CuA branches termi- eight branches. CuA long, pectinately branched. nating on posterior margin. CuP pectinately branched with few branches. Locality and horizon. Huangbanjigou Village, Chaomidian County, Beipiao City, Liaoning Province, Locality and horizon. Huangbanjigou Village, Cha- China. Yixian Formation, Early Cretaceous. omidian County, Beipiao City, Liaoning Province, China. Yixian Formation, Early Cretaceous. “Nymphites” sp. B Figs 18 and 19 Paratype CNU-NEU-LB2014003. Forewing preserved length 15.6 mm, preserved width 4.1 mm. Hind wing Diagnosis. Legs (at least hind) with tibial spurs; preserved length 13.1 mm, preserved width 4.1 mm. forewing costal crossveins seldom forked; hind wing Wings elongated. Wing markings present midway MP2 with four branches, CuA with at least five between Sc and R1, MP1 and MP2, along outer branches. gradate crossvein series area on forewing, and the area surrounding the fusion of Sc and R1 on both forewing Description. Forewing preserved length 23.7 mm, and hind wing. preserved width 9 mm. Hind wing preserved length Forewing costal crossveins seldom forked, but pro- 33.1 mm, preserved width 9.5 mm. gressively denser with more forks from pterostigma Compound eye medium-sized; antenna filiform; pro- area. Sc and R1 approximating basally, fused distally. thorax as wide as head; pterothorax dilated; legs slen- MA separated from Rs stem near basal one-third of der; tibiae with a pair of apical spurs. wing length. The first Rs branch separated from the Forewing costal crossveins seldom forked before Rs stem distally, almost midway along wing length. pterostigma area. CuA pectinately branched. MP forked before the basal one-fifth of wing length. Hind wing outer margin waved. Costal crossveins One oblique vein present between Rs stem and M stem only forked from pterostigma area. Sc and R1 closely right after the fork of MP. MP1 forked near termina- parallel. MA separated from the basal part of Rs stem, tion. MP2 pectinately branched with nine branches ter- but remote from wing base. Rs with seven branches. minating on outer margin. MP1 forked closely before termination. MP2 pecti- Hind wing costal crossveins not forked except from nately branched with four branches. CuA pectinately pterostigma to termination of Sc + R1. MA separated branched with at least five branches. C. Shi et al. / Cladistics 0 (2015) 1–36 21

(a)

(b)

(c)

(d)

Fig. 17. Wing venation of Sialium minor sp. nov. Paratype CNU-NEU-LB2014003. (a) Left forewing; (b) right forewing; (c) left hind wing; (d) right hind wing. 22 C. Shi et al. / Cladistics 0 (2015) 1–36

(a) (b)

Fig. 18. (a) Photograph and (b) line drawing of “Nymphites” sp. B. specimen CNU-NEU-NN2014003.

Material. CNU-NEU-NN2014003 versally considered to comprise the extant families Psychopsidae, Nymphidae, Myrmeleontidae, Ascala- Locality and horizon. Liutiaogou Village, phidae and Nemopteridae (e.g. MacLeod, 1964; Grim- Ningcheng County, Chifeng City, Inner Mongolia aldi and Engel, 2005; Winterton et al., 2010; Aspock€ Autonomous Region, China. Yixian Formation, et al., 2012), along with at least the extinct families Early Cretaceous. Palaeoleontidae, Araripeneuridae and Babinskaiidae (Yang et al., 2012; Makarkin et al., 2013b). The phy- Comments. This specimen is tentatively assigned to logenetic relationships among the various neuropteran the genus Nymphites based on the four branches of families are not universally accepted within Myrme- hind wing MP2 (referred to as MP by Shi et al., 2013) leontiformia (Aspock€ et al., 2001, 2012; Aspock,€ 2002; and the presence of tibial spurs. Due to the incomplete Winterton et al., 2010), with varying hypotheses of preservation of the specimen, there are not enough relationships among the families, differing mainly in significant observable characters to assign it to any the relative positions of Nemopteridae and Psychopsi- published species, or to formally name it as dae. Based solely on morphological characters of the representing a new species. wing venation, genitalia and larval mandible, some Species of Nymphites have been described from the authors place Nymphidae as the sister group of Myr- Middle Jurassic of China and the Late Jurassic of Ger- meleontidae + Ascalaphidae (Handlirsch, 1906–1908; many. This specimen could be the only representative Withycombe, 1925; Aspock€ et al., 2001, 2012; Aspock,€ of Nymphites from the Early Cretaceous, which indi- 2002; Beutel et al., 2010a, b). Yet the phylogenetic cates the genus might have survived for over 40 Myr analysis by Winterton et al. (2010) recovered Nymphi- in the Late Mesozoic. Nymphites is the earliest repre- dae as the sister group of Myrmeleontidae, Ascalaphi- sentative in Nymphinae, bearing a narrow costal area dae and Nemopteridae, and appears to be the best- and tibial spurs. Besides the unnamed Nymphites spe- supported hypothesis for placement for the family, cies described here, tibial spurs were also found on the based on both DNA sequence data and morphology Middle Jurassic species Nymphites bimaculatus Shi, of extant and extinct forms (e.g. New, 1984; Oswald, Makarkin et Ren, 2013, and an unnamed species 1998; Makarkin et al., 2013b). “Nymphites” sp. A (Shi et al., 2013). Phylogenetic relationships within Nymphidae

Discussion Nymphidae have been treated by some early authors as two separated families, Myiodactylidae (= Myiodac- Phylogenetic position of Nymphidae tylinae herein) and Nymphidae sensu stricto (s.s.) (= Nymphinae herein), with the former sometimes Myrmeleontiformia is a well-established and placed near Osmylidae (Handlirsch, 1906–1908; Withy- accepted clade within Neuroptera and is almost uni- combe, 1925; Tillyard, 1926; cf. Esben-Petersen, 1914; C. Shi et al. / Cladistics 0 (2015) 1–36 23

(a)

(b)

(c)

Fig. 19. Wing venation of “Nymphites” sp. B. specimen CNU-NEU-NN2014003. (a) Left forewing; (b) left hind wing; (c) right hind wing.

Adams, 1958) (note: there is striking superficial similar- phini (cf. Banks, 1913), Myiodactylini and Nymphyd- ity between green myiodactyline nymphids and green rini, but these have not been adopted by recent authors spilosmyline Osmylidae). Moreover, multiple plesio- (New, 1981, 1984). The validity of this distinction as morphic features in the wing venation of both families two families is questionable. Wing dimensions in the make differentiating incompletely preserved Mesozoic family are highly variable with numerous intermediate fossil taxa problematic, and their placement in particu- forms, especially in Myiodactylinae. Based on the ratio lar families is rarely a simple task (Lambkin, 1988; Ren data of forewing length to width measured in this and Engel, 2007). Myiodactylidae were supposed as the analysis alone (character 9), there is overlap between more “primitive” group, containing, for example, the Myiodactylinae and Nymphinae lineages; Osmylops in genera Myiodactylus and Osmylops. They were sepa- particular displays a wide range of morphometric rated from Nymphidae s.s. (i.e. Nymphes, Austronym- dimensions in wing shape. Presence or absence of tibial phes and Nesydrion) based on broad wing shape, broad spurs and costal space width are highly variable char- costal space and absence of tibial spurs. Nymphidae acters and are not sufficient to separate higher taxon were divided into three tribes by Navas (1922): Nym- groups reliably. For example, tibial spurs are present 24 C. Shi et al. / Cladistics 0 (2015) 1–36 or absent in closely related genera of Myrmeleontidae on hind legs, wing with narrow costal area and crossveins (New, 1985). Although the family displays a distinct present between branches of CuA, male genitalia with dichotomy morphologically, Nymphidae (inclusive of arms of gonarcus far separated medially and mediuncus Myiodactylidae) are strongly supported as a monophy- having large lateral lobes (Fig. 20a–d). All extant Nym- lum based on characters such as: bifid arolium, pres- phinae genera are now restricted to Australia although it ence of ScA bulge, presence of subcostal thyridiate is clear that the group was more widely distributed during crossveins, larva antennal flagellum distinctly thinner the Mesozoic than at present, with fossil genera known than pedicel, presence of a large single tooth midway from the Palaearctic and Nearctic (Ren and Engel, 2007; along the mandible, and presence of scoli on meso- and Archibald et al., 2009; Makarkin et al., 2013b). metathorax and abdomen. Other characters that easily Nesydrion was recovered as the sister genus to the diagnose adult Nymphidae include absence of ocelli on remaining Nymphinae genera and is characterized by the vertex and presence of wing trichosors. An ovirup- inconspicuous pterostigmata in both the forewing and tor is present in the egg of species in both Nymphinae the hind wing, infuscate spots present surrounding the and Myiodactylinae, a plesiomorphic character lost in distal crossvein between Sc + R1 and Rs on forewing more derived families of Myrmeleontiformia (New, or hind wing, forewing MP forked proximal to the 1989a). Thyridiate crossveins in the subcostal area are separation of MA, paramere with multiple small teeth considered here as a putative synapomorphy for the (Fig. 21d) and a paired gonapsis (Fig. 20n). Nesydrion family. is placed as the sister group of the remaining Nymphi- The earliest representatives of Nymphidae in the fos- nae genera and while the body is very similar to nym- sil record are from the Middle Jurassic of China (Ma- phine genera, the wing is very reminiscent of that karkin et al., 2013b; Shi et al., 2013), with the family found in myiodactyline genera (Fig. 3b). All other hypothesized to have originated during the Early extant Nymphinae genera have forewing MP forked Jurassic based on divergence time estimates of DNA distal to the separation of MA and, along with the sequence data (Winterton et al., 2010). Liminympha extinct genus Baissoleon, conspicuous pterostigmata in makarkini is the only Jurassic nymphid preserved with both wings as well (Figs 3a, f and 10). antennae, with antennae longer than half wing length The Cretaceous genus Baissoleon was originally (1:1) (Ren and Engel, 2007), an apomorphy also found described in Nymphitidae by Makarkin (1990a) and in Nymphes aperta; by comparison the antennae of includes two species with the addition of the new spe- most other extant nymphids are shorter. Liminympha cies described herein. The genus is distinct from other was recovered as the sister to the remaining Nymphi- Nymphinae genera as the costal space is not dilated dae and retains plesiomorphic characters such as a after the fusion of Sc and R1, forewing CuP is dichoto- recurrent humeral veinlet, a feature subsequently lost mously branched and one incomplete row of crossveins in most Nymphidae (Makarkin et al., 2013b). This is present between forewing CuA branches (Figs 10 and genus is also unique among Nymphidae in having no 11). Baissoleon is an intermediate genus in Nymphinae crossveins between the distal Rs branches (20:1). and, like Nesydrion, has a dichotomously branched The remaining Nymphidae (exclusive of Liminym- CuP in the hind wing (38:1). The remaining derived pha) form a characteristic dichotomy between Nym- Nymphinae have common characters such as multiple phinae and Myiodactylinae. Incongruously, while the rows of crossveins present between hind wing CuA family clearly originated during the Early Jurassic branches and hind wing CuP pectinately forked (38:0). based on both molecular and fossil data, Winterton The Cretaceous genus Sialium forms the next intermedi- et al. (2010) estimated the divergence of the subfami- ate clade with the Jurassic Nymphites, although with lies to be during the Early Palaeocene (with a range the latter rendering Sialium paraphyletic. Nymphites extending as far back as the late Early Cretaceous). can be reliably separated from Sialium based on the Based on the tree topology recovered here, with defini- presence of only a single row of cells between the tive Nymphinae genera (Baissoleon, Sialium and branches of CuA (24:1) and multiple forked crossveins Nymphites) placed as intermediate groups within the in the costal area (13:2); based on this we maintain the clade, Nymphinae and Myiodactylinae probably genera as separate at this time despite the paraphyly in diverged during the Middle Jurassic or Early Creta- the analysis. The extant sister genera Austronymphes ceous at the latest. and Nymphes both share the presence of numerous branches of forewing CuA (23:1). Austronymphes is Phylogenetic relationships within Nymphinae monotypic and is distinguishable based on forewing 2A and 3A forming a distinctive closed loop (Figs 2b and Nymphinae includes the extant genera Nesydrion, 3a). Nymphes is characterized by rose-coloured to Austronymphes and Nymphes, and Jurassic and Creta- brown markings in the apical area of both wings, and ceous fossil genera Baissoleon, Sialium and Nymphites. two rows of cells between forewing CuP and posterior They share characters such as tibial spurs present at least margin (Fig. 3f). C. Shi et al. / Cladistics 0 (2015) 1–36 25

(a) (b)

(c) (d) (e)

(f) (g) (h)

(i) (j) (k)

(l) (m) (n)

Fig. 20. Male genitalic structures, posteroventral view. (a–k) Mediuncus: (a) Nymphes myrmeleonoides Leach; (b) Austronymphes insularis Esben- Petersen; (c) Nesydrion fuscum Gerstaecker; (d) Nesydrion diaphanum Gerstaecker; (e) Myiodactylus osmyloides Brauer; (f) Norfolius howensis (Tillyard); (g) Osmylops ectoarticulatus Oswald; (h) Osmylops placidus (Gerstaecker); (i) Chasmoptera hutti (Westwood); (j) Psychopsis barnardi Tillyard; (k) Ithone fulva Tillyard. (l–n) Gonapsis: (l) Nymphes myrmeleonoides Leach; (m) Norfolius howensis (Tillyard); (n) Nesydrion fuscum Gerstaecker. Scale bar = 0.2 mm.

Most fossil nymphids have a similar elongated wing in Myiodactylinae are unique by paralleling the poster- shape to extant Nymphinae, so are easily placed in ior margin. In Nymphinae and a few Cretaceous that lineage. Costal space width varies widely in the nymphids (e.g. Cretonymphes, Sialium), there are mul- Middle Jurassic, being broad (e.g. Daonymphes)or tiple rows of crossveins between MP2 and CuA narrow (e.g. Liminympha). All fossil nymphids have branches, instead of a single row of crossveins found forewing MP dichotomously forked proximal to the in Jurassic and most Cretaceous fossil Nymphidae. separation of MA, which is the same as the extant Ne- The multiple branches of forewing CuA and many sydrion. In contrast, other extant nymphids have fore- rows of crossveins between them in Nymphes and wing MP forked distal to the separation of MA. Some Austronymphes are characteristics of Myrmeleontidae, of them have forewing MP forked nearly before termi- Ascalaphidae and Nemopteridae, too. nation, resulting in an inconspicuous MP1 and MP2, for example Austronymphes (Fig. 2b), Myiodactylus, Phylogenetic relationships within Myiodactylinae Umbranymphes and some species of Nymphes.In Jurassic nymphids, along with some genera from the Myiodactylinae includes the extant genera Umbra- Early Cretaceous, forewing veins MP2 and CuA have nymphes, Osmylops, Myiodactylus, Norfolius and Nym- a similar number of branches, like most Myiodactyli- phydrion, and the Cretaceous genus Spilonymphes. The nae and Nesydrion. The forewing veins MP2 and CuA extant Myiodactylinae are geographically more widely 26 C. Shi et al. / Cladistics 0 (2015) 1–36

(a) (b)

(c) (d)

(e) (f)

Fig. 21. Male genitalia, lateral view. (a) Nymphes myrmeleonoides Leach; (b) Nymphes aperta New; (c) Austronymphes insularis Esben-Petersen; (d) Nesydrion fuscum Gerstaecker; (e) Norfolius howensis (Tillyard); (f) Myiodactylus osmyloides Brauer. Scale bar = 1 mm. distributed than their sister clade, being found All three extant Nymphinae genera are highly dis- throughout New Guinea and Australia, including Nor- tinctive and easy to separate morphologically, but folk Island (New, 1981, 1987; Oswald, 1998). Like Myiodactylinae genera are very similar morphologi- Nymphinae, this group had a far more expansive his- cally with rather variable characters used to define tory, with Spilonymphes described here from Early genera previously, including width of the costal space Cretaceous deposits in China. All Myiodactylinae gen- base (Myiodactylus and Osmylops) and presence of sec- era share characters such as: absence of tibial spurs, ondary crossveins in the costal area of the forewing wing costal area broad (often with multiple forked forming multiple rows of cells (i.e. Umbranymphes, costal crossveins), no crossveins between forewing Norfolius). A probable artefact from the use of subjec- CuA branches; dorsal ends of gonarcus arms approxi- tive characters to define genera is the prevalence of mating each other (sometimes connected by a mem- monotypic genera based on distinctive autapomor- brane); mediuncus small, ball-like or trilobate ventrally phies. Winterton and Brooks (2002) found a similar with varying sized teeth and no obvious lateral lobes. scenario in the chrysopid subfamily Apochrysinae, C. Shi et al. / Cladistics 0 (2015) 1–36 27

(a) (b)

(c) (d)

(e)

Fig. 22. Male genitalia, lateral view (continued). (a) Osmylops sejunctus (Walker); (b) Osmylops placidus (Gerstaecker); (c) Chasmoptera hutti (Westwood); (d) Psychopsis barnardi Tillyard; (e) Ithone fulva Tillyard. Scale bar = 1 mm. with numerous monotypic genera based on highly var- features, such as parameres highly modified, presence iable characters rendering more generalized genera as of a gonapsis and paired membrane structures beneath paraphyletic; a phylogenetic analysis reduced the num- the mediuncus (Figs 20f, m and 21e). Nymphydrion is ber of genera from 13 to six. Three monotypic genera known only from a single, damaged female specimen, are found in this group, Umbranymphes, Nymphydrion and besides a distinctive venational peculiarity of the and Norfolius. Umbranymphes is endemic to New Gui- greatly elongated CuA, the genus is very similar to nea with a habitus very similar to Myiodactylus, but Myiodactylus and Norfolius. Nymphydrion is very simi- differentiated only by the presence of the secondary lar to Myiodactylus and is possibly synonymous, pend- crossveins in the costal area. It is recovered here with ing discovery of the male. Characteristics used by New weak support as the sister to Spilonymphes in a clade (1981) to define the genus (width of CuP space, i.e. sister to the remaining Myiodactylinae genera. Norfo- distance of CuP from posterior margin) have been lius (Figs 1c, 2a and 3e) is widely distributed and is shown to be not stable and the multiple species of My- defined based on both wings and male genitalic iodactylus described from northern Australia and New 28 C. Shi et al. / Cladistics 0 (2015) 1–36

Guinea since his redescription of the genus all exhibit the taxonomic part and nomenclature; Frederica Tur- a similar condition (New, 1987). Nymphydrion and co (Queensland Museum) and Beth Mantle (Australian Norfolius are weakly supported here as sister taxa in a National Insect Collection) for help with loans of clade sister to Myiodactylus. Osmylops is the most material and for providing images of types; and Jean species-rich genus of Nymphidae, consisting of 11 and Fred Hort, Anne-Marie McKinnon and CSIRO species (Oswald, 1998). Oswald (1997, 1998) estab- for use of their photographs of living nymphids and lished two species groups in this genus: the sejunctus larval drawings. Fossil specimens of the new species species group and armatus species group. The former are deposited in the Key Lab of Insect Evolution and is distinguished based on enlarged ectoproct with mul- Environmental Changes, College of Life Sciences, Cap- ticuspate processes, presence of entoprocessi and a pair ital Normal University, Beijing, China (CNU, Ren of elongated membranous processes beneath the para- Dong, Curator). This research was supported by the meres, and the mediuncus ventrally ball-like with teeth National Basic Research Program of China (973 Pro- (Figs 20g and 22a) (also found in Myiodactylus osmy- gram) (2012CB821906), the National Natural Science loides). The Osmylops armatus species group is less dis- Foundation of China (No. 41271063, 31230065, tinguishable than the sejunctus species group and is 41272006, 40972113), the Foundation for the Author not recovered as monophyletic in this analysis. Both of National Excellent Doctoral Dissertation of PR species included in this analysis from the O. armatus China (No. 201178), Great Wall Scholar and KEY species group have a distinctly small mediuncus with project of Beijing Municipal Commission of Education trilobate ventral processes (47:2) (Fig. 20h), but this Project (grants KZ201310028033), Starting Foundation character is also found homoplasiously in Norfolius. for Young Teachers of Sun Yat-sen University (No. The paraphyly of Osmylops, as well as the limited 32110-31101402), Program for Changjiang Scholars morphological synapomorphies supporting the mono- and Innovative Research Team in University phyly of most genera within Myiodactylinae, indicates (IRT13081) and by the United States National Science that this group of genera should be re-examined in Foundation (DEB 1144119). The Willi Hennig Society detail (with greater taxon sampling) and the generic facilitated the use of TNT and Winclada. The authors status of each assessed in a phylogenetic context. have no conflict of interest to declare.

References Conclusions Adams, P.A., 1958. Studies in the Neuroptera, with special reference The monophyly of Nymphidae, inclusive of Myiodac- to wing structure and evolution in the Osmyloidea. PhD tylidae, is confirmed here and supported by a suite of dissertation. Harvard University, Cambridge, MA, USA. Archibald, S.B., Makarkin, V.N., Ansorge, J., 2009. New fossil adult and larval morphological characters. Reflecting species of Nymphidae (Neuroptera) from the Eocene of North the historical dichotomy of nymphids into two distinct America and Europe. Zootaxa 2157, 59–68. lineages or types, the family is divided basally into two Aspock,€ U., 2002. Phylogeny of the Neuropterida (Insecta: – subfamilies, Nymphinae and Myiodactylinae. Inclusion Holometabola). Zool. Scr. 31, 51 55. Aspock,€ U., Aspock,€ H., 2008. Phylogenetic relevance of the genital of the numerous new genera and species of fossil Nym- sclerites of Neuropterida (Insecta: Holometabola). Syst. Entomol. phidae blurs this distinction somewhat, and while some 33, 97–127. Jurassic and Cretaceous genera can be placed in either Aspock,€ H., Aspock,€ U., Holzel,€ H., 1980. Die Neuropteren Europas. Vol. 2. Goecke and Evers, Krefeld, West lineage, others (e.g. Liminympha) represent more gener- Germany. alized stem taxa sister to the remaining Nymphidae. Aspock,€ U., Plant, J.D., Nemeschkal, H.L., 2001. Cladistic analysis Moreover, the spatial and temporal history of Nymphi- of Neuroptera and their systematic position within the dae is one now highlighted by extinction, obscuring Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Syst. Entomol. 26, 73–86. much of the evidence of vicariance and/or dispersal. The Aspock,€ U., Haring, E., Aspock,€ H., 2012. The phylogeny of the present-day fauna is restricted to the Australasian Neuropterida: long lasting and current controversies and region, but the presence of numerous extinct forms in challenges (Insecta: Endopterygota). Arthropod. Syst. Phylogeny 70, 119–129. deposits around the world of varying antiquity reflects a Banks, N., 1913. Synopses and descriptions of exotic Neuroptera. T. geographically broad, morphologically diverse and spe- Am. Entomol. Soc. 39, 201–242. cies-rich history of the family. Beutel, R.G., Friedrich, F., Aspock,€ U., 2010a. The larval head of Nevrorthidae and the phylogeny of Neuroptera (Insecta). Zool. J. Linn. Soc. 158, 533–562. Beutel, R.G., Zimmermann, D., Krauss, M., Randolf, S., Wipfler, Acknowledgements B., 2010b. Head morphology of Osmylus fulvicephalus (Osmylidae, Neuroptera) and its phylogenetic implications. Org. – Thanks are extended to Vladimir N. Makarkin (Far Divers. Evol. 10, 311 329. Brauer, F., 1866. Zusatze€ und Berichtigungen zu Hagen’s Eastern Branch of the Russian Academy of Sciences) Hemerobidarum Synopsis Synonymica und Beschreibung einer for helping preparing the draft manuscript, especially neuen Nymphiden-gattung: Myiodactylus osmyloides aus C. Shi et al. / Cladistics 0 (2015) 1–36 29

Australien. Verhandlungen der Kaiserlich-Koniglichen€ Kukalova-Peck, J., Lawrence, J.F., 2004. Relationships among Zoologisch-Botanischen Gesellschaft in Wien 16, 983–992. coleopteran suborders and major endoneopteran lineages: Carpenter, F.M., 1929. A Jurassic neuropteran from the lithographic evidence from hind wing characters. Eur. J. Entomol. 101, 95– limestone of Bavaria. Psyche 36, 190–194. 144. Comstock, J.H., 1918. The Wings of Insects. Comstock Publishing Lambkin, K.J., 1988. A re-examination of Lithosmylidia Riek from Co., Ithaca, NY. the Triassic of Queensland with notes on Mesozoic “osmylid- Engel, M.S., Grimaldi, D.A., 2008. Diverse Neuropterida in like” fossil Neuroptera (Insecta: Neuroptera). Mem. Queensl. Cretaceous amber, with particular reference to the paleofauna of Mus. 25, 445–458. Myanmar (Insecta). Nova Supplementa Entomologica 20, 1–86. Leach, W.E., 1814. Zoological Miscellany; Being Descriptions of Esben-Petersen, P., 1914. Australian Neuroptera. Part I. P. Linn. New, or Interesting Animals. Vol. 1. B. McMillan, London. Soc. N. S. W. 39, 635–645. Lewis, P.O., 2001. A likelihood approach to estimating phylogeny Ferris, G.F., 1940. The morphology of Plega signata (Hagen) from discrete morphological character data. Syst. Biol. 50, 913– (Neuroptera: Mantispidae). Microentomology 5, 33–56. 925. Gao, T., Shih, C.K., Rasnitsyn, A.P., Xu, X., Wang, S., Ren, D., MacLeod, E.G., 1964 Comparative morphological studies on the 2013. New transitional fleas from China highlighting diversity head capsule and cervix of larval Neuroptera (Insecta). PhD of Early Cretaceous ectoparasitic insects. Curr. Biol. 23, 1261– Thesis, Harvard University, Cambridge. 1266. Maddison, W.P., Maddison, D.R., 2011. Mesquite: A Modular Geinitz, F.E., 1884. Ueber die Fauna des Dobbertiner Lias. ZDGG System for Evolutionary Analysis. Version 2.75. http:// 36, 566–583. mesquiteproject.org. Geinitz, F.E., 1887. Beitrag zur Geologie Mecklenburgs. Archiv des Makarkin, V.N., 1990a. Baissoleon cretaceus gen. and sp. nov. Fossil Vereins der Freunde der Naturgeschichte in Mecklenburg 41, Neuroptera from the Lower Cretaceous of Baisa, East Siberia. 2. 143–216. Nymphitidae. Annales de la Societe Entomologique de France Gerstaecker, C.E.A., 1884 (1885). Zwei fernere Decaden (N.S.) 26, 125–126. Australischer Neuroptera Megaloptera. Mitt[h]eilungen aus dem Makarkin, V.N., 1990b. Novye setchatokrylye (Neuroptera) iz Naturwissenschaftlichen Verein fur€ Neu-Vorpommern und Rugen verkhnego mela Azii [New lacewings (Neuroptera) from the 16, 84–116. Upper Cretaceous of Asia]. In: Akimov, I.A. (Ed.), Novosti Goloboff, P.A., 1993. Estimating character weights during tree faunistiki i sistematiki. Sbornik nauchnyhk trudov [News of search. Cladistics 9, 83–91. faunistics and systematics. Collected scientific papers], Naukova Goloboff, P.A., Farris, J.S., Nixon, K.C., 2003. TNT (Tree Analysis Dumka Publ., Kiev, pp. 63–68. using New Technology) (BETA) Win. Menu Interface – Standard Makarkin, V.N., Yang, Q., Peng, Y.Y., Ren, D., 2012. A Version. Published by the authors, Tucuman, Argentina. comparative overview of the neuropteran assemblage of the Goloboff, P.A., Farris, J.S., Nixon, K.C., 2008. TNT, a free Lower Cretaceous Yixian Formation (China), with description of program for phylogenetic analysis. Cladistics 24, 774–786. a new genus of Psychopsidae (Insecta: Neuroptera). Cretaceous Grimaldi, D.A., Engel, M.S., 2005. Evolution of the Insects. Res. 35, 57–68. Cambridge University Press, Cambridge, UK. Makarkin, V.N., Yang, Q., Ren, D., 2013a. A new Cretaceous Haase, E., 1890. Bemerkungen zur Palaeontologie der Insecten. family of enigmatic two-winged lacewings (Neuroptera). Foss. Neues Jahrbuch fur€ Mineralogie, Geologie und Palaeontologie, Rec. 16, 67–75. Stuttgart 1890, 1–33. Makarkin, V.N., Yang, Q., Shi, C., Ren, D., 2013b. The presence of Hagen, H.A., 1854. Ueber die Neuropteren der Bernsteinfauna. the recurrent veinlet in the Middle Jurassic Nymphidae Verhandlungen des Zoologisch-Botanischen Vereins in Wien 4, (Neuroptera): a unique character condition in Myrmeleontoidea. 221–232. ZooKeys 325, 1–20. Hagen, H.A., 1856. Die im Bernstein befindlichen Neuropteren der Martins-Neto, R.G., 2005. New Neuroptera from Crato Formation, vorwelt bearbeitet von F. J. Pictet-Baraban [sic; for Pictet-del la Lower Cretaceous, Araripe Basin, northeast Brazil. GAEA 1, 5– Rive], und Dr. H. Hagen. In: Berendt, G.C. (Ed.), Die im 10. Bernstein befindlichen organischen reste der Vorwelt gesammelt, Martynova, O.M., 1949. Mesozoic lacewings (Neuroptera) and their in verbindung mit mehreren bearbeitet und herausgegeben von bearing on concepts of phylogeny and systematics of the order. Dr. Georg Carl Berendt. Bd. 2. Nicholaischen Buchhandlung, Trudy Paleon tologicheskogo Instituta 20, 150–170. Berlin, pp. 41–125. Menon, F., Martins-Neto, R.G., Martill, D.M., 2005. A new Lower Handlirsch, A., 1906–1908. Die fossilen Insekten und die Phylogenie Cretaceous nymphid (Insecta, Neuroptera, Nymphidae) from the der rezenten Formen. Ein Handbuch fur€ Palaontologen€ und Crato Formation of Brazil. GAEA 1, 11–15. Zoologen. W. Engelmann, Leipzig. Navas, L., 1921. Sur des Nevropteres nouveaux ou critiques. Handlirsch, A., 1939. Neue Untersuchungen uber€ die fossilen Quatrieme [IV] serie. Annales de la Societe Scientifique de Insekten mit Erganzungen€ und Nachtragen€ sowie Ausblicken auf Bruxelles, 41 (pt. 1), 130–137. phylogenetische, palaeogeographische und allgemein biologische Navas, L., 1922. Insectos nuevos o poco conocidos [I]. Memorias Probleme. II. Teil. zAnnalen des Naturhistorischen Museums in de la Real Academia de Ciencias y Artes de Barcelona, 17, Wien 49, 1–240. 383–400. Jarzembowski, E.A., 1993. A provisional checklist of fossil insects New, T.R., 1981. A revision of the Australian Nymphidae (Insecta: from the Purbeck Beds of Dorset. In: Proceedings of the Dorset Neuroptera). Aust. J. Zool. 29, 707–750. Natural History and Archaeological Society 114 (for 1992), 175– New, T.R., 1982. The larva of Nymphes Leach (Neuroptera, 179. Nymphidae). Neurop. Int. 2, 79–84. Jepson, J.E., Makarkin, V.N., Coram, R.A., 2012. Lacewings (Insecta: New, T.R., 1983a. Some early stages of Osmylops (Neuroptera: Neuroptera) from the Lower Cretaceous Purbeck Limestone Nymphidae). Syst. Entomol. 8, 121–126. Group of southern England. Cretaceous Res. 34, 31–47. New, T.R., 1983b. A revision of the Australian Osmylidae: Koch, L.E., 1967. The genus Chasmoptera (Neuroptera: Kempyninae (Insecta: Neuroptera). Aust. J. Zool. 31, 393–420. Nemopteridae), with the description of a new species from New, T.R., 1984. Intergeneric relationships in recent Nymphidae. In: Western Australia. Proc. Roy. Entomol. Soc. B 36, 137–146. Gepp, J.; Aspock,€ H.; Holzel,€ H. (Eds.), Progress in World’s Kruger,€ L., 1923. Neuroptera succinica baltica. Die im baltischen Neuropterology. Proceedings of the 1st International Symposium Bernstein eingeschlossenen Neuroptera des Westpreussischen on Neuropterology (22–26 September 1980, Graz, Austria). Graz, Provinzial-Museums (heute Museum fur€ Naturkunde und Austria, pp. 125–131. Vorgeschichte) in Danzig. Stettiner Entomologische Zeitung 84, New, T.R., 1985. A second species of Nymphidae (Neuroptera) from 68–92. Papua New Guinea. Neurop. Int. 3, 187–189. 30 C. Shi et al. / Cladistics 0 (2015) 1–36

New, T.R., 1987. Nymphidae (Insecta: Neuroptera) from New Stange, L.A., 1994. Reclassification of the New World antlion Guinea. Invertebr. Taxon. 1, 807–815. genera formerly included in the tribe Brachynemurini New, T.R., 1989a. An oviruptor in Nymphes Leach (Neuroptera: (Neuroptera: Myrmeleontidae). Insecta Mundi 8, 67–119. Nymphidae). Neurop. Int. 5, 239–240. Swofford, D.L., 1999. PAUP*. Phylogenetic Analysis Using New, T.R., 1989b. Planipennia, Lacewings. Handbuch der Zoologie, Parsimony (*and Other Methods), version 4.0b4a. Sinauer Vol. 4 (Arthropoda: Insecta), Part 30. Walter de Gruyter, Berlin. Associates, Sunderland, MA. New, T.R., Lambkin, K.J., 1989. The larva of Norfolius Tillyard, R.J., 1926. The Insects of Australia and New Zealand, 1st (Neuroptera: Nymphidae). Syst. Entomol. 14, 93–98. edn. Angus and Robertson, Sydney. Nixon, K.C., 2002. WinClada ver. 1.00.08. Published by the author, Tjeder, B., 1967. Neuroptera-Planipennia. The Lace-wings of Ithaca, NY. Southern Africa. 6. Family Nemopteridae. In: Hanstrom,€ B.; Oswald, J.D., 1993a. Phylogeny, taxonomy, and biogeography of Brinck, P.; Rudebec, G. (Eds.), South African Animal Life. Vol. extant silky lacewings (Insecta: Neuroptera: Psychopsidae). Mem. 13. Swedish Natural Science Research Council, Stockholm, pp. Am. Entomol. Soc. 40, iii+1–65. 290–501. Oswald, J.D., 1993b. Revision and cladistic analysis of the world Westwood, J.O., 1854. Contributions to fossil entomology. Quart. J. genera of the family Hemerobiidae (Insecta: Neuroptera). J. New Geol. Soc. London 10, 378–396. York Entomol. S. 101, 143–299. Whalley, P.E.S., 1988. Mesozoic Neuroptera and Raphidioptera Oswald, J.D., 1997. Review of the sejunctus species group of the (Insecta) in Britain. Bull. Br. Mus. Nat. Hist. (Geol.) 44, 45–63. split-footed Lacewing genus Osmylops Banks (Neuroptera: Winterton, S.L., Brooks, S.J., 2002. Phylogeny of the Apochrysine Nymphidae), with remarks on the functional morphology of green lacewings (Neuroptera: Chrysopidae: Apochrysinae). Ann. terminalic coupling. Aust. J. Entomol. 36, 351–358. Entomol. Soc. Am. 95, 16–28. Oswald, J.D., 1998. Osmylops Banks (Neuroptera: Nymphidae): Winterton, S.L., Makarkin, V.N., 2010. Phylogeny of moth generic review and revision of the armatus species group. J. lacewings and giant lacewings (Neuroptera: Ithonidae, Neuropterol. 1, 79–108. Polystoechotidae) using DNA sequence data, morphology and Ponomarenko, A.G., 1992. Setchatokrylye (Insecta, Neuroptera) iz fossils. Ann. Entomol. Soc. Am. 103, 511–522. nizhnego mela Zabaykal’ya [=Neuroptera (Insecta) from the Winterton, S.L., Hardy, N.B., Wiegmann, B.M., 2010. On wings of Lower Cretaceous of Transbaikalia]. Paleontol. Zh. 1992, 43–50. lace: phylogeny and Bayesian divergence time estimates of Rambur, J.P., 1842. Histoire Naturelle des Insectes, Nevropteres. Neuropterida (Insecta) based on morphological and molecular Librairie encyclopedique de Roret. Fain et Thunot, Paris. data. Syst. Entomol. 35, 349–378. Ren, D., 1998. Flower-associated Brachycera flies as fossil evidence Withycombe, C.L., 1925. Some aspects of the biology and for Jurassic angiosperm origins. Science 280, 85–88. morphology of the Neuroptera with special reference to the Ren, D., Engel, M.S., 2007. A split-footed lacewing and two immature stages and their possible phylogenetic significance. T. epiosmylines from the Jurassic of China (Neuroptera). Ann. Am. Entomol. Soc. 1924, 303–411. Zool. 57, 211–219. Yang, Q., Makarkin, V.N., Winterton, S.L., Khramov, A.V., Ren, Ren, D., Shih, C.K., Gao, T.P., Yao, Y.Z., Zhao, Y.Y., 2010. Silent D., 2012. A remarkable new family of Jurassic Insects Stories – Insect Fossil Treasures from Dinosaur Era of the (Neuroptera) with primitive wing venation and its phylogenetic Northeastern China. Science Press, Beijing. position in Neuropterida. PLoS One 7, e44762. Ronquist, F., Huelsenbeck, J.P., 2003. MrBayes 3: Bayesian Yao, Y.Z., Cai, W.Z., Xu, X., Shih, C.K., Engel, M.S., Zheng, X.T., phylogenetic inference under mixed models. Bioinformatics 19, Xhao, Y.Y., Ren, D., 2014. Blood-feeding true bugs in the Early 1572–1574. Cretaceous. Curr. Biol. 24, 1786–1792. Scudder, S.H., 1886. A review of mesozoic cockroaches. Mem. Boston Soc. Nat. Hist. 3, 439–485. Scudder, S.H., 1891. Index to the known fossil insects of the world Supporting Information including myriapods and arachnids. Bull. United States Geol. Survey 71, 1–744. Shi, C.F., Bethoux, O., Shih, C.K., Ren, D., 2012. Guyiling jianboni Additional Supporting Information may be found in gen. et sp.n., an antlion-like lacewing, illuminating homologies the online version of this article: and transformations in Neuroptera wing venation. Syst. Table S1. Morphological data matrix. – Entomol. 37, 617 631. Table S2. Molecular data matrix. Shi, C.F., Makarkin, V.N., Yang, Q., Archibald, S.B., Ren, D., 2013. New species of Nymphites Haase (Neuroptera: Nymphidae) from the Middle Jurassic of China, with a redescription of the type species of the genus. Zootaxa 3700, 393–410. C. Shi et al. / Cladistics 0 (2015) 1–36 31

Appendix 1. Checklist of extinct and extant Nymphidae Appendix 1. continued †Araripenymphes Menon, Martins-Neto et Martill, 2005 Araripenymphes seldeni Menon, Martins-Neto Osmylops monofoveatus Oswald, 1998 et Martill, 2005 Osmylops nesos Oswald, 1998 Austronymphes Esben-Petersen, 1914 Osmylops placidus (Gerstaecker, 1885) Austronymphes insularis Esben-Petersen, 1914 Osmylops sejunctus (Walker, 1853) †Pronymphes € †Baissoleon Makarkin, 1990a Kruger, 1923 Baissoleon cretaceous Makarkin, 1990a Pronymphes mengeana (Hagen in Pictet-Baraban et Hagen, Baissoleon similis sp. nov. 1856) †Cretonymphes Ponomarenko, 1992 Pronymphes hoffeinsorum Archibald, Makarkin et Ansorge, Cretonymphes baisensis Ponomarenko, 1992 2009 †Santananymphes †Dactylomyius Makarkin, 1990b Martins-Neto, 2005 Dactylomyius septentrionalis Makarkin, 1990b Santananymphes ponomarenkoi Martins-Neto, 2005 †Sialium †Daonymphes Makarkin, Yang, Shi et Ren, 2013 Westwood, 1854 Daonymphes bisulca Makarkin, Yang, Shi et Ren, 2013 Sialium minor sp. nov. †Elenchonymphes Engel et Grimaldi, 2008 Sialium sinicus sp. nov. Elenchonymphes electrica Engel et Grimaldi, 2008 Sialium sipylus Westwood, 1854 †Spilonymphes †Liminympha Ren et Engel, 2007 gen. nov. Liminympha makarkini Ren et Engel, 2007 Spilonymphes major gen. et sp. nov. Umbranymphes †Mesonymphes Carpenter, 1929 New, 1987 Mesonymphes apicalis Ponomarenko, 1992 Umbranymphes spinosus New, 1987 Mesonymphes hageni Carpenter, 1929 Mesonymphes rohdendorfi Panfilov in Dolin et al., 1980 Mesonymphes sibirica Ponomarenko, 1992 Myiodactylus Brauer, 1866 Appendix 2 Morphological character descriptions and Myiodactylus lyriformis New, 1987 character argumentation Myiodactylus maculatus New, 1986 Myiodactylus nebulosus McLachlan, 1877 Myiodactylus osmyloides Brauer, 1866 ADULT BODY (1–5) Myiodactylus paranebulosus New, 1987 Myiodactylus pubescens Banks, 1910 1. Antennae length Myiodactylus roseistigma Esben-Petersen, 1917 Myiodactylus sorongensis New, 1987 0. Shorter than half of forewing length; 1. Longer than half of Myiodactylus striatus New, 1982 forewing length. Nesydrion Gerstaecker, 1885 Comments. Antennal length varies considerably in Neuroptera, Nesydrion diaphanum Gerstaecker, 1885 from shorter than half of forewing length (e.g. Psychopsis, Psychopsi- Nesydrion fuscum Gerstaecker, 1885 dae), to longer than twice forewing length (e.g. Oviedus, Chrysopi- Nesydrion nigrinerve Esben-Petersen, 1914 dae). In Nymphidae, the apomorphic state is found in the earliest Nesydrion pallidum Banks, 1913 fossil nymphids (e.g. Liminympha), the sister group to the remaining Norfolius Navas, 1922 Nymphidae, as well the highly derived extant species Nymphes aperta. Norfolius howensis (Tillyard, 1917) All other nymphids examined in this analysis have short antennae. Nymphes Leach, 1814 Nymphes aperta New, 1982 2. Prothorax Nymphes modesta Gerstaecker, 1885 Nymphes myrmeleonoides Leach, 1814 0. Wider than long; 1. Longer than wide. Nymphes nigrescens New, 1982 Comments. The prothorax is usually short in Myrmeleontiformia Nymphes paramyrmeleonides New, 1982 (e.g. Psychopsidae, Nemopteridae and Ascalaphidae), but in Nym- †Nymphes georgei Archibald, Makarkin phidae the prothorax is slightly elongated, which is considered to be et Ansorge, 2009 an apomorphy in Neuroptera. An elongated prothorax is also found †Nymphites Haase, 1890 in Hemerobiiformia families such as some Mantispidae, Rhachibero- Nymphites priscus (Weyenbergh, 1869) thidae and Berothidae. In Mantispinae the prothorax has an elon- Nymphites bimaculatus Shi, Makarkin et Ren, 2013 gated tubular shape. Nymphydrion Banks, 1913 Nymphydrion delicatum Banks, 1913 3. Tibial spurs †Olindanymphes Martins-Neto, 2005 Olindanymphes makarkini Martins-Neto, 2005 0. Absent; 1. Present. Osmylops Banks, 1913 Comments. The presence of tibial spurs corresponds to the dichot- Osmylops armatus (McLachlan, 1867) omy of Myiodactylinae and Nymphinae; two groups that were ear- Osmylops bispinosus Oswald, 1998 lier considered as separate families (e.g. Handlirsch, 1906–1908; Osmylops clavatus Oswald, 1998 Withycombe, 1925; Tillyard, 1926). While tibial spurs are present in Osmylops crux Oswald, 1998 many lacewing families, they are absent in many others, so the Osmylops ectoarticulatus Oswald, 1997 polarity of such a character is globally plesiomorphic, but locally Osmylops halberdis Oswald, 1998 apomorphic, depending on outgroup selection. Here we treated the Osmylops hirsutus New, 1987 presence of tibial spurs as apomorphic, but find it is a plesiomorphic feature in Neuropterida. 32 C. Shi et al. / Cladistics 0 (2015) 1–36

4. First tarsal segment 8. Hind wing pterostigma colour 0. Elongated; 1. Short. 0. Inconspicuous; 1. Conspicuous. Comments. In Myrmeleontiformia, Psychopsidae and Nemopteri- Comments. See character 7. dae have legs with first tarsal segment much longer than the other tarsal segments, while all nymphids have legs with short, subequal 9. Forewing shape, ratio of forewing length to width tarsal segments. (gap: 2.4–2.6) 5. Arolium 0. Elongated, > 2.6; 1. Broad, < 2.4. Comments. Ratios of wing length to width, and costal width to 0. Single; 1. Split; 2. Absent. intraradial width were used to avoid the scaling effect from size dif- Comments. The presence of an arolium is considered to be plesio- ferences in different taxa. For each character, data of all taxa were morphic in the Myrmeleontiformia (Stange, 1994). The arolium is compared, and the most significant gap(s) were found to differentiate present in Ithonidae, a member of the sister group of the Myrme- character states. leontiformia, and the basal groups of Myrmeleontiformia, Psychopsi- dae and Nymphidae (Winterton et al., 2010). The arolium in Nymphidae is bifid, which is the synapomorphic character of the 10. Forewing costal space, ratio of costal width to family. intraradial width at MA separation level (gaps: 1.41–1.53, 2.5–2.8) 0. Broad, > 2.8; 1. Medium, 1.5–2.5; 2. Narrow, < 1.5. WINGS (6–38) Comments. To avoid the effect of varying width of costal space, the data were measured at the level of MA separation. The width of 6. Pattern of wing markings the costal space varies widely in Nymphidae. Significant differences were found between Nymphinae and Myiodactylinae. In Myiodac- 0. No markings: Ithone fulva, Chasmoptera hutti, Osmylops tylinae, the costal space of Myiodactylus and Nymphydrion is signifi- placidus, Osmylops armatus, Osmylops ectoarticulatus, Osmylops se- cantly broader than the other genera. junctus, Myiodactylus osmyloides, Myiodactylus pubescens, Baisso- leon similis, Austronymphes insularis; 1. Complex pattern: Psychopsis barnardi, Nemoptera coa; 2. Whole apical area: 11. Forewing ScA bulge Spilonymphes major, Sialium minor, Sialium sinicus, Nymphes mode- 0. Absent; 1. Present. sta, Nymphes aperta, Nymphes myrmeleonoides [Nymphes except Comments. ScA bulge is a vein-like thickening of the membrane for Nymphes nigrescens (see New, 1981)]; 3. Stripe along outer at the most basal costal area. Different from the real crossvein, it is gradate crossveins: Spilonymphes major, Sialium minor, Nymphites as thin as or slightly wider than the real crossvein/longitudinal vein, bimaculatus; 4. Apical spot: Nesydrion fuscum (both wings), Nesyd- but fainter and without setae. In Daonymphes, ScA bulge (referred rion nigrinerve (hind wing), Nesydrion diaphanum (forewing), to as ScA in Makarkin et al., 2013b) is present at the same position, Nymphydrion delicatum; 5. Red mark on posterior area: Spilonym- before the recurrent humeral veinlet. phes major, Norfolius howensis, Nymphydrion delicatum, Sialium minor. 12. Forewing humeral veinlet Comments. Compared with other lacewings, the wing markings in Nymphidae are relatively simple. There are three types of patterns in 0. Recurrent; 1. Simple. the recent nymphids: (i) coloured in the apical area of wings in Nym- Comments. The humeral veinlet is recurrent in Psychopsidae and phes, except for Nymphes nigrescens (New, 1981), which was used in Ithonidae, as well as Jurassic and Cretaceous nymphids (Liminym- key to species of Nymphes in New (1981); (ii) dark spot beneath the pha, Daonymphes, Sialium), whereas it is simple in extant nymphids. fusion of Sc and R1 in Nesydrion, but the spots on different wings in The humeral veinlet is simple in the more derived groups of Myrme- different species, which was used in key to species of Nesydrion in leontiformia (Myrmeleontidae, Ascalaphidae, Nemopteridae). The New (1981); (iii) red marks on the posterior area of hind wing in presence or absence of a humeral veinlet was used previously in phy- Norfolius. More complex wing markings typically associated with logenetic analyses of Neuroptera relationships by Winterton et al. crypsis are found only in fossil nymphids and never in recent taxa. (2010) and Yang et al. (2012). In Nymphites and Spilonymphes, a stripe along the outer gradate crossveins is present on the forewing, along with irregular markings. 13. Branching of forewing costal crossveins before The mark at the forewing apical area is also present in Sialium and – Spilonymphes. pterostigma area (gaps: 0, 9 13) 0. Simple; 1. Few forked, 1–10; 2. Many forked, > 12. 7. Forewing pterostigma colour Comments. The forewing costal crossveins are forked to varying extent among different genera of Nymphidae. Generally, taxa in My- 0. Inconspicuous; 1. Conspicuous. iodactylinae have more forked crossveins in the costal area, some- Comments. The pterostigma is considered plesiomorphic in Neu- times continuously forked, and with linking veins between them. roptera, but with conspicuous reduction in various families. Ptero- Myiodactylus, Norfolius, Nymphydrion and Nymphites have multiple stigmata are conspicuous in Ascalaphidae and Mantispidae, but forks in the forewing costal area. Genera in Nymphinae have fewer faint in Chrysopidae, Osmylidae and Myrmeleontidae (New, forked crossveins in the forewing costal area, and none with linking 1989b). Pterostigmata vary in different genera of Nymphidae, veins between them. Austronymphes, Baissoleon and Nymphes aperta sometimes even between forewing and hind wing in the same spe- have no forks in the forewing costal area. The number of forked cies (e.g. Myiodactylus osmyloides). Osmylops and Nesydrion have forewing costal crossveins varies among different species of Nym- inconspicuous pterostigmata. By contrast, Norfolius (red coloured), phes, Nesydrion and Osmylops, as well as Liminympha and Sialium. Nymphydrion, Nymphes (brown coloured), Austronymphes (rose col- Costal crossvein branching was used in the key to families of Neu- oured), and the Cretaceous Baissoleon have well-developed ptero- roptera in New (1989b), and in the phylogenetic analysis of Neurop- stigma. tera in Yang et al. (2012) (char. 21: subcostal veinlets). C. Shi et al. / Cladistics 0 (2015) 1–36 33

14. Forewing costal gradate series 20. Forewing, crossveins between distal Rs branches 0. Absent; 1. Present. 0. Present; 1. Absent. Comments. Forewing costal gradate series are not common in Comments. In Nymphidae, forewing crossveins between Rs Neuroptera, and are known in some extant Hemerobiidae, Ithoni- branches are irregularly arranged, but usually have a well-defined dae, Psychopsidae, Myrmeleontidae and Nymphidae and all of the outer gradate series. The outer gradate series is close to the outer Jurassic family Grammolingiidae. Forewing costal gradate series are margin in Nymphinae, while it is further from the wing margin in considered to be an apomorphy in Neuroptera. In Nymphidae, the Myiodactylinae. In the Jurassic Liminympha, crossveins between Myiodactylinae genera Umbranymphes and Norfolius have costal gra- distal Rs branches are absent, which is the apomorphy in Nym- date series in the forewing. Presence of a costal gradate series was phidae. used in the phylogenetic analysis of Hemerobiidae and Psychopsidae by Oswald (1993a,b). 21. Forewing MP branching 0. Dichotomously forked proximal to the separation of MA; 1. 15. Forewing subcostal thyridiate crossveins Dichotomously forked in the basal half of forewing, distal to the 0. Absent; 1. Short, thyridiate crossvein shorter than one-third of separation of MA; 2. Pectinately forked in the distal half of fore- subcostal width; 2. Medium, thyridiate crossvein length between one- wing. third and two-thirds of subcostal width; 3. Long, thyridiate crossvein Comments. This character contains two levels: MP branching pat- longer than two-thirds of subcostal width, almost reaching R1, but tern (dichotomous or pectinate) and fork position (proximal to the not touched. separation of MA, distal to the separation of MA but basal half of Comments. Subcostal thyridiate crossveins of varying length are forewing, or distal half of forewing). It could be inferred that in the present in all extant nymphids. This feature was described by family Nymphidae, forewing MP dichotomously forking is plesio- New (1981, 1984) and termed by Oswald (1997, 1998) but has morphic, which was found in all fossil nymphids and half of extant never been used as a diagnostic character for the family. Thyridi- nymphids. Correspondingly pectinately forking is apomorphic. ate crossveins originate on the Sc and extend into the subcostal Moreover, forewing MP forking proximal to the separation of MA space, but never fuse with R. The length of these thyridiate is considered to be plesiomorphic in Nymphidae, which is repre- crossveins varies among genera and sometimes within a genus. sented by all fossil nymphids and most species of the extant genus Moreover, the basal one or two crossveins may be significantly Nesydrion. Nesydrion is the sister genus to the remaining Nymphi- longer than the remaining crossveins. The presence of subcostal nae. Correspondingly, the forewing MP branching distal to the sepa- thyridiate crossveins are considered to be a synapomorphy of ration of MA (including states 1 and 2) is apomorphic, which is Nymphidae. present in the other extant nymphids.

16. Forewing costal area after the fusion of Sc and R1, 22. Forewing, crossveins between MP2 branches ratio of costal width at Sc+R1 last fork level with fusion 0. Many rows of crossveins; 1. One complete row of crossveins; 2. of Sc and R1 level (gap: 1.12–1.27) No crossvein, or an incomplete row of crossveins. 0. Narrow, < 1.2; 1. Broad, > 1.2. Comments. The Myiodactylinae genera have fewer crossveins between MP2 branches, while the extant Nymphinae have more Comments. In Myrmeleontiformia, the forewing costal area is crossveins between MP2 branches. Nymphes myrmeleonoides and all usually dilated distally (i.e. after the fusion of Sc and R1) with Nesydrion have more than one row of crossveins. All Myiodactylinae Sc + R1 terminated after wing apex. The Cretaceous Baissoleon is taxa except Osmylops ectoarticulatus have no crossveins or an incom- unique in having an equally narrow costal area in both wings, and is plete row of crossveins between the branches of MP2. an autapomorphy within Nymphidae. – 17. Forewing Rs origin, ratio of Rs origin (length from 23. Forewing CuA branches number (7 9; except wing base to Rs origin) to R1 relative length (length Chasmoptera: 8) from wing base to Sc + R1) (gap: 0.14–0.40) 0. Eight or fewer; 1. More than eight. 0. Distant from wing base, > 0.2; 1. Proximal to wing base, < 0.2. 24. Forewing, crossveins between CuA branches 18. Forewing MA separation, ratio of MA separation 0. Many rows of crossveins; 1. One (incomplete) row of crossve- (length from Rs origin to MA separation) to Rs relative ins; 2. Crossveins absent. length (length from Rs origin to Sc + R1) (gap: 0.25– Comments. Genera in Myiodactylinae have no crossveins between forewing CuA branches while extant Nymphinae taxa have multiple 0.29) rows of crossveins. Most fossil nymphids have a few crossveins 0. Proximal to origin of Rs, ≤ 1/4; 1. Distant from origin of Rs, between forewing CuA branches. > 1/4. Comments. Among extant Nymphidae, forewing MA separated 25. Forewing CuP space, ratio of CuP space width to from Rs close to Rs origin is found in Myiodactylinae genera, while intracubital space width (measured from the midpoint of remote from the origin of Rs in Nymphinae genera. The position of Cu fork to Cu fork) (gap: 1.60–1.75) forewing MA separation is variable in fossil genera. Some of them do not correspond to that of other extant nymphids within the same 0. Narrow, < 1.7; 1. Broad, > 1.7. subfamily. Comments. In Myiodactylinae, Spilonymphes and Umbranymphes are recovered as sister genera based on both having a wide CuP 19. Forewing Rs branch number space, a state also found in Nymphydrion. All fossil species of Nym- phinae have a wide CuP space, along with Nymphes modesta and > ≤ 0. 10; 1. 10. Nymphes myrmeleonoides. The width of CuP space has been fre- 34 C. Shi et al. / Cladistics 0 (2015) 1–36 quently used in the diagnosis of fossil Nymphidae genera (e.g. Ma- Coniopterygidae, Hemerobiidae, Ithonidae, Berothidae, Dilaridae karkin et al., 2013b; Shi et al., 2013). and Dipteromantispidae (Yang et al., 2012; Makarkin et al., 2013a). 26. Forewing CuP branching 0. Pectinately forked with multiple branches; 1. Dichotomously 34. Hind wing Rs origin, ratio of Rs origin (hind wing forked with few branches. base to Rs origin) to R1 relative length (hind wing base Comments. Dichotomously forked forewing CuP is only found in to the fusion of Sc and R1) (gap: 0.15–0.175) the Cretaceous genera Baissoleon and Olindanymphes. 0. Distant from wing base, > 0.15; 1. Proximal to wing base, ≤ 0.15; 2. Rs absent. 27. Forewing, crossveins between branches of CuP Comments. Most nymphids have hind wing Rs originating near 0. Absent; 1. Present. the wing base, with the exception of Myiodactylus pubescens, Nesyd- rion fuscum, Nesydrion diaphanum, Sialium minor and Austronymphes Comments. Nymphes is characterized by one row of crossveins insularis. In Neuroptera, Rs originating distant from the forewing between the branches of forewing CuP. base is a synapomorphy for the subclade of Myrmeleontiformia comprising Nemopteridae, Myrmeleontidae and Ascalaphidae (Win- 28. Forewing CuA fork position, ratio of CuA fork terton et al., 2010). (length from wing base to CuA fork) to wing length (gap: 0.36–0.42; except Ithone: 0.39) 35. Hind wing, crossveins between MP2 branches < > 0. Proximal to wing base, 0.4; 1. Distant from wing base, 0.4. 0. One (incomplete) row of crossveins; 1. More than one row of Comments. Forewing CuA of all Nymphinae is forked near the crossveins. wing base, while that of most Myiodactylinae is forked remote from Comments. All extant genera of Nymphinae share the character wing base, sometimes even in the distal half of forewing. state of multiple rows of crossveins present between the hind wing MP2 branches. 29. Forewing, ratio of CuP length to total wing length – (gap: 0.50 0.55) 36. Hind wing, crossveins between CuA branches > 0. Extending into apical half of wing, 0.5; 1. Ending in basal 0. Crossveins absent; 1. At least one row of crossveins. half of wing, ≤ 0.5. Comments. A subgroup within Nymphinae comprising Nymphites, Comments. All Nymphinae species have forewing CuP termi- Sialium, Austronymphes and Nymphes share the character state of at nating in the basal half of the wing, although length varies least one row of crossveins present between hind wing CuA widely among different species. The Osmylops armatus species branches, and forming at least two rows of cells between CuA and group and Myiodactylus osmyloides have a short CuP, while the posterior margin. other Myiodactylus species have CuP longer than half of fore- wing length. 37. Hind wing CuA, ratio of CuA length (length from 30. Forewing, ratio of 1A length to wing length (gap: hind wing base to CuA termination) to wing length (gap: – 0.18–0.20) 0.42 0.50) ≥ 0. Elongated, > 0.2; 1. Short, < 0.2. 0. Extending into apical half of wing, 0.5; 1. Ending in basal half of wing, < 0.5; 2. Hind wing CuA absent. Comments. Forewing 1A of all extant genera of Nymphinae are Comments. Hind wing CuA length is polymorphic throughout generally short, while all the other nymphids have an elongated 1A. Nymphidae. Most Nymphinae genera have a short CuA except for Nymphes modesta. In Myiodactylinae, Myiodactylus osmyloides and 31. Forewing, 1A fork the Osmylops armatus species group have a short CuA, while the 0. Dichotomous; 1. Pectinate. remaining Myiodactylinae have a long CuA. Comments. All Nymphinae have a dichotomously forked 1A. In Myiodactylinae, the subgroup comprising Norfolius, Nymphydrion, 38. Hind wing CuP branching Myiodactylus and the Osmylops sejunctus species group have pecti- 0. Pectinate; 1. Dichotomous; 2. Hind wing CuP absent. nately forked 1A. This feature was previously identified by Yang Comments. The hind wing CuP is dichotomously forked in multi- et al. (2012, char. 37: AA3 + 4 branching). ple genera of Nymphidae, and appears to be found in more basal groups. Derived taxa tend to have pectinate forking of CuP. While 32. Forewing 2A and 3A pectinate forking was considered the plesiomorphic state, the polar- 0. Separate along length; 1. Forming a closed loop. ity could easily be reversed, with pectinate forking of CuP occurring multiple times throughout Nymphidae, from a plesiomorphic dichot- Comments. The forewing veins 2A and 3A forming a closed loop omously forked condition. is unique to Austronymphes.

33. Hind wing shape MALE GENITALIA (39–61) 0. Elongated and highly narrowed; 1. Rounded, similar shape to forewing. 39. 9th tergite Comments. The hind wing being elongated is a synapomor- phy for Nemopteridae. Hind wing modifications, albeit reduc- 0. Ventral margin protruding posteriorly; 1. Both posterior and tion to brachyptery rather than extreme narrowing, is found in anterior margins protruding; 2. Anterior margin protruding. C. Shi et al. / Cladistics 0 (2015) 1–36 35

Comments. Most male nymphids 9th tergite ventral edge posteri- 47. Mediuncus medioventral process orly protruded (Fig. 21b). In Nesydrion, 9th tergite ventral edge 0. Absent; 1. Ball-like; 2. Trilobate; 3. Elongated with protruding anteriorly protruded. In Myiodactylus and Norfolius, 9th tergite ven- hooks. tral edge protruded posteriorly and anteriorly (Fig. 21e). Comments. The mediuncus of members of Nymphinae has an elongated ventral process, while the mediuncus in Myiodactylinae is 40. 9th sternite smaller with a ball-like or trilobate processes. 0. Posteriorly elongated; 1. Strongly sclerotized and modified; 2. Subtriangular, rounded; 3. Emarginate; 4. Squared. 48. Mediuncus ornamentation 0. Setae; 1. None; 2. Teeth-like processes. 41. Ectoproct shape Comments. The modification on the surface or process of mediun- 0. Posteriorly protruding; 1. Small and rounded; 2. Multi-lobed cus is highly variable in Nymphinae, but always with teeth in Myio- process. dactylinae. Comments. The multi-lobed process on the ectoproct (Fig. 22a) is a significant character of the Osmylops sejunctus species group, 49. Gonarcus shape which was also listed in the diagnosis of the species group by Oswald (1997). 0. Arms fused (singular arched structure), or arms proximate medially and connected by membrane; 1. Arms widely separated 42. Subanal position medially. Comments. The gonarcus in Nymphinae has the arms widely sep- 0. Absent; 1. Posterior to ectoproct; 2. Ventral face of ectoproct. arated medially, while the gonarcus in Myiodactylinae has arms clo- Comments. The subanal is placed posterior to the ectoproct sely proximate dorsomedially or connected. in Nymphinae, but ventral to ectoproct in Myiodactylinae (except Umbranymphes and Osmylops placidus). Within Myiodac- 50. Gonarcus position (dorsally) tylinae, Umbranymphes and Osmylops placidus are sister to the remaining Myiodactylinae and exhibit the plesiomorphic condi- 0. Posteriorly protruding from ectoproct; 1. Located in the poster- tion. ior half of ectoproct; 2. Located in the anterior half of ectoproct; 3. Located in the 9th tergite. Comments. The position of the gonarcus is highly variable within 43. Mediuncus lateral lobes most genera, except for the dorsal position in Nymphes and Nesydri- 0. Absent; 1. Large. on. Comments. The presence of enlarged lateral lobes on the mediun- cus is apomorphic for Nymphinae (Fig. 20a,c). 51. Gonarcus position (ventrally) 0. Located in the 9th tergite, or the lateral membrane area 44. Mediuncus between 9th tergite and sternite; 1. Located in posterior or ven- 0. Ventral to ectoproct; 1. Posteriorly protruding from ectoproct. tral to 9th sternite; 2. Located in between 8th tergite and ster- Comments. Generally the mediuncus in Nymphinae is large and nite; 3. Located in the 9th sternite; 4. Located in the 8th placed posterior to ectoproct (except Nesydrion diaphanum). The sternite. mediuncus in Myiodactylinae is small and placed ventral to ecto- Comments. See character 50. proct (except Umbranymphes). 52. Gonarcus width (lateral view) 45. Mediuncus orientation; angle of mediuncus relative 0. Broad, but tapered to ventral point; 1. Broad, but tapered to gonarcus to dorsal point; 2. Broad, but tapered ventrally and dorsally; 3. 0. Greater than 45°;1.0–45°. Equally narrow; 4. Narrow, but broad to ventral point; 5. Equally broad. Comments. This character is from the lateral view. The orienta- tion of mediuncus (observed laterally) is from the dorsal to the ven- tral. For the gonarcus in a vertical position, the orientation is from 53. Entoprocesses dorsal end to ventral end. For the gonarcus in a horizontal position, 0. Absent; 1. Present. the orientation is from the posterior end (assumed as dorsal) to the Comments. This structure has been termed previously as entopro- anterior end (assumed as ventral). When measuring, we adjusted to cessi (New, 1981) and extrahemigonarcal process (Oswald, 1997). merge the dorsal points of mediuncus and gonarcus, and measured The presence of entoprocesses (Fig. 22a) is unique for the Osmylops the angle between the orientations of the two structures, for 0–45°, sejunctus species group, and was listed as a diagnostic feature of this like Austronymphes insularis (Fig. 21c), Myiodactylus osmyloides; for species group by Oswald (1997). > 45°, like Norfolius. Although the mediuncus and gonarcus of Psychopsis appear parallel, when adjusting to merge the two dorsal points, the angle should be > 90°. So we coded Psychopsis as state 0. 54. Position of ventral end of gonarcus and paramere base 46. Mediuncus dorsal process 0. Touching; 1. Separated; 2. Overlapping. 0. Simple edge; 1. Single protrusion; 2. Paired protrusion; 3. Comments. The ventral ends of the gonarcus and paramere bases Triplet protrusion. are touching in Nymphinae. In Myiodactylinae, they are typically Comments. Generally the mediuncus in Nymphinae has diversely widely separated, with the exception of Osmylops placidus and Nor- modified top processes (except Nesydrion diaphanum). The mediun- folius howensis where the ventral ends of the gonarcus and bases of cus in Myiodactylinae lacks any modification. the parameres are overlapping. 36 C. Shi et al. / Cladistics 0 (2015) 1–36

55. Membranous area/process around paramere base Comments. Seven stemmata is plesiomorphic for Neuropterida, as in Nemopteridae. Six stemmata are found in Nymphidae. Secondary 0. Absent; 1. Small, covering paramere base; 2. Elongated, sepa- reduction in stemmata number occurred in Psychopsidae (five) and rated from paramere. in Ithonidae (zero) (Winterton et al., 2010). Comments. A membranous covering around the base of the para- mere is present in most genera of Myiodactylinae, as well as the Nymphinae genus Austronymphes. The Osmylops sejunctus species 63. Antenna length group has a membrane process beneath the paramere, and almost as 0. Longer than half of mandible; 1. Shorter than half of mandible. long as paramere, which is a synapomorphy for the Osmylops sejunc- Comments. Antennae of Nymphidae and Nemopteridae are tus species group (Fig. 22a) (Oswald, 1997). shorter than half of mandible length, while antennae of Ithonidae and Psychopsidae are almost equally long as the mandible. 56. Paramere dorsal process 0. Absent; 1. Midway along paramere; 2. Located in the distal 64. Antennal pedicel one-third of paramere. 0. Elongated, almost as long as flagellum; 1. Short, shorter than Comments. The dorsal process on the paramere is variable in two-thirds of flagellum. Nymphinae and polymorphic in some genera. In Myiodactylinae, Myiodactylus osmyloides and Norfolius howensis have a dorsal pro- cess on the distal one-third of paramere; Osmylops has no dorsal 65. Antennal flagellum width process. 0. As broad as pedicel; 1. Much narrower than pedicel. Comments. Antennal flagellum is distinctly thinner than the pedi- 57. Paramere apical modification cel in Nymphidae, while the antennal flagellum is equal in width to 0. Absent; 1. Setose; 2. Teeth-like processes. the pedicel in Ithonidae, Psychopsidae and Nemopteridae.

58. Shape of paramere apical process 66. Labial palp segments 0. Single point; 1. Two points; 2. Three elongated processes; 3. 0. Four; 1. Three; 2. Two. Three points. Comments. Labial palpi are four-segmented in Ithonidae and Psychopsidae, three-segmented in Nymphidae and two-segmented in 59. Gonapsis Nemopteridae. 0. Absent; 1. Single; 2. Paired. 67. Mandible tooth Comments. The presence of a gonapsis is not common in Nym- phidae. A simple gonapsis is found in Norfolius (Fig. 20m) and two 0. Absent; 1. Single large tooth midway along mandible. species of Nymphes (Fig. 20l); Nesydrion has paired gonapsis Comments. Presence of elongated teeth along inner edge of man- (Fig. 20n), which are a synapomorphy for the genus. dible is a synapomorphy of Nymphidae + (Myrmeleontidae + Ascal- aphidae) (Aspock€ et al., 2001; Winterton et al., 2010). Nymphidae 60. Membranous processes beneath mediuncus possess a single large tooth midway along the mandible. 0. Absent; 1. Present. 68. Scoli on meso- and metathorax and abdomen Comments. Membrane process beneath mediuncus is an autapo- morphy for Norfolius (Fig. 21e). 0. Absent; 1. Present. Comments. Presence of scoli on meso- and metathorax and abdo- 61.Gonosetae men is a synapomorphy for Nymphidae, Ascalaphidae and Myr- meleontidae. 0. Present; 1. Absent. Comments. Gonosetae are present in most Nemopteridae, the pres- Material examined ence of which is used to differentiate genera (Koch, 1967; Tjeder, 1967). Ithone fulva Tillyard, Nemoptera coa (Linnaeus), Chasmoptera hut- ti (Westwood), Psychopsis barnardi Tillyard, Osmylops ectoarticulatus Oswald, Osmylops armatus (McLachlan), Osmylops sejunctus LARVA (62–68) (Walker), Osmylops placidus (Gerstaecker), Myiodactylus osmyloides Brauer, Myiodactylus pubescens Banks, Norfolius howensis (Tillyard), Larval characters of Nymphidae were collected from Osmylops Nesydrion fuscum Gerstaecker, Nesydrion diaphanum Gerstaecker, (New, 1983a), Norfolius (New and Lambkin, 1989) and Nymphes Nesydrion nigrinerve Esben-Petersen, Austronymphes insularis Esben- (New, 1982). We assume these characters are conservative in Nym- Petersen, Nymphes aperta New, Nymphes myrmeleonoides Leach, phidae, including fossil and extant taxa. Nymphes modesta Gerstaecker, Nymphydrion delicatum Banks, Um- branymphes spinosus New, Liminympha makarkini Ren et Engel, Nymphites bimaculatus Shi, Makarkin et Ren, Sialium sinicus sp. 62. Stemmata number nov., Sialium minor sp. nov., Baissoleon similis sp. nov., Baissoleon 0. None; 1. Five; 2. Six; 3. Seven. cretaceous Makarkin, Spilonymphes major gen. et sp. nov.