Cladistics Cladistics 31 (2015) 455–490 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. 456 C. Shi et al. / Cladistics 31 (2015) 455–490 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 31 (2015) 455–490 457 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