JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 81(3), 2008, pp. 161–167 Aetheogrammatidae, A New Family of Lacewings from the Mesozoic of China (Neuroptera: Myrmeleontiformia)

1 2,3 DONG REN AND MICHAEL S. ENGEL

ABSTRACT: A family of extinct lacewings is described and figured from the Yixian Formation [Early Cretaceous (M.S.E.) or latest Jurassic (D.R.)] of Liaoning Province, China. Aetheogrammatidae Ren and Engel, new family, is established for Aetheogramma speciosa Ren and Engel, new genus and species, and is distinguished from other myrmeleontiform families. The enigmatic kalligrammatid genus Kallihemerobius Ren and Oswald is placed in a new subfamily, Kallihemerobiinae Ren and Engel. KEY WORDS: China, Mesozoic, Myrmeleontiformia, paleontology, Planipennia, taxonomy

The order Neuroptera (lacewings and their relatives) has a long and rich fossil history (Grimaldi and Engel, 2005). The order comprises, along with its near relatives the Megaloptera, Raphidioptera, and the extinct Protoneuroptera (Permoberothidae: sensu Grimaldi and Engel, 2005), some of the most archaic holometabolous insects. Despite their remarkable anatomical and biological diversity, particularly in larval habitats and forms, the Neuroptera are perhaps dwindling in overall numbers. During the past the order comprised a rich complex of families. While several of these families represent stem groups to more modern lineages, others existed alongside surviving families and truly represent entire branches of neuropteran diversity that have not persisted to the present (Grimaldi and Engel, 2005). Herein we describe a new genus and species of fossil lacewings, Aetheogramma speciosa, new genus and species, based on two rather well preserved specimens collected by Ren in the village of Huangbanjiegou, near the town of Shangyuan, 28 km southeast of Beipiao. The species possesses a remarkable combinationof features that exclude it from previously known families of Neuroptera, although it is not without some resemblance to the giants of the extinct family Kalligrammatidae. We consequently propose a new family for this enigmatic taxon and discuss its unique suite of venational attributes. The fossils described herein were recovered from the second member of the Yixian Formation exposed near the city of Beipiao in Liaoning Province, northeastern China (Ren et al., 1997). The dating of the Yixian Formation is of some legitimate controversy, with different scholars arguing for either an Early Cretaceous (e.g., Swisher et al., 1999, 2002; Yang et al., 2007) or Late Jurassic (e.g., Ren et al., 1995) age. The authors differ as to their position regarding this debate (DR agreeing with the older date, MSE agreeing with an Early Cretaceous age). The deposit has produced an abundance (more than 50,000 specimens excavated to date) of well

1 Department of Biology, Capital Normal University, 105 Xisanhuanbeilu, Haidan District, Beijing 100037, People’s Republic of China ([email protected]). 2 Division of Entomology (Paleoentomology), Natural History Museum, and Department of Ecology & Evolutionary Biology, 1501 Crestline Drive — Suite #140, University of Kansas, Lawrence, Kansas 66049-2811 ([email protected]). 3 To whom correspondence should be addressed. Accepted 1 May 2008; Revised 8 May 2008 E 2008 Kansas Entomological Society 162 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Fig. 1. Photomicrograph of holotype of Aetheogramma speciosa Ren and Engel, new genus and species (LB20010-1). preserved fossil insects from an array of orders, thereby providing a significant perspective on Mesozoic insect diversity. To date the diversity includes Ephemer- optera, Odonata, Orthoptera, Phasmatodea s.l. (i.e., Holophasmatodea), Dermap- tera, Hemiptera, Raphidioptera, Neuroptera, Coleoptera, Hymenoptera, Mecop- tera, and Diptera (e.g., Ren et al., 1995; Ren and Guo, 1996; Ren, 2002).

Systematic Paleontology

Suborder MYRMELEONTIFORMIA Henry AETHEOGRAMMATIDAE Ren and Engel, new family

TYPE GENUS: Aetheogramma Ren and Engel, new genus. DIAGNOSIS: Moderately large insects, with wing spans near 9 cm (Figs. 1–2). Wings oval, with rounded apices (Figs. 3–4), subequal in length, with broad fuscous bands running from costal margin to posterior margin (faintly preserved but evident in Figs. 1–2); pterostigma absent; trichosors and microtrichia absent; wings lacking conspicuous ‘‘eye-spot’’; crossveins numerous and regular over surface of wings (Figs. 3–4), not forming gradate series; principal longitudinal veins (Sc, R1, Rs, MA, MP) without twigging along wing margins (Figs. 1–4). Forewing without recurrent humeral vein; costal space moderately broad basally, tapering in width toward wing apex, all c-sc crossveins (costal veinlets) straight and simple (Fig. 3), forming single row of cells (without interlinking intercalary veins or crossveins); Sc paralleling R1 for entire wing length before terminating at C slightly prior to wing apex (Fig. 3), numerous and regular sc-r crossveins; R1 simple for most VOLUME 81, ISSUE 3 163

Fig. 2. Photomicrograph of holotype counterpart (LB20010-2) of Aetheogramma speciosa Ren and Engel, new genus and species. of length, branching once just before apical fourth of wing length, branches reaching wing margin slightly posterior to wing apex; Rs originating near wing base (Fig. 3), Rs branching in apical third of wing length, forming three principal branches, medial branch forking once; MA simple; MP branching pectinately, first branch at basal third of wing length, with at least four branches; CuA and CuP dichotomously branching; three anal veins present, 1A branching dichotomously, 2A simple, 3A apparently with two simple branches. Hind wing generally similar to forewing, slightly wider and with apex more broadly rounded (Fig. 4); venation similar to that of forewing except costal area narrower (Fig. 4), R with three branches reaching wing margin, 3A vestigialor perhaps even absent, and CuA, CuP, and 2A pectinate. COMMENTS: This family is most similar to the extinct giants of the family Kalligrammatidae, also known from the Mesozoic of Asia. Table 1 provides diagnostic features serving to distinguish Aetheogrammatidae from the two main branches of kalligrammatids. Kallihemerobius Ren and Oswald is an exceptionally bizarre genus of Kalligrammatidae and is perhaps sister to the remainder of the family. We have accordingly placed it in its own subfamily (vide Appendix and Table 1).

Aetheogramma Ren and Engel, new genus

TYPE SPECIES: Aetheogramma speciosa Ren and Engel, new species. DIAGNOSIS: As for the family (vide supra). 164 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Figs. 3–4. Wing venation of Aetheogramma speciosa Ren and Engel, new genus and species (wing markings omitted). 3. Forewing venation. 4. Hind wing venation.

ETYMOLOGY: The new genus-group name is a combination of the Greek words aethes (meaning, ‘‘unusual’’) and gramma (meaning, ‘‘mark’’ or ‘‘picture’’). The name is feminine.

Aetheogramma speciosa Ren and Engel, new species

DIAGNOSIS: As for the genus (vide supra). DESCRIPTION: The familial diagnosis is meant to simultaneously characterize the genus and species (ICZN, 1999: Arts. 13.4 and 13.5), with the following minor VOLUME 81, ISSUE 3 165

Table 1. Comparison of Aetheogrammatidae, Kalligrammatinae (5Kalligrammatidae s. str.), and the peculiar genus Kallihemerobius (Kallihemerobiinae: vide Appendix). Putatively apomorphic states boldfaced.

Kalligrammatinae Kallihemerobiinae Aetheogrammatidae

Remigial ‘‘eye’’ spot: present present absent Trichosors: absent present absent

Termination of Sc: into R1 into R1 at C R1*: simple pectinate branched apically Crossvenation: dense except margins** dense except margins dense including margins Recurrent humeral vein: present/absent*** present absent c-sc crossveins: forked forked simple

* The presence of ‘‘multiple’’ radial sectors results from the incorporation of Rs into R1 and thereby creates a more pectinate R1, a condition typical of Hemerobiidae and convergent in the enigmatic kalligrammatid genus Kallihemerobius. It is unclear whether the branching of R1 in Aetheogramma, however, is homologous with that seen in the aforementioned lineages as R1 is not pectinate but instead simply branched apically and quite distant from the origin of the main stem of Rs (vide etiam Discussion). ** Crossvenation is typically not present in area of marginal twigging and so the polarity of this trait is of some question as the loss of marginal crossvenation is linked to twigging. *** A couple of genera (Kalligramma and Oregramma) lack the recurrent humeral vein. This absence is likely an apomorphic feature within the family as these genera otherwise embody the broader suite of kalligrammatid characters (vide etiam Discussion). additions: Forewing length 45 mm, width 21 mm; hind wing length 44 mm, width 27 mm. HOLOTYPE: Holotype part (LB20010-1) and counterpart (LB20010-2); Yixian Formation, Huangbanjiegou Village, near Shangyuan Township, 28 km southeast of Biepiao City, Liaoning Province, China; deposited in the collection of the Department of Biology, Capital Normal University, Beijing, People’s Republic of China. ETYMOLOGY: The specific epithet is taken from the Latin term speciosus, meaning ‘‘splendid’’ or ‘‘beautiful.’’

Discussion The remarkable array of features embodied by A. speciosa is not found among any other lineage of myrmeleontiform Neuroptera, although there appears to be some affinity with the Kalligrammatidae. The high density of the crossveins in Aetheogrammatidae is putatively apomorphic. A more moderate degree of cross- venation is plesiomorphic for Neuroptera, although the extreme reduction seen in Coniopterygidae (Hemerobiiformia) is derived just as is the high density seen in the myrmeleontiform families Kalligrammatidae, Panfiloviidae, and Grammolingiidae. It is not immediately apparent whether or not such a feature unites these taxa as the assuredly related modern family Psychopsidae (and likely also Nymphidae) and other extinct families such as Osmylopsychopidae, Brongniartiellidae, and Nymphi- tidae exhibit more general crossvenation. As such, it is equally likely that the dense crossvenation of Aetheogrammatidae, Kalligrammatidae, Panfiloviidae, and Gram- molingiidae may be independently derived among subsets of these families. The recurrent humeral crossvein has been considered a primitive feature that has been lost repeatedly in various families in the course of neuropteran evolution (e.g., Barnard, 1981). Indeed, two genera within the Kalligrammatidae (Kalligramma 166 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Walther and Oregramma Ren), a family otherwise possessing a recurrent humeral vein, have lost this vein, putatively apomorphically. The absence of the recurrent humeral vein in Aetheogrammatidae is similarly interpreted as a derived feature. Nonetheless, the basalmost lineages of Neuroptera (e.g., Permoberothidae and others) lack a recurrent humeral crossvein, so this attribute is likely not a groundplan feature of the order, although it probably arose early in the course of neuropteran evolution and was then repeatedly lost in numerous lineages. Trichosors and branched costal veinlets are generally considered primitive features for many lineages of Neuroptera; therefore, the absence of trichosors and simple c-sc crossveins in A. speciosa are interpreted as apomorphic traits. In contrast, the absence of an ‘‘eye-spot’’ on the wings of Aetheogrammatidae is a primitive feature relative to Kalligrammatidae which generally have such markings. Additionally, the termination of Sc into C is likely primitive, whereas the fusion Sc with R1 is a derived feature among Neuroptera. The simple forewing MA and shape of the wing is reminiscent of some forewings of Nemopteridae, although the hind wings of this family are dramatically modified. For the present the family consists solely of A. speciosa, although Panfilov (1980) described a genus of Kalligrammatidae from the Late Jurassic of Karatau (southern Kazakhstan) that may be best classified in Aetheogrammatidae. The genus Kalligrammina Panfilov was proposed for a fragmentary portion of an isolated hind wing (Panfilov, 1980). Like Aetheogramma, the genus Kalligrammina possesses a single branch of R1 in the apical portion of the wing and greatly distant from the otherwise basal origin of the Rs stem. It is, therefore, possible that Kalligrammina is allied to Aetheogramma and should be transferred to Aetheogrammatidae. However, as noted, Kalligrammina is fragmentary, and in the absence of other definitive character information we prefer not to formally transfer the genus at this time. Certainly a great deal of cladistic work is needed for the Myrmeleontiformia. Fossils will assuredly be needed to elaborate the phylogeny of this group as the geological record documents a rich and now largely extinct history for the lineage, with a relatively few, apomorphic survivors. It is exactly for such groups that the incorporation of paleontological data is most crucial, as the unique character combinations provided by such extinct taxa are vital. Aetheogrammatidae is one such part of a mosaic that will help to bring together the rich tapestry of early neuropteran diversity and evolution.

Acknowledgments We are grateful to two anonymous reviewers who provided useful comments for improving the manuscript; and to Charles D. Michener for editorial assistance. This project was funded by the National Natural Science Foundation of China (30430100 to D.R.), the Beijing Natural Science Foundation (5082002 to D.R.), and the National Science Foundation (USA) (DEB-0542909 to M.S.E.). This is a contribution of the Division of Entomology, University of Kansas Natural History Museum.

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Appendix

Family KALLIGRAMMATIDAE Handlirsch KALLIHEMEROBIINAE Ren and Engel, new subfamily

TYPE GENUS: Kallihemerobius Ren and Oswald, 2002. DIAGNOSIS: Wings with recurrent humeral crossvein, trichosors, and ‘‘eye-spots’’; longitudinal veins with marginal twigging along wing margins, crossveins not present in area of marginal twigging. Forewing with c-sc crossveins forked; Sc terminating into R1 near wing apex; R1 pectinately branched (i.e., with multiple ‘‘Rs’’ veins).