JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 81(3), 2008, pp. 188–193 New from the Jiulongshan Formation of Inner Mongolia, China (Raphidioptera)

1 2 MICHAEL S. ENGEL AND DONG REN

ABSTRACT: Two new genera and of snakeflies (Raphidioptera) are described and figured from the Jiulongshan Formation of eastern Inner Mongolia, China. Ororaphidia megalocephala, new and species, and Styporaphidia magia, new genus and species, are both differentiated from other Mesozoic lineages. These constitute the oldest records of snakeflies from China. KEY WORDS: Raphidiomorpha, Neuropterida, Mesozoic, paleontology, , snakefly

The snakeflies (Raphidioptera) comprise six living and fossil families of distinctive neuropterid (Grimaldi and Engel, 2005). Today the order is restricted to the Holarctic and is represented solely by two families – and Inocelliidae – comprising the Neoraphidioptera. However, during the past the order was of global distribution and the habitat restrictions observed for modern snakeflies were clearly not present as species occurred in environments (e.g., warm tropics) that today exclude the lineage (Grimaldi and Engel, 2005). In addition, numerous families and species are regularly found in Jurassic and deposits suggesting that the order experienced its heyday during the Mesozoic and has subsequently dwindled in overall diversity. The diversity of fossil snakeflies was summarized by Engel (2002), with updates provided by Engel (2003), Aspo¨ck and Aspo¨ck (2004), Engel et al. (2006), de Flores and Nel (2006), and Perrichot and Engel (2007), and one overlooked species presented in Wang (1987). Herein we provide the description of two new Mesozoic snakeflies from the Jurassic Jiulongshan Formation of Inner Mongolia, China uncovered by Ren. These deposits have produced an abundance of fossils (e.g., Ren, 2002; Renand Oswald, 2002; Ren et al., 2002). The insect fauna of the Jiulongshan Formation has been summarized by Hong (1983) and Ren et al. (1995, 2002). Figures were prepared by D.R. using a Leica MZ12.5 stereomicroscope.

Systematic Paleontology Ororaphidia Engel and Ren, new genus TYPE SPECIES: Ororaphidia megalocephala Engel and Ren, new species. DIAGNOSIS: Head oval, slightly longer than wide, slightly longer than prothorax; ocelli present; coronal ecdysial cleavage lines (5Y-shaped ‘‘suture’’) present. Pronotum trapezoidal, lateral margins converging posteriorly such that anterior margin slightly wider than posterior margin, length about as long as combined lengths of meso- and metanotum. Wings slender, with relatively few crossveins in

1 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]). 2 College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidan District, Beijing 100037, People’s Republic of China ([email protected]). Accepted 13 April 2008; Revised 8 May 2008 E 2008 Kansas Entomological Society VOLUME 81, ISSUE 3 189 radial and medial fields; forewing Sc terminating into C beyond wing midlength; a single sc-r crossvein present basal of separation of Rs from stem of R; pterostigma well defined and elongate, with one pterostigmal crossvein in apical third; forewing MA originating slightly before tangent of MP forking and coalescent with stem of R for short distance; Rs and MA each with three terminal branches; MP with three medial cells; CuA connecting to M distal to separation of M from R, with a single terminal branch; 1A simple; hind wing MA originating from stem of R; hind wing MP with a single medial cell; 1mp-cua (‘‘arculus’’ vein) orthogonal to MP. ETYMOLOGY: The new genus-group name is a combination of the Greek word oros (meaning, ‘‘mountain’’) and Raphidia (first generic name of the Raphidioptera and stem for most names in the order). The name is feminine. COMMENTS: The new genus has a wing venation superficially similar to some Mesoraphidia, such as Mesoraphidia heteroneura Ren (Ren, 1997). However, Ororaphidia can be distinguished from these by the presence of a pterostigmal crossvein, a cup-a crossvein in the forewing, a single medial cell in the hind wing MP field, a shorter pronotum with lateral margins converging posteriorly, and shorter ovipositor.

Ororaphidia megalocephala Engel and Ren, new species (Figs. 1, 3–5) DIAGNOSIS: As for the genus (vide supra). DESCRIPTION: Female. Total body length (excluding ovipositor and antennae) 12 mm; forewing length 11.4 mm, width 4 mm. Compound eyes exophthalmic, positioned at anterior of head. Pterostigma dark brown, wing membrane otherwise hyaline. Legs, where evident, slender. Abdomen shorter than wings, parallel-sided, with six visible segments as preserved, each segment wider than long; ovipositor relatively short, apparently not surpassing wing apex. HOLOTYPE: Specimen No. NNS-200201 in the Department of Biology, Capital Normal University, Beijing, China; a complete well-preserved adult with wings. OCCURRENCE: Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Jurassic (Aalenian-Bajocian). ETYMOLOGY: The specific epithet is a combination of the Greek words megalos (meaning, ‘‘very large’’) and kephale (meaning, ‘‘head’’) and is a reference to the proportionally large size of the head capsule.

Styporaphidia Engel and Ren, new genus TYPE SPECIES: Styporaphidia magia Engel and Ren, new species. DIAGNOSIS: Head oval, longer than wide, longer than prothorax; ocelli present; coronal ecdysial cleavage lines (5Y-shaped ‘‘suture’’) not evident. Pronotum quadrate, shorter than combined lengths of meso- and metanotum. Wings slender, elongate, with relatively few crossveins in radial and medial fields; forewing Sc terminating into C beyond wing midlength; a single sc-r crossvein present basal separation of Rs from stem of R; pterostigma well defined and elongate, with two pterostigmal crossveins; forewing MA originating before tangent of MP forking and coalescent with stem of R for short distance; Rs and MA each with two terminal branches; MP with three medial cells; CuA connecting to M distal to separation of 190 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Figs. 1–2. Photomicrographs of Jurassic snakeflies from Inner Mongolia, China. 1. Holotype female of Ororaphidia megalocephala Engel and Ren, new genus and species (NNS-200201). 2. Holotype of Styporaphidia magia Engel and Ren, new genus and species (NNS-200202).

M from R, with a single terminal branch; 1A simple; hind wing MA originating from stem of R; hind wing MP with a single medial cell. ETYMOLOGY: The new genus-group name is a combination of the Greek word stypos (meaning, ‘‘stem’’ or ‘‘stump’’) and Raphidia (first generic name of the Raphidioptera and stem for most names in the order). The name is feminine. COMMENTS: The presence of two pterostigmal crossveins in the apical half of the pterostigma is a relatively uncommon feature among Mesozoic Raphidioptera and is typically known only in some baissopterids. The fossil described herein lacks the features of the primitive family Baissopteridae (e.g., the numerous crossveins in the radial and medial fields, concomitant larger number of medial cells), yet has this distinctive trait, a combination of attributes that makes the genus readily distinguishable from other Mesozoic snakeflies. VOLUME 81, ISSUE 3 191

Figs. 3–5. Line illustrations of Ororaphidia megalocephala Engel and Ren, new genus and species (NNS- 200201). 3. Outline of entire specimen (wing venation omitted). 4. Forewing venation. 5. Hind wing venation. Figures of wings share the same scale bar.

Styporaphidia magia Engel and Ren, new species (Figs. 2, 6–10) DIAGNOSIS: As for the genus (vide supra). DESCRIPTION: Sex indeterminate. Total body length (as preserved) 13 mm; forewing length 10 mm, width 4 mm. Compound eyes exophthalmic, positioned at anterior of head. Pterostigma dark brown, wing membrane otherwise hyaline. Legs, where evident, slender. Abdomen shorter than wings, parallel-sided, with eight preserved segments, each segment wider than long; abdominal apex not preserved. HOLOTYPE: Specimen No. NNS-200202 in the Department of Biology, Capital Normal University, Beijing, China; a complete well-preserved adult with wings. OCCURRENCE: Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; Jiulongshan Formation, Jurassic (Aalenian-Bajocian). ETYMOLOGY: The specific epithet is based on the Greek word mageia, meaning ‘‘magic’’.

Acknowledgments We are grateful to W. Wichard and R.J. Beckemeyer for comments on the manuscript. This research was partially supported by National Science Foundation 192 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Figs. 6–10. Line illustrations of Styporaphidia magia Engel and Ren, new genus and species (NNS- 200202). 6. Outline of entire specimen (wing venation omitted). 7. Left forewing venation. 8. Left hind wing venation. 9. Right forewing venation. 10. Right hind wing venation. Figures of wings share the same scale bar.

(USA) grant DEB-0542909 (to M.S.E.), National Natural Science Foundationof China grant 30430100 (to D.R.), Nature Science Foundation of Beijing grant 5082002 (to D.R.), and the Funding Project for Academic Human Resources under the jurisdiction of the Beijing Municipality (to. D.R.). This is a contribution of the Division of Entomology (Paleoentomology), University of Kansas Natural History Museum.

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