Zootaxa, New Fossil Prophalangopsidae (Orthoptera, Hagloidea

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New fossil Prophalangopsidae (Orthoptera, Hagloidea) from the Middle Jurassic of Inner Mongolia, China

JUNJIE GU, YUN-YUN ZHAO1 & DONG REN1 College of Life Science, Capital Normal University, Beijing 100048, China 1Corresponding author. E-mail: [email protected] or [email protected]

Abstract

Two new species attributed to the genus Sigmaboilus Fang, Zhang & Wang, 2007 are described from the Middle Jurassic Jiulongshan Formation of Daohugou Village, Inner Mongolia, China: Sigmaboilus fuscus sp. nov., S. peregrinus sp. nov. The diagnosis of the genus is revised.

Key words: Orthoptera, Prophalangopsidae, new species, Middle Jurassic, Daohugou, China

Introduction

Prophalangopsidae Kirby, 1906 contains one extant and five fossil subfamilies: Aboilinae Martynov, 1925 (Lower Jurassic-Upper Cretaceous, Siberia, Kazakhstan, Kirgyzstan, Mongolia, China, Japan, Germany; Martynov, 1925), Protaboilinae Gorochov, 1988 (Lower Jurassic, Middle Asia; Gorochov, 1988), Chifengiinae Hong, 1982 (Upper Jurassic-Lower Cretaceous, Siberia, China; Hong, 1982), Termitidiinae Zeuner, 1939 (Lower Cretaceous, England; Zeuner, 1939), Tettohaglinae Gorochov, 2003 (Lower Cretaceous, Siberia; Gorochov, 2003) and an extant subfamily, Prophalangopsinae Kirby, 1906 (India, China; Kirby, 1906). Aboilinae constitutes the most species-rich and diverse subfamily of all (Gorochov, 2003). Up to now, 19 genera and 46 species belonging to Aboilinae have been described (Deichmuller, 1886; Martynov, 1925; Sharov, 1962, 1968, Fujiama, 1976; Hong, 1982, 1983; Gorochov, 1988, 1990, 1996; Li et al., 2007; Fang et al., 2007). Recently, we recovered 24 fossil specimens of Aboilinae containing two new species: S. fuscus sp. nov., S. peregrinus sp. nov. Based on these well-preserved fossil specimens collected from the Middle Jurassic Jiulongshan Formation at Daohugou Village, Ningcheng County, Inner Mongolia, China, the diagnosis of Sigmaboilus Fang, Zhang & Wang, 2007 is revised and intraspecific forewing variation is briefly discussed. The age of the Jiulongshan Formation is still controversial, but most published biostratigraphic correlations and radiometric dates support a Middle Jurassic age (Chen et al. 2004; Ren et al. 1995; Ren et al. 2002; Tan & Ren 2002; Gao & Ren 2006).

Material and methods

All the type specimens of the new species are housed at the Key Lab of Insect Evolution & Environmental Changes, Capital Normal University, Beijing, China. The specimens were examined with a Leica MZ12.5 dissecting microscope and illustrated with the aid of a drawing tube attached to the microscope. Line drawings were made with CorelDraw 12 graphic software.

16 Accepted by D. Rentz: 17 Dec. 2008; published: 5 Feb. 2009 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

In order to establish a cladistic phylogeny of the Orthoptera, a new venation nomenclature and a new interpretation of the wing venation pattern have been proposed by Béthoux & Nel (2001, 2002). The venation nomenclature of Orthoptera remains a focus of discussion (Béthoux, 2007; Rasnitsyn, 2007). The wing venation nomenclature used in this paper is based on the interpretation of Béthoux & Nel (2001, 2002, 2005): ScA, anterior subcosta; ScP, posterior subcosta; RA, anterior radial; RP, posterior radial; MA, anterior media; MP, posterior media; CuA, anterior Cubitus; CuP1a, the anterior branch of first posterior Cubitus; CuP1b, the posterior branch of first posterior Cubitus; CuP2, the second posterior Cubitus; handle, a strong cross-vein appearing as a main vein, between origin of CuA + CuP1a and CuP1b.

Systematic Palaeontology

Order Orthoptera Olivier, 1789

Superfamily Hagloidea Handlirsch, 1906

Family Prophalangopsidae Kirby, 1906

Subfamily Aboilinae Martynov, 1925

Genus Sigmaboilus Fang, Zhang & Wang, 2007

Type species: Sigmaboilus gorochovi Fang, Zhang & Wang, 2007 Revised diagnosis: Forewing: Base of tegmen constricted; ScA long, mildly undulate, reaching anterior wing margin beyond midlength; ScP long with numerous branches ending in stem ScA; Cross-veins at the base of area between CuA+M and CuP sigmoidal; CuA and CuP1a remaining united as CuA+CuP1a for a short and appreciable distance . Hindwing: ScP and R with basal common stem, basal free part of R slightly curved; R diverging nearer wing base than in forewing, RP branched before first branch of RA; M bowed toward R and diverging before origin of RP. Species included: S. gorochovi Fang, Zhang & Wang, 2007, S. sinensis Fang, Zhang & Wang, 2007, S. fuscus sp. nov. S. peregrinus sp. nov. Remarks: The genus Sigmaboilus Fang, Zhang & Wang, 2007 was erected based on only a single forewing from the Middle Jurassic of Daohugou, Inner Mongolia, China. The diagnostic characteristics of the hind wing can be seen clearly from the material described here: ScP fused with R at base, separated before the point of origin MA.

Key to species of the genus Sigmaboilus based on tegmen

1. Whole tegmen fuscous...... 2 -. The coloration of tegmen formed irregular bands ...... 3 2. Second anterior branch of CuA+CuP1a forked and CuP1b straight ...... S. gorochovi Fang, Zhang & Wang, 2007 -. Second anterior branch of CuA+CuP1a pectinate and CuP1b curved...... S. fuscus sp. nov. 3. First and second anterior branch of CuA+CuP1a forked, the ‘handle’ forming a distinct angle with crossvein between the ‘handle’ and CuP1a...... S. sinensis Fang, Zhang & Wang, 2007 -. Second anterior branch of CuA+CuP1a dichotomously ramified, the ‘handle’ subparallel to each crossvein between the ‘handle’ and CuP1a...... S. peregrinus sp. nov.

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Sigmaboilus fuscus sp. nov. (Figs. 1–2)

Diagnosis: Forewing: Whole tegmen fuscous; precostal area approximately triangular; basal free part of CuA gently curved; second anterior branch of CuA+CuP1a pectinate with four branches; CuP1b slightly curved. Hindwing: ScP and R with a long basal common stem; R diverging nearer wing base than in forewing, RP branched before first branch of RA; M bowed towards R and diverging before origin of RP.

FIGURE 1. Sigmaboilus fuscus sp. nov., photograph of holotype, CNU-ORT-NN2008035.

FIGURE 2. Sigmaboilus fuscus sp. nov., line drawing of holotype, A. forewing; B. hindwing, CNU-ORT-NN2008035.

Description: Forewings; long and narrow; base of tegmen constricted. Preserved length about 30.5 mm, width 8.7 mm (opposite the fusion of ScA with anterior margin, holotype). Area between ScA and anterior margin approximately triangular and numerous ‘radial veinlets’ coupled with regular cross-veins at its base, loose branches and network of irregular cross-veins take up distal part. ScA long, mildly undulate, reaching

18 · Zootaxa 2004 © 2009 Magnolia Press GU ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. anterior wing margin slightly beyond first branch of RA. ScP apparently long, branches of ScP numerous and most of them end in stem ScA, straight and regular cross-veins between them; Area between ScP and R narrow. R straight and strong, R and M with basal common stem; R forking into RA and RP at about 1/3 basal tegminal length, RA posteriorly pectinate with 5 branches in preserved part, RP branched beyond first branch of RA, posteriorly pectinate at least with 4 branches; Area between R and M with straight and more or less parallel cross-veins in preserved part. M forked before the origin of RP, MA oriented towards RP at its origin, further gently curved and oriented toward posterior wing margin; Cross-veins at the base of area between CuA+M and CuP sigmoidal. Basal free part of CuA gently curved and slightly longer than basal free part of M; point of fusion of CuA and CuP1a beyond divergence of M; CuP1a twice the length of CuA before their fusion; CuA and CuP1a remaining united as CuA+CuP1a for a short and measurable distance, and exhibiting 6 terminal branches, first anterior branch of CuA+CuP1a oriented toward MP, beyond gently curved; second anterior branch of CuA+CuP1a pectinate with four branches; cross-veins between branches of CuA+CuP1a variable, some sigmoidal, some ramified; CuP1b slightly sigmoidal; CuP2 strong, sharply curved, probably Z- shaped, cross-veins between CuP2 and CuP1b curved. 1A strong and similar to CuP2 in shape, probably touching CuP2 at a point, 2A apparently S-shaped. Hindwing: Poorly preserved and largely hidden by forewing; Preserved length about 27.8 mm. Area between ScA and ScP very narrow, ScP and R with basal common stem, basal free part of R slightly curved towards M; R diverging nearer wing base than in forewing, RA posteriorly pectinate with 5 branches in preserved part; RP branched before first branch of RA and pectinate at least with 6 branches, cross-veins between RA and RP regular, areas between branches of RP with straight cross-veins in basal part and reticulated cross-veins in distal part; M bowed toward R and diverging before origin of RP, MA and MP gently curved and subparallel; Cubital and anal areas missing. Material: Holotype, male tegmen and poorly preserved hindwing, CNU-ORT-NN2008035; paratypes: 1, male tegmen: CNU-ORT-NN2008049. Etymology: The specific epithet derives from Latin ‘fuscus’, for its fuscous wing. Horizon and locality: Jiulongshan Formation, Middle Jurassic, Daohugou Village, Ningcheng County, Inner Mongolia, China. Remarks: The new species can be distinguished from the type species in following features on the forewing: Precostal area approximately triangular; branching mode of CuA+CuP1a different; basal free part of CuA slightly curved; CuP1b between ‘handle’ and posterior wing margin curved; and CuP1a twice the length of CuA before their fusion. It differs from Sigmaboilus sinensis in having a whole fuscous tegmen, and second anterior branch of CuA+CuP1a pectinate.

Sigmaboilus peregrinus sp. nov. (Figs. 3–8)

Diagnosis: Forewing: ScP long and ending on anterior margin at about the third fifth of the wing; CuA+CuP1a ramified with 4 to 6 terminal branches, first anterior branch of CuA+CuP1a simple, second anterior branch dichotomously ramified; and coloration formed irregular bands. Description: Forewings: 37.9mm/40mm preserved length (respectively left forewing/right forewing, holotype). Long and narrow, base of tegmen constricted. Precostal area between ScA and anterior margin very wide, with ‘radial veinlets’ and regular cross-veins at its base. ScA apparently long, mildly undulate; ScP long and ending on anterior margin at about the third fifth of the wing, branches of ScP numerous and most of them end in stem ScA. Area between ScP and R narrow and with regular cross-veins. R forking into RA and RP at about 1/3 basal tegminal length; RA pectinate with 6-8 terminal branches and RP with 6-7 terminal branches; RP branched beyond first branch of RA, but beyond second branch RA in left forewing of holotype. Area between R and M with oblique and more or less parallel cross-veins before mid-length, dense and network of regular cross-veins covering distal half; M forked before the origin of RP, MA oriented towards RP at its

JURASSIC PROPHALANGOPSIDAE FROM CHINA Zootaxa 2004 © 2009 Magnolia Press · 19 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. origin, beyond gently arched and oriented towards posterior wing margin; area between MA and MP with dense reticulated cross-veins in distal part, cross-veins at the base of area between CuA+M and CuP sigmoidal. CuP1a twice the length of CuA before their fusion. CuA and CuP1a remaining united as CuA+CuP1a for a short and appreciable distance, first anterior branch of CuA+CuP1a reaching posterior margin distal of the end of ScP; Second anterior branch dichotomously ramified with 2 to 4 branches. The ‘handle’ subparallel to each cross-vein between CuP1b and posterior branch of CuA+CuP1a. CuP1b curved, CuP1b between its base and the ‘handle’ straight, the ‘handle’ subparallel to each crossvein between the ‘handle’ and CuP1a; CuP2 very strong, sharply curved, clearly Z-shaped, cross-veins between CuP2 and CuP1b gently curved but not strong sigmoidal. 1A strong and similar to CuP2 in shape, convergent to CuP2 at about their midlength; 2A apparently S-shaped. The coloration spread over the whole tegmen and formed irregular bands. Material: Holotype, male tegmen, CNU-ORT-NN2008018; paratypes, 8, male tegmen: CNU-ORT- NN2008016/19/38/40/41PC/44PC/45/57; 13, female tegmen: CNU-ORT-NN2008017/34/43/47/48PC/51/53- 55/58-61. Etymology: The specific epithet derives from Latin ‘peregrinus’, for its special ‘handle’ vein. Horizon and locality: Jiulongshan Formation, Middle Jurassic, Daohugou Village, Ningcheng County, Inner Mongolia, China. Remarks: This species differs from the other species in the following features on the forewing: coloration formed irregular bands and spread over whole tegmen; first anterior branch of CuA+CuP1a simple and second anterior branch dichotomously ramified; and cross-veins between CuP2 and CuP1b gently curved but not sigmoidal.

Discussion

Based on the characters of revised diagnosis, the attribution of S. longus to the genus Sigmaboilus Fang, Zhang & Wang, 2007 is uncertain for the following reasons: (1) ScA extremely long almost reaching the same level of second branch of RA, (2) ScA is not distinctly undulate, (3) Cross-veins at the base of area between CuA+M and CuP straight, and (4) Coloration of tegmen formed relatively regular and dark bands. Variability of wing venation in orthopterids and difference between the left and right wings in the same individual have rarely been discussed. The species S. peregrinus described here shows some variability. Its right forewing differs from the left forewing in the following characters: (1) RP branched beyond first branch of RA (beyond second branch in left forewing). (2) More ScP branches and fewer CuA+CuP1a branches. (3) The ‘handle’ forms a special shape. Some paratypes of S. peregrinus sp. nov. differ from holotype in the following characters: specimens (CNU-ORT-NN2008016/19/37/48PC) have a long basal free part of CuA, about twice the length of basal free part of M, unlike in the holotype specimen (Fig. 3–A); specimens (CNU-ORT-NN2008040/41PC), have a basal free part of CuA which is slightly shorter or nearly the same length as the basal free part of M; the fifth branch of RP in specimen CNU-ORT-NN2008058 and the third branch of RP in specimen CNU-ORT- NN2008060 are forked, in other specimens they are simple; as a whole, the branch number of the second anterior branch of CuA+CuP1a is different, ranging between 4-6. Similar differences can be attributed to intraspecific variations in extant Orthoptera, but in fossils, it is almost impossible to differentiate specific and intraspecific variation. Therefore, we here prefer to assign these specimens (male and female, CNU-ORT- NN2008016/17/19/34/38/40/41PC/43/44PC/45/47/48PC/51/53-55/57/58-61) to S. peregrinus sp. nov. Thus, an appreciation of the intraspecific variability of the venation is very vital for identifying fossil orthopterids, especially when only isolated or fragmented wings are available.

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FIGURE 3. Sigmaboilus peregrinus sp. nov., A−B. ♂, photograph of holotype, A. part: CNU-ORT-NN2008018P; B. counterpart: CNU-ORT-NN2008018C. C−D. photograph of paratype, C. ♀, CNU-ORT-NN2008034. D. ♂, CNU-ORT- NN2008041P.

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FIGURES 4–7. Sigmaboilus peregrinus sp. nov., 4–5. line drawing of holotype, ♂, CNU-ORT-NN2008018: 4. left forewing. 5. right forewing. 6−7. line drawing of paratype, 6.♀, CNU-ORT-NN2008034. 7. ♂, CNU-ORT-NN2008041P.

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FIGURE 8. Sigmaboilus peregrinus sp. nov., additional material: ♂, A. CNU-ORT-NN2008016. B. CNU-ORT- NN2008019. C. CNU-ORT-NN2008040. D. CNU-ORT-NN2008045. G. CNU-ORT-NN2008044P. ♀, E. CNU-ORT- NN2008043. F. CNU-ORT-NN2008048P. H. CNU-ORT-NN2008061. All scale bars = 5 mm.

Acknowledgements

We are grateful to Dr. Shih ChungKun (College of Life Science, Capital Normal University) and Dr. Thomas A. Hegna (Dept. of Geology and Geophysics, Yale University), for their improvement of our manuscript. This research is supported by the National Natural Science Foundation of China (No.30430100, 40872022), the Nature Science Foundation of Beijing (No.5082002) and Scientific Research Key Program and PHR Project of Beijing Municipal Commission of Education.

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