Zootaxa, Revision of genera Quadraticossus ...

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ZOOTAXA

Revision of genera Quadraticossus, Martynovocossus and Fletcheriana (Insecta, Hemiptera) from the Middle Jurassic of China with description of a new species YING WANG1, LIN WANG2 & DONG REN1, 3 1

College of Life Science, Capital Normal University, Beijing 100048, China Beijing No.57 Junior High & High School, Beijing 100038, China 3 Corresponding author. E-mail: [email protected] 2

Abstract Four species, belonging to three known genera of Palaeontinidae (Hemiptera), are described from the Middle Jurassic of Daohugou Village, Inner Mongolia, China. They are Quadraticossus eumorphus sp. nov., Q. fangi Wang & Ren, 2007, Martynovocossus punctulosus (Wang & Ren, 2006) Wang & Zhang, 2008 and Fletcheriana colorata Wang et al, 2006. Based on these materials, a new species is erected; line drawing of hind wing of Q. fangi is reconstructed and the diagnoses of the genera Quadraticossus Wang & Ren, 2007, Martynovocossus (Martynov, 1931) Wang & Zhang, 2008 and Fletcheriana Evans, 1956 are revised. Key words: morphology, taxonomy, palaeontology, Cicadomorpha, Jurassic

Introduction Palaeontinidae is an extinct family of Hemiptera, which existed from the Triassic to the Cretaceous (Carpenter, 1992; Menon et al., 2005). So far, 53 species within 22 genera have been described based on fossils collected from the Mosozoic of China (Hong, 1982, 1983, 1984, 1986; Lin, 1992; Ren et al., 1995, 1998; Zhang, 1997; Wang B et al., 2006a–d, 2008; Wang Y & Ren, 2006, 2007a, b; Wang Y et al., 2007a, b). Many well-preserved insect fossils have been collected from the Middle Jurassic Jiulongshan Formation at Daohugou Village, Ningcheng County, Inner Mongolia in China (Tan & Ren, 2006; Yao et al., 2006; Liu et al., 2007). Based on new specimens collected from this locality and their unique characters, the diagnoses of Quadraticossus Wang and Ren, 2007, Martynovocossus (Martynov, 1931) Wang & Zhang, 2008 and Fletcheriana Evans, 1956 are revised.

Material and methods The specimens were examined by dissecting microscope (Type: LEICA MZ12.5) and illustrated with the aid of a drawing tube. The specimens studied here are deposited in the Key Lab of Insect Evolution and Environmental Change, College of Life Science, Capital Normal University, Beijing, China. The wing venation nomenclature used in this paper is based on the interpretation of Becker-Migdisova (1949). Systematic palaeontology

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Accepted by C. Dietric: 23 Jul. 2008; published: 22 Aug. 2008

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Order Hemiptera Linnaeus, 1758 Suborder Cicadomorpha Evans, 1946 Family Palaeontinidae Handlirsch, 1906 = Cicadomorphidae Evans, 1956 Genus Quadraticossus Wang & Ren, 2007 2007 Quadraticossus Wang Y & Ren, p. 41.

Type species. Quadraticossus fangi Wang & Ren, 2007 Revised diagnosis. Forewing triangular. Sc separated from R at wing base, and then fused with R, extending to near indentation and terminating on R1. Rs simple. M 4-branched, M4 sharply flexed after arising from M3+4. CuA connected with M4 by horizontal crossvein. Nodal line dividing discal cell into two parts, 2nd discal cell nearly rectangular. Hind wing small, with distinct costal indentation. Sc parallel to R at base, fused with R for short distance, terminating near indentation. M 3-branched, stem of M1+2 long, and M3+4 unbranched. Crossvein m3+4-cua short. Anal vein single. Species included. Q. fangi Wang & Ren, 2007; Q. longicaulis Wang & Ren, 2007; Q. eumorphus sp. nov. Remarks. This genus established by Wang Y & Ren (2007a) was based on a body with a pair of forewings and hind wings and 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: Sc and R separated at base, fused at point of origin of Rs, then Sc coalesced with R1+2 for short interval before extending to costal margin at indentation.

Quadraticossus eumorphus sp. nov. Figs.1, 2 Diagnosis. Hind wing with Sc extended to costal margin shortly beyond point of fusion with R, and Rs confluent with M1 for short interval. Description. Hind wing broad. Sc fused with R at branching point of R1+2 and Rs, then extended to costal margin. R1 arising from R1+2 at level of indentation, and nearly parallel with Sc but reaching costal margin slightly acclivitous. M 3-branched. M1 slightly extended upward after arising from M1+2 and downward after connecting with Rs for short interval. M3+4 unbranched. CuA gently bifurcating before level of indentation. A short crossvein m3+4-cua connected with M3+4 at level of M1+2 forks. CuP simple. Anal vein single. Hind wing 20 mm long; 12 mm wide. Etymology. The species epithet is a Latin eumorphus, beautiful, referring to the complete and beautiful features of the specimen. Material examined. Holotype: A single hind wing, No. CNU-H-NN2007006. Horizon and locality. Jiulongshan Formation, Middle Jurassic, Daohugou Village, Ningcheng County, Inner Mongolia in China. Remarks. This species is assigned to Quadraticossus Wang & Ren, 2007 by the main characters of hind wing, such as hind wing small, stem of M1+2 long, M3+4 unbranched and anal vein single. The new species can be distinguished from the type species in following features on the hind wing: Sc extended to costal margin soon after fused with R at one point, and Rs coalesced with M 1 for short interval.

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FIGURE 1. Quadraticossus eumorphus sp. nov., photograph of holotype, No. CNU-H-NN2007006.

FIGURE 2. Quadraticossus eumorphus sp. nov., line drawing of holotype, No. CNU-H-NN2007006.

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Quadraticossus fangi Wang & Ren, 2007 (Fig. 3) 2007 Quadraticossus Wang Y & Ren, p. 41. figure 1 (A–F)

Remarks. This species was erected based on both complete forewings and hind wings (Wang & Ren, 2007a). After re-examining part and counterpart of the holotype, we have reconstructed the hind wing of Q. fangi (Fig. 3). Sc and R are similar to those of Q. eumorphus: Sc parallel with R at base, fused with R at point of origin of Rs, then confluent with R1+2 for short interval, then extended to costal margin at indentation.

FIGURE 3. Quadraticossus fangi Wang and Ren, 2007, reconstruction based on hind wing of holotype, No, CNU-HNN2006093-1.

Genus Martynovocossus Wang & Zhang, 2008 1931 Pseudocossus Martynov; Martynov, p. 93–122. 1965 Pseudocossus Martynov; Becker-Migdisova and Wootton, p. 69–79. 1982 Pseudocossus Martynov; Kolosnitsyna, p. 13–25. 1992 Pseudocossus Martynov; Carpenter, p. 214–217. 1997 Pseudocossus Martynov ; Zhang, p. 312–323. 2006 Pseudocossus Martynov ; Wang Y et al., p. 289–293. 2008 Martynovocossus Wang & Zhang.; Wang B et al., p. 1–18.

Type species. Martynovocossus zemcuznicovi (Martynov, 1931) Wang & Zhang, 2008. Species included. Eight species: M. zemcuznicovi (Martynov, 1931) Wang & Zhang, 2008; M. turgaiensis



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(Becker-Migdisova & Wootton, 1965) Wang & Zhang, 2008 ; M. strenus (Zhang, 1997) Wang & Zhang, 2008; M. punctulosus (Wang & Ren, 2006) Wang & Zhang, 2008; M. bellus (Wang & Ren, 2006) Wang & Zhang, 2008; M. ancylivenius (Wang & Ren, 2006) Wang & Zhang, 2008; M. cheni Wang & Zhang, 2008; M. decorus Wang & Zhang, 2008. Revised diagnosis. Hind wing length 20-35mm; costal area wide; Sc terminating at costal margin slightly basad of indentation; Rs fused with M1 for short distance; M with four branches; M3+4 connected with CuA by short crossvein m-cua; CuA arising basally and bifurcating into CuA1 and CuA2; A with 2 branches, and A2 terminating at anal margin near wing base.

Martynovocossus punctulosus (Wang & Ren, 2006) Wang & Zhang, 2008 (Figs. 4, 5) 2006 Pseudocossus Martynov ; Wang Y et al., p. 289–293. figures 1, 3 2008 Martynovocossus Wang & Zhang.; Wang B et al., p. 1–18. figures 2–4

Revised diagnose. Hind wing with 2 anal veins, and A2 arising from A1+2 from wing base. Description. Hind wing ovoid, costal margin with distinct indentation. Sc parallel with R at base, fused with R beyond R1+2 origins, then separating and extending to costal margin. R1 arising from R1+2 slightly beyond indentation, then extended to costal margin. M 4-branched. Stem of M 2+3+4 short. M1 fused with Rs for short interval. M3+4 bifurcated into M3 and M4 at the same level of Rs fused with M1. CuA gently bifurcated into CuA1 and CuA2 before the point of M3+4 bifurcating. There is a short crossvein m3+4-cua. CuP simple. Anal vein with two branches. A2 arising from A1+2 at basal and terminating near the wing base. Hind wing 31 mm long; 19 mm wide. Material examined. A single hind wing, No.: CNU-H-NN2007005. Horizon and locality. Jiulongshan Formation, Middle Jurassic, Daohugou Village, Ningcheng County, Inner Mongolia in China. Remarks. This single hind wing has the color pattern and venation of other specimens of this species. The fossil examined here has the anal area complete, and the anal vein can be seen clearly.

FIGURE 4. Martynovocossus punctulosus (Wang & Ren, 2006) Wang & Zhang, 2008, photograph of No. CNU-HNN2007005.


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FIGURE 5. Martynovocossus punctulosus (Wang & Ren, 2006) Wang & Zhang, 2008, line drawing of No. CNU-HNN2007005.

Fletcheriana Evans, 1956 1956 Fletcheriana Evans, p. 224. 1976 Fletcheriana Evans; Riek, p. 809. 1997 Fletcheriana Evans; Zhang, p. 312. 2006 Fletcheriana Evans; Wang B et al., p. 115.

Type species. Fletcheriana triassica Evans, 1956 Revised diagnosis. Forewing with costal area broad; Sc lying alongside R basally, with several veinlets; Rs arising from R distant from wing base. Hind wing small, with distinct indentation; Sc nearly parallel to R from base, fusing with R1+2 beyond its origin from R and terminating at costal margin independently; Rs connected with M1 by crossvein r-m; M1 arising from M near or at origin of M3+4; short crossvein m3+4-cua present; anal vein with 2 branches. Species included. Five species: F. triassica Evans, 1956; F. magna Riek, 1976; F. jurassica Zhang, 1997; F. colorata Wang et al, 2006; F. minuta Wang et al, 2006. Remarks. This genus was erected by Evans (1956) based on incomplete forewings and hind wings from the Triassic of Brookvale Quarry, Australia. Recently, other species belonging to this genus from the Triassic of South Africa, and Jurassic of Xinjiang and Inner Mongolia, China were reported (Riek, 1976; Zhang, 1997; Wang B et al., 2006a). Among them, three species were described based on hind wings with the anal area REVISION OF THREE MESOZOIC PALAEONTINID GENERA


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missing. The fossils described here have a complete anal area, which enables us to supplement the generic diagnosis.

Fletcheriana colorata Wang, Zhang et Fang , 2006 (Figs.6, 7) 2006 Fletcheriana Evans; Wang B et al., p. 115.

Revised diagnosis. Hind wing rounded; Sc confluent with R for short distance and ending in costal margin before indentation; R1 dividing from R1+2 at the same level of indentation; M1 separating from M basad of M3+4 separation from M2; M3+4 connected with CuA by crossvein m-cua; A with 2 branches from base. Revised description. Hind wing rounded. Sc confluent with R1+2 after the latter arises from R, and for a short interval, terminating in costal margin before indentation. R1 arising from R1+2 at the same level of indentation, ending in costal margin just distal of indentation. Rs curved downwards when connected with M1 by short crossvein r-m at same level of indentation, and ending in apex of wing. M1 arising from M just basad of stem M3+4 separation from M2. M1 curved anteriorly at junction with r-m. M3+4 branching into M3 and M4 slightly beyond origin of R1+2 but before indentation. CuA bifurcated before M3+4 ramifications. CuP slightly arched. Anal vein slightly curved. Hind wing 26 mm long, 17 mm wide. Material examined. A single hind wing, with costal area missing, No. CNU-H-NN2006031.

FIGURE 6. Fletcheriana colorata Wang, Zhang et Fang, 2006, photograph of No. CNU-H-NN2006031.


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Horizon and locality. Jiulongshan Formation, Middle Jurassic, Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China.

FIGURE 7. Fletcheriana colorata Wang, Zhang et Fang, 2006, line drawing of No. CNU-H-NN2006031.

Acknowledgements We sincerely thank Dr. Shih ChungKun (College of Life Sciences, Capital Normal University) for his improvement of our manuscript. This research was supported by grants from the National Nature Science Foundation of China (No. 30430100) and the PHR (IHLB) Project of Beijing Municipal Commission of Education.

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