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Tettigarctidae (Insecta: Hemiptera: Cicadoidea) from the Middle Jurassic of Inner Mongolia, China
Geobios 42 (2009) 243–253
Original article
Tettigarctidae (Insecta: Hemiptera: Cicadoidea) from the Middle Jurassic of Inner Mongolia, China§ Tettigarctidae (Insecta : Hemiptera : Cicadoidea) du Jurassique moyen de Mongolie inte´rieure, Chine Bo Wang *, Haichun Zhang State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Road, 210008 Nanjing, Jiangsu, China Received 4 May 2008; accepted 23 September 2008 Available online 21 November 2008
Abstract Three new species of Tettigarctidae from the Middle Jurassic of Daohugou, China are described: Shuraboprosbole daohugouensis nov. sp., S. minuta nov. sp., and S. media nov. sp. The revised diagnostic characters of Shuraboprosbole Becker-Migdisova are provided based on wellpreserved whole-bodied fossil tettigarctids. So far, only four definitive species of Tettigarctidae are recorded from the Mesozoic of China. The pronotum concealing major of mesonotum seems to be an autapomorphy of Tettigarctinae. The ‘‘cicadellid-like tarsi’’ found in Cicadellidae and Brazilian Tettigarctidae are probably a parallel evolution. # 2008 Elsevier Masson SAS. All rights reserved. Résumé Nous décrivons ici trois nouvelles espèces de Tettigarctidae du Jurassique moyen de Daohugou, Chine : Shuraboprosbole daohugouensis nov. sp., S. minuta nov. sp. et S. media nov. sp. Les caractères diagnostiques de Shuraboprosbole Becker-Migdisova sont révisés à partir de spécimens fossiles complets bien préservés. À ce jour, seulement quatre espèces de Tettigarctidae sont connues dans le Mésozoïque de Chine. Le pronotum masquant principal du mesonotum semble être une autapomorphie des Tettigarctinae. La morphologie du tarse, proche de celle des Cicadellidae, a probablement évoluée de façon parallèle chez les Tettigarctidae brésiliens. # 2008 Elsevier Masson SAS. Tous droits réservés. Keywords: Tettigarctidae; Hemiptera; Taxonomy; New species; Middle Jurassic; Daohugou Mots clés : Tettigarctidae ; Hemiptera ; Taxonomie ; Espèces nouvelles ; Jurassique moyen ; Daohugou
1. Introduction The Tettigarctidae or hairy cicadas are a relict group of cicadas, consisting of two extant species found only in high altitude habitats of southeastern Australia (Moulds, 1990). They differ from true cicadas in features of the wing venation, head structure and in the presence of rudimentary tymbals in both sexes (Evans, 1941; Moulds, 2005). Unlike members of §
Corresponding editor: Gilles Escarguel. * Corresponding author. E-mail address: [email protected] (B. Wang).
the Cicadidae, the adults of living Tettigarctidae are nocturnal, hiding away from strong light during daylight hours (Evans, 1941). The fossil record of Tettigarctidae is relatively rich with more than 20 species within about 15 genera from the Mesozoic of Eurasia, Australia, Africa and South America (Becker-Migdisova, 1949; Shcherbakov, 1988; Hamilton, 1990; Nel, 1996; Nel et al., 1998; Menon, 2005; Moulds, 2005). The Late Triassic tettigarctids from Australia and South Africa most probably belong to other groups (Tillyard, 1919; Evans, 1956; Riek, 1976; Martins-Neto et al., 2003; Jell, 2004). The oldest tettigarctids were probably recorded from the latest Rhaetian of England (Whalley, 1983; Shcherbakov
0016-6995/$ – see front matter # 2008 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.geobios.2008.09.003
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and Popov, 2002). Recently, Cicadoprosbolidae has been considered as a synonym of Tettigarctidae within Cicadoidea (Shcherbakov, 1996; Nel et al., 1998; Menon, 2005). Therefore, the Tettigarctidae contains two subfamilies: Tettigarctinae Distant, 1905 and Cicadoprosbolinae BeckerMigdisova, 1947. Becker-Migdisova (1947) firstly separated Cicadoprosbolinae from the Tettigarctinae by forewing characters (see also Moulds, 2005: p. 426). So far, the unambiguous autapomorphies for Cicadoprosbolinae are based on the forewing (such as forewing elliptical with a broad costal area and clavus). However, some autapomorphies suggested by Becker-Migdisova (1947) also require revisions based on new material. Unfortunately, the revised diagnosis of Cicadoprosbolinae cannot be finished until revisions of all genera. Cicadoprosbolinae is believed to be a transitional unit between Prosbolidae and Cicadidae (Becker-Migdisova, 1947; Moulds, 2005). This opinion could affect the phylogeny of Cicadoidea. However, it is premature for discussing the phylogeny of Cicadoidea at the present time, because the limits of Tettigarctidae and Cicadoprosbolinae require further revisions and some available fossils require detailed, formal descriptions (such as Russian specimens). So far, there is no consensus on the relationships of four superfamilies (Cicadoidea, Cercopoidea, Membracoidea and Myerslopioidea) of Cicadomorpha (Dietrich et al., 2001; Szwedo et al., 2004; Cryan, 2005; Dietrich, 2005). The Mesozoic Tettigarctidae is therefore critical for examining the relationships among them. However, the definitive whole-bodied fossil tettigarctids were just previously reported from the Upper Jurassic of Karatau and Lower Cretaceous of Brazil (Hamilton, 1990; Shcherbakov and Popov, 2002; Menon, 2005), and furthermore most species of fossil Tettigarctidae are based on wing fragments. The Middle Jurassic Daohugou tettigarctids, including three new species within one genus, are described herein. They provide some information about the evolutionary processes of this important hemipteran family. 2. Material and methods Thirty-six specimens described herein were collected from the Middle Jurassic Daohugou deposits (418180 N, 1198130 E) in Ningcheng County, Chifeng City, Inner Mongolia of China. The fossils are preserved as impressions on the surface of grey tuffaceous siltstones. Specimens were examined dry and under alcohol, using a Nikon SMZ1000 stereomicroscope and drawings were made with the aid of a camera lucida. The photographs were prepared using a Nikon D100 digital camera, and the line drawings were readjusted on photographs using image-editing software (CorelDraw 13.0 and Adobe Photoshop CS). All specimens are deposited in the Nanjing Institute of Geology and Palaeontology (NIGP), Chinese Academy of Sciences. There is no consensus on the interpretation of vein nomenclature in Cicadomorpha. Correspondences between vein nomenclatures were provided by Nel et al. (1998). We tentatively followed the traditional terminologies with slight modifications (Wang et al., 2006).
3. Systematic palaeontology Order HEMIPTERA Linnaeus, 1758 Superfamily CICADOIDEA Latreille, 1802 Family TETTIGARCTIDAE Distant, 1905 Subfamily CICADOPROSBOLINAE Becker-Migdisova, 1947 Remarks: The generic classification of Mesozoic tettigarctids is mainly based on the forewing venation (Whalley, 1983; Nel, 1996). Only two isolated hindwings have been recorded from the Lower Cretaceous of Mongolia and China, respectively (Shcherbakov, 1986; Zhang, 1993). Some genera within Cicadoprosbolinae are known from only forewing fragments or body, such as Turutanovia and Liassocicada. Some of them have no clear diagnostic characters preserved, and cannot be compared with those of other genera. Therefore, any more refined study of relationships among these genera must wait until a recent taxonomic review of these taxa is available (Shcherbakov, in prep.). We follow the previously held concept for the Tettigarctidae and Cicadoprosbolinae (Moulds, 2005). Genus Shuraboprosbole Becker-Migdisova, 1949 Type species: Shuraboprosbole plachutai Becker-Migdisova, 1949; by original designation. Type horizon and locality: Lower–Middle Jurassic Sogul Formation; Sai-Sagul (Shurab 3), Tajikistan. Diagnosis: Head with inflated, declivous crown. Rostrum long, extending to or beyond hind coxae. Pronotum greatly expanded, with posterior half transversely rugose. Median length of pronotum three times that of vertex. Mesonotum partly exposed. Legs densely setose. Forewing elongate, with nodus at about basal 0.6 wing length; costal area wide; stem R + M branching at about basal 0.2 wing length; vein M fourbranched; vein R half or one third as long as stem R + M; vein RA two-branched; vein M fused with vein CuA for a distance; vein CuP ending at about midpoint of wing. Hindwing unknown. Ovipositor moderate, adpressed to pygofer. Included species: Four species: the type species S. plachutai, S. daohugouensis nov. sp., S. minuta nov. sp., and S. media nov. sp. Remarks: The diagnostic characters of body are based on S. daohugouensis nov. sp. and S. minuta nov. sp. This genus distinctly differs from others in possessing vein R much shorter than stem R + M, vein RA two-branched and vein M fused with vein CuA for a distance. Shuraboprosbole is similar to Turutanovia to some extent; however, the extensive comparison is impossible because the latter is just a broken forewing with distal part preserved. Key to species of the genus Shuraboprosbole: forewing: vein R bifurcates at the same level of vein M + CuA forking; vein A1 long: S. plachutai BeckerMigdisova, 1949; forewing: vein R bifurcates distal of vein M + CuA forking; vein A1 short and terminating at inner margin basal of stem R + M bifurcating.
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forewing: small (14–19 mm); costal margin convex at about middle of wing: S. minuta nov. sp.; forewing: not as above. forewing: large (29–35 mm); costal margin convex at about basal 0.3 wing length; vein RA curved posteriorly basal of nodal line; vein M bifurcates at basal 0.47 wing length: S. daohugouensis nov. sp.; forewing: medium-sized (about 25 mm); costal margin convex at basal 0.25 wing length; vein RP nearly straight basal of nodal line; vein M bifurcates at basal 0.39 wing length: S. media nov. sp. Shuraboprosbole daohugouensis nov. sp. Figs. 1–3(A, B) Holotype: NIGP149372a, b, female, almost complete adult in lateral aspect with two forewings preserved at top of the body. Paratypes: NIGP149373a, b, adult in ventral aspect with a forewing preserved on the right side of the body. Body obscured. Forewing with apex missing and basal part slightly destroyed. NIGP149374, female, almost complete adult in lateral aspect with two forewings preserved at top of the body. NIGP149375, incomplete adult in lateral aspect with forewings preserved over body; body obscured. Type locality: Daohugou Village, Chifeng City, Inner Mongolia, China. Stratum typicum: Middle Jurassic. Etymology: Specific epithet is after the type locality, Daohugou Village. Diagnosis: Forewing large, length 29–35 mm; costal margin convex at about basal 0.3 wing length; vein R bifurcating distal of vein M + CuA forking; vein RP curved posteriorly basal of nodal line; vein A1 short, terminating at inner margin basal of stem R + M bifurcation; wing membrane without distinct color patterns. Differs from S. plachutai in having a larger forewing with vein R bifurcating distal of vein M + CuA dividing, vein A1 shorter and wing membrane without distinct color patterns. Description: Holotype NIGP149372. Body dark, length about 26 mm. Ocelli invisible. Eyes large, rounded. Antennae slender and short, with four segments visible; scape thick and short; pedicel as long as first flagellomere; flagellum aristiform. Postclypeus strongly convex, with distinct transverse grooves (muscle attachments). Pronotum slightly wider posteriorly; anterior margin almost straight, lateral margins diverging posteriad, posterior margin distinctly concave medially. Fore femur thick, about 0.7 times as long as fore tibia. Forewing elongate, length: 29.4 mm, width: 10.8 mm, length/width ratio about 2.7. Costal margin convex at about basal 0.3 wing length, and costal area large. Nodal incision distinct at basal 0.6 wing length. Vein ScP fused with vein R + M at basal 0.1 wing length, branching from vein ScP + R at nodal line (basal 0.58 wing length) and terminating at nodus. Stem R + M slightly curved, connected with vein CuA at its base by a long, thin crossvein r + m-cua; stem R + M forked at basal 0.19 wing length. Vein R branching into veins RA and RP at basal 0.27 wing length. Vein R about half as long as stem R + M. Branch RA curved posteriorly, recurved at nodal line,
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and then branching into veins RA1 and RA2 at basal 0.65 wing length. Branch RA2 connected with vein RP by crossvein r at basal 0.7 wing length. Branch RP curved posteriorly basal of nodal line, and connected with branch M1 by crossvein r-m slightly basal of crossvein ir. Vein M curved posteriorly and fused with vein CuA for a short distance at basal 0.21 wing length, and then branching into veins M1+2 and M3+4 at basal 0.47 wing length. Vein M1+2 branching into veins M1 and M2 at basal 0.63 wing length. Vein M1 strongly curved anteriorly at crossvein r-m. Vein M2 connected with M3 by crossvein m at the same level of crossvein r-m. Vein M3+4 branching into veins M3 and M4 slightly basal of bifurcation of vein M1+2. Vein M3 strongly curved anteriorly at crossvein im. Vein M4 connected with branch CuA1 by crossvein m-cua at basal 0.6 wing length. Cubital triangle (Ct) narrow. Stem Cu bifurcating into veins CuA and CuP at wing base. Branch CuA curved anteriorly at fusion with vein M, recurved at nodal line, and then branching into veins CuA1 and CuA2 at basal 0.52 wing length. Vein CuA1 curved anteriorly at crossvein m-cua. Vein CuA2 very short. Vein CuP straight, ending at about midpoint of wing, forming a long clavus. Vein Pcu slightly curved. Vein A1 weak, curved anteriorly and terminating at inner margin basal of stem R + M bifurcating. Nodal line traceable as a crease along vein ScP, crossing veins M1+2 and M3+4 and then fused with veins CuA and CuA2. Abdominal segments 5–7 of similar size. Ovipositor ensiform, upcurved, adpressed to pygofer, extended just beneath anal tube. Paratype NIGP149373. Body length about 30 mm. Rostrum long, extending to hind coxae. Forewing length as preserved: 32 mm (estimated complete length about 35 mm), width about 12 mm, length/width ratio about 3. Venation in general as in holotype. Paratype NIGP149374. Body length about 27 mm. Abdomen length about 15 mm, five segments visible. Ovipositor upcurved, length: 5 mm. Forewing length: 34.1 mm, width: 11.8 mm, length/width ratio: 2.9. Venation in general as in holotype. Crossvein ir comparatively longer. Vein A1 obscured. Marginal membrane wider than vein and distinct at posterior margin. Paratype NIGP149375. Body length 27 mm. Abdomen pointed posteriorly, length: 12 mm, five segments visible. Hind femur: 5 mm and hind tibia: 8 mm. Forewing length as preserved: 31 mm (estimated complete length about 35 mm), width: 12.6 mm, length/width ratio about 2.8. Venation in general as in holotype. Shuraboprosbole minuta nov. sp. Figs. 3(C, D), 4 and 5 Holotype: NIGP149376a, b, male, almost complete adult in lateral aspect with two forewings preserved at top of the body. Paratypes: NIGP149377, NIGP149378, NIGP149379a, b, NIGP149380a, b, all female and complete adult in lateral aspect with two forewings preserved at top of the body. Other material: NIGP149381, NIGP149382, NIGP149383, all male and incomplete adult in lateral aspect with two forewings preserved at top of the body. NIGP149384a, b,
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Fig. 1. Shuraboprosbole daohugouensis nov. sp. Holotype NIGP149372. A, B. Photograph and illustration (same scale). Scale bar represents 10 mm. C. Enlarged antenna. Scale bar represents 1 mm.
NIGP149385a, b, NIGP149386, NIGP149387, NIGP149388, NIGP149389, NIGP149390, NIGP149391, NIGP149392, NIGP149393, NIGP149394, all female and incomplete adult in lateral aspect with two forewings preserved at top of the body. NIGP149395, NIGP149396, NIGP149397, all female and strongly deformed. NIGP149398, NIGP149399, NIGP149400, NIGP149401, NIGP149402, sex unknown and incomplete adult in lateral aspect with two forewings preserved at top of the body. NIGP149403, sex unknown and specimen distorted. NIGP149404, NIGP149405, NIGP149406, all almost complete forewings. Etymology: Specific epithet refers to the small size of the adult. Type locality: Daohugou Village, Chifeng City, Inner Mongolia, China. Stratum typicum: Middle Jurassic.
Diagnosis: Forewing small, length 14–19 mm; costal margin convex at about middle of wing; vein R bifurcating distal of vein M + CuA dividing; vein RP almost straight basal of nodal line; vein A1 short, terminating at inner margin basal of stem R + M bifurcating; wing membrane gray with lighter or darker narrow areas along veins. Differs from S. plachutai in possessing a smaller forewing with costal margin convex at about middle of wing, vein R bifurcating distal of vein M + CuA dividing, vein A1 shorter and different colour patterns; from S. daohugouensis in having a smaller forewing with costal margin convex at about middle of wing and distinct colour patterns. Description: Holotype NIGP149376. Body dark. Ocelli invisible. Eyes large, rounded or ovoid, slightly emarginated behind antennae. Antennae slender, short and two thirds as long as fore tibia, with four segments visible; scape thick and short; pedicel slightly shorter than first flagellomere; flagellum
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Fig. 2. Shuraboprosbole daohugouensis nov. sp. Paratypes. A, B. Photograph and illustration of the paratype NIGP149375 (same scale). C. Photograph of the paratype NIGP149374. D. Photograph of the forewing in paratype NIGP149374. Scale bars represent 10 mm.
aristiform. Postclypeus strongly convex, with distinct transverse grooves. Rostrum extending beyond hind coxae, nearly to apex of hind femur. Pronotum slightly wider posteriorly; anterior margin almost straight, lateral margins diverging posteriad, posterior margin distinctly concave medially. Mesonotum partly covered by wings, with distinctly separated, convex prescutum. Fore femur 1.2 times as long as hind femur, distinctly thicker than hind femur, unarmed. Fore tibia slightly longer than fore femur. Fore tarsus 0.4 times as long as fore tibia. Mid tibia 1.5 times as long as mid femur. Mid tarsus 0.3 times as long as mid tibia. Hind tibia slender, twice as long as femur, with two welldeveloped lateral spines on outer edge of tibia (longer than diameter of tibia, one at its 1/3, the other at about its 2/3) and two (or three?) apical teeth set on enlarged sepal base. Hind tarsus 0.4 times as long as hind tibia; first two tarsomeres with apical teeth (probable two teeth in each tarsomere); basitarsomere slightly longer than mid tarsomere; mid tarsomere 0.6 times as long as apical tarsomere. Tarsal claws well developed. Arolium invisible. Forewing elongate, with costal margin convex at about middle of wing. Nodal incision distinct at basal 0.65 wing length. Vein ScP invisible in wing base, dividing from ScP + RA at nodal line and ending at nodus. Stem R + M branching at basal 0.21 wing length. Vein R branching into veins RA and RP at basal 0.33 wing length. Branch RA curved anteriorly, recurved at nodal line, and then branching into veins
RA1 and RA2 at basal 0.68 wing length. Branch RA2 connected with vein RP by crossvein ir at basal 0.75 wing length. Branch RP sinuous, connected with branch M1 by crossvein r-m slightly basal of crossvein r. Vein M curved posteriorly and fused with vein CuA for a short distance at basal 0.23 wing length, and then branching into veins M1+2 and M3+4 at basal 0.4 wing length. Vein M1+2 branching into veins M1 and M2 at basal 0.69 wing length. Vein M2 connected with M3 by crossvein im just basal of crossvein r-m. Vein M3+4 branching into veins M3 and M4 basal of bifurcation of vein M1+2. Vein M3 curved anteriorly at crossvein m. Vein M4 connected with branch CuA1 by crossvein m-cua at basal 0.63 wing length. Stem Cu bifurcating into veins CuA and CuP at wing base. Branch CuA curved anteriorly at fusion with vein M, recurved at nodal line, and branching into veins CuA1 and CuA2 at basal 0.52 wing length. Vein CuA1 curved anteriorly at crossvein mcua. Vein CuA2 very short. Vein CuP straight. Vein Pcu slightly curved. Vein A1 weak, curved anteriorly and terminating at inner margin basal of stem R + M bifurcating. Nodal line traceable as a crease along vein ScP, crossing veins M1+2 and M3+4 and then fused with veins CuA and CuA2. Marginal membrane wider than vein and distinct at posterior margin. Wing membrane gray, with very narrow white bands along veins. Paratype NIGP149377. Hind tibia slender, with two welldeveloped lateral spines on outer edge of tibia (longer than diameter of tibia, one at its 1/3, the other at about its 2/3).
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Fig. 3. Comparison of forewings of Shuraboprosbole. A. S. daohugouensis nov. sp., NIGP149375. B. S. daohugouensis nov. sp., NIGP149374. C. S. minuta nov. sp., NIGP149405. D. S. minuta nov. sp., NIGP149378. E. S. media nov. sp., NIGP149407. F. S. plachutai Becker-Migdisova, 1949, redrawn from Becker-Migdisova (1949). Wings aligned horizontally with apex to the right. All to scale; scale bar represents 5 mm.
Abdomen pointed posteriorly. Ovipositor ensiform, upcurved, adpressed to pygofer, extended just below anal tube. Paratype NIGP149378. Forewing with vein M branching slightly basal of nodal line. Wing membrane gray, with very narrow dark bands along veins. Paratype NIGP149380. Hind tibia with three prominent lateral spines. Three spines longer than diameter of tibia; two on outer edge of tibia, one at its 1/3, the other at about its 2/3; one on inner edge of tibia, at about its midlength. Forewing with vein M branching slightly basal of nodal line. Wing membrane gray, with very narrow white bands along veins. NIGP149405. Venation in general as in holotype. Wing membrane gray, with very narrow white bands along veins. Measurements (mm): Male (N = 4), range given for available specimens with mean in parentheses. Body length:
13.5–14.1 (N = 2); vertex length: 0.9; rostrum length: 4.7; pronotum length: 2.7–3.4 (N = 2); forewing length: 14.8–16.5 (15.1; N = 4), width: 4.5–5.3 (4.9; N = 4), length/width ratio: 3.0–3.3 (3.1; N = 4); hind tibia length: 4.0–4.6 (N = 2). Female (N = 18). Body length: 14.1–15.9 (14.8; N = 9); vertex length: 1.0–1.2 (1.1; N = 7); pronotum length: 2.9–3.4 (3.2; N = 7); forewing length: 16.0–18.9 (17.6; N = 15), width: 5.5–6.2 (5.8; N = 15), length/width ratio: 2.9–3.2 (3.0; N = 15). Other forewings (N = 7). Forewing length: 15.2–18.2 (16.9), width: 4.8–6.2 (5.6), length/width ratio: 2.9–3.2 (3.0). Remarks: The length/width ratio of forewing and location of vein M branching are different among specimens. Two different colour types are detectable, but neither could be associated with any particular wing venation pattern. Furthermore, the consistent dimorphism in wing venation and colour
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Fig. 4. Shuraboprosbole minuta sp. nov. Holotype NIGP149376. A, B. Photograph and illustration (same scale). Scale bar represents 5 mm. C. Enlarged antenna. D, E. Photograph and illustration of enlarged hind tarsi (same scale). Scale bars represent 0.5 mm.
patterns was not detectable between the two sexes. Some of these intraspecific variations also occur in some fossil and living Cicadomorpha, such as palaeontinids and leafhoppers (Larsen and Walter, 2007; Wang et al., 2008). Therefore, it is reasonable to place these specimens into a single species. Shuraboprosbole media nov. sp. Figs. 3(E) and 6 Holotype: NIGP149407. Almost complete forewing with clavus slightly damaged. Type locality: Daohugou Village, Chifeng City, Inner Mongolia, China. Stratum typicum: Middle Jurassic.
Etymology: Specific epithet refers to the medium size of the forewing. Diagnosis: Forewing medium-sized, length about 25 mm; costal margin convex at basal 0.25 wing length; vein R bifurcating distal of vein M + CuA dividing; vein RP almost straight basal of nodal line; wing membrane without distinct colour patterns. It is similar to S. plachutai in the forewing shape (both with costal margin convex at basal 0.25 wing length), but differs from the latter in having a forewing with vein A1 shorter and wing membrane without distinct colour patterns. It is different from S. daohugouensis in having a smaller forewing with costal margin convex at basal 0.25 wing length, vein RP nearly straight basal of nodal line and stem M
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Fig. 5. Shuraboprosbole minuta sp. nov. Photographs. A. NIGP149377. B. NIGP149380. C. NIGP149379. D. NIGP149378. E. NIGP149385. F. NIGP149405. Scale bars represent 5 mm.
shorter. It also differs from S. minuta in having a larger forewing with costal margin convex at basal 0.25 wing length and wing membrane without distinct colour patterns. Description: Holotype NIGP149407. Forewing elongate, length: 25.8 mm, width: 10.1 mm, length/width ratio: 2.6. Costal margin convex at basal: 0.25 wing length, and costal area wide. Nodal incision distinct at about basal 0.6 wing length. Stem R + M branching at basal 0.17 wing length. Vein R branching into veins RA and RP at basal 0.27 wing length. Vein R about one third as long as stem R + M. Branch RA curved anteriorly, recurved at nodal line, and then branching into veins RA1 and RA2 at basal 0.68 wing length. Branch RA2 connected with vein RP by crossvein ir at basal 0.75 wing length. Branch RP almost straight basal of nodal line, connected with branch M1 by crossvein r-m slightly basal of crossvein ir. Vein M curved posteriorly and fused with vein CuA for a short distance at basal 0.18 wing length, and then branching into veins M1+2 and M3+4 at basal 0.39 wing length. Vein M1+2 branching into veins M1 and M2 at basal 0.67 wing length. Vein M1 strongly curved anteriorly at crossvein r-m. Vein M2 connected with M3
by crossvein im at the same level of crossvein r-m. Vein M3+4 branching into veins M3 and M4 slightly basal of bifurcation of vein M1+2. Vein M3 slightly curved anteriorly at crossvein im. Vein M4 connected with branch CuA1 by crossvein m-cua at about basal 0.6 wing length (Ct) narrow. Stem Cu bifurcating into veins CuA and CuP at wing base. Branch CuA curved anteriorly at fusion with vein M, recurved at nodal line, and branching into veins CuA1 and CuA2 at basal 0.49 wing length. Vein CuA1 strongly curved anteriorly at crossvein m-cua. Vein CuA2 very short. Vein CuP slightly curved, ending at about midpoint of wing, forming a long clavus. Vein Pcu sinuous. Nodal line traceable as a crease along vein Sc, crossing veins M1+2 and M3+4 and then fused with veins CuA and CuA2. Wing membrane gray. Marginal membrane wider than vein and distinct at posterior margin. 4. Discussion Up to now, 14 species within nine genera from China were placed in Tettigarctidae and all of them are restricted to the
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Fig. 6. Shuraboprosbole media nov. sp. Holotype NIGP149407. A, B. Photograph and illustration (same scale). Scale bar represents 5 mm.
Mesozoic. Most of them, however, are problematic in their systematic position. Protabanus chaoyangensis Hong, 1982, based on a specimen from the Middle Jurassic of Liaoning, was firstly referred to Tabanoidea (Diptera). Hamilton (1992) and Zhang (1993) considered it as a Tettigarctidae, but Nel (1996) excluded it from Cicadoidea. Based on the general appearance and forewing venation (Hong, 1982: Text-Fig. 150), it probably belongs to Cicadomorpha. However, judging from the original photograph of the impression (Hong, 1982: Pl. 38, Fig. 1), its forewing is obscure and broken in the middle and distal part, and hence its systematic position requires the reexamination of the type specimen. Luanpingia Hong, 1983, with three known species originally placed in Tettigarctidae, has been already referred to Procercopidae (Hamilton, 1992; Ren, 1995). Four specimens, respectively from South China and Xinjiang, were ever placed in Tettigarctidae: Lacunisbole ligonis Lin, 1986 from the Upper Triassic of Qujiang, Guangdong; Quadrisbole vieta Lin, 1986 from the Upper Triassic of Liuyang, Hunan; Quadrisbole stenis Lin, 1986 from the Lower Jurassic of Liuyang, Hunan; and Kerjieprosbole stigmatis Lin, 1992 from the Upper Triassic of Toksun, Xinjiang. Nel (1996) presented a detailed discussion regarding these four specimens; all of them are based on forewing fragments and require a further study. Sinocicadia shandongensis Hong, 1990 was erected mainly on the basis of a forewing from the Lower Cretaceous Laiyang Formation of Laiyang, Shandong. Based on the original description and illustration, neither distinct nodal line nor nodus is present, and vein M forks near wing apex. Therefore, it cannot
be included in Tettigarctidae, but its systematic position requires the reexamination of the type specimen (Nel, 1996). Sinocicadoprosbole hunjianensis Hong and Chang, 1993, from the Upper Triassic Beishan Formation of Hunjiang, Jilin, is based on a forewing with distal part missing and anal area destroyed. Judging from the photograph and illustration of the type specimen, the characteristics including absent nodal line and nodal incision, vein RA branching basal of middle of wing, vein CuA fused with stem R + M and vein CuA branching at the same level of vein M branching, exclude this species from Tettigarctidae, but probably attribute it to Prosbolidae. Last, Involuta perrara Zhang, 1993, certainly belonging to Tettigarctidae, is based on a complete hindwing from the Lower Cretaceous Fengjiashan Formation of Shangxian, Shanxi, China. Therefore, only four definitive species of Tettigarctidae are recorded from the Mesozoic of China. Based on our collections and recognitions, the large body with a greatly inflated frontoclypeus and a long rostrum indicates that early tettigarctids are probably primarily arboreal and xylem feeding. Considering the absence of angiosperms in the Daohugou flora, early tettigarctids probably live on some gymnosperms. So far, however, only few plant fossils from Daohugou have been studied in detail. Furthermore, plant fossils recorded from Daohugou (including ferns, cycads, ginkgoes and conifers) are also common in some other coeval biota. Therefore, little is known about the association between plants and cicadoprosbolines based on fossils from Daohugou. The pronotum exceedingly large and concealing major of mesonotum
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is common in extant Tettigarctinae and has been regarded as a diagnostic character of Tettigarctidae (Moulds, 2005). The pronotum of Cicadoprosbolinae is large, but not expanded over mesonotum (Figs. 1, 4 and 5; Hamilton, 1990). Therefore, the pronotum concealing major of mesonotum seems to be an autapomorphy of Tettigarctinae. Jurassic Tettigarctidae possess an elliptical forewing with broad costal area and clavus; whereas extant Tettigarctidae have elliptical forewing with narrow costal area and reduced clavus. This evolutionary trend is similar to that of Mesozoic Palaeontinidae, and probably also suggests an improvement of flight ability (Wang et al., in press). The earliest whole-bodied tettigarctids from the Middle Jurassic of Daohugou are slightly different from those from the Lower Cretaceous of Brazil by the detailed structure of hind legs. Hamilton (1990) showed that pretarsal structures of Architettix (Cicadoidea) are similar to those of Cercopoidea. But he did not suggest that Cicadoidea are closely related to Cercopoidea, and also did not give the reason. Because of the lack of detailed pretarsal structures in Daohugou specimens, we cannot decide whether this similarity is a parallel evolution between Architettix and Cercopoidea. Hamilton pointed out that Architettix compacta Hamilton, 1990 possessed an elongate hind basitarsomere with two rows of plantar setae, and furthermore he considered structures of ‘‘cicadellid-like tarsi’’ as synapomorphies linking the Membracoidea and Cicadoidea (Hamilton, 1990, 1996). However, Middle Jurassic Tettigarctidae show common cicadoid hind tarsi, in which the basitarsomere is short and unarmed. Therefore, the ‘‘cicadellidlike tarsi’’ found in Cicadellidae and Brazilian Tettigarctidae are probably a parallel evolution. Although almost distributed worldwide in the Jurassic and Early Cretaceous, the Tettigarctidae are very rare as fossils from the Late Cretaceous and Cenozoic. The scarcity of these insects in Cenozoic biotas is not a result of taphonomical bias, because singing cicadas occur in some Cenozoic deposits (Cockerell, 1906; Zhang and Zhang, 1990; Grimaldi and Engel, 2005). The decline of Tettigarctidae is probably due to changes in the vegetation and competitive displacement with singing cicadas. Acknowledgements We thank Huang Diying for his donation of some specimens, and K.G. Andrew Hamilton and André Nel for providing their literatures. Many thanks go to two anonymous reviewers for reviewing the manuscript and providing some constructive suggestions, and Gilles Escarguel for his helpful editorial comments. This research has been financially supported by the Major Basic Research Projects (2006CB806400) of MST of China, and the National Natural Science Foundation of China (40523004, 40632010, 40872015). References Becker-Migdisova, E.E., 1947. Cicadoprosbole sogutensis gen. nov. sp. nov., a transitional form between the Permian Prosbolidae and the recent Cicadidae. Doklady Akademii Nauk SSSR 55, 445–448 (in Russian).
Becker-Migdisova, E.E., 1949. Mesozoic Homoptera of Central Asia. Trudy Paleontologicheskogo Instituta 22, 1–20 (in Russian). Cockerell, T.D.A., 1906. A fossil cicada from Florissant, Colorado. Bulletin of the American Museum of Natural History 22, 457–458. Cryan, J.R., 2005. Molecular phylogeny of Cicadomorpha (Insecta: Hemiptera: Cicadoidea, Cercopoidea and Membracoidea): adding evidence to the controversy. Systematic Entomology 30, 563–574. Dietrich, C.H., 2005. Keys to the families of Cicadomorpha and subfamilies and tribes of Cicadellidae (Hemiptera: Auchenorrhyncha). Florida Entomologist 88, 502–517. Dietrich, C.H., Rakitov, R.A., Holmes, J.L., Black, W.C., 2001. Phylogeny of the major lineages of Membracoidea (Insecta, Hemiptera, Cicadomorpha) based on 28S rDNA sequences. Molecular Phylogenetics and Evolution 18, 293–305. Distant, W.L., 1905. Rhynchotal notes XXXIII. Annals and Magazine of Natural History 7, 1622–1635. Evans, J.W., 1941. The morphology of Tettigarcta tomentosa White (Homoptera, Cicadidae). Papers and Proceedings of the Royal Society of Tasmania 1940, 35–49. Evans, J.W., 1956. Palaeozoic and Mesozoic Hemiptera (Insecta). Australian Journal of Zoology 4, 165–258. Grimaldi, D., Engel, M.S., 2005. Evolution of the insects. Cambridge University Press, Cambridge. Hamilton, K.G.A., 1990. Homoptera. In: Grimaldi, D., (Ed.), Insects from the Santana Formation, Lower Cretaceous of Brazil. Bulletin of the American Museum of Natural History 195, pp. 82–122. Hamilton, K.G.A., 1992. Lower Cretaceous Homoptera from the Koonwarra fossil bed in Australia, with a new Superfamily and synopsis of Mesozoic Homoptera. Annals of the Entomology Society of America 84, 423–430. Hamilton, K.G.A., 1996. Cretaceous Homoptera from Brazil: implications for classification. In: Schaefer, C.W. (Ed.), Studies on Hemipteran Phylogeny. Proceedings, Thomas Say Publications in Entomology. Entomological Society of America, Lanham, pp. 89–110. Hong, Y.C., 1982. Mesozoic Fossil Insects of Jiuquan Basin in Gansu Province. Geological Publishing House, Beijing (in Chinese). Hong, Y.C., 1983. Middle Jurassic Fossil Insects in North China. Geological Publishing House, Beijing (in Chinese with English summary). Hong, Y.C., 1990. V. Insect fossils of Laiyang Formation. In: The Regional Geological Surveying Team, Shandong Bureau of Geology and Mineral Resources (Ed.), The Stratigraphy and Palaeontology of Laiyang Basin, Shandong Province. Geological Publishing House, Beijing, pp. 44–189 (in Chinese with English abstract). Hong, Y.C., Chang, J.P., 1993. V. Insects. In: Mi Jiarong, Zhang Chuanbo (Eds.), Late Triassic Stratigraphy, Paleontology and Paleogeography of the Northern Part of the Circum-Pacific Belt. China. Science Press, Beijing, pp. 186–188 (in Chinese). Jell, P.A., 2004. The fossil insects of Australia. Memoirs of the Queensland Museum 50, 1–124. Larsen, M.L., Walter, G.H., 2007. Intraspecific variation within Orosius argentatus Evans (Hemiptera: Cicadellidae): colour polymorphisms, morphometric analyses and host associations. Australian Journal of Entomology 46, 207–216. Lin, Q.B., 1986. Early Mesozoic fossil insect from the south China. Palaeontologica Sinica (Series B, 21). Science Press, Beijing, 1–112 (in Chinese with English summary). Lin, Q.B., 1992. Late Triassic insect fauna from Toksun. Xinjiang Acta Palaeontologica Sinica 31, 313–335 (in Chinese with English abstract). Martins-Neto, R.G., Gallego, O.F., Melchor, R.N., 2003. The Triassic insect fauna from South America (Argentina, Brazil and Chile): a checklist (except Blattoptera and Coleoptera) and descriptions of new taxa. Acta Zoologica Cracoviensia 46 (Suppl. Fossil Insects), 229–256. Menon, F., 2005. New record of Tettigarctidae (Insecta, Hemiptera, Cicadoidea) from the Lower Cretaceous of Brazil. Zootaxa 1087, 53–58. Moulds, M.S., 1990. Australian Cicadas. New South Wales University Press, Kensington. Moulds, M.S., 2005. An appraisal of the higher classification of cicadas (Hemiptera: Cicadoidea) with special reference to the Australian fauna. Records of the Australian Museum 57, 375–446.
B. Wang, H. Zhang / Geobios 42 (2009) 243–253 Nel, A., 1996. Un Tettigarctidae fossile du Lias Européen (Cicadomorpha, Cicadoidea, Tettigarctidae). EPHE 9, 83–94. Nel, A., Zarbout, M., Barale, G., Philippe, M., 1998. Liassotettigarcta africana sp. nov. (Auchenorrhyncha: Cicadoidea: Tettigarctidae), the first Mesozoic insect from Tunisia. European Journal of Entomology 95, 593–598. Ren, D., 1995. Systematic Palaeontology. Fossil insects. In: Ren, D., Lu, L.W., Guo, Z.G., Ji, S.A. (Eds.), Faunae and stratigraphy of Jurassic– Cretaceous in Beijing and the adjacent areas. Seismic Publishing House, Beijing, pp. 47–120, (in Chinese with English abstract). Riek, E.F., 1976. A new collection of insects from the Upper Triassic of South Africa. Annals of the Natal Museum 22, 791–820. Shcherbakov, D.E., 1986. Cicadina (= Auchenorrhyncha). In: Rasnitsyn, A.P. (Ed.), Insects in the Early Cretaceous Ecosystems of West Mongolia. Transactions of the Joint Soviet–Mongolian Palaeontological Expedition, vol. 28. Nauka, Moscow, pp. 47–57 (in Russian). Shcherbakov, D.E., 1988. New cicadas (Cicadina) from the later Mesozoic of Transbaikalia. Paleontological Journal 4, 52–63. Shcherbakov, D.E., 1996. Origin and evolution of the Auchenorrhyncha as shown in the fossil record. In: Schaefer, C.W. (Ed.), Studies on Hemipteran Phylogeny. Proceedings, Thomas Say Publications in Entomology. Entomological Society of America, Lanham, pp. 31–45. Shcherbakov, D.E., Popov, Y.A., 2002. Superorder Cimicidea Laicharting, 1781 order Hemiptera Linné, 1758. The bugs, cicadas, plantlice, scale insects, etc. In: Rasnitsyn, A.P., Quicke, D.L.J. (Eds.), History of Insects. Kluwer Academic Publisher, Dordrecht, pp. 152–155.
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Szwedo, J., Bourgoin, T., Lefebvre, F., 2004. Fossils planthoppers (Hemiptera: Fulgoromorpha) of the world. An annotated catalogue with notes on Hemiptera classification. Studio 1, Warszawa. Tillyard, R.J., 1919. Mesozoic insects of Queensland No. 7. Hemiptera Homoptera; with a note on the phylogeny of the suborder. Proceedings of the Linnaean Society 44, 857–896. Wang, B., Zhang, H.C., Fang, Y., 2006. Some Jurassic Palaeontinidae (Insecta, Hemiptera) from Daohugou, Inner Mongolia, China. Palaeoworld 15, 115–125. Wang, B., Zhang, H.C., Fang, Y., Zhang, Y.T., 2008. A revision of Palaeontinidae (Insecta: Hemiptera: Cicadomorpha) from the Jurassic of China with descriptions of new taxa and new combinations. Geological Journal 43, 1–18. Wang, B., Zhang, H.C., Szwedo, J. Jurassic Palaeontinidae from China and the higher systematics of Palaeontinoidea (Insecta: Hemiptera: Cicadomorpha). Palaeontology (in press). Whalley, P.E.S., 1983. A survey of recent and fossil cicadas (Insecta, Hemiptera–Homoptera) in Britain. Bulletin of the British Museum of Natural History (Geology) 37, 139–147. Zhang, J.F., 1993. A contribution to the knowledge of insects from the late Mesozoic in southern Shaanxi and Henan provinces, China. Palaeoworld 3, 49–56 (in Chinese with English summary). Zhang, J.F., Zhang, X.Y., 1990. Fossil insects of cicada (Homoptera) and true bugs (Heteroptera) from Shanwang, Shandong. Acta Palaeontologica Sinica 29, 337–348.
Original article
Tettigarctidae (Insecta: Hemiptera: Cicadoidea) from the Middle Jurassic of Inner Mongolia, China§ Tettigarctidae (Insecta : Hemiptera : Cicadoidea) du Jurassique moyen de Mongolie inte´rieure, Chine Bo Wang *, Haichun Zhang State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Road, 210008 Nanjing, Jiangsu, China Received 4 May 2008; accepted 23 September 2008 Available online 21 November 2008
Abstract Three new species of Tettigarctidae from the Middle Jurassic of Daohugou, China are described: Shuraboprosbole daohugouensis nov. sp., S. minuta nov. sp., and S. media nov. sp. The revised diagnostic characters of Shuraboprosbole Becker-Migdisova are provided based on wellpreserved whole-bodied fossil tettigarctids. So far, only four definitive species of Tettigarctidae are recorded from the Mesozoic of China. The pronotum concealing major of mesonotum seems to be an autapomorphy of Tettigarctinae. The ‘‘cicadellid-like tarsi’’ found in Cicadellidae and Brazilian Tettigarctidae are probably a parallel evolution. # 2008 Elsevier Masson SAS. All rights reserved. Résumé Nous décrivons ici trois nouvelles espèces de Tettigarctidae du Jurassique moyen de Daohugou, Chine : Shuraboprosbole daohugouensis nov. sp., S. minuta nov. sp. et S. media nov. sp. Les caractères diagnostiques de Shuraboprosbole Becker-Migdisova sont révisés à partir de spécimens fossiles complets bien préservés. À ce jour, seulement quatre espèces de Tettigarctidae sont connues dans le Mésozoïque de Chine. Le pronotum masquant principal du mesonotum semble être une autapomorphie des Tettigarctinae. La morphologie du tarse, proche de celle des Cicadellidae, a probablement évoluée de façon parallèle chez les Tettigarctidae brésiliens. # 2008 Elsevier Masson SAS. Tous droits réservés. Keywords: Tettigarctidae; Hemiptera; Taxonomy; New species; Middle Jurassic; Daohugou Mots clés : Tettigarctidae ; Hemiptera ; Taxonomie ; Espèces nouvelles ; Jurassique moyen ; Daohugou
1. Introduction The Tettigarctidae or hairy cicadas are a relict group of cicadas, consisting of two extant species found only in high altitude habitats of southeastern Australia (Moulds, 1990). They differ from true cicadas in features of the wing venation, head structure and in the presence of rudimentary tymbals in both sexes (Evans, 1941; Moulds, 2005). Unlike members of §
Corresponding editor: Gilles Escarguel. * Corresponding author. E-mail address: [email protected] (B. Wang).
the Cicadidae, the adults of living Tettigarctidae are nocturnal, hiding away from strong light during daylight hours (Evans, 1941). The fossil record of Tettigarctidae is relatively rich with more than 20 species within about 15 genera from the Mesozoic of Eurasia, Australia, Africa and South America (Becker-Migdisova, 1949; Shcherbakov, 1988; Hamilton, 1990; Nel, 1996; Nel et al., 1998; Menon, 2005; Moulds, 2005). The Late Triassic tettigarctids from Australia and South Africa most probably belong to other groups (Tillyard, 1919; Evans, 1956; Riek, 1976; Martins-Neto et al., 2003; Jell, 2004). The oldest tettigarctids were probably recorded from the latest Rhaetian of England (Whalley, 1983; Shcherbakov
0016-6995/$ – see front matter # 2008 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.geobios.2008.09.003
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and Popov, 2002). Recently, Cicadoprosbolidae has been considered as a synonym of Tettigarctidae within Cicadoidea (Shcherbakov, 1996; Nel et al., 1998; Menon, 2005). Therefore, the Tettigarctidae contains two subfamilies: Tettigarctinae Distant, 1905 and Cicadoprosbolinae BeckerMigdisova, 1947. Becker-Migdisova (1947) firstly separated Cicadoprosbolinae from the Tettigarctinae by forewing characters (see also Moulds, 2005: p. 426). So far, the unambiguous autapomorphies for Cicadoprosbolinae are based on the forewing (such as forewing elliptical with a broad costal area and clavus). However, some autapomorphies suggested by Becker-Migdisova (1947) also require revisions based on new material. Unfortunately, the revised diagnosis of Cicadoprosbolinae cannot be finished until revisions of all genera. Cicadoprosbolinae is believed to be a transitional unit between Prosbolidae and Cicadidae (Becker-Migdisova, 1947; Moulds, 2005). This opinion could affect the phylogeny of Cicadoidea. However, it is premature for discussing the phylogeny of Cicadoidea at the present time, because the limits of Tettigarctidae and Cicadoprosbolinae require further revisions and some available fossils require detailed, formal descriptions (such as Russian specimens). So far, there is no consensus on the relationships of four superfamilies (Cicadoidea, Cercopoidea, Membracoidea and Myerslopioidea) of Cicadomorpha (Dietrich et al., 2001; Szwedo et al., 2004; Cryan, 2005; Dietrich, 2005). The Mesozoic Tettigarctidae is therefore critical for examining the relationships among them. However, the definitive whole-bodied fossil tettigarctids were just previously reported from the Upper Jurassic of Karatau and Lower Cretaceous of Brazil (Hamilton, 1990; Shcherbakov and Popov, 2002; Menon, 2005), and furthermore most species of fossil Tettigarctidae are based on wing fragments. The Middle Jurassic Daohugou tettigarctids, including three new species within one genus, are described herein. They provide some information about the evolutionary processes of this important hemipteran family. 2. Material and methods Thirty-six specimens described herein were collected from the Middle Jurassic Daohugou deposits (418180 N, 1198130 E) in Ningcheng County, Chifeng City, Inner Mongolia of China. The fossils are preserved as impressions on the surface of grey tuffaceous siltstones. Specimens were examined dry and under alcohol, using a Nikon SMZ1000 stereomicroscope and drawings were made with the aid of a camera lucida. The photographs were prepared using a Nikon D100 digital camera, and the line drawings were readjusted on photographs using image-editing software (CorelDraw 13.0 and Adobe Photoshop CS). All specimens are deposited in the Nanjing Institute of Geology and Palaeontology (NIGP), Chinese Academy of Sciences. There is no consensus on the interpretation of vein nomenclature in Cicadomorpha. Correspondences between vein nomenclatures were provided by Nel et al. (1998). We tentatively followed the traditional terminologies with slight modifications (Wang et al., 2006).
3. Systematic palaeontology Order HEMIPTERA Linnaeus, 1758 Superfamily CICADOIDEA Latreille, 1802 Family TETTIGARCTIDAE Distant, 1905 Subfamily CICADOPROSBOLINAE Becker-Migdisova, 1947 Remarks: The generic classification of Mesozoic tettigarctids is mainly based on the forewing venation (Whalley, 1983; Nel, 1996). Only two isolated hindwings have been recorded from the Lower Cretaceous of Mongolia and China, respectively (Shcherbakov, 1986; Zhang, 1993). Some genera within Cicadoprosbolinae are known from only forewing fragments or body, such as Turutanovia and Liassocicada. Some of them have no clear diagnostic characters preserved, and cannot be compared with those of other genera. Therefore, any more refined study of relationships among these genera must wait until a recent taxonomic review of these taxa is available (Shcherbakov, in prep.). We follow the previously held concept for the Tettigarctidae and Cicadoprosbolinae (Moulds, 2005). Genus Shuraboprosbole Becker-Migdisova, 1949 Type species: Shuraboprosbole plachutai Becker-Migdisova, 1949; by original designation. Type horizon and locality: Lower–Middle Jurassic Sogul Formation; Sai-Sagul (Shurab 3), Tajikistan. Diagnosis: Head with inflated, declivous crown. Rostrum long, extending to or beyond hind coxae. Pronotum greatly expanded, with posterior half transversely rugose. Median length of pronotum three times that of vertex. Mesonotum partly exposed. Legs densely setose. Forewing elongate, with nodus at about basal 0.6 wing length; costal area wide; stem R + M branching at about basal 0.2 wing length; vein M fourbranched; vein R half or one third as long as stem R + M; vein RA two-branched; vein M fused with vein CuA for a distance; vein CuP ending at about midpoint of wing. Hindwing unknown. Ovipositor moderate, adpressed to pygofer. Included species: Four species: the type species S. plachutai, S. daohugouensis nov. sp., S. minuta nov. sp., and S. media nov. sp. Remarks: The diagnostic characters of body are based on S. daohugouensis nov. sp. and S. minuta nov. sp. This genus distinctly differs from others in possessing vein R much shorter than stem R + M, vein RA two-branched and vein M fused with vein CuA for a distance. Shuraboprosbole is similar to Turutanovia to some extent; however, the extensive comparison is impossible because the latter is just a broken forewing with distal part preserved. Key to species of the genus Shuraboprosbole: forewing: vein R bifurcates at the same level of vein M + CuA forking; vein A1 long: S. plachutai BeckerMigdisova, 1949; forewing: vein R bifurcates distal of vein M + CuA forking; vein A1 short and terminating at inner margin basal of stem R + M bifurcating.
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forewing: small (14–19 mm); costal margin convex at about middle of wing: S. minuta nov. sp.; forewing: not as above. forewing: large (29–35 mm); costal margin convex at about basal 0.3 wing length; vein RA curved posteriorly basal of nodal line; vein M bifurcates at basal 0.47 wing length: S. daohugouensis nov. sp.; forewing: medium-sized (about 25 mm); costal margin convex at basal 0.25 wing length; vein RP nearly straight basal of nodal line; vein M bifurcates at basal 0.39 wing length: S. media nov. sp. Shuraboprosbole daohugouensis nov. sp. Figs. 1–3(A, B) Holotype: NIGP149372a, b, female, almost complete adult in lateral aspect with two forewings preserved at top of the body. Paratypes: NIGP149373a, b, adult in ventral aspect with a forewing preserved on the right side of the body. Body obscured. Forewing with apex missing and basal part slightly destroyed. NIGP149374, female, almost complete adult in lateral aspect with two forewings preserved at top of the body. NIGP149375, incomplete adult in lateral aspect with forewings preserved over body; body obscured. Type locality: Daohugou Village, Chifeng City, Inner Mongolia, China. Stratum typicum: Middle Jurassic. Etymology: Specific epithet is after the type locality, Daohugou Village. Diagnosis: Forewing large, length 29–35 mm; costal margin convex at about basal 0.3 wing length; vein R bifurcating distal of vein M + CuA forking; vein RP curved posteriorly basal of nodal line; vein A1 short, terminating at inner margin basal of stem R + M bifurcation; wing membrane without distinct color patterns. Differs from S. plachutai in having a larger forewing with vein R bifurcating distal of vein M + CuA dividing, vein A1 shorter and wing membrane without distinct color patterns. Description: Holotype NIGP149372. Body dark, length about 26 mm. Ocelli invisible. Eyes large, rounded. Antennae slender and short, with four segments visible; scape thick and short; pedicel as long as first flagellomere; flagellum aristiform. Postclypeus strongly convex, with distinct transverse grooves (muscle attachments). Pronotum slightly wider posteriorly; anterior margin almost straight, lateral margins diverging posteriad, posterior margin distinctly concave medially. Fore femur thick, about 0.7 times as long as fore tibia. Forewing elongate, length: 29.4 mm, width: 10.8 mm, length/width ratio about 2.7. Costal margin convex at about basal 0.3 wing length, and costal area large. Nodal incision distinct at basal 0.6 wing length. Vein ScP fused with vein R + M at basal 0.1 wing length, branching from vein ScP + R at nodal line (basal 0.58 wing length) and terminating at nodus. Stem R + M slightly curved, connected with vein CuA at its base by a long, thin crossvein r + m-cua; stem R + M forked at basal 0.19 wing length. Vein R branching into veins RA and RP at basal 0.27 wing length. Vein R about half as long as stem R + M. Branch RA curved posteriorly, recurved at nodal line,
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and then branching into veins RA1 and RA2 at basal 0.65 wing length. Branch RA2 connected with vein RP by crossvein r at basal 0.7 wing length. Branch RP curved posteriorly basal of nodal line, and connected with branch M1 by crossvein r-m slightly basal of crossvein ir. Vein M curved posteriorly and fused with vein CuA for a short distance at basal 0.21 wing length, and then branching into veins M1+2 and M3+4 at basal 0.47 wing length. Vein M1+2 branching into veins M1 and M2 at basal 0.63 wing length. Vein M1 strongly curved anteriorly at crossvein r-m. Vein M2 connected with M3 by crossvein m at the same level of crossvein r-m. Vein M3+4 branching into veins M3 and M4 slightly basal of bifurcation of vein M1+2. Vein M3 strongly curved anteriorly at crossvein im. Vein M4 connected with branch CuA1 by crossvein m-cua at basal 0.6 wing length. Cubital triangle (Ct) narrow. Stem Cu bifurcating into veins CuA and CuP at wing base. Branch CuA curved anteriorly at fusion with vein M, recurved at nodal line, and then branching into veins CuA1 and CuA2 at basal 0.52 wing length. Vein CuA1 curved anteriorly at crossvein m-cua. Vein CuA2 very short. Vein CuP straight, ending at about midpoint of wing, forming a long clavus. Vein Pcu slightly curved. Vein A1 weak, curved anteriorly and terminating at inner margin basal of stem R + M bifurcating. Nodal line traceable as a crease along vein ScP, crossing veins M1+2 and M3+4 and then fused with veins CuA and CuA2. Abdominal segments 5–7 of similar size. Ovipositor ensiform, upcurved, adpressed to pygofer, extended just beneath anal tube. Paratype NIGP149373. Body length about 30 mm. Rostrum long, extending to hind coxae. Forewing length as preserved: 32 mm (estimated complete length about 35 mm), width about 12 mm, length/width ratio about 3. Venation in general as in holotype. Paratype NIGP149374. Body length about 27 mm. Abdomen length about 15 mm, five segments visible. Ovipositor upcurved, length: 5 mm. Forewing length: 34.1 mm, width: 11.8 mm, length/width ratio: 2.9. Venation in general as in holotype. Crossvein ir comparatively longer. Vein A1 obscured. Marginal membrane wider than vein and distinct at posterior margin. Paratype NIGP149375. Body length 27 mm. Abdomen pointed posteriorly, length: 12 mm, five segments visible. Hind femur: 5 mm and hind tibia: 8 mm. Forewing length as preserved: 31 mm (estimated complete length about 35 mm), width: 12.6 mm, length/width ratio about 2.8. Venation in general as in holotype. Shuraboprosbole minuta nov. sp. Figs. 3(C, D), 4 and 5 Holotype: NIGP149376a, b, male, almost complete adult in lateral aspect with two forewings preserved at top of the body. Paratypes: NIGP149377, NIGP149378, NIGP149379a, b, NIGP149380a, b, all female and complete adult in lateral aspect with two forewings preserved at top of the body. Other material: NIGP149381, NIGP149382, NIGP149383, all male and incomplete adult in lateral aspect with two forewings preserved at top of the body. NIGP149384a, b,
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Fig. 1. Shuraboprosbole daohugouensis nov. sp. Holotype NIGP149372. A, B. Photograph and illustration (same scale). Scale bar represents 10 mm. C. Enlarged antenna. Scale bar represents 1 mm.
NIGP149385a, b, NIGP149386, NIGP149387, NIGP149388, NIGP149389, NIGP149390, NIGP149391, NIGP149392, NIGP149393, NIGP149394, all female and incomplete adult in lateral aspect with two forewings preserved at top of the body. NIGP149395, NIGP149396, NIGP149397, all female and strongly deformed. NIGP149398, NIGP149399, NIGP149400, NIGP149401, NIGP149402, sex unknown and incomplete adult in lateral aspect with two forewings preserved at top of the body. NIGP149403, sex unknown and specimen distorted. NIGP149404, NIGP149405, NIGP149406, all almost complete forewings. Etymology: Specific epithet refers to the small size of the adult. Type locality: Daohugou Village, Chifeng City, Inner Mongolia, China. Stratum typicum: Middle Jurassic.
Diagnosis: Forewing small, length 14–19 mm; costal margin convex at about middle of wing; vein R bifurcating distal of vein M + CuA dividing; vein RP almost straight basal of nodal line; vein A1 short, terminating at inner margin basal of stem R + M bifurcating; wing membrane gray with lighter or darker narrow areas along veins. Differs from S. plachutai in possessing a smaller forewing with costal margin convex at about middle of wing, vein R bifurcating distal of vein M + CuA dividing, vein A1 shorter and different colour patterns; from S. daohugouensis in having a smaller forewing with costal margin convex at about middle of wing and distinct colour patterns. Description: Holotype NIGP149376. Body dark. Ocelli invisible. Eyes large, rounded or ovoid, slightly emarginated behind antennae. Antennae slender, short and two thirds as long as fore tibia, with four segments visible; scape thick and short; pedicel slightly shorter than first flagellomere; flagellum
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Fig. 2. Shuraboprosbole daohugouensis nov. sp. Paratypes. A, B. Photograph and illustration of the paratype NIGP149375 (same scale). C. Photograph of the paratype NIGP149374. D. Photograph of the forewing in paratype NIGP149374. Scale bars represent 10 mm.
aristiform. Postclypeus strongly convex, with distinct transverse grooves. Rostrum extending beyond hind coxae, nearly to apex of hind femur. Pronotum slightly wider posteriorly; anterior margin almost straight, lateral margins diverging posteriad, posterior margin distinctly concave medially. Mesonotum partly covered by wings, with distinctly separated, convex prescutum. Fore femur 1.2 times as long as hind femur, distinctly thicker than hind femur, unarmed. Fore tibia slightly longer than fore femur. Fore tarsus 0.4 times as long as fore tibia. Mid tibia 1.5 times as long as mid femur. Mid tarsus 0.3 times as long as mid tibia. Hind tibia slender, twice as long as femur, with two welldeveloped lateral spines on outer edge of tibia (longer than diameter of tibia, one at its 1/3, the other at about its 2/3) and two (or three?) apical teeth set on enlarged sepal base. Hind tarsus 0.4 times as long as hind tibia; first two tarsomeres with apical teeth (probable two teeth in each tarsomere); basitarsomere slightly longer than mid tarsomere; mid tarsomere 0.6 times as long as apical tarsomere. Tarsal claws well developed. Arolium invisible. Forewing elongate, with costal margin convex at about middle of wing. Nodal incision distinct at basal 0.65 wing length. Vein ScP invisible in wing base, dividing from ScP + RA at nodal line and ending at nodus. Stem R + M branching at basal 0.21 wing length. Vein R branching into veins RA and RP at basal 0.33 wing length. Branch RA curved anteriorly, recurved at nodal line, and then branching into veins
RA1 and RA2 at basal 0.68 wing length. Branch RA2 connected with vein RP by crossvein ir at basal 0.75 wing length. Branch RP sinuous, connected with branch M1 by crossvein r-m slightly basal of crossvein r. Vein M curved posteriorly and fused with vein CuA for a short distance at basal 0.23 wing length, and then branching into veins M1+2 and M3+4 at basal 0.4 wing length. Vein M1+2 branching into veins M1 and M2 at basal 0.69 wing length. Vein M2 connected with M3 by crossvein im just basal of crossvein r-m. Vein M3+4 branching into veins M3 and M4 basal of bifurcation of vein M1+2. Vein M3 curved anteriorly at crossvein m. Vein M4 connected with branch CuA1 by crossvein m-cua at basal 0.63 wing length. Stem Cu bifurcating into veins CuA and CuP at wing base. Branch CuA curved anteriorly at fusion with vein M, recurved at nodal line, and branching into veins CuA1 and CuA2 at basal 0.52 wing length. Vein CuA1 curved anteriorly at crossvein mcua. Vein CuA2 very short. Vein CuP straight. Vein Pcu slightly curved. Vein A1 weak, curved anteriorly and terminating at inner margin basal of stem R + M bifurcating. Nodal line traceable as a crease along vein ScP, crossing veins M1+2 and M3+4 and then fused with veins CuA and CuA2. Marginal membrane wider than vein and distinct at posterior margin. Wing membrane gray, with very narrow white bands along veins. Paratype NIGP149377. Hind tibia slender, with two welldeveloped lateral spines on outer edge of tibia (longer than diameter of tibia, one at its 1/3, the other at about its 2/3).
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Fig. 3. Comparison of forewings of Shuraboprosbole. A. S. daohugouensis nov. sp., NIGP149375. B. S. daohugouensis nov. sp., NIGP149374. C. S. minuta nov. sp., NIGP149405. D. S. minuta nov. sp., NIGP149378. E. S. media nov. sp., NIGP149407. F. S. plachutai Becker-Migdisova, 1949, redrawn from Becker-Migdisova (1949). Wings aligned horizontally with apex to the right. All to scale; scale bar represents 5 mm.
Abdomen pointed posteriorly. Ovipositor ensiform, upcurved, adpressed to pygofer, extended just below anal tube. Paratype NIGP149378. Forewing with vein M branching slightly basal of nodal line. Wing membrane gray, with very narrow dark bands along veins. Paratype NIGP149380. Hind tibia with three prominent lateral spines. Three spines longer than diameter of tibia; two on outer edge of tibia, one at its 1/3, the other at about its 2/3; one on inner edge of tibia, at about its midlength. Forewing with vein M branching slightly basal of nodal line. Wing membrane gray, with very narrow white bands along veins. NIGP149405. Venation in general as in holotype. Wing membrane gray, with very narrow white bands along veins. Measurements (mm): Male (N = 4), range given for available specimens with mean in parentheses. Body length:
13.5–14.1 (N = 2); vertex length: 0.9; rostrum length: 4.7; pronotum length: 2.7–3.4 (N = 2); forewing length: 14.8–16.5 (15.1; N = 4), width: 4.5–5.3 (4.9; N = 4), length/width ratio: 3.0–3.3 (3.1; N = 4); hind tibia length: 4.0–4.6 (N = 2). Female (N = 18). Body length: 14.1–15.9 (14.8; N = 9); vertex length: 1.0–1.2 (1.1; N = 7); pronotum length: 2.9–3.4 (3.2; N = 7); forewing length: 16.0–18.9 (17.6; N = 15), width: 5.5–6.2 (5.8; N = 15), length/width ratio: 2.9–3.2 (3.0; N = 15). Other forewings (N = 7). Forewing length: 15.2–18.2 (16.9), width: 4.8–6.2 (5.6), length/width ratio: 2.9–3.2 (3.0). Remarks: The length/width ratio of forewing and location of vein M branching are different among specimens. Two different colour types are detectable, but neither could be associated with any particular wing venation pattern. Furthermore, the consistent dimorphism in wing venation and colour
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Fig. 4. Shuraboprosbole minuta sp. nov. Holotype NIGP149376. A, B. Photograph and illustration (same scale). Scale bar represents 5 mm. C. Enlarged antenna. D, E. Photograph and illustration of enlarged hind tarsi (same scale). Scale bars represent 0.5 mm.
patterns was not detectable between the two sexes. Some of these intraspecific variations also occur in some fossil and living Cicadomorpha, such as palaeontinids and leafhoppers (Larsen and Walter, 2007; Wang et al., 2008). Therefore, it is reasonable to place these specimens into a single species. Shuraboprosbole media nov. sp. Figs. 3(E) and 6 Holotype: NIGP149407. Almost complete forewing with clavus slightly damaged. Type locality: Daohugou Village, Chifeng City, Inner Mongolia, China. Stratum typicum: Middle Jurassic.
Etymology: Specific epithet refers to the medium size of the forewing. Diagnosis: Forewing medium-sized, length about 25 mm; costal margin convex at basal 0.25 wing length; vein R bifurcating distal of vein M + CuA dividing; vein RP almost straight basal of nodal line; wing membrane without distinct colour patterns. It is similar to S. plachutai in the forewing shape (both with costal margin convex at basal 0.25 wing length), but differs from the latter in having a forewing with vein A1 shorter and wing membrane without distinct colour patterns. It is different from S. daohugouensis in having a smaller forewing with costal margin convex at basal 0.25 wing length, vein RP nearly straight basal of nodal line and stem M
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Fig. 5. Shuraboprosbole minuta sp. nov. Photographs. A. NIGP149377. B. NIGP149380. C. NIGP149379. D. NIGP149378. E. NIGP149385. F. NIGP149405. Scale bars represent 5 mm.
shorter. It also differs from S. minuta in having a larger forewing with costal margin convex at basal 0.25 wing length and wing membrane without distinct colour patterns. Description: Holotype NIGP149407. Forewing elongate, length: 25.8 mm, width: 10.1 mm, length/width ratio: 2.6. Costal margin convex at basal: 0.25 wing length, and costal area wide. Nodal incision distinct at about basal 0.6 wing length. Stem R + M branching at basal 0.17 wing length. Vein R branching into veins RA and RP at basal 0.27 wing length. Vein R about one third as long as stem R + M. Branch RA curved anteriorly, recurved at nodal line, and then branching into veins RA1 and RA2 at basal 0.68 wing length. Branch RA2 connected with vein RP by crossvein ir at basal 0.75 wing length. Branch RP almost straight basal of nodal line, connected with branch M1 by crossvein r-m slightly basal of crossvein ir. Vein M curved posteriorly and fused with vein CuA for a short distance at basal 0.18 wing length, and then branching into veins M1+2 and M3+4 at basal 0.39 wing length. Vein M1+2 branching into veins M1 and M2 at basal 0.67 wing length. Vein M1 strongly curved anteriorly at crossvein r-m. Vein M2 connected with M3
by crossvein im at the same level of crossvein r-m. Vein M3+4 branching into veins M3 and M4 slightly basal of bifurcation of vein M1+2. Vein M3 slightly curved anteriorly at crossvein im. Vein M4 connected with branch CuA1 by crossvein m-cua at about basal 0.6 wing length (Ct) narrow. Stem Cu bifurcating into veins CuA and CuP at wing base. Branch CuA curved anteriorly at fusion with vein M, recurved at nodal line, and branching into veins CuA1 and CuA2 at basal 0.49 wing length. Vein CuA1 strongly curved anteriorly at crossvein m-cua. Vein CuA2 very short. Vein CuP slightly curved, ending at about midpoint of wing, forming a long clavus. Vein Pcu sinuous. Nodal line traceable as a crease along vein Sc, crossing veins M1+2 and M3+4 and then fused with veins CuA and CuA2. Wing membrane gray. Marginal membrane wider than vein and distinct at posterior margin. 4. Discussion Up to now, 14 species within nine genera from China were placed in Tettigarctidae and all of them are restricted to the
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Fig. 6. Shuraboprosbole media nov. sp. Holotype NIGP149407. A, B. Photograph and illustration (same scale). Scale bar represents 5 mm.
Mesozoic. Most of them, however, are problematic in their systematic position. Protabanus chaoyangensis Hong, 1982, based on a specimen from the Middle Jurassic of Liaoning, was firstly referred to Tabanoidea (Diptera). Hamilton (1992) and Zhang (1993) considered it as a Tettigarctidae, but Nel (1996) excluded it from Cicadoidea. Based on the general appearance and forewing venation (Hong, 1982: Text-Fig. 150), it probably belongs to Cicadomorpha. However, judging from the original photograph of the impression (Hong, 1982: Pl. 38, Fig. 1), its forewing is obscure and broken in the middle and distal part, and hence its systematic position requires the reexamination of the type specimen. Luanpingia Hong, 1983, with three known species originally placed in Tettigarctidae, has been already referred to Procercopidae (Hamilton, 1992; Ren, 1995). Four specimens, respectively from South China and Xinjiang, were ever placed in Tettigarctidae: Lacunisbole ligonis Lin, 1986 from the Upper Triassic of Qujiang, Guangdong; Quadrisbole vieta Lin, 1986 from the Upper Triassic of Liuyang, Hunan; Quadrisbole stenis Lin, 1986 from the Lower Jurassic of Liuyang, Hunan; and Kerjieprosbole stigmatis Lin, 1992 from the Upper Triassic of Toksun, Xinjiang. Nel (1996) presented a detailed discussion regarding these four specimens; all of them are based on forewing fragments and require a further study. Sinocicadia shandongensis Hong, 1990 was erected mainly on the basis of a forewing from the Lower Cretaceous Laiyang Formation of Laiyang, Shandong. Based on the original description and illustration, neither distinct nodal line nor nodus is present, and vein M forks near wing apex. Therefore, it cannot
be included in Tettigarctidae, but its systematic position requires the reexamination of the type specimen (Nel, 1996). Sinocicadoprosbole hunjianensis Hong and Chang, 1993, from the Upper Triassic Beishan Formation of Hunjiang, Jilin, is based on a forewing with distal part missing and anal area destroyed. Judging from the photograph and illustration of the type specimen, the characteristics including absent nodal line and nodal incision, vein RA branching basal of middle of wing, vein CuA fused with stem R + M and vein CuA branching at the same level of vein M branching, exclude this species from Tettigarctidae, but probably attribute it to Prosbolidae. Last, Involuta perrara Zhang, 1993, certainly belonging to Tettigarctidae, is based on a complete hindwing from the Lower Cretaceous Fengjiashan Formation of Shangxian, Shanxi, China. Therefore, only four definitive species of Tettigarctidae are recorded from the Mesozoic of China. Based on our collections and recognitions, the large body with a greatly inflated frontoclypeus and a long rostrum indicates that early tettigarctids are probably primarily arboreal and xylem feeding. Considering the absence of angiosperms in the Daohugou flora, early tettigarctids probably live on some gymnosperms. So far, however, only few plant fossils from Daohugou have been studied in detail. Furthermore, plant fossils recorded from Daohugou (including ferns, cycads, ginkgoes and conifers) are also common in some other coeval biota. Therefore, little is known about the association between plants and cicadoprosbolines based on fossils from Daohugou. The pronotum exceedingly large and concealing major of mesonotum
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is common in extant Tettigarctinae and has been regarded as a diagnostic character of Tettigarctidae (Moulds, 2005). The pronotum of Cicadoprosbolinae is large, but not expanded over mesonotum (Figs. 1, 4 and 5; Hamilton, 1990). Therefore, the pronotum concealing major of mesonotum seems to be an autapomorphy of Tettigarctinae. Jurassic Tettigarctidae possess an elliptical forewing with broad costal area and clavus; whereas extant Tettigarctidae have elliptical forewing with narrow costal area and reduced clavus. This evolutionary trend is similar to that of Mesozoic Palaeontinidae, and probably also suggests an improvement of flight ability (Wang et al., in press). The earliest whole-bodied tettigarctids from the Middle Jurassic of Daohugou are slightly different from those from the Lower Cretaceous of Brazil by the detailed structure of hind legs. Hamilton (1990) showed that pretarsal structures of Architettix (Cicadoidea) are similar to those of Cercopoidea. But he did not suggest that Cicadoidea are closely related to Cercopoidea, and also did not give the reason. Because of the lack of detailed pretarsal structures in Daohugou specimens, we cannot decide whether this similarity is a parallel evolution between Architettix and Cercopoidea. Hamilton pointed out that Architettix compacta Hamilton, 1990 possessed an elongate hind basitarsomere with two rows of plantar setae, and furthermore he considered structures of ‘‘cicadellid-like tarsi’’ as synapomorphies linking the Membracoidea and Cicadoidea (Hamilton, 1990, 1996). However, Middle Jurassic Tettigarctidae show common cicadoid hind tarsi, in which the basitarsomere is short and unarmed. Therefore, the ‘‘cicadellidlike tarsi’’ found in Cicadellidae and Brazilian Tettigarctidae are probably a parallel evolution. Although almost distributed worldwide in the Jurassic and Early Cretaceous, the Tettigarctidae are very rare as fossils from the Late Cretaceous and Cenozoic. The scarcity of these insects in Cenozoic biotas is not a result of taphonomical bias, because singing cicadas occur in some Cenozoic deposits (Cockerell, 1906; Zhang and Zhang, 1990; Grimaldi and Engel, 2005). The decline of Tettigarctidae is probably due to changes in the vegetation and competitive displacement with singing cicadas. Acknowledgements We thank Huang Diying for his donation of some specimens, and K.G. Andrew Hamilton and André Nel for providing their literatures. Many thanks go to two anonymous reviewers for reviewing the manuscript and providing some constructive suggestions, and Gilles Escarguel for his helpful editorial comments. This research has been financially supported by the Major Basic Research Projects (2006CB806400) of MST of China, and the National Natural Science Foundation of China (40523004, 40632010, 40872015). References Becker-Migdisova, E.E., 1947. Cicadoprosbole sogutensis gen. nov. sp. nov., a transitional form between the Permian Prosbolidae and the recent Cicadidae. Doklady Akademii Nauk SSSR 55, 445–448 (in Russian).
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