Revision of the genus Sinochlora Tinkham (Orthoptera: Tettigoniidae, Phaneropterinae)

The genus Sinochlora Tinkham [type species Sinochlora longifissa (Matsumura and Shiraki)] (Orthoptera: Tettigoniidae) is revised. Six new species, S. nonspinosa sp. nov., S. trapezialis sp. nov., S. tibetensis sp. nov., S. mesominora sp. nov., S. retrolateralis sp. nov., S. aequalis sp. nov., and the male of S. hainanensis are described. S. gracilisulcula Shi and Zheng, 1996 and S. kiangsuensis Tinkham, 1945 are synonymized with S. szechwanensis Tinkham, 1945. Holochlora voluptaria Carl, 1914 is transferred to the genus Sinochlora. The significance of each taxonomic character is discussed. Characteristics of the male epiproct of most species are firstly studied in detail and considered most important and relied on to distinguish species. Some known species are redescribed based on the characters of the male epiproct and male stridulatory area. The map of geographical distribution with GIS is drawn, and the possible patterns of distribution are discussed. A key to the 13 known species in the world is provided.


Introduction
provided a valuable work on Phaneropterinae in China, in which he established the genus Sinochlora, including five species from China (type species Sinochlora kwangtungensis Tinkham, 1945). Sinochlora is most closely related to Holochlora and distinguished by the following synapomorphies: white and black tegminal costal vein, black femoral spine, male tenth abdominal tergum produced into a pair of forcipate lateral processes and a central one, male subgenital plate from moderately to very strongly arcuately reflexed, and female ovipositor with the upper valvulae strongly truncated at the apex.
Later, Shi and Zheng (1996) described one species, S. gracilisulcula, from Sichuan Province, China. In Kang's master's dissertation (1987), he indicated that Holochlora longifissa should be transferred to the genus Sinochlora, H. longifissa and S. kwangtungensis are junior synonyms of S. longifissa, and S. kwangtungensis is mistakenly identified. Liu

Diagnosis
Male. Tegmen with stridulatory area elongate, left tegmen with stridulatory vein distinctly coarse, strongly elevated and massive above the wing plane, and right tegmen with mirror indistinct, only having a small transparent area (Figures 1, 2). Stridulatory file on underside of left tegmen with teeth regularly arranged, becoming gradually smaller from the middle to both sides ( Figure  Female. Ovipositor equally broad throughout, axe-shaped, dorsal margin with apical part truncated and serrated, ventral margin with apical part spiniferous ( Figures 4F, 6G, 9D-G).

Notes
We consider that Holochlora voluptaria Carl, 1914 should be transferred to the genus Sinochlora for its following characters: black and white tegminal costal vein, strong black femoral spines, male tenth abdominal tergum possessing a straight crassus median process and two small lateral lobes (which is similar to S. sinensis Tinkham and S. nonspinosa sp. n.), male epiproct developed ( Figure 5E), male subgenital plate with lateral margin convex after basal constriction ( Figure 5F), and female ovipositor with apical half of dorsal margin truncated, finely serrated ( Figure 5G).

Discussion
The genus Sinochlora is most closely related to the genus Holochlora in the tribe Holochlorini. Most similarities and differences between the two genera have been discussed in detail in Tinkham's work (1945). Here we just mention features of the male stridulatory apparatus and male epiproct. These two genera share the following characters: male stridulatory vein on male left tegmen coarse, strongly elevated and massive above the wing plane, teeth of stridulatory file on the underside of left tegmen regularly densely arranged and becoming smaller proximally, and mirror on the right tegmen indistinct.     2. Central process of male tenth abdominal tergum much more elongate, and lateral processes almost indistinct ( Figure 4A, B, G, H); male subgenital plate split from approximately apical half ( Figure 4E, M) . . . . . . . . . . . . 3 -Central process of male tenth abdominal tergum slightly shorter, and lateral processes distinct ( Figure 5A, B, E); male subgenital plate variable. . . . . 5 3. Central process of male tenth abdominal tergum possessing distal half strongly deflexed and with apical margin truncate or slightly clavate ( Figure 10F). Male epiproct short, triangular, thickened, with a spine recurved upward at apex ( Figure 10G). Female subgenital plate large, base wider than the apex, closely quadrangular with a deeper triangular notch on posterior margin. Guizhou Province, China ( Figure 10H) . . . . Sinochlora stylosa Shi and Chang, 2004 -Distal part of central process of male tenth abdominal tergum somewhat deflexed with apical margin emarginate to different extent ( Figure 4A    -Size much larger (length of male: tegmina 56.0 mm, posterior femora 34.0 mm) (Carl 1914). Male tenth abdominal tergum with lateral lobe rather big ( Figure 5E). Male subgenital plate split from about basal third, with lateral margins strongly convex after the basal constriction ( Figure 5F). North Vietnam . . . . . . . . . Sinochlora voluptaria (Carl, 1914) comb. n.
6. Male tenth abdominal tergum with each lateral lobe rather long and strongly inclined downwards, the median process large, gradually constricted toward both swollen lateral apices ( Figure 11D). Male epiproct smaller, and just bearing three small round lobes at apex, among which the middle one is distinctly smaller than both the lateral ones ( Figure 11F). Female subgenital plate large, widely triangular, lateral margins approximately straight, and with a small broad shallow triangular notch at the apex ( Figure 5I).  Figure 6A, B). Male epiproct approximately trapezoid, much longer than wide, and with each posterior lateral corner extending into a long upcurved sharp spine ( Figure 6C, D). Female subgenital plate large, broadly triangular, with strongly convex lateral margins and with an acute triangularly incised apex ( Figure 6H). Hainan Island, China . . . . . . . . . . . . . . . Sinochlora hainanensis Tinkham, 1945 -Lateral lobes of male tenth abdominal tergum not strongly diverging as above.
Male epiproct and female subgenital plate not as above . . . . . . . . 8 8. Central process of male tenth abdominal tergum large and elevated with its posterior margin approximating the apices of the lateral arms ( Figure 7A, B). Male epiproct smaller, slightly wider than long, and bearing a pair of relatively short spine-like lateral processes at apex, which are projected dorsad, notch between trapezoid ( Figure 7C, D). Female subgenital plate short and very broadly triangular, with a broad shallow triangular notch at the apex ( Figure 7F). China, Korea, Japan . . . . . Sinochlora longifissa (Matsumura and Shiraki, 1908) -Male epiproct not as above. Female subgenital plate triangular, longer than wide 9 9. Male epiproct equilaterally triangular, dorsally concave towards the inside, with a pair of black sharp spines directing dorso-caudad, between which is triangular ( Figure

Redescription
Male. Stridulatory file on underside of left tegmen with about 50 teeth ( Figure 3A). Epiproct approximately long triangular, dorsally concave towards the inside, without long black heavy bristles in the ventral surface; basal half area semicircular, then abruptly narrowed; apical third very narrow, subparallel-sided, and armed with a pair of very small spine-like processes projected dorsad only at apex, notch between triangular ( Figure 4C, D).
Color. Deep foliage-green. Costal vein white and black, and each tegmen with a black spot at the base of the green subcostal vein.
( Figures 1C, D Figure 3B). Male tenth abdominal tergum with central process strongly produced, distal half strongly deflexed, apical margin emarginated, and lateral processes indistinct ( Figure 4G-I). Male epiproct approximately trapeziform, slightly wider than long, with dorsal surface strongly concave towards the inside symmetrically, without long black heavy bristles in the ventral surface; basal margin broadly concave, lateral margin gradually constricted towards the apex, and each posterior lateral corner extending into an upcurved sharp spine. Male cerci rather long, incurved, and conical ( Figure 4J-L). Male subgenital plate slightly upcurved, split from apical half, and with small styli ( Figure 4M).
Color. Deep foliage-green. Costal vein white and black, and each tegmen with a black spot at the base of the green subcostal vein.

Measurements of male (mm)
Length of body 29.0, length to tip of wing 55.5, length of pronotum 6.3, length of tegmen 43.5, largest width of tegmen 8.5, length of hind wing 48.5, length of anterior femur 8.5, length of median femur 11.0, length of posterior femur 29.0.

Remarks
In having elongated deflexed central process and indistinct lateral lobes of male tenth abdominal tergum, the new species closely resembles S. stylosa from Guizhou Province in southwestern China. It differs from S. stylosa by the male epiproct being trapezoid with each lateral apex formed into an upcurved spine (not triangular with apex formed into a spine).

China.
Sinochlora nonspinosa sp. n. (Figures 1E, F Figure 3C). Male tenth abdominal tergum with central process produced, gradually narrowed, apical margin rounded, and each lateral one distinctly present as a small round lobe ( Figure 5A, B). Male epiproct triangular, dorsally strongly concaved entad, and with long black heavy bristles in apical part of ventral surface, and apical part upcurved, gradually tapering into an obtuse finger-shaped process at apex. Male cerci rather long, incurved, and conical ( Figure 5C). Male subgenital plate strongly upcurved, split from apical third, and with small styli ( Figure 5D).
Color. Deep foliage-green. Costal vein white and black, each tegmen with a black spot at the base of the green subcostal vein.

Measurements of male (mm)
Length of body 25.0, length to tip of wing 52.0, length of pronotum 5.5, length of tegmen 42.0, largest width of tegmen 7.5, length of hind wing 47.0, length of anterior femur 7.0, length of median femur 10.5, length of posterior femur 27.0.

Remarks
The new species closely resembles S. voluptaria in male tenth abdominal tergum with central process strongly produced far beyond the somewhat distinct lateral lobes, but distinctly distinguished from the latter in the small size, the shape of the male subgenital plate and male epiproct triangular with apex obtuse and upcurved.

Distribution
China.

Description
Female (holotype). Size larger for the genus (length of tegmen: male 55 mm, female 56.0-57.0 mm; length of posterior femur: male 33.5 mm, female 36.0 mm). Form, coloration, fastigum verticis, and pronotum also similar to the congenera. Tegmen with Rs bifurcate and given off slightly after the middle, and R stem also with three other lateral branches. Female tenth abdominal tergum approximately trapeziform, with apical margin slightly obtuse. Cerci short, conical, suddenly sharpened into a spine at apex. Subgenital plate triangular, wider than long, with slightly convex lateral margin and obtuse emarginated apical margin ( Figure 5I). Ovipositor similar to other congener ( Figure 5J).
Male. Stridulatory file on underside of left tegmen with about 74 stridulatory teeth ( Figure 11A). Tenth abdominal tergum with each lateral lobe rather long and strongly inclined downwards, the median process large, declined from base to center on the dorsal surface, gradually constricted toward both swollen lateral apices, and extending beyond the apices of the lateral lobes ( Figure 11C, D). Epiproct smaller, approximately trapezoid, slightly wider than long, dorsally concave entad, with numerous black bristles in the ventral surface, and just bearing three small round lobes at apex, among which the middle one is distinctly smaller than both the lateral ones ( Figure 11F, G). Male subgenital plate with apical two-fifths incised, and the forked processes gently recurved ( Figure 11H).
Color. Deep foliage-green. Costal vein white and black, each tegmen with a black spot at the base of the green subcostal vein. Apical half lateral lobes of male tenth abdominal tergum black. Male epiproct marginalized black lateral stripes, and also with black half apical part.

Remarks
The new species differs from other Sinochlora species by the larger size, structure of male tenth abdominal tergum, male epiproct, and female widely triangular subgenital plate. It slightly resembles S. longifissa in the shape of female subgenital plate, but differs by the larger size, male abdominal apex and the female subgenital plate with obtuse emarginated apical margin (not angular emarginated). It resembles S. hainanensis in size but differs by the male abdominal apex and the shape of female subgenital plate.

First description of male (nov.)
Form and size large for the genus (male length of tegmen: male 48.5 mm; length of posterior femur: male 31.5 mm). Stridulatory file on underside of left tegmen with about 55 teeth ( Figure 3D). Male tenth abdominal tergum strongly produced, distal quarter deflexed, split into two approximately triangular strongly diverging lateral lobes ( Figure 6A, B). Male epiproct approximately trapezoid, much longer than wide, dorsal surface strongly concaved symmetrically, and without long black bristles in the ventral surface; basal margin broadly concave, lateral margin strongly constricted towards the apex, and with each posterior lateral corner extending into a long upcurved sharp spine ( Figure 6C, D). Male cerci rather long, incurved, and conical. Male subgenital plate slightly upcurved, widest at basal third part, then strongly tapering, split from apical third, and both subparallel lateral lobes with minute styli ( Figure 6F).
Color. Deep foliage-green. Costal vein white and black, each tegmen with a black spot at the base of the green subcostal vein.

Variation
One specimen has triangular subgenital plate with straight lateral margin and emaginated apical margin, which is very different from the original description of Sinochlora hainanensis.

Remarks
The species is distinctly distinguished by male tenth abdominal tergum strongly produced, distal quarter deflexed, split into two approximately triangular strongly diverging lateral lobes, and male epiproct approximately conversely trapezoid, much longer than wide, and each posterior lateral corner extending into a long upcurved sharp spine, and without long black bristles in the ventral surface.

Redescription
Male. Stridulatory file on underside of left tegmen with about 10 undeveloped teeth in the basal part, and 36 normal teeth in the other part ( Figure 3E). Tenth abdominal tergum with each lateral lobe short and forcipate, the median process large, elevated, and with its posterior margin approximating the apices of the lateral lobes ( Figure 7A, B). Epiproct smaller, slightly wider than long, dorsally concave entad, without black bristles in the ventral surface, and bearing a pair of relatively short spine-like lateral processes at apex, which are projected dorsad, notch between trapezoid ( Figure 7C, D). Male subgenital plate with apical fourth incised, and the forked processes gently recurved ( Figure 7E).
Female. Subgenital plate short, triangular, wider than long, and with a shallow broad triangular notch at the apex ( Figure 7F).
Color. Deep foliage-green. Costal vein white and black, each tegmen with a black spot at the base of the green subcostal vein.

Redescription
Male. Stridulatory file on underside of left tegmen with 10 undeveloped teeth at the basal part and about 64 normal teeth in the remaining part ( Figure 3F, J-M). Epiproct bigger, longer, dorsally concave entad, with long black heavy bristles in the ventral surface, and apical area with a pair of relatively longer spine-like lateral processes which are projected dorsad beyond the median tumid area of the tenth abdominal tergum, notch between cambered ( Figure 8G-R).

Discussion
The type specimens of S. gracilisulcula have been damaged. The original description of S. gracilisulcula just differs from that of S. szechwanensis in the former having the following apomorphies: small distance between the lateral lobes of male tenth abdominal plate, female subgenital plate slightly broader than long, and with a distinct obtuse-angular notch at the base and basal lateral margins distinctly enlarged. After carefully examining a damaged female paratype and the material from O'Mei Mt., Sichuan Province at hand, we found that these specimens somewhat vary to a different extent regarding male tenth abdominal tergum and shape of female subgenital plate. Thus as concerning male tenth abdominal tergum and female subgenital plate, we consider that there lies much morphological variation within the geographical population in O'Mei Mt., Sichuan Province; and the apomorphies mentioned about S. gracilisulcula are just not beyond the variation. So we consider S. gracilisulcula just as a junior synonym of S. szechwanensis.
When carefully examining specimens from nine different geographical populations: Sichuan Province, Hunan Province, Hubei Province, Henan Province, Anhui Province, Jiangxi Province, Jiangsu Province, Zhejiang Province, and Guangxi Province, we can trace all transitive forms of the different character states-male tenth abdominal tergum dorsally with or without a middle elongate sulcus, and slightly varied female subgenital plate between S. kiangsuensis and S. szechwanensis. More importantly, male epiprocts of different geographical populations are nearly similar in structure, differing only by size of protuberance at the base of the ventral surface, of which all transitive forms with different size also can be traced ( Figure 8G-R). The other characters including male stridulatory file, male tenth subgenital plate and male cerci are of slight or no difference. Thus we also consider S. kiangsuensis as a junior synonym of S. szechwanensis.

Distribution
Sinochlora szechwanensis Tinkham is distributed over the Chinese district of the Oriental Region.

Description
Male (holotype). Stridulatory file on the underside of left tegmen with about 68 teeth ( Figure 3G). Tenth abdominal tergum produced backwards, basal half triangularly sulcate in the middle, lateral forcipate processes gently decurved in the apical half, and apical margin approximately flat in the middle ( Figure 9A, B). Epriproct dorsally concaved, with black bristles in the middle and both lateral sides in ventral surface, basal part widest, gradually constricted and shaped into three upcurved spines at apex, among which the middle one is smaller than the lateral ones, directing caudad at the base, and curved craniad apically, and both lateral symmetrical ones directing caudad ( Figure 9D-G). Cerci conical, rather long, slender, and slightly curved. Subgenital plate broad at the base and very deeply cleft at the basal third, and the long arms strongly recurved and terminated with minute apical styli ( Figure 9C).
Color. Brownish green. Costal vein white and black, each tegmen with a black spot at the base of the green subcostal vein.

Measurements of male (mm)
Length of body 28.0, length to tip of wing 58.0, length of pronotum 6.5, length of tegmen 46.5, largest width of tegmen 9.0, length of hind wing 51.5, length of anterior femur 8.0, length of median femur 12.0, length of posterior femur 29.0.

Remarks
The new species most closely resembles S. trispinosa except that its male epiproct has a smaller middle spine than two lateral ones at apex.

Distribution
China.

Redescription
Male. Stridulatory file on the underside of left tegmen with about 63 teeth ( Figure 3I). Lateral forcipate processes of the tenth abdominal tergum decurved in the apical area, central process slightly produced with a rather broad central groove ( Figure 9H, I). Epiproct symmetrical, equilaterally triangular in the whole view, dorsally concave entad, with long black heavy bristles in the basal ventral surface, and apical part with three spines directing dorso-caudad, among which the middle one longer than two lateral ones, notches between triangular ( Figure 9J-L).
Color. Green. Costal vein white and black, each tegmen with a black spot at the base of the green subcostal vein.

Measurements of male (mm)
Length of body 29.0, length to tip of wing 58.0, pronotum 6.0, length of tegmen 47.0, largest width of tegmen 9.5, length of hind wing 52.0, length of anterior femur 8.0, length of median femur 11.5, length of posterior femur 27.0.

Remarks
The species is recognized by the male epiproct providing three spines directing dorsocaudad at apex, among which the middle one is longer than the two lateral ones.
( Figures 2G, H Figure 3H). Dorsal surface of undivided part possessing a converse triangular longitudinal groove in the middle, the lateral forcipate processes decurved in the apical area, and apical margin emarginated in the middle ( Figure 10A, B). Cerci longer and coniform. Epiproct symmetrical, equilaterally triangular in the whole view, dorsally concave towards the inside, with long black heavy bristles on the basal ventral surface, and apical area with three prongs at apex, among which the middle longest, both symmetrical lateral shorter and diverging forth, and notches between triangular ( Figure 10C-E). Subgenital plate deeply split for over apical one-half, lateral lobes upcurved beyond the tenth abdominal tergum, and with small styli.
Color. Deep foliage-green. Costal vein white and black, each tegmen with a black spot at the base of the green subcostal vein.

Measurements of male (mm)
Length of body 30.0, length to tip of wing 58.0, pronotum 7.0, length of tegmen 47.0, largest width of tegmen 9.5, length of hind wing 52.0, length of anterior femur 7.5, length of median femur 10.0, length of posterior femur 29.5.

Remarks
The new species is similar to S. szechwanensis, S. mesominora, S. trispinosa in the size, form, tegmen, tenth abdominal tergum, and subgenital plate, but differs by apical part of the epiproct shaped into three large spines, among which the larger middle one is straightly produced dorso-caudad, and two lateral ones diverging forth.

Description
Male (holotype). Stridulatory file on the underside of left tegmen with about 51 teeth ( Figure 11B). Tenth abdominal tergum produced backwards, apical half elliptically sulcate in the middle, lateral forcipate processes gently decurved in the apical half, and apical margin approximately flat in the middle ( Figure 11I, J). Epriproct dorsally concaved, with little bristles in the middle and both lateral sides in ventral surface, basal part widest, gradually constricted and shaped into three upcurved spines at apex, among which the middle one is approximately as long as lateral ones, directing dorsad ( Figure 11K, L). Cerci conical, rather long, slender, and slightly curved. Subgenital plate broad at the base and very deeply cleft at the basal third, and the long arms strongly recurved and terminated with minute apical styli.
Female. Subgenital plate short, triangular, wider than long, and with a rather deep broad rounded notch at the apex ( Figure 11N).
Color. Green. Costal vein white and black, each tegmen with a black spot at the base of the green subcostal vein. Female ovipositor with apical third brown.

Remarks
The new species is distinguished from other species in the genus by the male epiproct bearing three spines, of which the middle one is approximately as long as the lateral ones, and the female subgenital plate with broad rounded notch at apex.

Distribution
China.

Biogeography
The genus is mostly distributed in the Oriental Region (Map 1). The origin of the genus is uncertain, although the majority of species exist in the tropical region today. There are two possible patterns. The more probable one is that the genus is of a tropical origin, from there it respectively spread over the Himalayan Region towards the northwest and Palaearctic Region (including China, Korea, and Japan) towards the northeast; the other is that the genus is of Palaearctic origin, from which it spread and diversified towards the Oriental Region.
The five species are distributed in the Guangxi Province, southwest of China, adjacent to Vietnam, from where another species, Sinochlora voluptaria, was described. The type species S. longifissa is widely distributes in China, Korea, and Japan. S. sinensis and S. szechwanensis are widely distributed in the south of China. S. tibetensis is endemic to Tibet and S. hainanensis to Hainan Island.