Revision of the Oriental spittlebug genera Baibarana Matsumura and Telogmometopius Jacobi (Hemiptera: Cercopidae: Rhinaulacini)

The Oriental spittlebug genera, Baibarana Matsumura and Telogmometopius Jacobi (Hemiptera: Auchenorrhyncha: Cercopidae), previously included in Cosmoscartini and Bandusiini, respectively, are transferred to Rhinaulacini and revised to include five and nine species, respectively. The species recognized are: Baibarana bicolor sp. nov. (Vietnam), B. longispina sp. nov. (Laos and Thailand), B. rufofasciata sp. nov. (Vietnam), B. sinuata sp. nov. (Vietnam), B. uchidai Matsumura (Taiwan), Telogmometopius angulatus sp. nov. (Malaysia: northern Borneo), T. bicarinatus sp. nov. (northeastern India), T. bifasciatus sp. nov. (Malaysia: northern Borneo), T. carinilabratus (Chou and Wu) (from Baibarana Matsumura), comb. nov. (southwestern China: Yunnan; Tibet), T. himalayensis sp. nov. (northeastern India: east Himalayas, southwestern China: Tibet), T. obsoletus Jacobi, 1921 (southern and central China: Hubei, Gansu, Sichuan, Guizhou, ?Jiangxi), T. rangoonensis sp. nov. (Burma), T. sabahensis sp. nov. (Malaysia: Sabah), and T. sarawakensis sp. nov. (Malaysia: Sarawak). Descriptions and keys to genera and species are provided together with structural illustrations. Scanning electron micrographs of the antennal sensilla, rostral apex, and male genitalia of the two genera, are provided for the first time. A lectotype is designated for Telogmometopius obsoletus Jacobi.


Introduction
Cercopid spittlebugs (Hemiptera: Auchenorrhyncha) form the largest group of plant xylem sap-feeding insects in the world. Although the New World fauna has been recently revised (Carvalho and Webb 2006), the cercopid family Cercopidae (sensu Metcalf, 1961) in the Old World remains one of the most neglected Auchenorrhyncha groups at all levels. Descriptions of species and higher taxa are scattered in the older taxonomic literature and generally lack illustrations of the diagnostic male genitalia, and therefore often make species 4. Hind wing with cross vein m-cu meeting Cu1a after the Cu1a/Cu1b bifurcation. Basal half of Cu1b strongly arched against Cu2 (Liang 1993); aedeagal shaft robust (Liang and Webb, 2002); subgenital plates with a strongly incurved apical process (Liang 1993) . . . . . . . . . . . . Paracercopis Schmidt -Hind wing with cross vein m-cu meeting Cu1a before the Cu1a/Cu1b bifurcation. Basal half of Cu1b straight and remote from Cu2 ( Figure 8); aedeagal shaft elongate, or, if robust, subgenital plates without a strongly incurved apical process 5 5. Hind wing with bifurcation of Cu1a/Cu1b near m-cu cross vein; aedeagal shaft robust (Liang 1994a

Description
Small, finely pubescent spittlebugs, length (from apex of vertex to tip of fore wings): 3.8-5.0 mm. General colour grey to brown, vertex and pronotum usually olivaceous yellow; fore wings sometimes with pale granules.
Head relatively broad; roundly or angularly produced anteriorly, narrow in profile. Vertex flat, sloping forward, posterior margin concave; ocelli very close to each other, situated halfway between anterior and posterior margins. Antennae with pedicel distinctly longer than scape, flagellar base expanded and subglobose, with one peg-like basiconic sensillum on apical part; ultrastructure not examined. Postclypeus weakly convex, in males upper half variably compressed laterally and longitudinally keel-like medially; lateral areas with distinct muscle impressions (striations) dorsally. Females with postclypeus more bulbose with lateral areas weakly compressed and medial longitudinal region flatter and smoother and muscle impressions fainter. Rostrum extending slightly beyond middle coxae. Pronotum hexagonal, broader than long, centrally and posteriorly convex, sloping to depressed vertex; anterior margin broadly arched anteriorly, anterior lateral margins relatively short, nearly half to one-third of posterior lateral margin, posterior margin mesially arched anteriorly. Scutellum as long as broad at base, disc shallowly depressed, apex sloping downward caudally. Fore wings about 2.5 times as long as broad, surface sometimes with numerous granules, some of them confluent, forming ridges; Rs branched apically; veins indistinct or sometimes distinct on apical area resulting in obscure or prominent apical cells, respectively. Hind wings with four elongate, closed, apical cells, Cu1 branched, Cu1a and Cu1b on relatively long stalk, cross vein m-cu meeting Cu1a well before branch of Cu1a/Cu1b. Hind tibiae with one lateral spur slightly distad of mid-length. Male pregenital segment very short, much shorter than pygofer in ventral view.
Male genitalia with pygofer in lateral view protruding anteriorly on upper half and posteriorly on lower half, wider dorsally than ventrally, anterior margin somewhat Sshaped, posterior margin sinuate and slightly angular submesially. Anal segments and anal style short and small. Subgenital plates not fused to pygofer, comprising an expanded basal part with an apical spine-like process. Lateral plates usually very elongate and large in lateral view, distinctly protruding posterodorsally above subgenital plates and nearly as long as the upper genital styles in lateral view. Styles relatively elongate, positioned above lateral plates in lateral view; produced dorsally slightly distad of mid-length or club-like (in sinuata); pilose over apical half. Aedeagal shaft in lateral view very elongate, cylindrical, extreme basal part produced ventrally between subgenital plates then curved posterodorsally and strongly anterodorsally, subapically expanded with extreme apex usually with a fine, posteroventrally directed hook; gonopore subapical on dorsal surface.

Remarks
Baibarana was placed in the tribe Cosmoscartini Schmidt by Metcalf (1961). Cosmoscartini is characterized by the strongly bulbose frons, hind tibiae with two lateral spurs, and male genitalia with subgenital plates short, without an apical spine-like process, without lateral plates, and aedeagal shaft short and not strongly curved. We here transfer Baibarana to the tribe Rhinaulacini based on the following characters: postclypeus weakly bulbous; hind tibiae with one lateral spur; subgenital plates with an expanded basal region and a spine-like apical process; lateral plates present; and aedeagal shaft elongate, cylindrical, and strongly curved.
Baibarana can be distinguished by its relatively broad head ( Figure 1) and from most other Rhinaulacini (especially those from the Oriental and Australian Regions) by the postclypeus with its upper part usually keel-like medially and its lower part relatively broad, flat and smooth. In other genera the postclypeus is medially sulcate, or keel-like throughout length in Telogmometopius. In the male genitalia Baibarana is very similar to Sounama (see Liang and Webb 2002, Figures 58-94).
In common with most spittlebug groups, currently no biological data are available for Baibarana species. Collecting data show that adults can be collected in almost all seasons and certain species, e.g. B. longispina, can be attracted and collected at light (see collecting data below).

Distribution
Oriental Region, including Taiwan, Vietnam, Laos, and Thailand.  Male genitalia with subgenital plates  with expanded basal region broad in ventral view, apical spine-like process short and stout, directed anterodorsally. Lateral plates ( Figure 41) with apical margin broad in lateral view, moderately protruding beyond pygofer. Styles ( Figure 43) with a dorsal subapical process and a ridge extending ventrally from near base of process. Aedeagus as in Figure 44.

Distribution
Vietnam.

Remarks
This species can be distinguished by its contrasting colour pattern and its basally broad subgenital plates with short apical process. It differs from the very similar B. uchidai from Taiwan, by its smaller size and narrower subgenital plates basally and shorter aedeagal shaft with more weakly arched basal part.

Baibarana longispina sp. nov. (Figures 26, 56-62)
Description Small species, length: " 3.8-4.0 mm, R 4.0 mm. Vertex and pronotum pale ochraceous; ocelli reddish; callous areas near posterior margins of vertex and on anterior area of pronotum suffused with brown, sometimes posterior half of pronotum suffused with brown; scutellum and fore wings dark brown or pale fuscous; head and thorax ventrally, basal segment of rostrum, fore and middle femora basally and abdomen, brown; apical segment of rostrum, apex of fore and middle femora, fore and middle tibiae and tarsi, hind legs, metapleura, metasterna, and veins on apical area of fore wings ochraceous.
Fore wings with distinct apical cells.
Male genitalia with subgenital plates (Figures 56-58) with expanded basal region short and narrow, apical spine-like process very elongate, greater than 2.5 times length of expanded basal region, directed posterodorsally in lateral view. Lateral plates (Figures 58,  61) large, broadly rounded apically. Styles (Figures 59, 60) terminating in a small, dorsally directed bifurcate process. Aedeagus as in Figure 62.

Remarks
This species can be distinguished by its elongate apical process of the subgenital plate and bifurcate apical process of the style.
Fore wings with distinct apical cells.

Remarks
This species can be very easily distinguished by the two transverse rufous fasciae on the vertex and pronotum, ochraceous apical veins of the fore wings and by the male genitalia, particularly by the short apical process of the subgenital plates.
Vertex and pronotum olivaceous yellow, posterior half of pronotum with pale brown suffusion; face and sternum, apical segment of rostrum, scutellum, leg, fore wings and abdomen brown; basal segment of rostrum, apical veins of fore wings, metapleura, metasterna, apex of fore and middle femora, fore and middle tibiae and tarsi and hind legs, ochraceous.
Fore wings with distinct apical cells.

Distribution
Vietnam.

Remarks
This species can be distinguished by the shape of its subgenital plates and styles. Lectotype ", designated by Komatsu and Hayashi 1998, p 170, Taiwan (HU) [examined].

Description
Length: " 4.8-5.0 mm, R 4.6-5.0 mm. Vertex and pronotum olivaceous yellow; ocelli red. Face ochraceous with antennal sockets, antennal pedicel and upper region of frons brown, the latter sometimes with a narrow, transverse brown or fuscous fascia; scutellum, thorax beneath, and legs ochraceous, sometimes with brown suffusion. Fore wings brown, sometimes with costal and apical area and subapical transverse band, ochraceous. Abdomen brown.
Fore wings densely granulate, some granules confluent forming ridges; veins indistinct. Male genitalia with subgenital plates (Figures 45-47) with expanded basal region nearly triangular in lateroventral view; apical spine-like process very short and stout, directed anterodorsally, with a laminate, finger-like process along basal half of inner margin. Lateral plates (Figure 47) short and broad in lateral view, moderately posteriorly protruded beyond posterior margin of pygofer. Styles (Figures 46, 47) with bifurcate apex, outer fork smaller, laminate, strongly sclerotized and hook-shaped and inner fork larger, somewhat papilliform. Aedeagus as in Figure 50.

Remarks
This species is very similar to B. bicolor but can be distinguished from the latter by its distinctly larger size and in the male genitalia by their broader subgenital plates and more elongate aedeagal shaft with a more broadly arched basal part. Telogmometopius Jacobi 1921, p 43;Wu 1935, p 48;Schulze, Kü kenthal and Heider 1938, p 3406;Neave 1940, p 417;Lallemand 1949, p 56, 57;Metcalf 1961, p 327. Type species: Telogmometopius obsoletus Jacobi, 1921, by original designation and monotypy.

Description
Small to medium-sized, slender, finely pubescent spittlebugs, length (from apex of vertex to tip of fore wings): " 5.0-8.4 mm, R 5.3-8.5 mm. General colour grey brown or brown; vertex and pronotum usually ochraceous, probably olivaceous yellow in life.
External features as in Baibarana with the following differences. Head (Figures 2, 9) relatively narrow, broad in profile; posterior margin of vertex more or less straight; ocelli situated slightly basad of mid-distance from anterior to posterior margin of vertex; postclypeus (Figures 4, 10, 11) strongly compressed laterally and medially longitudinally keel-like. Ultrastructure of antennal pedicel: very sparsely setose and sculptured with scalelike pattern, apex hollowed with inner surface strongly spiralled ( Figure 12); flagellum base nearly smooth, sensory organs in subglobose base including one long blunt, peg-like, basiconic sensillum located in a broad, shallow pit adjacent to a group of about five, small, coeloconic sensilla on rim (Figures 12, 13). Rostrum tip (Figures 17, 18) consisting of two lateral lobes separated by a dorsal stylet groove, each lateral lobe with about 12 peg sensilla, numerous short cuticular teeth and some divided acanthae near inner edge. Fore wings with relatively distinct venation. Male pregenital segment very short, much shorter than pygofer in ventral view.
Male genitalia as in Baibarana with the following differences. Lateral plates very small. Styles strongly expanded over distal half in lateral aspect with apex excavate with two lobes, outer lobe with two small, dorsally directed teeth on outer dorsal edge. Aedeagal shaft with basal part horizontal, thereafter strongly and narrowly curved anteriorly, with an apical or dorsal subapical, anteriorly directed, sinuate or coiled, filamentous process.

Biology
In common with most spittlebug genera, no biological data are currently available for species of Telogmometopius. Species of the genus may be associated with bamboos (see collecting data under T. sabahensis). Collecting data show that adults can be collected in all seasons and certain species, e.g. T. obsoletus and T. himalayensis, can be attracted and collected at light (see collecting data below).

Remarks
The monobasic genus Telogmometopius Jacobi was placed by Lallemand (1949) and Metcalf (1961) in the Afrotropical tribe Bandusiini Lallemand, together with the Afrotropical genus Bandusia Stål. We here transfer Telogmometopius to the tribe Rhinaulacini based on the following characters: postclypeus weakly bulbous; hind tibiae with one lateral spur; subgenital plates with an expanded basal region and a spine-like apical process; lateral plates present; aedeagal shaft elongate, cylindrical, and strongly curved; distribution and type of antennal sensilla (see Telogmometopius can be distinguished from other Oriental cercopid genera by its strongly laterally compressed postclypeus, with a median carina, keel-like in ventral view (Figures 10, 11) and in the male genitalia by the small lateral plates and very elongate aedeagus with the basal part horizontal and strongly and narrowly curved anteriorly with an apical or subapical filamentous process.
Key to the species of Telogmometopius 2. Postclypeus with lower two-thirds of median keel distinctly dilated laterally resulting in a diamond-shaped, flattened, median area; subgenital plates ( Figure 71)

Description
Length: " 5.2-5.4 mm, R 5.3-5.6 mm. Vertex, pronotum, and scutellum pale stramineous (possibly olivaceous yellow in live specimens); ocelli pale red; ocelli, callous areas on posterior area of vertex and anterior area of pronotum, posterior half of pronotum and middle area of scutellum usually with brown suffusion; head and thorax ventrally ochraceous, basal antennal segments, postclypeus laterally, fore-and mesothorax beneath and legs sometimes suffused with brown; fore wings grey brown, with an indistinct, oblique, pale white transverse fascia before middle, the costal area behind the pale white fascia fuscous, apical margin fuscous; abdomen dark brown, posterior margin ochraceous.
Postclypeus with medial keel very sharp. Male genitalia with subgenital plates (Figures 91-93) with expanded basal region elongate and apical spine-like process short and robust, about half length of expanded basal region, directed dorsally at a nearly 90u angle in lateral view. Style as in Figure 92. Aedeagal shaft ( Figure 92) laterally compressed, in lateral view relatively short and broad with a very fine, anterodorsally directed filamentous process, arising subapically from dorsal margin of an expanded membranous apex.

Remarks
This species can be distinguished by its relatively short and stout aedeagal shaft with the apex expanded and with a filamentous process arising subapically from the dorsal margin and by its stout subgenital plates with a dorsally directed apical process.

Description
Small and slender species, length: " 5.4 mm, R 5.3 mm. Vertex and pronotum pale ochraceous; ocelli suffused with reddish; callous areas on posterior area of vertex and anterior area of pronotum brown; head ventrally brown, median keel of postclypeus (including its lower expanded area), anteclypeus and rostrum ochraceous; antennae with pedicle and expanded flagellar base fuscous; thorax ventrally and legs ochraceous with brown suffusion; fore wings pale stramineous with an indistinct, narrow, obliquely transverse, pale white fascia, before middle; area behind transverse pale white fascia and apical outer margin dark brown; abdomen brown or dark brown with posterior margins ochraceous.
Postclypeus with lower two-thirds of median keel distinctly dilated laterally resulting in a diamond-shaped, flattened, median region, laterally carinate.
Male genitalia with subgenital plates (Figures 69-71) with expanded basal region small and apical spine-like process very long and straight, about three times length of expanded basal region. Style as in Figures 72 and 73. Aedeagal shaft (Figure 70) very elongate with distal third tapered into an apical, sinuate, filamentous process, apex curved posterodorsally.

Distribution
Northeastern India.

Remarks
This species can be distinguished externally by its postclypeus with the lower two-thirds of the median keel distinctly dilated laterally, resulting in a diamond-shaped, slightly sulcate, median area bordered laterally with a carina. It can be distinguished in the male genitalia by its subgenital plates having a small expanded basal region and a very elongate apical spinelike process.

Description
Relatively small species, length: " 5.2 mm. General colour brown; ocelli suffused with red; antennal scape, pedicel, and expanded flagellar base, upper margin of frons, and mesosterna, fuscous; fore wings pale, with two transverse fuscous fasciae: one before middle and the other subapical.
Postclypeus strongly compressed laterally and median keel on postclypeus very fine and sharp.
Male genitalia with pygofer ( Figure 83) in lateral view with posterior margin rounded; subgenital plates ( Figure 84) with basal expanded region narrow and slender, nearly triangular in ventral view, base broad, tapered from base to apex; apical spine-like process relatively stout, tapered from base to apex, posterodorsally directed in lateral view; genital styles not examined (missing). Aedeagal shaft (Figure 85) very long and slender with an elongate, anteriorly directed filamentous process arising from dorsal margin of an expanded apex, in lateral view.
Non-type material examined. One female specimen from Sabah, Mt Kinabalu (BMNH) may belong to this species.

Remarks
This species is similar to rangoonensis but has the subgenital plate tapering more gradually and the aedeagal shaft more elongate. The styles and one subgenital plate of the unique type are missing. With respect to the aedeagus its process is now detached and lying within the pygofer (drawn re-associated in Figure 85). In consequence it is not clear if the lightly sclerotized apical part (basad of the process), as figured, is complete. Despite these deficiencies, sufficient critical parts of the male genitalia remain to enable adequate identification.

Description
Length: " 7.5-8.4 mm, R 7.5-8.5 mm. Vertex and pronotum ochraceous; callous areas on posterior area of vertex and anterior area of pronotum brown, posterior area (excluding posterior margin) of pronotum brown; scutellum brown with apex and basal lateral angles ochraceous; face brown with median keel of frons and rostrum ochraceous, sometimes rostrum with apical segment suffused with brown; thorax ventrally brown with mesosterna, metapleura, and metasterna ochraceous; legs brown with apex of fore and middle femora and base of fore and middle tibiae, and hind tibiae and tarsi ochraceous; fore wings brown, apical part and costal margin in middle much darker; abdomen brown with segmental posterior margins ochraceous. Sometimes body beneath almost uniformly brown, except metapleura and metasterna of thorax and hind tibiae and tarsi, ochraceous.
Postclypeus with median keel moderately sharp. Male genitalia with subgenital plates (Figures 74-76) with expanded basal region very elongate, evenly tapered from base to apex, joined basally; apical spine-like process short and fine, half length of expanded basal region. Style as in Figure 78. Aedeagal shaft ( Figure 79) extremely elongate, apical part distinctly filamentous and coiled.

Remarks
This species is the largest of the genus. Its male genitalia are similar to T. himalayensis but the aedeagal shaft is slightly narrower in lateral view.

Description
Length: " 5.4-5.5 mm, R 5.6 mm. Vertex and pronotum ochraceous, callous areas on posterior area of vertex and anterior area of pronotum brown; scutellum ochraceous, central foveate area and two callous areas on lateral basal angles brown. Head ventrally brown, postclypeus with median keel and the areas between lateral striate fasciae on lateral compressed areas ochraceous. Thorax ventrally brown, fore-and mesosterna, metapleura, and metasterna ochraceous. Legs brown, fore femora with extreme base ochraceous; posterior tibiae (excluding base) and tarsi of hind legs ochraceous; rostrum ochraceous. Fore wings grey brown, apical area and costal marginal area after middle fuscous. Abdomen brown with posterior segmental margin, pygofer and genitalic segment ochraceous.
Postclypeus with median keel very sharp. Male genitalia similar to T. carinilabratus but aedeagus ( Figure 81) slightly more robust with apical filamentous process sinuate rather than coiled. Non-type material examined. The following specimens may belong to this species. China, Tibet: 3"", 3RR (1R collected at

Remarks
This species is very similar externally to T. carinilabratus but can be distinguished by its smaller size and small differences of the aedeagus as noted in its description. A series of specimens from China, Tibet (SIE; IZCAS), may be this species but individuals are slightly larger.

Description
Length: " 5.4-5.6 mm, R 5.6-5.8 mm. Vertex, pronotum, and scutellum ( Figure 36) pale ochraceous; ocelli with red or brown suffusion; callous areas on vertex and pronotum, posterior area of pronotum and central foveate area of scutellum usually with brown suffusion; antennal sockets, basal antennal segments and anterior margin of frons in ventral view brown, forming a distinct, transverse, brown fascia in ventral view; rostrum ochraceous; thorax ventrally ochraceous, mesosterna brown; fore and middle legs brown, extreme base of fore and middle femora ochraceous, hind legs ochraceous; fore wings grey brown, sparkled with ochraceous, with an indistinct, narrow, obliquely transverse, pale white fascia before middle; abdomen brown, posterior segmental margin and genitalic segment ochraceous.
Postclypeus with medial keel moderately sharp. Male genitalia with subgenital plates (Figures 14-16, 98-100) closely approximated, with expanded basal region moderately long and oval-shaped; apical spine-like process short and fine, distinctly shorter than expanded basal region. Style as in Figure 102. Aedeagal shaft (Figure 103) relatively short and broad in lateral view, compressed laterally, distal two-thirds evenly broad in lateral view; with an anterodorsally directed, membranous, sinuate filamentous process arising subapically from upper margin.

Remarks
This species is similar to sabahensis but can be distinguished by the subgenital plates with the apical spine-like process shorter and finer and the aedeagal shaft slightly more robust in lateral view. We have not been able to verify the distribution of the species from Jiangxi [Kiangsi], China (Wu 1935;Metcalf 1961).

Description
Small species, length: " 5.0 mm. General colour pale stramineous; ocelli suffused with red; callous areas on posterior area of vertex and anterior area of pronotum pale brown; pronotum with posterior area pale brown; body ventrally ochraceous.
Postclypeus with median keel very sharp. Vertex with a faint, fine, median longitudinal carina.
Male genitalia with subgenital plates (Figures 86-88) with expanded basal region moderately large and apical spine-like process elongate and fine, slightly longer than expanded basal region. Style as in Figure 90. Aedeagal shaft ( Figure 87) relatively short and stout, with upper anteriorly directed part slightly arched in lateral view, apex expanded and slightly upturned, with an elongate, anterodorsally directed, sinuate, filamentous process arising from apical upper margin.

Remarks
This species is similar to bifasciatus but has the subgenital plates tapering more abruptly and the aedeagal shaft shorter.

Remarks
This species can be distinguished by the caliper-like subgenital plates and looped apical filamentous process of the aedeagus.