Revision of Pseudoparicana ( Hemiptera : Fulgoromorpha : Tropiduchidae ) with description of a new species

The planthopper genus Pseudoparicana Melichar, 1914 is revised. Three previously known species, P. curvifera (Distant, 1907), P. sanguinifrons Muir, 1931 and P. tepida Melichar, 1914 are re-described and P. analoga Wang & Malenovský sp. n. is described as new. Lectotypes are designated for P. curvifera and P. tepida. An identification key and illustrations of diagnostic characters, including male and female genitalia, are provided for all taxa and the systematic position of this genus and the asymmetry of the genitalia of the species are briefly discussed. As far as known, the distribution of Pseudoparicana is restricted to New Guinea and the Aru Islands. 365 * Corresponding author; e-mail: liangap@ioz.ac.cn Type species. Paricana curvifera Distant, 1907, by original designation. Included species. P. curvifera (Distant, 1907), P. tepida Melichar, 1914, P. sanguinifrons Muir, 1931 and P. analoga Wang & Malenovský sp. n.


INTRODUCTION
The planthopper genus Pseudoparicana (Hemiptera: Fulgoromorpha: Tropiduchidae) was established by Melichar (1914) for Paricana curvifera Distant, 1907 and Pseudoparicana tepida Melichar, 1914. Muir (1931) added a third species, P. sanguinifrons Muir, 1931.The genus has been recorded from Indonesia (Aru Islands) and New Guinea (Melichar, 1914;Muir, 1931).Unfortunately, the original descriptions of all three species lack illustrations and clear diagnostic information, particularly on the morphology of the male and female genitalia, which are crucial for their reliable identification.Moreover, Melichar's drawings (1914) of a dorsal view of the head and fore wing of P. tepida are inaccurate.Pseudoparicana was included in the tribe Paricanini by Melichar (1914) but transferred to the tribe Tropiduchini by Fennah (1982).Members of the Tropiduchini mainly occur in the Oriental and Afrotropical regions, including the western Pacific (Bourgoin, 2012).
Specimens of Pseudoparicana are rare in collections.While sorting and identifying Tropiduchidae in material from the California Academy of Sciences, San Francisco and the Moravian Museum, Brno, we found a new species of Pseudoparicana collected in Papua New Guinea.This new species and a revision of the type material of the previously described species enabled us to re-describe the genus and for the first time describe in detail with illustrations the male and female genitalia.We also provide a key for identification of the four currently known species and additional illustrations of the habitus, head, pronotum and wings.According to Article 74 of ICZN (1999), lec-totypes for P. curvifera and P. tepida are designated to stabilize the nomenclature in the genus.

Description
Coloration.General colour of body is straw-colored to brownish yellow.Fore wings (Figs 2A-F, 3D) hyaline, marked with two more or less narrow transverse brown bands, one situated sub-basally, the other along the nodal line (in P. tepida, band along nodal line is indistinct).
Head and thorax.Head in dorsal view (Figs 2A-C, E, 3B) short and broad, produced in front of eyes approximately for a half the length of an eye, with apex broadly convex; in lateral view (Figs 2D, F, 3C) evenly and broadly rounded down to base of frons, vertex and frons not separated, lateral carinae continuous.Vertex (Figs 2A-C, E, 3B) about 2.75 times as broad at base than long at midline, and slightly longer than the length of pronotum at midline (1.25 : 1); lateral margins ridged and gently convex; posterior margin ridged and concave; anterior part of vertex smoothly convex; posterior part of vertex distinctly depressed, lateral margins of depression incurving anteriorly and meeting medially to form a nearly straight or slightly convex line; vertex without median carina.Frons (Figs 2G-J, 3A) longer than broad, widest in lower portion, usually medially swollen and shiny, disc between median swollen portion and lateral carinae depressed in shallow grooves on each side, converging down to clypeus (in P. tepida, frons is regularly convex, lacking median swelling and depressed grooves); lateral margins of frons distinctly convex adjacent to clypeus; frontoclypeal suture obtusely angled, concave.Clypeus convex, medially swollen in continuation with the median swollen area on frons and lacks lateral carinae.Labium long, reaching between hind coxae, apical segment distinctly longer than broad.Compound eyes oval.Lateral ocelli distinct.Antenna with a short ring-like scape; pedicel cylindrical, covered with many micro-setae extending to base of pedicel.
Distribution.Species of Pseudoparicana are currently known from the Aru Islands (administrative part of Indonesia) and New Guinea (Fig. 1).E-F).Posterior margin of male pygofer nearly straight in left lateral view (Fig. 4B), produced into a broadly obtuse lobe dorsally in right lateral view (Fig. 4C).Gonostyli, in left lateral view, with obtusely rounded apex (Fig. 4B
Coloration.General colour of body pale yellow.Clypeus and median swollen portion of frons orange to red.Posterior lateral corners of vertex, eyes, disc of pronotum, anterior portion of mesonotum and dorsum of abdomen all suffused with red.Mesonotum with trans-verse suture black.Mesopleura with large orange stripes, metapleura reddish black.Fore wings hyaline, marked with two narrow dark brown transverse bands sub-basally and on nodal line; apical margin narrowly bordered dark brown in posterior third; scutellar border red.Hind wings hyaline, veins dark brown, anal area infuscated apically.Legs yellow, hind coxae, trochanters, femora, fore tibia, and tarsi all infuscated dark brown; apices of all spines black.Freshly emerged specimens uniformly pale yellow.
Male genitalia.Pygofer asymmetrical, narrow and high in lateral view; in left view (Fig. 3G), wider ventrally than dorsally, with posterior margin dorsally produced into a blunt, tooth-like lobe directed dorso-caudad (Figs 3F-G); in right lateral view (Fig. 3H), posterior margin regularly convex; in ventral view (Fig. 3I), posterior margin of pygofer with right (dextral) part with a U-shaped excavation to accommodate gonostyli.Anal tube (Figs 3F-H) short; in lateral view, expanded and decurved in distal half from the axis of the basal half; in dorsal view, basal part distinctly narrowed, expanded toward apex, apical margin truncate.Gonostyli (Figs 3F-I) fused together, distinctly curved along medial axis and forming a sub-cylindrical structure, joined by membrane to pygofer at base; in left lateral view (Fig. 3G), spade-like, with truncate apex and a relatively long curved, hook-like and strongly sclerotized process arising on apical half on left side of gonostyli and directed dorsocephalad (Fig. 3I); in right lateral view (Fig. 3H), gonostyli broad, without process on right side laterally.Aedeagus situated left of gonostyli (Figs 3F-I), asymmetrical, tube-like; in lateral view (Figs 3G, K), left distal part abruptly expanded into a sub-spherical lobe oriented dorsad; in dorsal view, apex on right lateral side produced into a broad triangular process directed laterad (Figs 3F, I-J); endosoma with two narrow apical processes: one spine-like, situated proximally and directed dorso-mesiad and another sinuate, denticulate along its margin, situated distally and directed dorso-laterad (Figs 3J-K); phallotreme split-like, running along the apex of aedeagus, forming a distinct longitudinal furrow.
Female genitalia.Unknown.Etymology.From the adjective analogus, Latinized form of Ancient Greek άνάλογος = correspondent, like, similar, compa-rable.The new species so named because in general appearance it is similar to P. curvifera (Distant, 1907).

Remarks. P. analoga
Wang & Malenovský sp.n. is similar to P. curvifera externally but can be distinguished 372 from the latter by the shape of the male pygofer, gonostyli and aedeagus as indicated in the corresponding key couplet above.

Description
Size.Male.Total length (from apex of vertex to tip of fore wings): 7.5 mm; body length (from apex of vertex to tip of anal tube): 5.0 mm; fore wing length: 6.2 mm.Female.Total length: 8.1 mm; fore wing length: 6.8 mm.
Coloration, head and thorax as in P. analoga Wang & Malenovský sp.n.
Male genitalia.Pygofer asymmetrical, narrow and high in lateral view; in left view (Fig. 4B), wider ventrally than dorsally, posterior margin nearly straight, truncate; in right lateral view (Fig. 4C), with posterior margin dorsally produced into a broadly obtuse lobe directed dorsocaudad; in ventral view (Fig. 4D), posterior margin of pygofer with U-shaped depression on right (dextral) part to accommodate gonostyli.Anal tube (Figs 4A-C) short; in lateral view, slightly expanded and decurved in distal half from the axis of the basal half; in dorsal view, basal part narrowed, expanded toward apex, apical margin rounded.Gonostyli (Figs 4B-D) fused together, distinctly curved along medial axis and forming a sub-cylindrical structure, joined by a membrane to base of pygofer; in left lateral view (Fig. 4B), narrow at base, with obtuse apex and a relatively long curved, hook-like and strongly sclerotized process arising on apical half on left side of gonostyli and directed dorso-cephalad; in right lateral view (Fig. 4C  Female genitalia.Gonoplacs with 11 strongly sclerotized teeth along apical margin arranged in two rows ( Figs 4G-H).Gonapophyses VIII strongly sclerotized with 7 blunt teeth on dorsal margin; base of gonapophyses VIII asymmetrical in ventral view: left part produced into two small, tooth-like lobes, right part expanding into one relatively long lobe (Fig. 4H).Sternite VII with posterior margin shallowly sinuate, shallowly notched in obtuse angle at middle in ventral view (Fig. 4H).
Remarks.P. curvifera is similar to P. analoga Wang & Malenovský sp.n. externally, but can be distinguished from the latter by the shape of the male pygofer, gonostyli and aedeagus (see the key above).The female of P. curvifera differs from P. sanguinifrons and P. tepida in the shape of the base of the gonapophyses VIII.
A comparison with the corresponding type material revealed that one male and one female specimen from "Fiume Purari" deposited in L. Melichar collection (MMBC) and cited as Pseudoparicana curvifera by Melichar (1914) and Wilson & Malenovský (2007) were misidentified and belong to P. sanguinifrons.
Coloration.General colour of body straw yellow.Frons (Figs 2H-I) uniformly yellow or the swollen middle portion reddish.Clypeus dark brown to black.Posterior tip of mesonotum narrowly and mesoscutellum entirely dark brown to black.Fore wings (Fig. 2B) hyaline with two well-delimited and relatively broad transverse dark brown bands, one situated sub-basally, the other along nodal line; apical margin of fore wing widely bordered dark brown in posterior third (dark coloration of membrane extending from node round wing apex nearly to clavus); scutellar border red.Legs yellowish or light brownish, coxae dark brown to black, femora ventrally, fore tibia entirely, mid and hind tibiae basally and tarsi all infuscated dark brown; apices of all spines on legs black.Abdomen light ochreous to reddish brown basally, darker brown in the mid-dorsal portion, two apical sternites dark brown to black.
Male genitalia.Pygofer asymmetrical; in left lateral view (Fig. 5A), narrow and high, wider ventrally than dorsally, with posterior margin produced into a large, sharply triangular tooth on dorsal third, directed dorsocaudad; in right lateral view (Fig. 5B), pygofer narrow, with posterior margin regularly convex in dorsal part; in ventral view (Fig. 5D), posterior margin of pygofer with right (dextral) part widely excavated to accommodate gonostyli.Anal tube (Xth segment; Figs 5A-C) short, expanded and slightly decurved in distal half from the axis of the basal half in lateral view; apical margin irregularly rounded in dorsal view.Gonostyli (Figs 5A-B, D) fused together, distinctly curved along medial axis and forming a sub-cylindrical structure, with a rounded, strongly curved apex in left lateral (Fig. 5A) and ventral view (Fig. 5D), and a relatively long curved, hook-like and strongly sclerotized process arising on apical half on left side of gonostyli and directed dorso-cephalad (Figs 5A, C-D); in right lateral view (Fig. 5B), gonostyli broadly truncate at apex, without process on lateral side.Aedeagus (Figs 5A, E-F) situated left of gonostyli, asymmetrical, tubular-like; in left lateral view, with apical part nearly parallel-sided, slightly curved dorsad; right lateral side of aedeagus with two large processes arising from aedeagus shaft medially: in right lateral view, one is long and runs along ventral margin of aedagus, blunt at apex, the other small, tooth-like, with sharp apex, oriented dorsad; endosoma with a vermicular twisted apical process on left side, directed obliquely cephalad (Fig. 5F).Phallotreme split-like, forming a longitudinal furrow.
Remarks.P. sanguinifrons can be differentiated from the related species by its slightly larger size, extended dark markings on the fore wings, presence of a sharp triangular tooth on the left posterior margin of male pygofer and the structure of the gonostyli and aedeagus.The female differs from P. curvifera and P. tepida also in the shape of the base of gonapophyses VIII.
Coloration.General colour of body (Figs 2C, F) brownish yellow.Frons, clypeus (Fig. 2J), vertex, pronotum and major portions of mesonotum and scutellum uniformly dirty yellow to brownish yellow; posterior fifth of mesonotum light brown; mesoscutellum basally narrowly dark brown.Pleural sclerites of thorax marked with dark brown.Fore wings hyaline, marked with a not clearly delimited narrow transverse dark brown band subbasally; nodal line and node dark brown but dark coloration restricted to veins, hardly extending into membrane in adjacent cells; apical margin of forewing clear, lacking dark markings; a small basal portion of scutellar border red.Legs yellowish or light brownish, coxae dark brown to black, hind femora and tarsi infuscated dark brown; apices of all spines on legs black.Abdomen light ochreous, reddish brown in the mid-dorsal portion, two apical sternites dark brown to black.
Male genitalia.Pygofer asymmetrical; in left lateral view (Fig. 7A), narrow and high, slightly wider ventrally than dorsally, posterior margin regularly convex in dorsal half; in right lateral view (Fig. 7B), pygofer broad, concave medially, produced into a broadly rounded lobe dorsally; in ventral view (Fig. 7D), posterior margin of pygofer with left (sinistral) part widely excavated to accommodate gonostyli.Anal tube very short, in lateral view (Figs 7A-B) nearly triangular; in dorsal view (Fig. 7C), round, with apical margin slightly concave.Gonostyli fused together, distinctly curved along medial axis and forming a sub-cylindrical structure; in left lateral view (Fig. 7A), with a broadly rounded apex; in dorsal view (Fig. 7C), with a sclerotized tooth on inner side of left dorsal margin medially and a relatively long curved, hook-like and strongly sclerotized process arising on apical half on right side of gonostyli and directed dorsocephalad; in right lateral view (Fig. 7B), gonostyli distinctly curved and apically rounded.Aedeagus situated right of gonostyli (Figs 7A-D), asymmetrical, tubularlike, shaft bearing a small sclerotized tooth-like apical process dorsally (Figs 7C, E-F); in right lateral view (Fig. 7E), apex of aedeagus broadly truncate; endosoma with a large hook-like apical process largely surpassing aedeagus apex (Figs 7E-F).Phallotreme running along the apex of aedeagus, forming a distinct, wide opening.
Female genitalia.Gonoplacs with 10 strongly sclerotized teeth arranged in two rows along apical margin.Gonapophyses VIII strongly sclerotized with 6-8 blunt teeth on dorsal and three on ventral margin; base of gonapophyses VIII asymmetrical in ventral view: left part with broadly rounded apex, right part distinctly sinuate subapically (Fig. 6E).Sternite VII with posterior margin shallowly and broadly V-shaped, indistinctly notched at middle in ventral view (Fig. 6E).Posterior vagina relatively small, irregularly heart-shaped in ventral view (Fig. 6E).Bursa copulatrix membranous, bladder-like with surface covered in numerous small sclerotized cuticular spines, sharply triangular or bifid at apex.Distribution.South-eastern New Guinea (Fig. 1).

Remarks. P. tepida differs from all other
Pseudoparicana species in its regularly convex frons lacking a median swelling and lateral grooves, the absence of a distinct dark brown band extending into the membrane along the nodal line on the fore wings, a more evenly rounded fore wing apex lacking dark brown pigmentation, and the structure of the male genitalia, including its chirality, the shape of the pygofer, the presence of a sclerotized tooth on the left latero-dorsal margin of the gonostyli, and the form of the aedeagus and its processes.The female differs from P. curvifera and P. sanguinifrons in the shapes of the posterior margin of sternite VII and the base of gonapophyses VIII and from P. curvifera also in the smaller posterior vagina and microsculpture on the bursa copulatrix.
Melichar's drawing of a dorsal view of the head of P. tepida (Melichar, 1914: 113, Fig. 27) is wrong in showing a median carina on the vertex.In fact, the three syntype specimens examined do not have such a carina.
One female paralectotype of P. tepida is parasitized by a female of Strepsiptera, tentatively identified as Halictophagus sp.Halictophagus Curtis, 1831 is a large cosmopolitan genus comprising 97 species associated with various families of Auchenorrhyncha (Kathirithamby & Moya-Raygoza, 2000;Kathirithamby, 2012).Two species of Halictophagus are recorded from New Guinea: H. ancylophallus Kifune & Hirashima, 1989and H. eurycephalus Kifune & Hirashima, 1989(Kifune & Hirashima, 1989;Kathirithamby, 2012).More material, particularly males, is needed for a precise identification of the parasite.Remarks.The above cited females collected in northwestern and northern New Guinea (Fig. 1) are similar to P. analoga Wang & Malenovský, sp.n. and P. curvifera in general habitus, particularly the fore wing pattern with two well-distinct, narrow, transverse brown bands.The base of gonapophyses VIII is broadly irregularly rounded and lacks distinct tooth-like lobes as in P. curvifera.The identification of these specimens is uncertain -they may belong to P. analoga Wang & Malenovský, sp.n. (for which no female has been collected at the type locality) or to one or two additional, undescribed species.More material, particularly of the males, is needed for identification.

Systematic position and diagnosis of the genus
In the absence of a phylogenetic analysis of Tropiduchidae, which would test homologies of morphological characters, the systematic position of Pseudoparicana within the family remains enigmatic.Melichar (1914) included it in the tribe Paricanini, defined by him mainly by the elongate forewing with the nodal line situated basad of the middle, the apical part longer than the corium and the presence of short erect setae on veins.Fennah (1982) transferred Pseudoparicana to the Tropiduchini, apparently based e.g. on the asymmetrical, fused gonostyli shared with some twenty other genera.In general habitus, Pseudoparicana is similar to Paricana Walker, 1857 (Paricanini) and Dichoneura Lethierry, 1890 (transferred from Delphacidae to Tropiduchidae: Cyphoceratopini by Fennah, 1982).
Pseudoparicana represents a distinct taxon within Tropiduchidae which can be diagnosed e.g. by forewing venation.The vein CuA forks at the nodal line that does not include the apex of the clavus.The free basal part of CuA2 is short, curved before merging into the postclaval margin of the tegmen.This fusion results in cell C5 being delimited by icu, CuA1 and the postclaval margin.This might represent an apomorphy for the genus, while a similar condition is approached, but probably independently due to tegmina reduction, in some brachypterous tropiduchid genera such as Durium Stål, 1861 and Neommatissus Muir, 1913. C5 is also apically open and closed by the postclaval margin in Trypetimorpha Costa, 1862 and Ommatissus Fieber, 1875.In Paricana and Dichoneura the transverse veinlet uniting CuA2 and the postclaval margin is present as in most other tropiduchids, and the branch CuA2 does not fuse with the postclaval margin, reaching the margin as a normally developed terminal.

Chirality of male genitalia
Asymmetry of genitalia might have important implications for understanding genital evolution and sexual selection (Eberhard, 2010).It is common in several different groups of insects and a consequence of convergent evolution from a symmetric situation (Huber et al., 2007;Huber, 2010;Schilthiuzen, 2007).In planthoppers, it has also been mentioned in Delphacidae, which exhibit almost exclusively an asymmetric morphology (Guglielmino & Bückle, 2010).In Tropiduchidae, asymmetry is widely found in the male aedeagus, periandrium, pygofer and gonostylus, and in the female gonocoxae VIII and sternite VIII (Wang et al., 2009).Asymmetry of genital structures (genital styles) was mentioned as a feature of the tribe Tropiduchini by Fennah (1982).In Pseudoparicana, asymmetry concerns the male aedeagus, pygofer and gonostyli (Figs 3F-K, 4A-F, 5A-F, 7A-F), and the female gonocoxae VIII (Figs 4G, 6D-E).
Interestingly, in this genus, left-or right-handed asymmetry (chirality) of the male genitalia occurs: the male genitalia of P. analoga, P. curvifera and P. sanguinifrons are "left-handed" while that of P. tepida are "righthanded".Within the Tropiduchidae, the chirality of the male genitalia seems also to be a diagnostic character at the species level in some other genera, e.g.Daradax Walker, 1857, Tropiduchus Stål, 1854, Vanua Kirkaldy, 1906and Varma Distant, 1906.However, in some Delphacidae, chiral dimorphism (right or left anti-symmetry of aedeagus) can occur within a single species (Guglielmino & Bückle, 2010).
of the Czech Republic (grant no.MK000094862, awarded to the Moravian Museum, Brno).

Fig. 1 .
Fig. 1.Geographical distribution of Pseudoparicana species in New Guinea and adjacent islands.
), gonostyli broad, without process on right side laterally, apex truncate.Aedeagus situated left of gonostyli (Figs 4A-D), asymmetrical, tube-like; in left lateral view (Figs 4B, E), distal part slightly expanded; in dorsal view (Figs 4A, F), sub-apical part on right side with two spine-like processes, directed laterad and dorsad, respectively (Figs4A, E-F); apex on left dorsal side with one large twisted process, directed ventrolaterad; endosoma with a large spine-like process; phallotreme (Figs4A, E-F) trumpet-like, running along the apex of aedeagus, forming a distinct, wide opening.