Journal of Natural History, 2002, 36, 601–616

New taxaof Vizcayinae ( :Auchenorrhynch a:), includinga remarkablenew genus from China

AI-PING LIANG Department ofEntomology,Institute ofZoology, Chinese Academy of Sciences,19 Zhongguancun Lu, Beijing 100080, P.R. China; e-mail: [email protected]

(Accepted10 November 2000 )

Neovizcayasinica gen.and sp. nov., the second genus of thedelphacid subfamily Vizcayinae,is described and illustrated from Yunnan, south-west China. A key tothe nine species of Vizcaya (theother genus of the subfamily) is provided, includingthe following four new species, which are described and illustrated: V. aschei sp.nov. (south India), V.latifrons sp.nov. ( Taiwan), V.longispinosa sp. nov.(south-west China: south Yunnan) and V.lombokensis sp.nov. ( Indonesia: Lombok).The new genus Neovizcaya isof special taxonomic and phylogenetic interestas it di Œers from Vizcaya inmany characters. Based on thesedi Œerences andfrom studies of theantennal sensilla, a revisedde® nition of thesubfamily is given.Micrographs of the antennae, egg, rostral apex and pretarsus of the subfamilyare provided for the ® rsttime.

Keywords: Asia,Delphacidae, Vizcayinae, new genus, new species, ultra- structure,antennal sensilla, egg, rostrum, pretarsus.

Introduction The subfamily Vizcayinae( Hemiptera: Delphacidae) wasestablished byAsche (1990)for the genus Vizcaya Muir with ®vespecies from southern India and South- EastAsia. Vizcayinae is of considerable interest within the Delphacidae asit is regarded asaphylogenetic link between the Asiracinae and the remaining delphacids. While sorting and identifying Delphacidae from materialin the Collections of the Institute of Zoology,Chinese Academy of Sciences, Beijing( IZCAS),aseries of bizarre specimens, collected atKunming, Yunnan, south-west China,were found that represent aremarkable newgenus and species of the Vizcayinae.The newgenus Neovizcaya gen. nov. diŒers from Vizcaya in manycharacters, necessitating changes in the de® nition of the subfamily Vizcayinae.In addition, four newspecies of the Vizcaya werealso found from southwestern China(southern Yunnan), Taiwan, Lombok and southern India. These discoveries havegreatly broadened our know- ledgeon the morphology, phylogenyand biogeography of the Vizcayinae.No information is currently availableon the biologyof these although most

Journalof Natural History ISSN0022-2933 print /ISSN1464-5262 online Ñ 2002Taylor & FrancisLtd http://www.tandf.co.uk /journals DOI:10.1080 /00222930110062327 602 A.-P. Liang delphacids feed from the sap of Monocotyledones from which they feed close to the soil.

Materials andmethods The specimens studied in the course of this work aredeposited in the following institutions whose names areabbreviated in the text asfollows: BMNH, The Natural History Museum (formerly British Museum (Natural History)), London, UK; BPBM,Bernice P.Bishop Museum, Honolulu, Hawaii,USA; CAS, California Academy of Sciences, San Francisco, CA,USA; HUS, Laboratory of Systematic Entomology, Hokkaido University, Sapporo, Japan;IZCAS, Institute of Zoology, Chinese Academy of Sciences, Beijing,P.R. China;MNHU, Museum fu È r Naturkunde der Humboldt-UniversitaÈt, Berlin,Germany. For SEM studies, dry pinned specimens wereemployed. Specimens wereplaced and cleaned in 10%KOH for 20min, then washed in distilled watertwice, mounted on aluminium stubs bydouble-sided sticky tape, air-dried atroom temperature, and coated with gold-palladium using asputter coater. Observations weremade with a Zeiss DSM 950or aJEOL5200LV scanning electron microscope, operated at acceleratingvoltages of 10±25kV. Morphological terminology follows that of Asche (1985,1990 )and Lewisand Marshall (1970)and Marshall and Lewis( 1971)(antennal sensilla).

Checklistof Vizcayinae VizcayinaeAsche, 1990 Neovizcaya gen. nov. sinica sp. nov.Ð SWChina( Yunnan). Vizcaya Muir, 1917 adornata Asche, 1990ÐIndonesia (Sulawesi). aschei sp. nov.Ð south India (Madras). bakeri Muir, 1917ÐPhilippines (Luzon). latifrons sp. nov.Ð Taiwan. lombokensis sp. nov.Ð Indonesia (Lombok). longispinosa sp. nov.Ð SW China( Yunnan). orea Asche, 1990ÐIndonesia (Sumatra), Thailand( Doi Suthep, Chiangmai), Vietnam( BanMe Thuot). piccola Asche, 1990ÐMalaysia(Sarawak). vindaloa Asche, 1990Ðsouth India (Travancore).

SubfamilyVIZCAYINAE Asche VizcayinaeAsche, 1990: 157. Type genus: Vizcaya Muir,1917: 351. Includedgenera. Vizcaya Muir (nine species), Neovizcaya gen.nov. (one species). Distribution. South India, south-west Chinaand South-East Asia. Remarks. The taxonomic position of Vizcayinaeis problematical. Asche’s (1990)phylogenetic analysisof the Delphacidae indicated that the Vizcayinaeformed alink between the Asiracinae and the remaining Delphacids (Kelisiinae, Stenocraninae, Plesiodelphacinae and Delphacinae). However,based on characters of the nymph, Emeljanov( 1995)downgraded the Vizcayinaeto atribe of Delphacinae. Emeljanov( 1995)also questioned the polarity of anautapomorphy Vizcayinaedelphacids 603 proposed byAsche (1990)Ðthe arrangement of teeth on the tip of the basalsegment of the metatarsi (see below). In the current work, Asche’s(1990)subfamily placement of Vizcayinaeis followed pending further work on the group. Vizcayinaecan be distinguished from allother delphacid subfamilies bythe following combination of characters (from Asche, 1990with newcharacters in italics): vertexwith aninverted `v’-shaped carina(® gure 44);antennal scape and pedicel veryor moderately elongate, scape compressed (® gures 10,15, 46, 49 )[similar to Asiracinae];antennal pedicel with numerous sensory plaque organs, irregularly arrangedover the whole surface (® gures 10,15, 16 )[similarto Asiracinae:Ugyopini and some Delphacinae]; sensoryplaque organs on pedicel surface with relatively few (8±12)denticlesand relatively few (17±28 )digitateprocesses (® gures 11,17 ); dent- icles relativelylarge and broad, compressed and curved inwardly, nearly pyramidical withacute apex, outer edge curved and distinctly keel-like, thedepressed surface with oneto three indistinctlongitudinal ridges (® gures 11,17 )[similarto Asiracini (see Shih and Yang,1996; Liang, 1998 )]; basal pit of Xagellumtraversed bya cuticular spinoseprocess (® gure 14);metatibial spur conical inshape, with 5±12teeth (includ- ingapical tooth) on inner margin(® gures 18,20 )[similarto Kelisiinae, Plesiodelphacinae and some Delphacini];apicalpecten row of the 1st metatarsomere with the median spine smaller than the remainder and displaced proximally(arrowed in ®gure 53);forewings elongate with veins prominent and thickly covered with granules each bearinga long seta [similarto Asiracinae];2nd abdominal tergite of maledrumming organwith adeep central depression (arrowed in ®gure 19). One of the characters used byAsche (1990)to support the monophyly of the Vizcayinaeis absent in Neovizcaya.This is the smoothly rounded fore marginof the head in Vizcaya which is strongly compressed in Neovizcaya. The two Vizcayinaegenera can be distinguished bythe characters noted in table 1.

Descriptionof taxa

Neovizcaya gen. nov. Typespecies: Neovizcayasinica sp. nov. Small,slender, somewhat dorsoventrally ¯attened delphacid. Head relativelylarge and ¯at,longer and narrower than pronotum, gradually tapered from base to apexin lateralaspect. Vertexlong, mediallyabout 1.6 3 longer than broad atbase, about 1.4 3 length of pronotum, distinctly extending in front of eyes,broad atbase and converginganteriorly, apexsomewhat broad, posterior marginnearly straight, lateralmargins laminately carinate and graduallyconverging from base to apexand divergingat mid-length to lateralmargins of frons, disk of vertexmuch depressed, sloping to apex,with anindistinct `Y’-shaped ridge on posterior half. Frons large,smooth, apexrounded in ventralaspect, gradually broadened from upper to lower part, widest slightlyabove frontoclypeal suture, about 1.43 higher than maximum width, upper part much depressed, lower part distinctly convex,lateral margins carinate, without mediallongitudinal carina. Postclypeus much narrower than frons, convexand smooth, lateralmargins slightly carinate,without medialcarina. Anteclypeus convex,without lateraland medial carinae.Rostrum long, reachingbeyond hind trochanters. Compound eyesin lateral viewelongate, kidney-shaped, distinctly incised mediallyabove antennal base from inferior marginto about one-half height of eye.Antennae relativelylong (but much 604 A.-P. Liang

Figs 1± 9. Dorsalhabitus of Neovizcaya and Vizcaya species:( 1,2) N. sinica ( ,paratypes, China:Yunnan ( IZCAS));(3,4 ) V.longispinosa ( ,paratypes,China: Y unnan (IZCAS));(5) V.latifrons ( ,holotype,T aiwan( HUS));(6) V.lombokensis ( , holotype,Lombok ( HUS));(7) aschei ( ,holotype,S. India:Madras (CAS ));(8,9 ) V. vindaloa Asche ( ,S.India:Madras (CAS )). Vizcayinaedelphacids 605

Figs 10± 18. Externalmorphology of Vizcayinae(SEM ).(10±14 ) Neovizcayasinica sp. nov., (10)leftantenna of male, frontal view; ( 11)asensoryplaque organ on pedicel; ( 12) pittedcuticular process of a plaqueorgan; ( 13)swollen¯ agellarbase; ( 14)topopening ofswollen ¯ agellarbase, dorsal view; ( 15±18) Vizcayalongispinosa sp.nov., ( 15)right antennaof male, frontal view; ( 16)partof pedicel; ( 17)asensoryplaque organ on pedicel;( 18)distalend of metatibia and basal part of metatarsomereI. shorter in comparison with Vizcaya)with apexof 2nd segment reachingonly base of forewings in respose; 1st segment shorter than median length of vertex,depressed, distinctly concaveon inner side; 2nd segment terete, about 1.7±1.8 3 longer than 1st,with more than 20sensory ®leds irregularlydistributed over distal surface; both 1st and 2nd segments with numerous sturdy bristles on surface. Pronotum relatively large,shorter than head, distinctly separated from head; disk convex,strongly tricarinate, anterior marginnearly straight, posterior marginangularly incurved; strongly sloping lateradand curved down, extreme lateralareas strongly foliaceous and somewhat re¯exed. Mesonotum with anterior marginprojected under posterior marginof pronotum, disk relatively¯ atand strongly tricarinate, sloping laterad, caudaltip concaveand faintlytransversely striate basally.Forewings (® gure 36) elongate,nearly parallel-sided, surpassing abdomen byabout one-third of total length and 3.9 3 longer than maximum width, widest slightlydistad of nodal line, 606 A.-P. Liang

Figs 19± 27. Externalmorphology of Vizcayinae (SEM ).(19±23) Neovizcayasinica sp. nov., (19)malemetathorax and abdominal segments I± III, dorsal view; ( 20)distalend of metatibiaand basal part of metatarsomere I; (21)egg;( 22)pretarsusof hind leg; ( 23) apexof rostrum; ( 24±27 ) Vizcayalongispinosa sp.nov., ( 24)apexof rostrum; ( 25±27 ) pretarsusof hind leg. narrowest atbasal one-third; veins prominent and thickly covered with granules eachwith along seta; costal membrane without transverse veins,Sc and Rand Cu1aand Cu1bbranching near mid-length of wingwith outer and inner anteapical cells relativelylarge; subapical transverse nodal line relativelystraight. Hindwings wider than forewings, venation asin ®gure 39.Legs moderately elongate and slender, fore and middle coxaelong, smooth and somewhat concavein ventralview, fore femora and tibiae normal, not foliaceously dilated; hind tibiae with two lateralteeth, Vizcayinaedelphacids 607

Figs 28± 39. Neovizcayasinica sp.nov. ( 28)maleholotype; ( 29)head,ventral view; ( 30) head,lateral view; ( 31)distalend of metatibia and metatarsus; ( 32)malegenitalic capsule,ventrocaudal view; ( 33)aedeagus,dorsal view; ( 34)secondabdominal apodeme ofmale, caudal view; ( 35)rightparameres, ventrolateral view; ( 36)leftforewing; ( 37) femalegenitalia, ventral view; ( 38)malepygofer, lateral view; ( 39)righthindwing. the basalone near the base, verysmall and indistinct, the distal one beyond the middle prominent, metatibial spur conical, with ®veteeth (including apicaltooth) on inner margin(® gures 20,31 ),metatarsomere Ilong, about 1.1 3 longer than metatarsomeres II 1 III, black-tipped spines atend of hind tibiae and metatarsomeres Iand IInumbering 5,5 and 4,respectively; pretarsus with one long seta on each unguis and one pairof relativelylong setae on arolium (® gure 22). Drumming organsexually dimorphic: in females asin Vizcaya;inmalessecond abdominal tergite dilated, convex,with ashallowoval depression centrally(arrowed in ®gure 19);metapostnotum with apair of veryslender apodemes directed caudad; 2nd abdominal sternite with apairof elongate apodemes directed dorsocaudad (® gure 34). 608 A.-P. Liang

Figs 40± 53. Externalmorphology of Vizcaya species.( 40,41 ) V.longispinosa sp.nov., ( 40) head,ventral view; ( 41)head,lateral view; ( 42±48 ) V.latifrons sp.nov., ( 42)head, ventralview; ( 43)head,lateral view; ( 44)vertex,dorsal view; ( 45)femalegenitalia, ventralview; ( 46)leftantenna, frontal view; ( 47)leftforewing; ( 48)metatibiaand metatarsus;( 49±51 ) V. vindaloa Asche,( 49)antenna;( 50)leftforewing; ( 51)metatibia andmetatarsus; ( 52,53 ) V.lombokensis sp.nov., ( 52)leftforewing; ( 53)metatibiaand metatarsus.

Malegenitalia with pygofer (® gures 32,38 )in lateralview trapezoid, in caudal viewcircular, ventrallyabout 1.3 3 longer than dorsally,laterodorsal edges strongly produced caudad in lateralview, without teeth on dorsal margin,laterocaudal Vizcayinaedelphacids 609

Table1. Comparisons between Vizcaya and Neovizcaya.

Vizcaya Neovizcaya

Body size Large,body length 4.9± 6.5 mm Small, body length 3.8± 4.8 mm Body form Subcylindrical Dorsoventrallycompressed Head Narrow,a littleprotruding in Very broad, well protrudingin frontof eyes frontof eyes Vertex Normal Stronglydorsoventrally compressed Vertexcarinae Very® neandweak, without `Y’ -Strongand distinct, with an shapedridge on posterior half indistinct `Y’ -shapedridge on posteriorhalf Frons Narrow Very broad Antenna Extremelyelongate, apex of Moderatelyelongate, apex of pedicelreaching basal one-third pedicel reaching base of toone-half of the costal margin forewings in respose offorewings in respose Pronotumand Narrow,disk arched, lateral Broad, relatively ¯ at,lateral mesonotum marginsnormal; median and margins laminate and re¯ exed; lateralcarinae very weak and median and lateral carinae very indistinct strongand very distinct Number of 6± 12 5 metatibialspur spines Egg Withoutring-like hatching cap at With ring-like hatching cap at apex apex Distribution S.India,SW China( Yunnan),SW China( Yunnan) SEAsia,Philippines ( Luzon)and Indonesia(Sumatra, Sulawesi, Lombok)

marginsbroadly rounded to diaphragm,median processes on ventrocaudal margin small,acute, widelyseparated; ventralangles of pygofer acute, tooth-like in lateral view,distance between ventralangles great; anal tube short and broad, lateroapical anglesproduced into short, ventrallydirected process. Anal style smalland short. Parameres (® gure 35)in ventralview arched, basallyexpanded and broad in inner side, graduallytapering to apex,apex with averysmall laterally recurving hooked barb, nearlybeak-shaped in lateralview. Aedeagus (® gure 33)with shaft elongate, slender, slightlydepressed ventrodorsally, graduallytapered from base to apex,with one elongate,sinuate process directed ventrallyand two relativelyshort and acute spinose processes apicallyon right side, and one short and acute spinose process subapicallyon left side. Femalegenitalia as in ®gure 37.Eggs (® gure 21)much smaller than those of Vizcaya with distinct ring-like hatching capat apexÐ Asche (1990:175 )reported that no ring-like hatching capwas seen in the eggsof Vizcaya. Etymology. The generic name is derived from Greek neo-,new,and the existing genus name Vizcaya.Gender: feminine. Includedspecies and distribution. One species; south-west China( Yunnan).

Neovizcayasinica sp. nov. (® gures 1,2, 10± 14, 19± 23, 28± 39 ) 610 A.-P. Liang

Length (from apexof vertexto tip of forewings): 3.8± 4.2 mm; 4.5± 4.8 mm; length of 1st 1 2nd antennal segments 1.2±1.3mm. Vertexpale yellowish, posterior areasometimes with brownish su Œusion, lateral carinaebrown; frons, postclypeus, anteclypeus, genaeand lora paleyellowish, anterior marginand upper part of lateralmargins of frons, sides in front of and abovecompound eyes(® gure 30),and lora in female, brown; antennae brown, distal half of 2nd segment blackish brown, bristles paleyellowish brown; rostrum pale yellowish,apex blackish brown; pronotum dark brown or blackish brown, lateral areaspale yellowish, disc sometimes yellowishbrown; mesonotum shining blackish brown; forewings blackish brown, costa yellowishbrown, abroad oblique transverse fascianear base, anelongate longitudinal subapical costal spot which is posteriorly crossed bya blackish brown fascia,and asmallirregular spot near apexof clavus whitish and subhyaline; granualeson veins brown, setae on veins paleyellowish brown; hindwings hyalinewith veins brownish; thorax beneath paleyellowish, lateral areasof meso- and metapleurae dark brown; legspale yellowish, claws and metatar- somere III, base of coxae,base and distal two-thirds of fore tibiae,base of middle tibiae,inner side of hind femora and base of hind tibiae brown or su Œused with brown, tips of lateraland apicalteeth on hind tibiae and tarsi, and tips of lateral and apicalteeth of metatibial spurs black; abdomen brown or dark brown, posterior segmental margins,ventrolateral areasof tergites, and acentral longitudinal band on sternites yellowishbrown or reddish brown; female genitaliawith valvifersVIII and lateralgonapophyses IXyellowish,with brownish tinge. Maleand female genitaliaas in generic description (® gures 32,33, 35, 37, 38 ). Etymology. This newspecies is named for its occurrence inChina. Materialexamined. Holotype , China:Yunnan Province, Kunming, Heilongtan, 1900m, 24August 1958( Y.-R. Zhang)(IZCAS). Paratypes: China: 17 , 8 ,same dataas holotype (15 , 6 in IZCAS; 2 , 2 in BMNH; 1 , 1 in MNHU ). Distribution. South-west China( Yunnan).

Genus Vizcaya Muir Vizcaya Muir,1917: 351; Metcalf, 1943: 68; Asche, 1990: 158. Type species: Vizcayabakeri Muir,1917, by monotypy.

Vizcaya comprises averyuniform group of delphacids which canbe easily distinguished bythe following combination of characters: body slender in appearance with legsand forewings long; antennae with scape and pedicel both strongly elongate, scape slightlyor welldilated, pedicel terete with numerous sensory ®elds, which are irregularlyarranged over the whole surface (® gures 10,15, 16 );legsvery elongate and slender, hind legspine formula 5:5:4,metatibial spur conical in shape, with 6±12 teeth (including apicaltooth) on inner margin;head smalland narrow, trans- ition of vertexto frons smooth and rounded; head and pronotum predominantly ochraceous with vertex,upper half of frons, postclypeus and lorashining black, forewings (® gures 47,50 )usuallywith alarge,distinct fuscous patch atapical area beyond nodal line (excluding costal area)and with or without ashort fuscous stripe which branches from the fuscous patch and runs to costal margin,clavus and basal costal areausually marked with fuscous; abdomen usuallyblackish. For details of the antennal sensilla,egg, rostrum and pretarsus (shown for the ®rst time) (see ®gures 10±17,21 ±27). Vizcayinaedelphacids 611

Includedspecies and distribution. Nine species; S.India, Philippines (Luzon), Indonesia (Sulawesi, Lombok (new record)),Malaysia(Sarawak), Thailand, Vietnam,SW China(south Yunnan) (new record),Taiwan(new record).

Keyto species of Vizcaya 1Firstantennal segment well dilated,relatively broad and short (® gures 15, 46 )..2 ±Firstantennal segment slightly dilated, relatively slender and elongate (® gure 49 )..5

2Anteclypeusrelatively broad basally (® gure 42 );Taiwan. ... latifrons sp. nov. ±Anteclypeusrelatively narrow basally (® gure 40 )...... 3

3Aedeaguswith apical left spinose process elongate and slender, reaching almost one- halflength of theca (® gure 59 );SWChina( Yunnan). ... longispinosa sp. nov. ±Aedeaguswith apical left spinose process relatively short and stout, reaching almost one-thirdlength of theca...... 4

4Forewingswith apical fuscous patch relatively elongate, without fuscous stripe at apical costalarea (see Asche, 1990: ® gure24 );aedeagalshaft with apical spinose processes relativelyshort and slender (see Asche, 1990: ® gure31 );metatibialspur with six (left)1 eight(right )teeth;Philippines ( Luzon)...... bakeri Muir ±Forewingswith apical fuscous patch relatively short, with short fuscous stripe at apical costalarea (see Asche, 1990: ® gure33 );aedeagalshaft with apical spinose processes relativelyelongate and broad (see Asche, 1990: ® gure43 );metatibialspur with 9± 11 teeth;Indonesia ( Sumatra),Thailand ( DoiSuthep, Chiangmai ),Vietnam( BanMe Thuot)...... orea Asche

5Costalarea of forewings with a longitudinalfuscous stripe ...... 6 ±Costalarea of forewings hyaline ...... 7

6Forewingswith apical fuscous patch relatively elongate and broad, fuscous stripe at apicalcostal area short, costal area with narrow longitudinal fuscous stripe, clavus withone complete elongate fuscous stripe (see Asche, 1990: ® gure7 );metatibialspur withseven teeth; Indonesia (Sulawesi )...... adornata Asche ±Forewingswith apical fuscous patch small and short, with fuscous stripe at apical costalarea very long, costal area with broad longitudinal fuscous stripe connecting theapical fuscous patch, clavus with two disjunct patches: one basal and another subapical(see Asche, 1990: ® gure46 );metatibialspur with six teeth; Malaysia (Sarawak)...... piccola Asche

7Bodyextremely elongate (® gure 6 );forewingswith apical fuscous patch very narrow andslender, occurring less than one-third of the apical area, clavus without any brown suŒusion(® gure 52 );aedeaguswith two short apical processes, equal in length, expandedsubapically and acute apically (® gure 74 );rightmetatibial spur with six teeth;Indonesia ( Lombok)...... lombokensis sp. nov. ±Bodymoderately elongate; forewings with apical fuscous patch relatively large occur- ringmore than one-third of the apical area, clavus with distinct brown su Œusion; aedeaguswith two or three narrow apical processes of unequal length (® gure 64 ); metatibialspur with nine or more teeth ...... 8

8Forewingswith apical fuscous patch relatively broad and large (® gures 8, 9, 50 ); pygoferwith laterodorsal edges with one tooth subapically on dorsalmargin (® gure 66 ); aedeagalshaft with three stout apical spinose processes (® gure 69 );metatibialspur with9 (left) 1 12(right )teeth;south India ( Travancore). ... vindaloa Asche ±Forewingswith apical fuscous patch narrow and slender (® gure 7 );pygoferwith laterodorsaledges without teeth subapically on dorsal margin (® gure 61 );aedeagal shaftwith two very slender apical spinose processes (® gure 64 );metatibialspur with nineteeth; south India ( Madras)...... aschei sp. nov. 612 A.-P. Liang

Vizcayaaschei sp. nov. (® gures 7,60± 64 ) Length: 6.0 mm. Externallysimilar to V. vindaloa Asche (see description of vindaloa given by Asche, 1990:174 )but forewings (® gure 7)with the apicalfuscous patch verynarrow, slender, evenlybroad and nearlyC-shaped, the apicalmargin and the apicalinner marginhyaline, and the fuscous longitudinal stripe in clavusbetween common stem of analveins and inner marginvery short, smalland nearlytriangular and metatibial spur with nine teeth (including apicaltooth) on inner margin. Malegenitalia with pygofer (® gures 60,61 )short and high, long ventrallyand short dorsallyin lateralaspect, laterodorsal edges relativelybroadly produced caudad, without teeth subapicallyon dorsal margin;median projection of ventrocau- dalmargin (® gure 63)moderately long, slightlyV-shaped distallyin ventralaspect; analtube short, broad, somewhat rounded, and apicalmargin incised mediallyin dorsal view,anal styles veryshort, smalland slender (® gure 62);parameres (® gure 60)relativelynarrow and slender, nearlyS-shaped inlateralaspect, distinctly broad subbasally,apical one-third evenlybroad, apexslightly acute; aedeagus (® gure 64)with shaft long and slender, depressed ventrodorsally, with two slender, ventrallydirected, spinose processes apically,one relativelylong, reachingnear mid- length of the shaft, the other short. Etymology. This newspecies is named in honour of Dr Manfred Asche, Museum fuÈ rNaturkunde der Humboldt-UniversitaÈtzuBerlin,Germany for his very valuablecontribution to the world delphacid systematics and for whose pioneering work on the Vizcayinaehas made this work possible. Distribution. Southern India. Materialexamined. Holotype , South India:Madras State, AnamalaiHills, Cinchona, 1067m, May1959 ( P.S.Nathan) (CAS). Paratype: South India: 1 , Madras State, NilgiriHills, Devala,675 m, May1961 ( P.S.Nathan) (CAS). Remarks. This species is similarto another Indian species, V. vindaloa, but can be distinguished bythe characters noted in the abovedescription.

Vizcayalatifrons sp. nov. (® gures 5,42± 48 ) Female. Length: 6.7mm. Externallysimilar to V. orea (see description of orea givenby Asche, 1990:168 ) but discalareas of pro- and mesonotum more ochraceous, anteclypeus relatively

Figs 54± 74. Malegenitalia of Vizcaya species.( 54±59 ) V.longispinosa sp.nov., ( 54)genital capsule,ventrocaudal view; ( 55)pygofer,lateral view; ( 56)parameres,ventral view; (57)analsegment, dorsal view; ( 58)medianprojection of ventrocaudal margin of pygofer,ventral view; ( 59)aedeagus,left dorsolateral view; ( 60±64 ) V. aschei sp. nov., (60)genitalcapsule, ventrocaudal view; ( 61)pygofer,lateral view; ( 62)analsegment, dorsalview; ( 63)medianprojection of ventrocaudal margin of pygofer, ventral view; (64)aedeagus,dorsal view; ( 65±69 ) V. vindaloa Asche,( 65)genitalcapsule, ventrocau- dalview; ( 66)pygofer,lateral view; ( 67)analsegment, dorsal view; ( 68)median projectionof ventrocaudalmargin of pygofer, ventral view; ( 69)aedeagus,dorsal view; (70± 74) V.lombokensis sp.nov., ( 70)genitalcapsule, ventrocaudal view; ( 71)pygofer, lateralview; ( 72)analsegment, dorsal view; ( 73)medianprojection of ventrocaudal marginof pygofer, ventral view; ( 74)aedeagus,dorsal view. Vizcayinaedelphacids 613 614 A.-P. Liang broad basally,forewings with the brown longitudinal stripe in clavusbetween common stem of analveins and inner marginof wingshort and distinct only in basalone-half, and metatibial spur with seven teeth (including apicaltooth) on inner margin. Male. Unknown. Etymology. This species is named for its relativelybroad frons. Materialexamined. Holotype , Taiwan:[underside] Horisha [in Japanese], 30April [19]07;Matsumura (HUS). Distribution. Taiwan.This is the ®rst report of Vizcaya species from Taiwan. Remarks. This species canbe distinguished from allother known species in the genus byits 1st antennal segment being welldilated and relativelybroad and short, pro- and mesonotum with disc areasmore ochraceous, relativelybroad anteclypeus, forewings with broad fuscous patch atapical area and short fuscous stripe atapical costal areaand metatibial spur with seven teeth (including apicaltooth) on inner margin. Basedon the antennae this species forms agroup with longispinosa sp. nov., bakeri Muir and orea Asche, allhaving well dilated and relativelybroad and short 1st segment. The discovery of amaleand study of the malegenitalia of this species areparticularly desirable to determine its relationship to other species.

Vizcayalombokensis sp. nov. (® gures 6,52, 53, 70± 74 )

Length: 5.7 mm. Externalcharacters and colour asin V. adornata Asche (see description of adornata givenby Asche, 1990:160 )but di Œers in its smaller size, narrower body, base and basallateral margins of vertexand central and posterior areasof pronotum paleochraceous, forewings with apicalfuscous patch greatlyreduced, verysmall, narrow and slender, occurring over less than one-third of the apicalarea, the fuscous stripe atapical costal areavery long, clavusand costal marginwithout distinct fuscous suŒusion (® gure 52)and the right metatibial spur with sixteeth (including the apicaltooth) (left hind legmissing ). Malegenitalia with pygofer (® gures 70,71 )distinctly long ventrallyand very short dorsallyin lateralaspect, laterodorsal edges distinctly produced caudad, with one distinct, smalltooth subapicallyon dorsal margin;median projection of ventro- caudalmargin of pygofer narrrow, nearlyV-shaped distallyin ventralaspect (® gure 73);paramere (® gure 70)relativelyevenly broad, graduallytapered from subbasal part to apex;aedeagus (® gure 74)with shaft expanded mesially,with two short, equal-lengthed, relativelybroad, ventrallydirected, apicalspinose processes, both processes expanded subapicallywith apexacute. Etymology. This species is named for its occurrence in Lombok. Materialexamined. Holotype , Indonesia:Lombok, Sapit, 2000ft, April 1896( H.Fruhstorfer), [Matsumura’shandwriting] Hygops [sic] curvfascia det. Matsumura (HUS). Distribution. Indonesia (Lombok). This is the ®rst report of Vizcaya species from Lombok. Remarks. This species di Œers from other species byits more slender body (® gure 6).In other respects it is similarto V. adornata Asche but diŒers in the characters noted in the abovedescription. Vizcayinaedelphacids 615

Vizcayalongispinosa sp. nov. (® gures 3,4, 15 ±18,24 ±27,40, 41, 54 ±59) Length: 6.4 mm; 6.5± 6.7 mm Externallysimilar to V. orea Asche (see description of orea givenby Asche, 1990:168 ).Metatibial spur with 8±11 teeth (including apicaltooth) on inner margin (number of teeth varyingamong individuals and on left and right legs). Malegenitalia with pygofer (® gures 54,55 )in lateralview trapezoid, in caudal viewcircular, ventrallyabout 1.5 3 longer than dorsally,laterodorsal edges relatively broadly produced caudad, with three smallteeth subapicallyon dorsal margin; distance between ventralangles relatively small; median projection of ventrocaudal marginrelatively long, comparativelynarrow, distallywith nearlyU-shaped incision in ventralview (® gure 58);analtube relativelylarge and broad, expanded laterad mesially,lateroapical angles not produced into ventrallydirected processes; anal style slender; parameres (® gure 56)relativelybroad, sinuous and slightlyS-shaped in lateralaspect, graduallytapering from base to apex,apical part acute in lateral view;aedeagus (® gure 59)with shaft elongate,slender, depressed ventrodorsally, arched in lateralaspect, with two long, slender, ventrallydirected, spinose processes apicallyon left and dorsal side: left process long, longer than one-half total length of theca, dorsal spine base bifurcate. Etymology. This species is named for its relativelylong apicalspinose processes on the aedeagus. Materialexamined. Holotype , China:Yunnan, Xishuangbanna, Menga, 1050±1080 m, 6June 1958(S.-Y. Wang)(IZCAS). Paratypes: China: 7 , Yunnan, Xishuangbanna, Yunjinghong, 850m, 25June 1958( L.-Y. Zheng)(5 in IZCAS; 2 in BMNH ). Distribution. South-west China( Yunnan). This is the ®rst report of Vizcaya species from mainland China. Remarks. This species issimilarto V. orea but diŒers in havingthe metatibial spur with 8±11 teeth (including apicaltooth) on inner marginand the aedeaguswith relativelyslender apicalspinose processes with the left one long, longer than one- half total length of theca.

Vizcayavindaloa Asche (® gures 8,9, 49± 51, 65± 69 ) Vizcayavindaloa Asche,1990: 174, ® gures47± 49. Holotype ,India (BMNH)[examined]. This species wasdescribed and illustrated byAsche (1990)from asinglefemale specimen taken atsouth India. An additional maleand female specimen from southern India werefound during this work. The maledi Œers from the female in its smaller size (6.2mm) and in havingits ®rst antennal segment slightlymore elongate (compare ®gure 49with Asche, 1990:® gure 48).The second antennal segment (missing from the type described byAsche), isshown in ®gures 8,9, 49. Malegenitalia with pygofer (® gures 65,66 )with laterodorsal edges with one smalland short tooth subapicallyon dorsal margin;median projection of ventrocau- dalmargin (® gure 68)short, and relativelynarrow, nearlyV-shaped distallyin ventralaspect; parameres (® gure 65)distinctly broad subbasally,tapered to relatively broad apex;aedeagus (® gure 69)with three spinose processes subapically,the longest one extending nearlyto middle of shaft, alittle expanded beyond middle and then 616 Vizcayinaedelphacids tapered to acute apex,the middle-lengthed one expanded subapicallywith acute apex,the shortest one acute. Materialexamined. Holotype , India:Thekkadi, PeriyarDam, Travancore, 6±10 May[ 19]37[underside] B.M.-C. M.Expdn. to South India. April± May1937; [red label ] Holotype Vizcayavindaloa n. sp. det. M.Asche 1990( BMNH). South India: 1 , 1 ,Madras State, AnamalaiHills, Cinchona, 1067m, May1959 (P.S.Nathan); 1 [abdomen missing],Cherangode, NilgiriHills, 3500ft, October 1950( P.S.Nathan) (CAS). Distribution. South India. Remarks. This species is similarto another Indian species, V. aschei (see remarks under aschei).

Acknowledgements Iamgrateful to the following individuals and institutions for loans of specimens or access to collections: Mr DavidPreston, Drs Godon Nishida and Scott Miller (BPBM),Dr Norman Penny (CAS)and Drs MasaakiSuwa and Masahiro Ohara (HUS).Ithank Ms PelingFong-Melville, Interdepartmental Laboratory,American Museum of Natural History, NewYork and Mr Mike Turner, University of Wales, CardiŒ,UK, for assistance with the scanning electron microscopy. Iwish also to thank Mr Mick Webb (BMNH)and Dr Manfred Asche, Museum fu È rNaturkunde der Humboldt-UniversitaÈtzu Berlin,Germany, for reviewingthe manuscript. The work on which this paper is based wassupported bythe National Natural Science Foundation of Chinagrant numbers 39925006and 39770115,the Chinese Academy of Sciences BiologicalInnovation Fund C2999084,and the Theodore Roosevelt Memorial Fund, Postdoctoral Fellowship Program, American Museum of Natural History, NewY ork.

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