The Delphacid Genus Sogatella and Related Groups: a Revision with Special Reference to Rice-Associated Species (Hornoptera: Fulgoroidea)

Systematic Entomology (1990) 15, 1-42

The delphacid genus Sogatella and related groups: a revision with special reference to rice-associated species (Hornoptera: Fulgoroidea)

MANFRED ASCHE and MICHAEL R. WILSON* Fachbereich Biologie der Philipps Universitat, Marburg, West Germany, and *CAB International Institute of Entomology, c/o Department of Entomology, British Museum (Natural History), London, U.K.

ABSTRACT. The delphacid planthopper genus Sogatella Fennah is rede- fined and a key provided to males of the fourteen included species. The type-species of Sogatodes Fennah, S.molinus Fennah is considered to be a Sogatella species and consequently Sogatodes becomes a junior subjective synonym of Sogarella. The remaining species of Sogatodes are transferred to Tagosodes gen.n. (type species T.cubanus (Crawford) comb.n.), Latis- tria Huang et al. or Sogatellana Kuoh. A key is provided to distinguish the four genera. A check list of species in each genus is given. Several species are important rice pests and information on biology and pest status is sum- marized.

Introduction species of Sogatodes, S.molinus, belongs to the genus Sogatella, Consequently, Sogatodes must Several delphacid (planthopper) species are im- be synonymized with Sogatella and subsequently portant pests of cereal crops such as maize, there is a need to establish at least one genus in wheat and rice. Some species currently placed in which to place the remaining species. This study the genera Sogatella Fennah and Sogatodes Fen- has been undertaken to resolve the situation and nah, are serious pests of rice in Asia and south also to provide names for use in a handbook for and central America by virtue of the effects of the identification of rice planthoppers currently their direct feeding and by their role as vectors of under preparation by M. R. Wilson and M. F. virus disease (e.g. Ou, 1985, and see references Claridge. in Wilson & Claridge, 1985). There have been Material studied is deposited in the following considerable difficulties in the identification of collections denoted by the abbreviations given: species in these genera since their separation is AMNH: American Museum of Natural History, mainly based on small differences in the male New York, U.S.A.; BMNH: British Museum genitalia. However, these difficulties in identifi- (Natural History), London, U.K.; BPBM: Ber- cation have been compounded by taxonomic nice P. Bishop Museum, Honolulu, U.S.A.; problems. In Asia it appears that many names KU: Snow Entomological Collection, Univer- have been applied to a small number of species. sity of Kansas, Lawrence, U.S.A.;LV: R. Lin- Also, Okada (1977) realized that names given to navuori collection, Raisio, Somersoja, Finland; Sogutella species in Japan might not concur with LOB: L. O’Brien collection, Tallahassee, their usage elsewhere. We now find that the type Florida, U.S.A.; MA: M. Asche collection, Correspondence: Dr M. Asche, FB-Zoologie der Marburg, West Germany; MAK: Zoologisches Philipps Universitat, Postfach 1929, Marburg, D- Forshungsinstitut und Museum Alexander 3550, West Germany. Koenig, Bonn, West Germany; NCIP; National 1 2 Manfred Asche and Michael R. Wilson

Insect Collection, Pretoria, South Africa; NM: fused with S.kolophon. In addition to this the Naturhistorisches Museum, Basel, Switzerland; species re-illustrated by Fennah (1963a) as MNHN: Museum National d’Histoire Naturelle, S.longifurcifera strongly resembles the species Paris, France; USNM: United States National Japanese researchers call S.panicicola in various Museum, Washington, U.S.A.; UTS: Theron characters.’ Dr R. Kisimoto kindly sent speci- Collection, University of Stellenbosch, Stellen- mens from Japan and examination of his speci- bosch, South Africa. mens confirmed that Okada’s opinion was In the section ‘Distribution and material indeed correct. examined’ countries from which specimens have A number of Sogatella species have been de- been examined are indicated by *. scribed since 1963. For our study considerably more specimens were available than were examined by Fennah, including several type specimens that he did not see. The results of our Taxonomic history study appear to move us back towards the broader species concept of Muir & Giffard. This The species now placed in Sogatella are among is not entirely so, as eight of their ten synonyms some of the most widely distributed and of S.furcifera are considered valid species. The taxonomically difficult of any Delphacidae. It is major changes to Sogatella outlined in this paper not intended here to provide a full history of concern the status of S.vibix (Haupt) and Sogatella and its species, but some information is S.kolophon (Kirkaldy) in which eight and seven given as background to the problem. The type synonyms respectively are newly proposed. We species, S.furcifera (Horvhth) is listed in Met- regard both species as having a wide distribution calf‘s (1943) catalogue with ten synonyms and and the majority of synonyms have arisen from three subspecies, following Muir & Giffard descriptions of new nominal species based on (1924). Those authors, realizing the difficulties slight morphological differences without proper of identification, wrote ‘The authors cannot pro- appreciation of variation within and between test too strongly against erecting species in this populations or without reference to other de- group on color or even slight structural charac- scribed species at all. The analytical approach ters. Such species not only lead to confusion, but adopted in this study is, by necessity, mor- lead to totally wrong conceptions as to relation- phological in emphasis and centred on charac- ship and geographical distribution’. Unfor- ters of the male genitalia. Few biological data tunately, until Fennah’s (1963a) study, no stable are available to support species separation in generic concept had been established for what most cases. However, a study of the courtship- he called the ‘species-complex known as Sogata songs of the three Asian Sogatella species has furcifera’. By that time furcifera had been in- been made by Uhm et al. (1982). They dem- cluded by various authors in six different genera onstrated species-specific differences in both (Delphux Fabr., Liburnia Stil, Sogata Distant, males and females of S.furcifera, S. vibix (given Megumelus Fieber, Delphacodes Fieber and as S.panicicola (Ishihara)) and S. kolophon Chloriona Fieber). As an interim measure until (given as S. longifurcifera Esaki & Ishihara). ‘a more critical assessment of its relationships These data lend support to our morphological could be made’ Fennah (1956a) established concept, especially since we recognize only these Sogatella as a subgenus of Chloriona. In his later three species occurring sympatrically through- study (Fennah, 1963a) almost all the synonyms out Asia. It would be worthwhile to study the included under S.&rc#era by Muir & Giffard (1%4) acoustic repertoire of Sogatella species in Africa were raised from synonymy and a number of where thirteen species are found, many of them new species were described. Fennah’s study has, sympatric. for 25 years, been regarded as definitive, but the Sogatella belongs to a group of genera that difficulties expressed in our introduction have have a slender body shape, narrow vertex and necessitated that Sogatella should be ‘revisited’. frons and a white or pale yellow longitudinal Okada (1 977 : 10) expresses one of the problems stripe from the head across the pro- and clearly. In discussing S.longifurcifera Esaki & mesanotum (Fig. 1). This group contains Latis- Ishihara, he writes ‘. . . S.longifurcifera referred triu, Matutinus (recently revised by Fennah, 1972), to by Japanese researchers is possibly being con- Sogatella, Sogatelha and Tagosodes gen.n. These The delphacid genus Sogatella and related groups 3

FIG. 1. Sogatella furcifera (Horvath), habitus, 6 from Sulawesi; scale line: 0.5 mm. 4 Manfred Asche and Michael R. Wilson genera may mainly be separated by characters of cubanus and T.oryzicolus as rice, Echinochloa the male genitalia. A key is given below. and other Gramincae. Xfurcifera is a serious rice pest in Japan but appears unable to overwinter there and under- goes long-distance migration (with Nilaparvata Distribution of Sogatella, Tagosodes, lugens (351) the brown planthopper) each year Latristria and Sogatellana species from southern China (Kisimoto, 1976, 1987). (Names here and below in ‘Biology’ and ‘Economic aspects’ are as used in accordance Economic aspects of Sogatella and with the synonymies and new combinations Tagosodes species proposed later in the paper.) Sogatella species are found throughout the S.furcifera is second only to Nilaparvata lugens sub-tropical and tropical regions of the world. (Stll) as a rice pest in Asia where it damages the They are concentrated in Africa where thirteen crop by direct feeding. Other species such as species are found. Two species, S.kolophon and S. nigeriensis and S. vibix are frequently found on S,rnolina, arc found in the Nearctic and Neotrop- rice (e.g. Ammar, 1977; Ammaretal., 1980) but ical regions. With our revised concept only three are not considered to be important pests at pres- species, S.furcifera, S.kolophon and S. vibix, are ent. Tagosodes pusanus (Distant) is also found found in Asia, the Pacific and Australasia. These commonly on rice in Asia. At present none of are the most widely distributed species; S.kolo- these species are known to spread virus diseases phon is found throughout the entire range of the to rice. However, Tagosodes oryticolus and genus, while S.vibix is found in Asia and the T.cubanus are well-known vectors of Hoja Ethiopian region. S.furcifera is common through- blanca virus of rice in south and central out Asia and extending to the Middle East. America, where SO% of the yield may be lost Tagosodes species, are also found throughout to this disease, .and as such are considered the tropics but with more species recognized important pest species in that region (King & from the Nearctic region than with Sogatella. Saunders, 1984). S.vibix is the vector of maize Latistria and Sogatellana species are found only rough dwarf virus in the middle east (Harpaz, in the old world tropics. 1966) and (as S.1ongifurcifera) is reported as a vector of Maize Sterile Stunt in Australia (Grebcr, 1982). S.kolophon is the vector of Biology and host plants Digitaria striate virus in Australia (Greber, 1979). Julia & Mariau (1982) reported that These genera are entirely grass-associated but S. kolophon andTagosodes cubanus (as little evidence is available for specificity. Sogatodes) were the vectors of dry bud rot of Mochida & Okada (1971) list the host plants of coconuts in West Africa. The causative agent of Japanese Delphacidae, including Sogatella the disease is not known. species. Unfortunately they fail to differentiate adequately between casual host records and true Key to genera (males only) host plants. However, for S.furcifera a considerable list of Gramineae is given from which Anal segment with two pairs of processes (e.g. Figs nymphs were reared to adult under experimen- 136, 137) ...... Sogatellana tal conditions. Lee & Kwon (1980) also give an extensive list of host plants of S.Jurcift.ra. Ammar Anal segment with one pair of processes (e.g. Figs 21, 112, 126) ...... 2 (1977) reared S. vibix in Egypt, on wheat experimental conditions for eight successive genera- Mediodorsal margin of diaphragm of genital seg- tions. The cultures originated from rice and ment forming a broad U-shape (Fig. 20), aedeagus other Gramineae. Ammar et at. (1980) also compressed, twisted, with two rows of spines ...... Sogatella reared S.nigeriensis (recorded as S.furcifera) under similar conditions and from the same ori- Diaphragm of genital segment different, aedeagus ginal hosts. In central America, King & Saun- tubular, with irregular teeth and/or teeth in rows ders (1984) record the host plants of Tagosodes ...... 3 The delphacid genus Sogatella and related groups 5

Mediodorsal margin of diaphragm forming a con- sinuate, after its basal third it is bent dorsad, spicuous caudodorsad directed, shoe-shaped pro- with the tip curved ventrad, slightly compressed tuberance; anal segment with rather long sinuate and twisted tapering to the apex. Two rows of processes which originate subdistally; relatively robust species with acute angled transition of frons small teeth are present ascending from the ven- to vertex ...... Macutinus trodorsal third on both sides to the dorsal third. The phallotreme is situated subapically on the Diaphragm of genital segment and processes of left side. anal segment different, small and slender species with transition of vertex to frons either rounded or 3. The parameres are diverging, in most of the slightly acute-angled ...... 4 species tapering to apex and distally bifurcated.

Diaphragm of genital segment rnediodorsally with a sinuate protrusion (Figs 124, 129); parameres, Synonymy of Sogatodes with Sogatella slender (Figs 125, 130), almost reaching the laterodorsal angles of the genital segment Fennah (1963a) established the genus ...... Latistria Sogatodes to accommodate the species Dic- ranotropis cubanus Crawford, Sogata biangulata Diaphragm medio-dorsally with either a raised T- Muir, Sogata orizicola Muir, Megamelus ap- shaped area or a distinct rectangular protusion (Figs 92,101,104), or with inverse W-shaped cent- proximatus Crawford, Sogata brazilensis Muir, ral projection (Fig. 108). Parameres variously Sogata nautica Muir, Sogata anomala Muir, shaped but not long and slender Sogata dorsolineata Beamer, Liburnia al- ...... Tagosodes gen.n. bolineosa Fowler and Sogatodes molinus. Un- fortunately, by an oversight no species was designated as type species. Subsequently, Sogatella Fennah S.molinus was designated as type species (Fen- nah, 1963b). Chloriona (Sogatella) Fennah, 1956a: 471. Type Several other species have since been transfer- species (by original designation), Delphax fur- red to Sogatodes newly described in that cifera Horvith, 1899: 372. or genus (see list under Tagosodes). For reasons Sogatodes Fennah, 1963a: 71. Type species (by given later in this paper S.molinus is regarded as subsequent designation by Fennah, 1963b: a Sogatella species and accordingly all species 166), Sogatodes molinus Fennah, 1963a: 72. currently placed in Sogatodes have to be placed Syn.n. in other genera. The genus Tagosodes is de- Diagnosis. Small and slender delphacids. scribed below to accomodate the bulk of the Macropterous males about 2.5-3.5 mm, mac- species while others are transferred to Latistria, ropterous females about 3.0-4.00 mm. Vertex and Sogatellana. rounded onto frons, anterior cell of vertex long, reaching to apex of vertex or bent towards frons. Coloration. Males with a whitish longitudinal stripe across vertex and the middle portion of Checklist of Sogatella species and pro- and mesonotum; lateral portions of pro- sub-species and mesonotum brown or black. Females distinctly lighter than males, yellow to orange, More detailed synonyms are given under each stramineous or light brown. species. Sogatella can be distinguished from other ex- Sogatella albofimbriata (Muir) ternally similar genera by a combination of =Sogata rhodesi Muir characters of the male genitalia as follows. We Sogatella camptistylis Fennah consider these three characters to be constitutive Sogatella capensis (Muir) characters for Sogatella: Sogatella colorata colorata (Distant) 1. The dorsal margin of the diaphragm in the Sogatella colorata nigrior Fennah middle posseses two cone-shaped processes Sogatella furcifera (HorvBth) which are medially connected by a dorsally =Soguta distincta Distant slightly concave tuberosity, altogether forming a =Sogata pallescens Distant broad U-shaped structure. =Sogata kyusyuensis Matsumura & Ishihara 2. The adeagus is moderately long, slightly syn.n. 6 Manfred Asche and Michael R. Wilson

FIGS 2-14. Sogutella spp.; left paramere, maximum view, scale line: 0.1 mm; 2-8: S.furciferu-group; 2, S.furciferu (Horvhth), specimen from Belau Is.; 3, S.nigeriensis (Muir), specimen from Nigeria: 4, S.cumptistylis Fennah, specimen from Uganda; 5, S.cupensis (Muir), specimen from S. Africa (Cape Pro- vince); 6, S.yei Linnavuori, holotype, Sudan (Equatoria); 7, S.munetho Fennah, holotype, Zimbabwe; 8, S.petux Fennah, specimen from Tanzania (Zanzibar). 9-12: Xkolophon-group; 9, Xkolophon (Kirkaldy), paratype, Australia (Queensland); 10, S.molina (Fennah), paratype, Mexico; 11, Xnigrigenis (Jacobi), specimen from Uganda; 12, S.vibix (Haupt), specimen from Israel. 13-14. S.albofimbriuta-group; 13, S.al- bofimbriatu (Muir), specimen from Ivory Coast; 14, Xkrugeri (Muir), specimen from S. Africa (Natal). The delphacid genus Sogatella and related groups I

=Sogata tandojamensis Qadri & Mirza nomen Key to ZQBatella species (males only) nudum 1. OutP/2nd inner angle of the apex of parameres Sogatella kolophon (Kirkaldy) aba'.kequally produced or the outer one rounded, =S. kolophon atlantica Fennah syn.n. .~-tcalmargin shallowly concave or sinuate (S.fur- $era-group, Figs 2-8) ...... 2 = Opiconsiva insularis Distant syn.n. =Sogata meridiana Beamer syn.n. Parameres distinctly bifurcate distally. inner api- = Opiconsiva balteata Distant syn.n. cal process rod-like, outer one strongly produced .# 'l =Sogatella chenhea Kuoh syn.n. / (S.kolophon-group) or parameres thin and elong- = Opiconsiva derdicta distant syn.n. .>- J ate, inner distal process strongly reduced or ab- * *' sent (S.albofmbriuru-group) ...... 8 =Delphacbdes elegantissima lshihara syn.n. =Sogatella nebris Fennah syn.n. 2. Face with frons and clypeus and genae dark brown Sogatella krugeri (Muir) c0mb.n. (carinae may be stramineous or pale yellow) . 3 Sogatella manetho Fennah - Sogatella molina (Fennah) c0mb.n. Frons and clypeus stramineous or pale yellow, genae dark brown ...... 5 Sogatella nigrigenis (Jacobi) Sogatella nigeriensis nigeriensis (Muir) 3. Apex of clavus in forewing with dark brown mark- =Sogatella nigeriensis troilos Fennah syn.n. ing ...... Sogatella petax Fennah - Apex of clavus without marking; parameres with Sogatella vibix (Haupt) apical margin undulate, shallowly concave, outer =Sogatella catoptron Fennah syn.n. corner forming a pointed process (Fig. 5) =Sogatella diachenhea Kuoh syn.n...... capensis =Delphacodes dogensis Ishihara syn.n. 4. Parameres strongly dilated at base with apex rela- = Delphacodes longifurcifera Esaki & tively small, almost equally bifurcate (e.g. Fig. 2). Ishihara syn.n. diaphragm mediodorsally broadly U-shaped (Fig. =Liburnia matsumurana Metcalf syn.n. 20) ...... furcifera =Sogatella parakolophon Linnavuori syn.n. - Sogatella yei Linnavuori Parameres not dilated at base, apical part distally about as broad as base (Fig. 3), U-shaped process Species transferred from Sogatella: of diaphragm rather narrow ...... nigeriensis Sogatella hedai Kuoh, 1977 to Toya Distant: Toya hedai (Kuoh) c0mb.n. 5. Parameres with apical margin distinctly concave, outer apical corner forming a pointed edge or pro- Remarks: The type-material of this species jection (Figs 4, 6) ...... 6 could not be studied. The transfer is made on the basis of the drawings of the male genitalia. They - Parameres with apical margin shallowly concave, appear very similar to the type species of Toya, outer apical corner rounded (Figs 7,8) 7 T.attenuata Distant. 6. Outer apical corner of parameres less produced Sogatella fulva Yang 1989:192 to Toya: than inner forming a short, pointed edge; inner Toya fulva (Yang) c0mb.n. corner strongly produced to a long r cess (Fig. 6) ...... Sogatella lima Yang 1989:199 to Toya Toya lima (Yang) c0mb.n. - Parameres with outer apical corner more strongly Sogatella timaea Fennah, 1969b to produced than inner for Tagosodes gen.n.(see below). curved process (Fig. 4) . . Sogatella wallacei (Muir & Giffard, 1924) to 7. Outer apical corner of parameres broadly Tagosodes gen.n.(see below). rounded, not distinctly produced, inner apical [Mochida & Okada (1971) and Lee & Kwon corner forming a bill-sha (1980) include Delphacodes terryi and Sogata ...... petax sirokata in Sogatella. Neither of these species is a - Outer apical corner of parameres projected to a Sogatella species under the criteria established rounded process, apical margin of parameres al- here. They are currently placed, respectively in most straight, inner apical corn Toya (where terryi is a synonym of Toya tuber- like (Fig. 7) ...... culosa) and in Paracorbulo.] 8. Parameres distinctly bifurcate distally, inner apical process rod-like, outer one strongly produced (S.kolophon-group. Figs 9-12) ...... 9 8 Manfred Asche und Michael R. Wilson

- Parameres thin, elongate, inner apical process - Head with dark brown genae (e.g. Fig. 72) . 11 strongly reduced or absent (S.albofirnbriata- group, Figs 13-14) ...... 13 11. Frons with area between the carinae fuscous or dark brown (Figs 58, 60); forewing with apex of 9. Head with frons, clypeus, genae and anterior part clavus with brown marking (Figs 61,62) of vertex shiny dark brown ...... colorutu ...... molina

- - Head otherwise coloured ...... 10 Frons with area between the carinae stramineous or pale yellow. apex of clavus without brown 10. Frons, clypeus and genae pale yellow or stramin- marking ...... 12 eous, in some specimens a slight brown suffusion at lower area of frons and around the ocelli (Figs 12. Parameres with outer apical corner produced in a 35-38 ...... kolophon medially distinct broadened process (e.g. Fig. 12) ...... vibix

FIGS 15-18. Sogufellufurciferu (Horvath); 15-17: coloration of head and thorax, 6 specimen from Borneo (Sarawak), scale line: 0.5 mm;15, head and thorax, lateral view; 16, head, frontal view; 17, head left lateral view; 18, left forewing, macropterous 6,Sulawesi, scale line 0.5 mm. The delphacid genus Sogatella and related groups 9

- Outer apical corner of parameres strongly pro- ing a pointed process) and in the longer proces- duced in a long, slender process which tapers con- ses of the anal segment. tinuously to apex (Fig. 11) ...... nigrigenis Remarks. S.capensis seems to be closely re- 13. Parameres with broad base, then narrowing to lated to S.nigeriensis, possibly also to S.carnptis- truncate apex (Fig. 14) ...... krugeri tylis Fennah and S.manetho Fennah (see below), the latter two differing in the lighter coloration. - Parameres narrowing conically to apex, to small Distribution and material examined. SOUTH edge subapically on inner margin (Fig. 13) ...... albofimbriata AFRICA: Cape Province (Muir, 1929b: 212; Fennah, 1958b: 203; Fennah, 1963a: 57), Natal, SW-Africa (Muir, 1929b: 212; Fennah, 1963a: Sogatella furcifera-group (seven species) 57). Additional specimens from South Africa* (BMNH, NICP, UTS). S.camptistylis, S.cupensis, S.furcifera, S.manetho, s. nigeriensis, S.petax, S.yei

Diagnosis. Male genitalia with outer and inner Sogatella furcifera (Horvath) (Figs 1, 2, apical angles of parameres about equally pro- 15-34) duced, or with the outer one rounded, apical margin shallowly ancave or sinuate (see Figs 2-8). Delphax furcifera HorvBth, 1899: 372, syntypes (6 0)JAPAN (Hokkaido), not examined. Sogata distincta Distant, 1912: 191, Lectotype Sogatella camptistylis Fennah (Fig. 4) d, designated by Fennah, 1963a: 50, INDIA, (BMNH), examined. Syn. by Muir, 1919: 7. Sogatella camptistyh Fennah, 1%3a: 70, Holotype Sogata pallescens Distant, 1912: 192, Lectotype 8.UGANDA (BMNH), examined. 6, designated by Fennah, 1963a: 50, INDIA, (BMNH), examined. Syn. by Muir, 1919: 7. Diagnosis. Stramineous with light face, males Sogata kyusyuensis Matsumura & Ishihara, with dark genae (externally resembling S. vibir). 1945: 65, Holotype d, JAPAN (Kyushu Uni- Male genitalia similar to those of S.yei, but also versity, Fukuoka, Japan). Syn.n. S.capensis and S.rnanetho, but differing from Sogata tandojamensis Qadri & Mirza, 1960: 115, these species in the shape of the parameres (dis- nomen nudum, PAKISTAN, not examined. tally shallowly concave, outer tip pronounced and slightly recurved basad). Diagnosis. Males with dark frons, clypeus and Disrribution. Atrica: Uganda. genae. Tegmina with dark or fuscous marking at Only known from the holotype and a second tip of clavus, which in females may be less de- male from Uganda (BMNH). veloped or missing. This species can readily be distinguished from the other Sogatella species by structures of the Sogatella capensis (Muir) (Fig. 5) male genitalia; the parameres strongly dilated at base, apex relatively small, almost equally bifur- Sogata furciferu capensis Muir. 1929b: 212. cate (Figs 2,22,25-34). Lectotype 6,designated Fennah, 1Y63a: 57, Remarks. The synonymy of S. kyusyuensis S. AFRICA (Cape Province) (BMNH) with S.furcifera is based on the examination of examined. the holotype of S.kyusyuensis, a brachypterous male. The name, S.tandojamensis (Qadri & Diagnosis. Dark fuscous, externally similar to Mirza, 1960) is invalid based on article 13 S.furcifera and S.nigeriensis. Male with shiny (availability of name) of the code of Zoological dark-brown frons, clypeus and genae. Male Nomenclature 1985. The species is, in any case, genitalia similar to those of S.nigeriensis, but dif- clearly identical with S.furcifera, based on the fering in the shape of the mediodorsal process of rather poor illustrations. the diaphragm which is broad U-shaped in S.furcifera shows a wide range of intraspecific S.capensis, in the tip of the parameres (apically variation in several characters such as intensity less incised, more truncate, outer corner form- and extent of coloration and genital structures 10 Manfred Asche and Michael R. Wilson

, ,

FIGS 19-24, Sogatella ficrcifcra (Horvjth) 3 genitalia. specimen from India. scale lincs 0 1 mm: 19. genitalia in repose, ventocaudal view; 20, diaphragm of genital segment caudal view; 21, genitalia without genital segment, left lateral view; 22, left paramere maximum view; 23. aedeagus right lateral view: 24. aedeagus ventrocaudal view. The delphacid genus Sogatella and related groups 11

(e.g. parameres) even within the same popula- 1987: 195), VIETNAM* (Fennah, 1978: 221). tion (Figs 25-34). In some specimens variation Pacific region: FIJI IS.: Viti Levu* (Muir, in the shape of the left and right parameres of the 1917: 328; Fennah, 1950: 42) (BPBM). MIC- same individual was observed (Fig. 34). The de- RONESIA: Belau Is. (=Palau Is.)* (BPBM), gree of intraspecific variation proved to be about Guam* (BPBM), Marshall Is.* (BPBM), the same in all populations studied from various Ponape Is. (all Fennah, 1956b: 116). RYUKYU localities (Figs 25-34). However, Fennah IS.* (Clausen, 1931: 45) (BPBM). VANUATU (1963a) discussed the status of the two In- (=NEW HEBRIDES)* (BPBM), dian taxa S.distincta and S.pallescens Fennah Australian region: AUSTRALIA: Queens- (1Y63a) and considered they might be consid- land* (Fennah, 1965b: 47) (BPBM, MA), ered as geographical subspecies of the typical Northern Territory (MA). S.furcifera. Our study of large numbers (>500 Remarks. The western limits of the distribu- specimens) of S.furcifera specimens from India tion of S.furcifera are still unclear. We have not and many other parts of the Oriental Region and seen a single specimen of this species from Af- Australia has not revealed any evidence for the rica, Europe or the New World. All specimens geographical concentration of any particular from these regions which had been previously morphological or colour configurations which identified as S.furcifera proved to be other would support the idea of subspecific differentia- species. Records from Europe (e.g. Oshanin, tion in India. 1907: 315), N. Africa (e.g. Matsumura, 1910b: Distribution and material examined. Wide- 17; Ammar et al., lY80) and tropical Africa spread in the Eastern Palaearctic, the Oriental (e.g. Matsumura, 1910b: 17) concern either Region, the Western Pacific and Australia. S.nigeriensis (Muir) specimens examined from Numerous specimens have been examined Egypt, Canary Islands, Madeira) or Svibix from various localities in all parts of the Haupt (for instance the records from Sicily, distribution. Yugoslavia). Records of S.furcifera from the Palaearctic region: JAPAN: Hokkaido* New World countries (Muir & Giffard, 1924: 13; (BMNH, BPBM), Honshu, Kyushu, Shikoku Muir, 1926a: 26) in most cases concern S.molina (e.g. Ishihara, 1949: 63). KOREA* (Nakayama, (Fennah) or S.kolophon (Kirkaldy) (see below). 1930: 181; Lee & Kwon, 1976: 60; 1977: 103; The western-most populations of true S.furci- 1980: 31). U.S.S.R.: Maritime Territory (Vil- fera we could examine originate from Pakistan baste, 1968: 21; Anufriev & Emeljanov, 1988: and Saudi Arabia. Its closest relative, 361), Kurile Is. (Anufriev, 1977: 863), Mongolia S.nigeriensis, is widely distributed in Africa and (Dlabola, 1967: 212; Emeljanov, 1982: 72). occurs sympatrically with Xfurcifera in Saudi Oriental region: CAMBODIA* (BPBM). Arabia. However, no transition zone (hybrid belt CHINA (e.g. Matsumura, 1910a: 17), Fennah, or cline) between these two species has been found. 1956a: 472, Kuoh et al., 1983: 277). FUKIEN* The distribution of S.furcifera and S.nigeriensis (BPBM). HONG KONG' (BPBM). INDIA* appears to overlap in the Middle East. (e.g. Matsumura, 1910b: 17), INDONESIA: Amboyna, Ceram (Muir, 1917: 328), Java* (BMNH, BPBM), Kalimantan* (BMNH, Sogatella manetho Fennah (Fig. 7) BPBM), Sulawesi* (BMNH, MA), Sumatra* (Muir, 1923: 174) (BMNH, BPBM). LAOS* Sogatella manerho Fennah. 1963a: 66. Holotype (BPBM). MALAYSIA: Malay Peninsula* 8,ZIMBABWE (BMNH). examined. (BPBM) (Kathirithamby, 1981: 15), Sabah' (BPBM), Sarawak* (BMNH, BPBM). Diagnosis. Light coloured with pale yellow NEPAL* (BMNH). PAKISTAN* (Dlabola, face. Males with slightly darkened genae. The 1971: 381) (LOB). PHILIPPINE IS.* Luzon male genitalia resemble S.capensis, but differ in (e.g. Muir, 1917: 328) (BMNH, BPBM), Min- the rather short and apically almost blunt danao (BPBM), Palawan (BPBM). SAUDI aedeagus, the blunt outer apical process of the ARABIA* (NM). SRI LANKA (CEYLON) parameres (Fig. 7) and in the shape and length of (Melichar, 1903: 104; Fennah, 1975: 94), the processes of the anal segment. TAIWAN* (e.g. Matsumura, 1910a: 17) Remarks. The configuration of the genitalia (BMNH, BPBM). THAILAND (Hongsaprug, suggests that S.manetho is the light-coloured 12 Manfred Asche und MichueI R. Wilson

FIGS 25-34. SoRotclln firrcifero (Horvith). intraspecific variation in pnrnmcrcs. scalc line 0.1 mm: 2.5. Japan (Hokkaido) d 1. 26. same population 62; 27, same population 83; 28. Belau Is. (Bebelthuap I.;); Fiji Is. (Viti Levu I.) 30. Borneo (Sarawak);31, Philippine Is. (Luzon); 32. India; 33. Saudi Arabia; 34. Au- stralia (Queensland). left-right variation in the same individual. The delphacid genus Sogatella and related groups 13 equivalent to S.capensis, possibly a close rela- Saudi Arabia (see above) lends support to our tive. S.manetho might replace S. capensis geo- view that they are distinct species without any graphically in northeast South Africa. S.petax morphological transition. may have evolved from Xmanetho-like forms Dismhuion and mterkd examined. S.nigeriemis since it shows similarly-shaped rounded outer is widespread in the Ethiopian region the south- apical corners of the parameres. western portion of the Palaearctic (e.g. Canary Distribution and material examined. Southern islands, Madeira, Egypt, Israel) and Madagas- Africa. At present known only from the type locality car. Numerous specimens from Africa have in Zimbabwe. Additional material examined: been examined. paratypes from the type locality (BMNH). Palaearctic region: AZORES (e.g. Lindberg, 1Y54a: 8). CANARY IS.* (Linnavuori, IY73:~ log), EGYPT* (Linnavuori, 1969: 208; Fennah, Sogatella nigeriensis Muir (Fig. 3) 1963a: 61) (BMNH). ISRAEL* (MA), MAD- EIRA* (e.g. Lindberg, 1961) (BMNH). OMAN* Megamelus furcifer (sic) nigeriensis Muir, 1920: (BMNH). SAUDI ARABIA* (MNHB). 143, Holotype 6, NIGERlA (BMNH), Ethiopian region: ALDABRA* (BMNH). examined. ETHIOPIA* (Fennah, 1969b: 56), GUINEA Sogatella nigeriensis troilos Fennah, 1963a: 62, (Fennah 19.58~:490). ILES GLORIEUSES* Holotype 8, MADAGASCAR (BMNH), (Fennah, 1963a: 61). IVORY COAST* (Asche, examined. Syn.n. 1988: 208). KENYA* (BMNH). MADAGAS- CAR* (Fennah, 1963a: 61) (BMNH). MALI Diagnosis. S.nigeriensis is a dark species ex- (Fennah, 1958c: 490). NIGER (Fennah, 1963a: ternally closely resembling S.furcifera, but also 60). NIGERIA* (Muir, 1920: 143; Medler, similar to S.capensis. Males possess a dark 19801, SENEGAL (Fennah, 1961: 311), brown frons and genae. Tegmina with fuscous SOUTH AFRICA: Cape Province, Pondoland, marking towards the end of clavus. In the male Natal (Muir, 1926b, as S.furcifera) (BMNH). genitalia the U-shaped process at the dorsal mar- SUDAN: Blue Nile, Darfur, Equatoria, Kordo- gin of the diaphragm rather narrow; parameres fan, Upper Nile (Linnavuori, 1973: 108). TAN- comparatively short, inner base moderately pro- ZANIA* (BMNH). UGANDA* (BMNH). duced (not dilated lobe-like as in S.furcifera), ZAIRE* (BMNH) distal part similar to that in S.furcifera, but outer process more strongly developed (Fig. 3). S. nigeriensis can be distinguished from S. capensis by the dark marking in the apex of the clavus Sogatella petax Fennah (Fig. 8) (absent in S.capensis). The male genitalia have been figured by Asche (1988: 208). Sogatella petax Fennah, 1963a: 68, Holotype 8, Remarks. Fennah (1963a) based the sub- EGYPT (BMNH), examined. species s.nigeriensis troilos from Madagascar and the Iles Glorieuses on slight differences in Diagnosis. S.petax is light stramineous in col- the coloration of the tegirna between continental our, males with dark genae, externally resembl- African populations and the island specimens. ing S.vibix. It is readily distinguishable from all In our study we could find lighter coloured ‘is- other Sogatella species by the shape of the para- land-forms’ also in populations from mainland meres which are distally sinuate and with the Africa, and vice versa fuscous ‘continental- outer apical angle rounded and not distinctly forms’ in Malagasian populations. Accordingly produced laterad, the inner apical angle almost we cannot accept the validity of S.nigeriensis beak-like and produced mediad (Fig. 8). troilos as a separate geographic subspecies. Remarks. Within the S.furcifera-group S.nigeriensis is in all characters very similar to S.petax seems somewhat isolated by the entirely S.furciferu, and it is most likely its closest rela- rounded outer apical angle of the parameres. Of tive geographically replacing S.furcifera in the all species included in the S.furcifera-group it Ethiopian Region. They do appear to overlap in might be closest related to S.manetho which dis- the Middle East to some degree. The syntopic plays a blunt outer apical angle of the parameres occurrence of S. furcifera and S.nigeriensis in (see Fig. 7). 14 Manfred Asche and Michael R. Wilson

Distribution and material examined. Appa- Sogatella yei Linnavuori (Fig. 6) rently widespread at least in the eastern part of Africa and parts of the Middle East. Sogatella yei Linnavuori, 1973: 109, Holotype Palaearctic region: EGYPT* (Fennah, 1963a: 8,SUDAN (Equatoria), (AMNH), examined. 70; Linnavuori, 1964: 341) (BMNH). JOR- DAN* (Fennah, 1963a: 70) (BMNH.) Diagnosis. Xyei externally resembles S.vibix Ethiopian region: SUDAN: Equatoria (Lin- (Haupt) but differs from this and all other navuori, 1973: 109), TANZANIA* (BMNH), Sogatella species in the shape of the male NIGERIA* (Medler, 1980: 62) (BMNH). genitalia. The parameres have a very short

___I

- FIGS 15-19 Sogntella kolophon (Kirkaldy): 15-78. colour patterns of head and thorny. fi specimen from Australia (Queensland),(Que scale lines: 0.5 mm; d 1, head and thorax dorsal vlew; 35, d , head and thorax dorsal view; 36, d 1 head frontal view; 37. d 1, head frontal view; 38, 82,samc population, head frontal view 39. left forewing. macropternup 6. Australia (Queen4and). Kdk line 0 5 mm The delphacid genus Sogatella and related groups 15

FIGS 40-45. Sogatella kolopphon (Kirkaldy), 6 genitalia, specimen from Australia (Queensland), scale line 0.1 mm; 40, genitalia in repose ventrocaudal view; 41, diaphragm of genital segment caudal view: 42, genitalia without genital segment, left lateral view; 43, left paramere maximum view; 44, aedeagus right lateral view; 45, aedeagus ventrocaudal view. 16 Munfred Asche and Michael R. Wilson pointed outer apical corner and a long, rod-like Sogatella colorata nigrior Fennah inner process (Fig. 6). Sogutella colorntu nigrior Fennah, 1963a: 64. Remarks. Within the S.furcifera-group S.yei is Holotype 6, MAURITIUS (BMNH), very similar to S.camptisty1i.s from Uganda in the examined. shape of the apical part of the parameres. It possesses a less produced outer apical process and a Diagnosis. Fennah separated the Mauritian more strongly developed inner apical process populations from the nominate of S.colorata than found in S.camptistylis. These two species mainly by darker coloration of the postclypeus appear to be closely related. and by a yellow rather than castaneous colora- Distribution and material examined. Only tion of the first antenna1 segment. known from the holotype collected in Sudan Remarks. We have not seen enough material (Equatoria). of both taxa to be able to judge the range of intraspecific variation. Provisionally we follow Fennah and regard the Mauritian population as a separate subspecies. Sogatella kolophon-group (five species) Distribution and material examined. Mauritius (Fennah, 1963a: 64). Holotype only, examined. S.colorata, S. kolophon, S.molina, S.nigrigenis, S.vihix Soga tella kolophon (Ki r ka Idy) The group is defined by the male genitalia in (Figs 9, 35-56) which the parameres are distally bifurcate with the apical margin shallowly or deeply concave, Delphax kolophon Kirkaldy, 1907: 157, inncr angle short and stick-like, and the outcr Holotype 6,AUSTRALIA, (Queensland), angle strongly produced (Figs 9-12). (BPBM), examined. Sogatella kolophon atlanticn Fennah, 1963a: 58, Holotype 6, ST HELENA (BMNH) examined. Syn.n Sogatella colorata (Distant) Opiconsiva insularis Distant, 1917: 303, Lec- totype 6,designated by Fennah, 1963a: 60, 0picon.viva colorata Distant, 1917: 301, Lec- SEYCHELLES IS. (BMNH), examined. totype 6, designated by Fennah, 1963a: 63, Syn.n. SEYCHELLES IS. (BMNH), examined. Sogatella kolophon insularis: Fennah, 1963a: 59. Sogutellu coloruta: Fennah, l9h3a: 62. Sogata meridiana Beamer, 1952: 111, Holotype 6 U.S.A. (Florida), (KU), examined. Syn.n. Diagnosis. S.colorata differs from all other Sogatella kolophon meridiana: Fennah, 1963a: Sogatella species by the homogeneous shiny cas- 59. taneous or dark brown coloration of frons, Opiconsiva balteata Distant (in part), 1917: 302, genae, anterior portion of vertex, pronotum and Lectotype 6,designated by Fennah, 1963a: mesonotum (except for a fine white stripe in 64,SEYCHELLES IS. (BMNH), examined. middle line). The male genitalia are very similar Syn.n. to those of S.kolophon. Sogatella belateata: Fennah, 1963a: 64. Remarks. S.colorata appears to be closely Sogatella chenhea Kuoh, 1977: 440, Holotype 6, related to S.kolophon, both species occurring CHINA (Kwangtung), (Anhwei Agric. Col- sympatrically. As we have not seen any kind of lege), not examined. Syn.n. transition in the coloration of the two taxa we as- Opiconsiva derelicta Distant, 1917: 303, sume that S.colorata represents a good species Holotype 6,SEYCHELLES IS. (BMNH), which might have derived from S. kolophon-like examined. Syn.n ancestors that have undergone adaptive radia- Sogatella derelicta Fennah, 1963a: 62. tion on the Seychelles Islands. Delphacodes elegantissima Ishihara, 1952: 45, Distribution and material examined. Known Holotype 6,JAPAN, (Shikoku) (Ehime only from the Seychelles Is.; Lectotype and Univ., Matsuyama, Japan), not examined other specimcns examined (BMNH). Syn.n. The delphacid genus Sogatella and related groups 17

46 47

50 51

FIGS 46-56. Sogarella kolophon (Kirkaldy), intraspecific variation in parameres, scale line 0.1 mm; 46, Sulawesi. 6 1; 47. Sulawesi. same population. 62;48, Sulawesi, same population. 63;49. Sulawesi. same population, 64; 50. Australia (Queensland); Japan (Hokkaido); 52, Pitcairn I.; 53, Galapagos Is. (Santa Cruz I.); 54, Mexico; 55, S. Africa (Pondoland), paratype of Xnehris Fennah; 56, Philippine Is. (Luzon I.), left-right variation in the same individual. 18 Manfred Asche and Michael R. Wilson

Sogatella elegantissirnu Fennah, 1963a: 76. The specific synonomies proposed here of Sogatella nebris Fennah, 1963a: 67, Holotype d , 0.balteata, 0.derelicta and S.nebris with SOUTH AFRICA (Pondoland) (BMNH), S.kolophon are based on the examination of examined. Syn.n. type-material. The synonymy of Schenhea and D.elegantissima we deduce from the original de- Diagnosis. S.kolophon is small and slender in scriptions and drawings. The characters given general appearance. In colour it is light yellow to there do not separate these species from the pale stramineous; males with yellow face and range of intraspecific variation of S. kolophon. In genae, in some specimens a small brown or red the case of S.chenhea, all Chinese specimens of spot around or below the ocelli and a brown the S. kolophon-group we have examined mark in the lower half of the frons (Figs 35-38); proved to be either S.kolophon or S.vibix tegmina hyaline, in apical half with a grey or (Haupt), the latter being clearly distinct by other light brown suffusion of varying extent (Fig. 39). characters (see below). Male genitalia. Laterodorsal angles of the gen- Within the S. kolophon-group the species ital segment slightly produced, mostly bent seems closely related to the African species mediad; outer apical angle of the parameres S.nigrigenis (Jacobi), the Seychelles species strongly produced, more or less continuously S.colorata, and the New World species S.molina tapering to its apex, in some specimens slightly (Fennah) by the shape of the male genitalia. sinuate, not distinctly dilated in middle part; Distribution and material examined. Widely inner angle of parameres relatively short and distributed in Australia, the Oriental Region, stout; apical margin of parameres shallowly the Pacific, the Ethiopian Region, the Atlantic concave (Figs 9,43,46-56). Aedeagus compara- Islands, the New World and the eastern tively short, sinuate, apically not acutely taper- Palaearctic. It is most commonly found in the ing, tip in most specimens blunt, left row with tropics. 15-22 teeth, right row with 5-8 teeth (>lo0 Numerous specimens (>300) have been specimens examined) (Figs 42, 44, 45). examined from various localities in all parts of S.kolophon can be separated from all other the distribution. species of the genus by a combination of charac- Palaearctic region: AZORES (e .g. Lindberg, ters such as coloration of head and wings (it is the 1960: 3). CANARY IS.* (MA). JAPAN* (Ishi- only species with light genae in males), the shape hara, 1952: 45, as Delphacodes elegantissirnu) of the parameres and the apically blunt (BPBM). KOREA* (BMNH). aedeagus. Ethiopian region: CAPE VERDE IS.* Remarks. S. kolophon is the most widely dis- (Lindberg, 1958: 151, as Sogata furcifera ‘Rasse tributed of all Sogatella species. There is varia- colophon’ (sic); Fennah, 1963a: 59) (BMNH). tion in the intensity of coloration of head (see IVORY COAST* (Julia & Mariau 1982: 517; Figs 36.38) and tegmina and in the structures ot Asche, 1988: 206) (MNHP, MA). NIGERIA* the male genitalia e.g. parameres (see Figs 46- (Medler, 1980: 62). SOUTH AFRICA: Natal, 56). Fcnnah (1963a) used the shape of the caudal Pondoland, Transkei, Zululand (Fennah, margin of the genital segment in lateral view, 1963a: 68, as S.nebris), Swaziland* (UST). especially the form of the laterodorsal angles, to Malagasian region: MAURITIUS, Iles discriminate the subspecies S. kolophon adan- Glorieuses (Fennah, 1964: 140, as S.kolophon tica, S.kolophon insularis and S.kolophon insularis). RODRIGUES I.* (BMNH) meridinnu.. However, in examining large num- Oriental region: CAMBODIA* (BPBM). bers of S. kolophon specimens from various CHINA: Anhwei, Fukien* (BMNH, BPBM), localities we found that the outline of the caudal Kwangsi, Kwangtung, Yunnan (Kuoh, 1977: margin is highly variable, even within the same 440. as J.chrnheu. Kuoh et al.. IY83: 116). population. Moreover, the shape of this struc- HONG KONG* (BPBM). INDIA* (LOB). ture is strongly affected by shrinking during the INDONESIA: J;Iv;I* (BPBM). Kalimantan* drying process of the specimen. In all the charac- (BPBM). Sulawesi (BMNH. MA), Sumatra* ters examined the three subspecies lay well (BPBM). LAOS” (BPBM). MALAYSIA: within the range of variation of Australian, Malay Peninsula* (BPBM), Sabah* (BPBM), Oriental and Pacific populations and we cannot Sarawak* (BPBM). PAPUA NEW GUINEA* see any reason to retain the concept of the geo- (BPBM). PHILIPPINE IS.: Luzon* (Muir, graphical subspecies. 1916: 385), Mindanao* (BPBM) Palawan* The delphacid genus Sogatella and related groups 19

FIGS 57 63.Sn,qotc/(n rndirta (Fcnnah): 57-60. coloration of head and thorax. 3 spccimcnsfrom Bermuda Is.. scale lines: 0.5 mm; 57. 61. head and thorax dorsal view; 58. 8 I. head frontal view; 59. 6 1, left lateral view; 60. 62. same population. head frontal view. 61-62? coloration of forewings, macropterous males. wale line 0.1 mm: 61. Bermuda Is.: A?. Mexico. paratypc. 30 Man,frtvi Asche and Michael R. Wilson

(BPBM). SEYCHELLES IS.* (Distant, 1917: Sogatella molina (Fennah) c0mb.n. 302-303, as Opiconsiva insularis, 0.balteata (Figs 10,57-69) partim, 0.derelicta; Fennah, 1963a: 60: 62: 64). SRI LANKA (CEYLON) (Fennah, 1963a: 58). Sogatodes molinus Fennah, 1963a: 72, Holotype THAILAND* (BPBM). VIETNAM* (Fennah, 6,MEXICO, (BMNH), examined. 1978: 16) (BPBM). Pacific region: BELAU IS. (=PALAU)* Diagnosis. Externally, S.molina appears (Fennah, 1956b: 117; 1971a: 573) (BPBM). somewhat similar to S.furcifera. The vertex and BONIN IS.* (Fennah, 1956b: 116; 1971a: 572) middle carina of frons white or pale yellow; (BPBM). FIJI IS.: Viti Levu* (Kirkaldy, 1907: genae and compartments of frons light brown, in 157) (BPBM). GALAPAGOS IS.* (Fennah, specimens from Bermuda less intense (Figs 57- 1967b: 76) (BPBM). GUAM" (Fennah, 1956b: 60). Tegmina with a distinct light brown or in 117) (BPBM). HAWAII IS.* (Fennah, 1963a: most specimens dark brown spot in this postcrior 58) (BPBM). MANGAREVA IS.* (Fennah, angle of the clavus (Figs 61.62); tegmina hyaline 1958a: 194-195) (BPBM). MARQUESAS IS." or slightly infumed, from apex of clavus to apex (Fennah, 19SXa: 194-195). MARSHALL IS.* of tegmina a brown band-like suffusion (in speci- (Fennah, 1956b: 117) (BPBM) ENIWETOK mens from Mexico and Florida less developed or ATOLL (Japtan 1.) (Fennah, 1971a: 573), Gil- almost missing (see Figs 61,62), at apex area be- bert Is. (Makarei I.) (Fennah. 1971a: 573). MIC- tween the ends of the veins in some specimens RONESIA: Caroline atolls* (Fennah, 1956b: lighter or hyaline. Male genitalia (Figs 63-69) 117) (BPBM). Ponhpei Is. (Ponape)* (Fennah, resembling those of S.kolophon, but outer distal 1956b: 117) (BPBM), Truk Is. (Ton I.)* (Fen- process of parameres longer and straight; nah, 1056b: 117) (BPBM), Yap Is. (Map I.)* aedeagus with an acute tip (blunt in (Fennah, 1956b: 117; 1971a: 573) (BPBM). S.kolophon). Externally the species can be NEW CALEDONIA* (Fennah, 1969a: 3.5) distinguished from other Sogatella species by (BPBM). NORTHERN MARIANA IS.: the coloration of the head as mentioned above Saipan I. (Fennah, 1956b: 116) (BPBM) PIT- combined with the presence of a brown spot at CAIRN I.* (Fennah. 1958: 194-195) (BPBM). claval apex. SOCIETY IS.: Raiatea I.* (Fcnnah, 1058a: Remarks. S. molinus is transferred here from 194-195) (BPBM). SOLOMON IS.: Guadal- Sogatodes to Sogatella for the following reasons: canal I.* (BMNH, BPBM), Malaita 1. (BPBM), Fcnnah (1963a) discriminated Sogatodes from New Georgia Is.* (BPBM). TONGA IS.* (Fen- the genera Sogatella and Matutinus mainly by nah, 1967a: 46) (BPBM, MA). WESTERN the form of the ovipositor and by the carination SAMOA: Savaii I.* (MA), Upolu I.* (MA) of the thorax. However, re-examination of the Australian region: AUSTRALIA: Queens- holotype of the type-species Sogatodes molinus land* (Kirkaldy, 1907: 157; Fennah, 196Sb: 47) revealed the presence of the complete set of (BPBM, MA), Northern Territory', Kakadu characters that we consider define the genus NP (MA). Sogatella. Moreover, it appears that the shape of Nearctic region: BERMUDA IS.* (BMNH). the dorsal margin of the ovipositor and its denta- USA.: Florida*, Texas. Georgia (Beamer. 1952: tion varies in other species included in Sogatodes 111, as Sogata meridiana) (KU). and even in closely related Sogatella species, and Neotropical region: EQUADOR* (BPBM). is a rather weak character. Therefore we con- JAMAICA, MONTSERRAT, SANTA LUCIA, sider Sogatodes molinus congeneric with VENEZUELA (all Fennah. IYSY: 25')). Sogurellu. Consequently. the genus Sogarodes GUYANA, MEXICO" (both Fennah. 1963a: falls in synonymy with Sogarella as discussed 59) (USNM). ahovc. Atlantic ocean: ST HELENA I.* (Fennah, Within the S.kolophon-group S.molina re- 1963a: 59; 1976: 266; as S. kolophon atlantica and sembles S. kolophon, S.nigrigenis from Africa 1976: 267, as S.ne6ris) (BMNH). and S.colorata from the Seychelles in the shape of the male genitalia. Distribution and material examined. S.molina is exclusively a New World species. MEXICO* (Fennah. 1963a: 73) (BMNH). CAYMAN IS. The delphacid genus Sogatella and related groups 21

FIGS 63-69. Sogarr/la rnolirra (Fennah). 3 genitalia. scale line 0. t mm: 63-68. Mexico. paratype, 69. specimen from Bermuda Is.; 63, genitalia in repose ventrocaudal view; 64, diaphragm of genital segment caudal view; 65, genitalia without genital segment left lateral view; 66. left paramere maximum view; 67, aedeagus (Mexico) right lateral view; 68. aedeagus (Mexico) ventrodcaudal view; 69. aedeagus (Bermuda) right lateral view. 22 Manfred Aschc and Michael R. Wilson

(Fennah, 1971b: 321). U.S.A.: Florida* Sogatella diachenhea Kuoh, 1977: 441, Holotype (USNM). BERMUDA IS.* (BMNH). 8,CHINA (Shantung) (Anhwei Agric. Col- lege), not examined. Syn.n. Delphacodes dogensis Ishihara, 1952: 47, Holotype 6,JAPAN (Shikoku) (Ehime Univ., Sogatella nigrigenis (Jacobi) (Fig. 11) Matsuyama, Japan), not examined. Syn.n. Delphacodes 1ongifircife.a Esaki & Ishihara, 1947: Delphax nigrigenis Jacobi, 1917: 530, KENYA. 41, Holotype 6,JAPAN (Kyushu) (Kyushu Type material assumed deposited in either Univ., Fukuoka, Japan), examined. Syn.n. Humboldt Museum, Berlin or Dresden Sogatella longifurcifera Fennah, 1963a: 53. Museum, not examined. Liburnia matsumurana Metcalf, 1943: 364, Sogatella nigrigenis Fennah 1963a: 55. ITALY (Sicily), not examined. [given as nom.nov. for Delphax furcata Matsumura, Diagnosis. In external appearance similar to 1910b: 34 (nec Delphax furcata Provancher, S. vibix in possessing dark genae but differing in 1872)] Syn.n. the shape of the parameres. Male genitalia al- Sogatella matsumurana: Nast, 1975: 2. most as in S.kolophon, but laterodorsal angles of Delphacodes panicicola Ishihara, 1949: 51, genital segment broadly rounded and the tip of Holotype 6,JAPAN (Honshu) (Ehime the aedeagus acutely tapering (as in S.furcifera). Univ., Matsuyama, Japan), examined, syn. Outer distal process of the parameres continu- with S.longifurcifera by Ding etal. (1981). ously tapering, straight. Stramineous, genae and Sogatella panicicola Fennah, 1963a: 76. abdomen dark brown. Sogatella parakolophon Linnavuori, 1973: 108, Remark$. bennah (IY63a) apparently did not Holotype 8,SUDAN (Equatoria), (AMNH), examine the type material of S.nigrigenis but his examined. Syn.n. interpretation of the species based on specimens from Uganda has been followed. Within the Diagnosis. In coloration S.vibix is stramin- S. kolophon-group the relationships of this cous or pale yellow. Males have dark brown species are unclear: it resembles S. vibix in the genae similar to S.nigrigenis. S. vibix differs from coloration of head and tegmina, and S. kolophon other Sogatella species in the shape of the male in the shape of the parameres, also in other genitalia. The parameres have the outer process characters it combines the configuration of these of the apical birfurcation dilating from base to two species middle then tapering to apex with dorsal margin Distribution and material examined. Only forming a blunt angle (Figs 12,77, 81-91). known at present from East Africa: KENYA* Remarks. S. vibix individuals vary in size, col- (Jacobi, 1917: 530), UGANDA* (Fennah, oration and genital structures to a very similar 1963a: 56) (BMNH), SUDAN: Blue Nile, extent in all populations studied (see variation of Equatoria (Linnavuori, 1973: 108) (AMNH). parameres Figs 84-91) The synonymies of S. catoptron, S.parakolophon and D.longifurcifera (this species was damaged) and the con- specificity of D.panicicola are based on the Sogatella vibix (Haupt) (Figs 12, 70-91) examination of the holotypes. the synonymy of S.matsumurana on the study of topotypic Liburnia vibix Haupt, 1927: 13, Lectotype 6, material from Sicily. The synonymies of paralectotype P (here designated), ISRAEL S.diachenhea and D.dogensis we deduce from (collection Haupt, Jena, East Germany), the original descriptions and illustrations. The examined. figure of S.diachenhea (Kuoh, 1977: 442, Fig. 2) Sogatella suezensis Linnavuori, 1964: 341, mis- shows the lateral view of the head with the genac identification [not suezensis Matsumura 1910. pale rather than dark coloured as is typical for Nast , 1975 transferred Delphax suezensis to S. vibix. The shape of the parameres figured by Toya]. Kuoh and the fact that in almost all populations Sogatella catoptron Fennah, 1963a: 54, studied (including some from China) lighter col- Holotype d, ISRAEL (BMNH), examined. oured specimens could be found, convinces us Syn.n. that this species is synonymous with S.vibix. The delphacid genus Sogatella and related groups 23

Externally S. vibir resembles S.nigrigenis; Azores (Lindberg, 1941: 26; 1954a: 8; 1960: 33) however, at present closer relationship to any Madeira (Lindberg, 1961: 62), the Canary Is. particular species of the S.kolophon group can- (Lindberg, 1954b: 184) and the Capeverde Is. not be assessed. (Lindberg, 1958: 152) concern either S.kolophon Distribution and material examined. S.vibix is or Xnigeriensis. widely distributed ifi the Palaearctic region, the Numerous specimens (>400) from various Lthiopian Kegion, the Oriental Kegion, Aus- localities in all parts of the species occurrence tralia and the Western Pacific. It is absent in the have been examined. New World. The records of S.vibir from the Palaearctic region: AFGHANISTAN'

FIGS 70-73. Sogarella vibix (Haupt); 70-72. coloration of head and thorax, 6 specimen from Israel, scale line 0.5 mm;70, head and thorax, dorsal view; 71, head frontalview; 72, head left lateral view. 73, leftfore- wing, macropterous d from Israel, scale line 0.5 mm. 24 Manfred Asche and Michael R. Wilson

(Dlabola, 1971: 381; 1972: 203) (MAK). CY- 1977: 442; 1983: 117, as S.diachenhea), Anhui* PRUS (Lindberg. 1948: 126). EGYPT* (Lin- (BMNH), Fukien+ (BPBM). INDIA',' (LOB). navuori, 1Y64: 341; IY6Y: 2U8, as Colligypona INDONESIA: Ceres* (BPBM), Irian Jaya* (Sogatella) suezensis; 1964, loc. cit. as C. (BPBM), Java* (BPBM) Kalimantan* (Sogarella) caroptron: Fennah, 1963a: 55, as (BPBM), Sulawesi* (BMNH, MA), Sumatra' S.catoptron. GREECE* (Dlabola, 1Y7Ya: 25, as (BPBM). LAOS* (BPBM). PAKISTAN* Sogarella suezensis; Asche & Remane, 1982: (Dlabola, 1971: 381) (LOB). PHILIPPINE IS.*: 263: Asche & Hoch, 1982: 60; Drosopoulos, Luzon*, Mindanao*, Palawan* (BPBM). SIN- Asche & Hoch, 1986: 32). IRAN (Dlabola, GAPORE* (BPBM). THAILAND* (BPBM). 1960: 5; 1971: 381). IRAQ* (CLV). ISRAEL* TAIWAN (Fennah. 1963a: 55 as Xkongifurcif- (Haupt, 1927: 13; Linnavuori, 1962: 26; Fennah, eru. VIETNAM (Fennah, 1978: 222 as S.lon- 1963a: 52; 55, as S.catoptron. ITALY: Sicily* gifurcifera). (Oshanin, 1907: 315, as Delphax furcifera; Mat- Pacific region: BONIN IS. (Fennah, 1971: sumura, 1910b: 34 as Delphax furcata; Haupt, 574, as SJongifurcifera). FIJI IS.*: Viti Levu I. 1924: 229, as D.furcata; Servadei, 1958: 9; 1967: (I male paratype of D.kolophon, misidentifica- 82, as Liburnia matsumuriana (sic); Nast, 1975: tion, BPBM). NEW CALEDONIA (Fennah. 2, as S.matsumurana). JAPAN: Kyushu* (Esaki IY6Y: 34, as S.fongifurcifera). RYUKYU 1S.* & Ishihara, 1947: 47 and Ishihara, 1949: SO, as (BPBM). SOLOMON IS.: Choiseul I.* D.longifurcifera), Shikoku (Ishihara, 1952: 47. (BPBM) Guadalcanal I.* (BMNH, BPBM), as Delphacodes dogensis: 45, as D.panicicola), Malaita I. (BPBM), New Georgia Is* (BPBM), Honshu* (Ishihara, 1949: 51 and 1952: 45, as Santa Isabel I.* (BPBM). TONGA IS.: Vavau D.panicico/a).JORDAN (Fennah. 1963a: 55, as I.* (Fennah, 1967a: 46, as S.longifurcifera) S.catoprron; Dlabola, 1965: 427). KOREA" (BPBM), Tongatapu I. (MA). VANUATU: Es- . (Lee & Kwon, 1977: 103 and 1980: 32, as S.lon- piritu Santo I. (BPBM). gifurcifera and S.panicicola) . LE B ANON * Australian region: AUSTRALIA: Queens- (Dlabola, lY65: 427; Asche & Hoch, 1Y86: 18). land* (Fennah, 196Sb: 47, as Xlongifurcifera MONGOLIA (Dlabola, 1967: 212). MOROCCO* (BPBM, MA), Northern Territory* (MA). (Lindberg, 1963: 32). TURKEY: Anatolia* (Dlabola, 1971: 381; 1981: 155; Lodos & Kal- kandelen. 1980: 112, as Toya suezensis; Asche, Soga tella albofimbriata-g ro up (two 1982: 83) (MA). SAUDI ARABIA (Dlabola, species) 1979b: 117, as Sogatella suezensis. U.S.S.R.: Ukraine, Krim (Logvinenko, 1970: 628; 1975: S.albofimbriata, S.krugeri 127, as Sogatella suezensis), Maritime Territory (Vilbaste, 1968: 21, as S.longifurcifera: 22, as Diagnosis. Male genitalia with parameres S.panicicola; Anufriev & Emeljanov, 1988: 361, slender to apex, elongate, in repose almost as S.longifurcifera. YUGOSLAVIA (Jankovic. reaching the dorsolateral angles of the genital 1975: 123, as Sogata vibix; Jankovic & Papovic, segment and the inner distal process almost ab- 1981: 129, as Sogatella suezensis. sent or only forming a very small edge (Figs 13, Ethiopian region: SUDAN: Northern Pro- 14). vince, Blue Nile, Upper Nile, Kordofan, Bharel Ghazal. Equatoria' (Linnavuori. 1973: 108, as S.suezen.sis and as S.parakolophon). Sogatella albofimbriata (Muir) (Fig. 13) Oriental region: BISMARK IS.: New Ireland I.* (BPBM), New Britain I.* (BPBM). Sogata albofimbriata Muir, 1926b: 33, Holotype BOUGAINVILLE I.* (BPBM). CAMBO- d, SOUTH AFRICA (Cape Province), DIA* (BPBM). CHINA: Shantung (Kuoh, (BMNH), examined.

FIGS 74-80. Sogarella vibix (Haupt), 6 genitalia, scale line: 0.1 mm; 74-79, specimen from India, 80, lectotype d from Israel; 74, genitalia in repose ventrocaudal view; 75. diaphragm of genital segment caudal view; 76, genitalia without genital segment, left lateral view; 77, left paramere, maximum view; 78, aedeagus (India), right lateral view; 79, aedeagus (India), ventrocaudal view; 80, aedeagus (Israel), right lateral view. The delphacid genus Sogatella and related groups 25 26 Manfred Asche and Michael R. Wilson

Chloriona albofimbriata: Fennah, 1958b: 204. Sogatella krugeri (Muir) c0mb.n. (Fig. 14) Chloriona (Sogutella) albofimbriata Fennah, 1958~:490. Sogata krugeri Muir, 1929b: 209, Holotype 6, Sogatella albofimbriata Fennah, 1964: 141. SOUTH AFRICA (Cape Province) Sogutodes albofimbriatus Medler, 1980: 62. (BMNH), examined. Sogata rhodesi Muir, 1929b: 208, Holotype 6, SOUTH AFRICA (Natal) (BMNH), Diagnosis. Externally S.krugeri is very similar examined. Synonymized by Asche. 1988: 206. to S.albofimbriuta, but somewhat smaller. It differs from all other Sogatella species by the struc- Diagnosis. Xalbojimbriata is a small and slender ture of the male genitalia and the shape of the Sogatella species. Genae dark brown, frons vertex. The laterodorsal angles of the genital stramineous or light brown, median carina over- segment not produced, but broadly rounded, layed by a broad white stripe which continues angles not bent mediad, the lateral arms of the over all parts of vertex; lateral portions of pro- U-shaped diaphragm process moderately long. notum pale yellow. Tegmina hyaline, slightly in- Parameres with a broad base then narrowing to fumed, a broad brown longitudinal band from apex, almost straight, outer margin slightly base to apex extending from inner margin to sinuate, apically truncate, inner angle of apex media. Differing from all other Sogatella species slightly produced mediad (Fig. 14). Aedeagus by the structures of the male genitalia: twisted with an oblique row of about 20 very tine Laterodorsal margin of genital segment slightly teeth on the left side, from right to ventral side a produced and rounded, edges slightly bent short row of 0-4 teeth mediad; lateral processes of the dorsal margin of Vertex very narrow and acutely produced (1.7 the diaphragm rather long, directed dorsad; pa- times longer in mid-line than broad at base), rameres broad at base, conically narrowing to separated from frons by a weak transverse apex, subapically on the inner margin a distinct carina. Vertex white, area of frons, genae and edge (Fig. 13); aedeagus as long as parameres, lateral portion of pronotum dark brown, post- on left side a row of numerous minute teeth, on and anteclypeus pale yellow, carinae of frons right side a row of about 12-15 teeth. male white or pale yellow. Tegmina similar to S.al- genitalia figured in Asche (1988). bofimbriata with a broad longitudinal brown suf- Remarks. As discussed in Asche (1988), it is fusion covering the inner half. difficult to decide evolutionary trends in the pa- Remarks. The systematic position of S. krugeri rameres of the genus, i.e. whether the S.al- is somewhat difficult. By virtue of the coloration bofimbriutu-like form marks the beginning or and the male genitalia it is placed easily into the the endpoint for the forked parameres of the Sogutella concept and within it is clearly related S.kolophon or the S.furcifera type. Moreover, to S.albofimbriata. However, S.krugeri differs similarly elongate parameres are also found in from other Sogatella species by the acutely pro- some non-Sogatella species, e.g. in some Toyu duced head and the vertex separated from the species. Within Sogutella, S.albofimbriata is ap- frons by a transverse carina. In this character it parently closely related to S. krugeri. resembles Delphax eupompe Kirkaldy (here Distribution and material examined. Ethio- placed in Latrisrria see below), but in this species pian region and Mauritius. SOUTH AFRICA: the diaphragm structure is not of the Sogatella Cape Province* (Muir, 1926b: 33; 1929b: 213), form and the phallotreme is subapically on the Zululand (Muir, 1929: 213 as S.rhodesi,Fennah, right side and not subapically left as in all 1958b: 204) (BMNH), Swaziland* (UST). Sogatella species (see Figs 124-133). MAURITIUS* (Fennah, 1% 141) (Bh4M-I). Distribution and material examined. AL- SENEGAL (Fennah, 195%: 4W). NIGERIA" DABRA* (BMNH). TANZANIA* (BMNH). (Medler, 1980: 62). IVORY COAST* (Asche, 1988: SOUTH AFRICA: Cape province* (Muir, 2%) (M,MA. BMNH). OMAN* (BMNH). 1929: 209), Natal' (UTS). The delphacid genus Sogatella and related groups 27

81 82

85 87

FIGS 81-91. Sogatella vibix (Haupt), intraspecificvariationof parameres, scale line 0.1 mm; 81, Israel, lectotype; 82, Israel, 61; 83, Israel, same population 62; 84, Israel, same population, 83; 85, Egypt; 86, Sudan (Equatoria), holotype of Xparakolophon Linnavuori; 87, Japan (Hokkaido); 88, Philippine Is. (Luzon); 89, Solomon Is. (Guadalcanal); 90 Tonga Is. (Tongatapu I.); 91, Saudi Arabia, left-right variation in the same variation. 28 Manfred Asche and Michael R. Wilson

FIGS Y2-Y5. Tugosodes anomulus (Muir) d genitalia, holotype from Brazil, genitalia mounted in balsam on slide, scale line 0.1 mm; 92, diaphragm of genital segment, caudal view; 93, left paramere, left lateral view; 94, genitalia without genital segment and parameres, left lateral view; 95, aedeagus right lateral view. FIGS 96-100. Tugosodes cubanus (Crawford). d genitalia, specimen from Ivory Coast, scale line 0.1 mm; 96, diaphragm of genital segment, caudal view; 97, left paramere maximum view; 98, aedeagus, left lateral view; 99, aedeagus right lateral view; 100, aedeagus, ventrocaudal view. The delphacid genus Sogatella and related groups 29

FIGS 101 -103. Tugosodrs nuuticu (Muir). 6 genitalia. paratype from Guyana, genitalia mounted in balsam on slide, scale line 0.1 mm; 101, diaphragm of genital segment, caudal view; 102, left paramere, maximum view; 103, genitalia without genital segment and parameres, left lateral view (basal parts of aedeagus and connective missing). FIGS 104-107. Tugosodes nicias (Fennah). 6 genitalia, specimen from Papua New Guinea, scale line 0.1 mm; 104, diaphragm of genital segment caudal view; 105, left paramere, maximum view; 106, genitalia without genital segment, left lateral view; 107, aedeagus, right lateral view. 30 Manfred Asche and Michael R. Wilson

110 108

FIGS 108-114. Tagosodes orizicolus (Muir). 6 genitalia, scale line 0.1 mm; 109, holotype from Guyana, all others paratypes; 108. diaphragm of genital segment, caudal view; 109. left paramere, maximum view (undissected holotype); 110, left paramere, maximum view (paratype 1); 111, left paramere, slightly bent caudad (paratype 1); 112, genitalia without genital segment, caudal view (paratype 1); 113, tip of aedeagus (paratype 2); 114, aedeagus right lateral view (paratype 1).

FIGS 115-119. Tagosodespusanus (Distant), 6 genitalia, specimen from Sulawesi, scale line 0.1 mm; 115. diaphragm of genital segment caudal view; 116, left paramere maximum view; 117, genitalia without genital segment lateral view; 118, aedeagus right lateral view; 119, aedeagus, dorsal view. FIGS 120-123. Tagosodes wallacei (Muir & Giffard), 6 genitalia, paratype from Cuba, mounted in balsam on slide, scale line 0.1 mm; 120. diaphragm of genital segment, caudal view; 121, left paramere, maximum view; 122, genitalia without genital segment and parameres, left, lateral view (basal part of the aedeagus and connective missing, position of processes of anal segment probably due to embedding in balsam); 123, aedeagus, right lateral view. The delphacid genus Sogarella and related groups 31 32 Manfred Asche and Michael R. Wilson

TagosWres gen.n. pressed, if at all; from its base slightly bent dorsad; teeth irregular and/or forming rows; phallo- Type species: Dicranotropis cubanus Crawford, treme apically or subapically on left or right 1914: 595. sidc.

Description. Small delphacids, about 3-4 mm Diagnosis. Tagosodes species resemble in length (including tegmen). In external charac- Sogatella species in coloration and external ap- ters similar to Sogatella species. pearance but differ in the structure of the male Vertex slender, almost parallel sided, about genitalia. In particular the diaphragm never 1.4-1.7 times longer in middle line than at base; forms a broad U-shape as in Sogatella but dis- posterior compartments shallowly concave with plays various different configurations as detailed median carina weak or absent; anterior cell long above. The aedeagus is much less compressed, if and slender, anteriorly ending at tip of vertex or at all, compared with Sogatella; the teeth may be reaching down onto frons; vertex and frons in rows but in most species additional irregularly forming a slightly acute angle, transition of ver- spaced teeth occur; the aedeagus is never twisted tex to frons mostly rounded. Frons high and as found in Sogatella but often forms a simple, slender, about 2 times or a little more as high as more or less straight tube. Tagusodes may be maximum width; widest at or shortly before the separated from Matutinus and Latistria mainly frontoclypeal suture; median carina prominent, by the shape of the diaphragm of the genital seg- especially at the transition of vertex to frons. ment (forming a shoe-shaped structure in Post- and anteclypeus with distinct median Matutinus and a sinuate bilobed cross plate in carina. Rostrum attaining the hindcoxae. An- Latistria. Tagosodes species can be readily sepa- tennae: Second joint about 1.8-2 times longer rated from Sogatellana species by having only than tirst; arrangement ot antennd sensory tield one pair of anal segment processes rather than 16/7. Male drumming organ with elongate two in Sogatellana. apodcrnes of the second abdominal sternite di- Remarks. Several species groups are recog- rected dorsad. Post-tibal spur foliaceous with nized within Tagosodes based on the form of the numerous minute teeth on its hind margin similar to diaphragm structure and some of these groups that of Sogatella species. Proportions of the hind may be monophyletic and may later be found to tarsi, tegmina and wings as in Sogatelfa species. have generic status. Further work is necessary to Coloration. Along the middle line from the re-describe some of the species included below anterior cell of vertex to the caudal tip of the and further re-arrangement cannot be ruled out. mesonoturn a white band, the pro- and mesonotum parts laterad of that band brown or black. Male genitalia. Genital segment ring-like, in List of included species (alphabetical caudal view subcircular or oval; laterodorsal order) angles not excessively produced; central part of diaphragm raised, mediodorsal margin either Tagosodes albifacies (Caldwell) c0mb.n. caudodorsad produced in a T-shaped (Figs 92, Sogata albifacies Caldwell, 1951, in Caldwell 104) or triangular (Fig. 96) area, in a distinct rec- & Martorell 1951: 173 (Puerto Rico) tangular (Fig. 101) or in a U- or W-shaped pro- Tagosodes albolineosus (Fowler) c0mb.n. trusion (Fig. 108), or central part bulbous oval or Liburnia albolineosus Fowler, 1905: 135 ridged and with numerous minute teeth (Figs (Mexico). 115, 120); opening for the parameres of broad Tagosodes anomalm (Muir) c0mb.n. trapezoid shape (e.g. Fig. 96). Anal segment (Figs 92-95) with two spinose processes originating at the Sogata anomala Muir, 1926a: 27, Holotype 8, laterodorsal angles on the ventral side, processes (Brazil), (BPBM), examined. medially approaching each other, in lateral view Tagosodes approximatus (Crawford) c0mb.n. slightly curving ventrad. Parameres variously Megamelus approximata Crawford 1914: 622 shaped, with broad base, slender middle section (Nicaragua). and dilated apical area; mostly diverging from Tagosodes baina (Ding & Kuoh) c0mb.n. base, then at least inner apical angle converging Himeunka baina Ding & Kuoh, 1981, in Kuoh to middle. Aedeagus tubular, only slightly com- et al., 1981: 24 (China). The delphacid genus Sogatella and related groups 33

Tagosodes biangulatus (Muir) c0mb.n. Chloriona (Sogatella) panda Fennah, 1958c: Sogata biangulata Muir, 1929a: 84 (Argentina). 491 (Ivory Coast), syn. mentioned in Fennah, Tagosodes candiope (Fennah) c0mb.n. 1969b: 56. Sogatodes candiope Fennah, 1975: 97 (Sri Remarks. T.cubanus is found in south and Lanka). central America and also in West Africa. Male Tagosodes cubanus (Crawford) c0mb.n. (type genitalia illustrated by Asche (1988). species) (Figs 96-100) Tagosodes dorsolineatus (Beamer) c0mb.n. Dicranotropis cubanus Crawford, 1914: 595 Sogata dorsolineatus Beamer, 1952: 112 (Cuba). (U.S.A.: Texas). Megamelus jlavolineatus Muir 1920: 143 Tagosodes elpenor (Fennah c0mb.n. (Nigeria). (mentioned by Muir & Giffard, Sogatodes elpenor Fennah, 1964: 139 1924: 12, syn. by Wolcott, 1923: 273). (Mauritius). Delphacodes pallidivitta Fennah, 1945: 433, Tagosodes fzaviceps (Muir) comb. n. Holotype 6 (USNM), paratype 6 6 (Trinidad) SogataJlaviceps Muir, 1929b: 210, Holotype 6 (BMNH), examined. Syn.n.

126

128

FIGS 124-128. Latistria eupompe (Kirkaldy) d genitalia, specimen from Australia (New South Wales), compared with holotype 6,scale line 0.1 rnm; 124, diaphragm of genital segment, caudal view; 125, left paramere, maximum view; 126, genitalia without genital segment and parameres, left lateral view; 127, aedeagus, right lateral view; 128, aedeagus, ventrocaudal view. 34 Manfred Asche and Michael R. Wilson

(South Africa: Zululand) (BMNH), examined. Remarks. Male genitalia illustrated by Asche Tagosodes incanus (Yang) c0mb.n. (1988). Sogatodes incanus Yang, 1989: 176, Holotype Tagosodes nauticus (Muir) c0mb.n. 6 (Taiwan) not examined. (Figs 101-103) Tagosodes melicerta (Fennah) c0mb.n. Sogata nautica Muir, 1926a: 26, Holotype 6 Chloriona (Sogatella) melicerta Fennah, (Guyana) (BPBM), examined. 1958c: 490 Holotype 6 (Portugese Guinea) Tagosodes nicias (Fennah) c0mb.n. (MNHN), examined. (Figs 104-107) Sogatodes neomphalus Asche, 1988: 209, Sogatodes nicias Fennah, 1965b; 45 (Papua Holotype 6 (Ivory Coast) (MNHN) examined. New Guinea). Syn.n. Tagosodes orizicolus (Muir) c0mb.n. (Figs 108-114)

FIGS 129-133. Latistria placitus (Van Duzee), male genitalia, specimen from Belau Is.: Babelthuap I., scale bar 0.1 mm; 129, diaphragm of genital segment caudal view: 130, left paramere maximum view; 131, genitalia without genital segment and parameres left lateral view; 132, aedeagus right lateral view; 133, aedeagus ventrocaudal view. The delphacid genus Sogatella and related groups 35

Sogata orizicola Muir, 1926a: 27 (Guyana). Tagosodes pusanus (Distant) c0mb.n. Sogata brazilensis Muir, 1926a: 26 (Brazil), (Figs 115-119) syn. by Fennah 1965a: 215. Sogatapusana distant, 1912: 191 (India). Remarks. It appears that the species presently Kelisia fieberi Muir, 1917: 331 (Philippines), known as S.orizicola may consist of several closely syn. by Fennah, 1971a: 574. related species that may be found in South Unkana formosella Matsumura, 1935: 72 America. Further work is necessary on this topic. (Taiwan), syn. by Fennah, 1971a: 574.

FIGS 134-138. Sogatellanu geranor (Kirkaldy), 8 genitalia, holotype from Australia (Queensland), scale line 0.1 mm; 134, diaphragm ofgenital segment, caudal view; 135, left paramere, maximum view; 136, anal segment, ventrocaudal view; 137, genitalia without genital segment, left lateral view; 138, aedeagus dorsal view. 36 Manfred Asche and Michael R. Wilson

Sogata striatus Qadri & Mirza, 1960: 117 assimilis Yang was described from just one spec- (Pakistan) nomen nudum. imen differing from pusanus only in proportions Himeunka chibana Tian & Kuoh, 1981, in of the frons and coloration of the tegmina. We Kuoh et al., 1981: 193 (China). Syn.n. regard this as merely variation in this widespread Sogatodes assirnilis Yang 1989: 178 (Taiwan). species. Syn.n. Tagosodes sternalis (Distant) c0mb.n. Remarks. The name Sogata striatus Qadri & Sogata sternalis Distant, 1916: 139 (Sri Mirza is considered invalid under article 13 of Lanka). the code of Zoological Nomenclature (1985). Tagosodes timaea (Fennah) c0mb.n. Based on the drawings in Qadri & Mirza (1960) Sogatella timaea Fennah, 1969b: 56 (Sudan). the species is, in any case, identical with Tagosodes wallacei (Muir I% Giffard) c0mb.n. T.pusanus.The synonymy of Himeunka chibana (Figs 120-123) is based on the drawings and description given Sogata wallacei Muir & Giffard, 1924: 13 by Tian & Kuoh in Kuoh eta1 (1981). Sogatodes (Guyana).

FIGS 139-144. Sogatellanu quadrispinosa (Muir), 6 genitalia, holotype and another non-type specimen from Singapore, scale line 0. I mm; 139, diaphragm of genital segment, caudal view (holotype); 140, left paramere, maximum view (holotype); 141, anal segment, ventrocaudal view (non-type specimen); 142, genitalia without genital segment, left lateral view, tip of aedeagus broken and missing (holotype); 143, aedeagus, left lateral view (non-type specimen); 144, aedeagus, right lateral view (non-type specimen). The rlelphacid genus Sogatella and related groups 37

Latristria Huang et al. Latistria fuscipennis Huang & Ding Latistria fuscipennis Huang & Ding, 1980, in Latistria Huang et al., 1980: 166; type species Huang etal., 1980: 167 (China). L.testacea Huang et al., 1980: 166 by original Latistria placitus (Van Duzee) designation. (Figs 129-133) Diagnosis. Latristria species externally resem- Sogata placitus Van Duzee, 1937: 120 ble small slender Sogatella species, especially the (Caroline Is.). Xalbofimbriata group with anteriorly a Chloriona (Sogatella) euterpe Fennah, 1956b: somewhat acute vertex. However, Latistria differs 118 (Mariana Is.), syn. by Fennah, 1969a: 21. Sogatodes eupompe Yang, 1989: 170 misident. from Sogatella and all other genera displaying Latistria testacea Huang KL Ding the white longitudinal stripe across the vertex, pro- and mesonotum in the shape of the diap- Latistria testacea Huang & Ding, 1980, in hragm which mediodorsally forms a broad plate- Huang etal., 1980: 166 (China). like protusion with rounded lobe-like lateral edges (Figs 124, 129). The parameres are long Sogatellana Kuoh and slender with tapering or truncate apex, parameres in repose almost reaching the laterodorsal Sogatellana Kuoh in Huang et al., 1980: 169. corners of the genital segment. Type species S.marginuta Kuoh by original Remarks. A similarly shaped diaphragm in designation. combination with elongate parameres is found in some species currently placed in Toya Distant; Diagnosis. Resembles Sogatella in coloration however, these species lack the white dorsal and body proportions. Differs from all other re- stripe and they differ considerably in bodily lated genera of the Sogatellu-group by the pos- proportions. Further studies are necessary to session of two pairs of processes on the ventral further elucidate relationships in these groups of side of the anal segment (Figs 136, 137, 141, species. 142). Included species (alphabetical order) Remarks. The possession of the two pairs of anal segment processes is apparently also found Latistria eupompe (Kirkaldy) c0mb.n. in the type species of Himeunka. If so the two (Figs 124-128) genera may be synonymous. Delphux eupompe Kirkaldy 1907: 162 (Aus- tralia, Fiji) Delphux ochrius Kirkaldy 1907: 157 (Au- Included species (alphabetical order) stralia, Fiji). Syn.n. Sogatodes infestus Yang, 1989: 172. Syn.n. Sogatellana costata Ding Remarks. This species is clearly placed in Sogatellana costuta Ding, 1985: 432 (China). Latistria by virtue of the form of the male Sogatellanafusca Tian & Ding genitalia with the tapering apex to the para- Sogatellana fusca Tian & Ding, 1980, in meres. It is possible that one of the listed Huang et al., 1980: 170 (China: Jiangsi). Chinese species will be a synonym of either of Sogatellana geranor (Kirkaldy) c0mb.n. L.eupompe or L.placitus (see below). Delphax (Figs 134-137) ochrias was described from brachypterous Delphax geranor Kirkaldy, 1907: 158 (Aus- females. Studies in Sulawesi and Australia have tralia). shown that the short-winged females of Delphax sponsa Kirkaldy, 1907: 148, Holotype L.eupompe are of different colour to the males; 0 (Australia) (BPBM), examined. Syn.n. they are stramineous and resemble in all charac- Sogutelluna quadrispinosa (Muir) c0mb.n. ters the holotype female of Delphax ochrias. On {(Figs139-144) this basis the synonymy is proposed here. The Sogata 4-spinosu Muir, 1919: 526, Holotype d , synonymy of S.infestus Yang is made on the (Singapore) (BPBM), examined. basis of the original drawings. Sogatellana marginata Kuoh Latistria flavotestacea Kuoh Sogatellana marginata Kuoh, 1980, in Huang Latistriajlavotestacea Kuoh, 1980, in Huang et et al., 1980: 169 (China: Guangdong). al., 1980: 168 (China). Sogatellana semicirculara Yang 38 Manfred Asche and Michael R. Wilson

Sogatellana semicirculara Yang, 1989: 196 References (Taiwan). Sogatellana yunnanana (Ding) c0mb.n. Ammar. E.D. (1977) Biology of the planthopper Himeunka yunnanana Ding, 1985: 431 Sogatella vibiv (Haupt) in Giza, Egypt (Horn. Del- phacidae). Deutsche Entomologische Zeitschrift, 24, (China: Yunnan). 151- 158. Ammar, E.D., Lamie, 0. & Khodeir, LA. (1980) Biology of the Planthoppcr Sogatella furcifera Horv. Himeunka Matsumura & lshihara in Egypt (Horn. Delphacidae). Deutsche Entomo- logische Zeitschrift, 27, (l-3), 21-27. Himeunka Matsumura & Ishihara, 1945: 70. Anufriev. G.A. (1977) Delphacids (Homoptera, Au- Type species H.tateyamaella Matsumura by chenorrhyncha) of the Kurile Islands fauna. Zoologischeskii Zhurnal, 56, (6).855-869. subsequent designation. Anufriev, G.A. & Emeljanov, A.F. (1988) Keys tothe Insects of the Far East of USSR, in 6 volumes. Remarks. Kuoh et al. (1981) revised Hemiptera, pp. 1-472. Himeunka with descriptions of two new Chinese Asehe, M. (1982) Beitragc ziir Delphaciden-Fauna species, H.baina and H.yunnanana, in addition der Tiirkei (Anatolien) (Homoptera Cicadina Del- phacidae). Marburger Entomologische Publika- to a re-description of what he considered to be tionen, 1, (7), 71-98. H.tateyamaella. One of the Chinese species Asche, M. (1988) Delphacidae from Cote d’lvoire (H.baina) does not display the 4 processes of the (Homoptera Fulgoroidea). Revue FranGaise d’En- anal segment and is transferred to Sogatellana tomologie (N.S.),10, (2), 151-231. (see above). The other species, H.yunnanana, Asche, M. & Hoch, H. (1982) Beitrage ziir Dclphaci- den-Fauna Griechenlands I1 (Homoptera Cicadina possesses all essential characters of Sogatellana Delphacidae). Marburner Entomolonische Pub- I and is transferred to this genus. Unfortunately likaiionen, 1,’ (7), 37-70: we have not seen any specimens of H.tateyamaella Asche. M. & Hoch. H. (1986) Cixiidae and Del- but the illustrations given by Kuoh et al. (1981) phacidae (Homoptera-Fulgoroidea) from Leba- non-a preliminary synopsis. Proceedings ofthe 2nd clearly represent H.tateyamaella a a Sogatellana Inirrnationul Congress concerning the Rhynchota species by virtue of the two pairs of anal segment Fauna of Balkan and Adjacent Regions, pp. 8-13. processes in the male genitalia. If this is the rase Asehe, M. & Remane, R. (1982) Beitrlge ziir Del- then Sogatellana becomes a junior synonym of phaciden-Fauna Grieehcnlands l (Homoptera Himeunka and the .species listed above would Cicadina Delphacidae) . Marburger Entomologische Publikationen, 1, (6), 231-290. become Himeunka species. This situation will Beamer, R.H. (1952) One old and five new species of not be resolved until the type material of Delphacine Fulgorids (Homoptera: Fulgoridae). H.tateyarnaella can be compared with Sogatel- Journal of the Kansas Entomological Society, 25, lana species. 111-115. Caldwell, 1,s. & Martorell, L.F. (1951) Review of the Auchenorrhynchous Hornoptera of Puerto Rieo, Acknowledgments Part 11. The Fulgoroidea except Kinnaridae. Jour- nal of the Agricultural University, Puerto Rico, 34, (2), 133-269, i-x. We should like to thank the following for their Crawford, D.L. (1914) A contribution toward a help; Dr R. Kisimoto for his interest in the work, monograph of the Hornopterous insects of the fam- for sending specimens from Japan and for assist- ily Delphacidae of North and South America. Pro- ance in arranging loans of type material, Dr Y. ceedings of the United States National Museum, Washington, 46,557-640. Hirashima (Kyushu University, Japan), Dr T. Ding Jin-Hua (1985) Descriptions of three new species Ishihara and Dr M. Miyatake (Ehime Univer- of Delphaeidae (Homoptera). Acta Entomologica sity, Japan) for the loan of specimens. Dr S. W. Sinica, 28, (4). 430-433. Wilson (Central Missouri State University, Ding, J.H. etal. (1981) [A revision of scientific name on Chinese Sogatella panicicola (Ishihara) U.S.A.), T. J. Henry (USNM) and R. Schuh (Homoptera: Aeropodidae (sic))]. Kunchong (AMNH) also arranged the loan of specimens. Zhishi, 18, (3), 142. [In Chinese]. Dr G. Nishida (BPBM) for help in allowing Distant, W.L. (1912) Descriptions of new genera and examination of specimens in the Bishop species of Oriental Homoptera. Annals and Museum. M. D. Webb and W. R. Dolling Maxazine of Natural History, (8),9, 181-194. Distant. W.L. (1916) Rhynchota. Homoptera: Ap- (BMNH) made useful comments on the manus- pendix. The Fauna of British India, including cript and H. Hoch also read the manuscript and Ceylon and Burma, Vol. 6, pp. 1-148. Taylor & helped with the preparation of the figures. Francis, London. The delphacid genus Sogatella and related groups 39

Distant, W.L. (1917) Rhynchota. Part ii: Suborder Fennah, R.G. (1Y58a) Fulgoroidea of South-Eastern Homoptera. The Percy Sladen Trust Expedition to Polynesia. Transactions of the Royal Entornological the Indian Ocean in 1905, under the leadership of Society of London, 110,117-220. Mr. J. Stanley Gardiner, M.A. The Transactions of Fennah, R.G. (1958b) Hemiptera. Homoptera. Ful- the Linnean Society of London, Second Series, goroidea. South African Animal Life. Results of the Zoology 17,273-322. Lund University Expedition in 1950-1951, 5, 193- Dlabola, J. (1960) Iranische Zikaden (Homoptera, 211. Auchenorrhyncha). Stuttgarter Beitrage ziir Natur- Fennah, R.G. (1958~)Fulgoroidea from West Africa. kunde, 41,l-24. Bulletin de l’lnstitut Franfaises de Afrique Noire, 20, Dlabola, J. (1965) Jorhanische Zikaden (Homoptera Ser. A, 460-538. Auchenorrhyncha). Acta Entomologica Musei Na- Fennah, R.G. (1959) Delphacidae from the Lower tionalis Pragae, 36,419-450. Antilles (Homoptera: Fulgoroidea). Bulletin of the Dlabola, J. (1967) Ergebnisse der 2. mongolisch- British Museum (National History), Entomology, 8, tschechoslovakischen entomologisch-botanischen 245-265. Expedition in der Mongolei. Nr. 12: Reisebericht, Fennah, R.G. (1961) Le Parc National de Niokolo- Lokalitateniibersicht und Bearbeitung der gesam- Koba. XXXIII. Homoptera Fulgoroidea. Mkmoires melten Zikaden (Homopt. Auchenorrh.). Acta de l‘institut Franfaises de I’Afrique Noire, 62, 305- Faunistica Entomologica Musei Nationalis Pragae, 320. 12,207-230. Fennah, R.G. (1963a) The Delphacid species-com- Dlabola. J. (1971) Taxonomische und chorologische plex known as Sogata furcifera (Horvath) (Homop- Erganzungen der Zikadenfauna von Anatolien, tera: Fulgoroidea). Bulletin of Entomological Re- Iran, Afghanistan und Pakistan (Homoptera. Au- search, 54,45-79. chenorrhyncha). Acta Entomologica Bohemo- Fennah, R.G. (1963b) Erratum toFennah 1963a. Bul- slovaca, 68, (6), 377-396. letin of Entomological Research, 54, 166. Dlabola, J. (1972) Beitrage ziir Kenntnis der Fauna Fennah. R.G. (1963~)New Delphacidae (Homoptera: Afghanistans. Homoptera Auchenorrhyncha. Acta Fulgoroidea) from South America and West Africa. Musei Moraviae, 56157 (1971 -72). 189-248. Annals and Magazine of Natural History, 13, (9, Dlabola, J. (1979a) Chorologische Erganzungen zur 689-700. Zikadenfauna des Mittelmeergebietes (Homopt- Fennah, R.G. (1964) Delphacidae from Madagascar era- Auchenorrhyncha). Sbornik Narodniho Muzea and the Mascarene Islands (Homoptera: Ful- Praze (B), 33, (1-2), 1977 (1979), 21-40. goroidea). Transactions of the Royal Entomological Dlabola, J. (1979b) Insects of Saudi-Arabia. Homop- Society of London, 116, (7), 131-150. tera. Fauna of Saudi Arabia, 1,115-139. Fennah, R.G. (1965a) Sogatodes brasilensis (Muir), a Dlabola, J. (1981) Ergebnisse der tschechos- new synonym of S. orizicola (Muir) (Fulgoroidea, lowakische-iranischen entomologischen Expedition Delphacidae). Bulletin of Entomological Research, nach dem Iran (1970 und 1973) (Mit Angaben iiber 56, (2), 215-217. einige Sammelresultate in Anatolien) Homoptera: I:ennah, R.G. (1965b) Delphacidae from Australia Auchenorrhyncha (11. Teil). Acta Entomologica and New Zealand (Homoptera: Fulgoroidea). Bul- Musei Nationalis Pragae, 40, 127-31 1. letin of the British Museum (Natural History), En- Drospoulos, S., Asche. M. & Hoch, H. (1986) Con- tomology, 17, (1) ,3-59. tribution to the planthopper-fauna of Greece Fennah, R.G. (1967a) New species and records of Ful- (Homoptera Auchenorrhyncha Fulgoromorpha goroidea (Homoptera) from Samoa and Tonga. Delphacidae). Annales de l’lnstitut Phyto- Pacijic Insects, 9, (l), 29-72. pathologique Benaki (N.S.) (1983) (1986), 14, 1948. Fennah, R.G. (1967b) Fulgoroidea from the Emeljanov. A.F. (1982) [Fulgoroidea (Homoptera) Galapagos Archipelago. Proceedings of the Califor- collected in the Mongolian People’s Republic by the nia Academy of Sciences, Fourth Series, 35, (4), 53- entomofaunistic group of the Soviet Mongolian 102. complex biological expedition in 1970-1975.1 In- Fennah, R.G. (1969a) Fulgoroidea (Homoptera) from sects of Mongolia, 8, 69-122 [In Russian.] New Caledonia and the Loyalty Islands. Pacific Zn- Esaki, T. 19 Ishihara, T. (1947) Species nova vel minus sects Monographs, 21. 1-116. cognita Araeopidarum Japonaricum (Hemiptera). Fennah, R.G. (1969b) Delphacidae (Homoptera: Ful- Mushi, 17,39-42. goroidea) from the Sudan Republic. Acta En- Fennah, R.G. (1945) The Fulgoroidea, or Lanternflies tomologica Fennica, 26, 1-78. of Trinidad and adjacent parts of South America. Fennah, R.G. (1971a) Homoptera: Fulgoroidea Sup- Proceedings of the United States National Museum, plement. B. P. Bishop Museum, Insects of Mic- 95,411-520. ronesia, 6, (8), 563-609. Fennah, R.G. (1950) Fulgoroideaof Fiji. B.P. Bishop Fennah, R.G. 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