SYSTEMATICS Placement of the Faltala Oman Group (: Cicadellidae: ) with Descriptions of Three New Species

1 2 J. N. ZAHNISER AND M. D. WEBB

Ann. Entomol. Soc. Am. 97(4): 667Ð674 (2004) ABSTRACT Three new species of the Faltala Oman leafhopper group, Kramerana junina n. sp. and K. adusta n. sp. from the Andes Mountains of Peru and Clorindaia brasileira n. sp. from the Serra da Mesa of Brazil, are described. The tribal placement of the group and the high incidence of brachyptery and reduced (or absent) ocelli in the group are discussed. A revised key to the Faltala group genera and additional characters for Kramerana are provided.

KEY WORDS Deltocephalinae, , Hecalini, Neotropical, new species

OMAN (1938) DESCRIBED THE Faltala based on one Placement of the Faltala Group species from Argentina. Since then, four additional LinnavuoriÕs (1959) placement of the Faltala group closely related genera (Kramerana, Virganana, and in Athysanini was based on the Y-shaped connective, Aequicephalus described by DeLong and Tham- a character that cannot be considered a synapomor- bimuttu [1973], and Clorindaia described by Linna- phy of this poorly deÞned tribe, as members of several vuori[1975a]) and 12 further specieshave been de- deltocephaline tribes, indeed most, possess a similar scribed. Linnavuori and DeLong (1977) placed all Y-shaped connective. Later, Linnavuori (1975) also genera in the Faltala group of the Athysanini (Delto- noted the similarity in cephalic structure of Faltala and cephalinae). Linnavuori and DeLong (1977) gave a Clorindaia to Hecalini (Deltocephalinae) (see be- diagnosis of this group and a key to genera, and low). Recent morphological phylogenetic analyses of Blocker and Fang (1992) provided a useful synopsis of (Knight and Webb 1993) and Deltoceph- the included species and their distributions. In the alinae and related subfamilies (Zahniser and Dietrich, current study, two new species of Kramerana and one unpublished data) revealed several characters, espe- new species of Clorindaia are described, and newly cially of the female genitalia, that are useful in deÞning recorded details of the leg chaetotaxy and female some tribes and higher groups. In their analysis, Zah- genitalia for Kramerana are provided. The tribal place- niser and Dietrich recovered a clade of grass-special- ment of the Faltala group and the high incidence of izing , including the deltocephaline tribes brachyptery and reduced (or absent) ocelliinthe Hecalini, Stenometopiini (ϭStirellini), and Macros- group are discussed. In the course of identifying the telini sensu Knight and Webb in part (Balclutha species described in this work, we found that some of Kirkaldy) and the subfamilies Arrugadinae, Eupelici- the characters provided in the key to genera by Lin- nae (Eupelicini, Paradorydiini, Dorycephalini, and navuori and DeLong (1977) do not easily distinguish Listrophorini), and Drakensbergeninae, based largely the genera. A simpliÞed key is provided that reßects on the following features of the female genitalia (with previous generic concepts, adds some newly discov- some reversals, as indicated): 1) Þrst valvulae with a ered characters, and discards some ambiguous char- deÞnitely delimited apicoventral sculptured area, sub- acters. All of these genera are apparently closely re- triangular in shape (Figs. 12 and 30; reversed in Eu- lated, and as more species are identiÞed, some may be pelicini and Drakensbergeninae); 2) Þrst valvulae dor- sal sculpturing pattern maculose or scale-like, with the found to be paraphyletic. maculae or scales not overlapping and clearly sepa- rated from one another (Fig. 11) (reversed in Eupeli- cini); 3) Þrst valvulae dorsal sculpturing submarginal (Figs. 11, 12, and 30) (also present in Deltocephalinae: 1 Department of Entomology, 320 Morrill Hall, University of Chiasmusini and Doraturini); 4) second valvulae lack- Illinois at Urbana/Champaign, Urbana, IL 61801 (e-mail address: ing large dorsal teeth (Figs. 10, 28, and 33) (reversed [email protected]). in Eupelicini, Drakensbergeninae, and Dorycephalini 2 Department of Entomology, The Natural History Museum, Crom- well Road, London, SW7 5BD, United Kingdom (e-mail address: [ Kouchakewitch]); 5) pygofer macro- [email protected]). setae absent or reduced in number.

0013-8746/04/0667Ð0674$04.00/0 ᭧ 2004 Entomological Society of America 668 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 4

Figs. 1–5. K. junina n. sp. (1) Male, dorsal habitus. (2) Female, same. (3) Female, lateral habitus. (4) Nymph, dorsal habitus. (5) Male, face. All scale bars ϭ 1.0 mm.

All the Faltala group taxa have the characters listed lini and, in combination with an elongate connective, above and, together with some hecalines, have the the following were also removed: Tenucephalus De- head produced anteriorly; the anterior margin of head Long, Onura Oman, Acrolithus Freytag and Ma, and foliacious, carinate, or transversely striate; the tergum Egenus Oman. All of these taxa were transferred by with several longitudinal stripes; and strong sexual Hamilton to sensu Hamilton (1975). Of dimorphism in head length and brachyptery (see be- these genera, Hecullus and Tenucephalus have been low). However, although the inclusion of the Faltala examined in the current study (Hecullus sp.: Figs. 33 group in Hecalini would Þt in with this wider concept and 34) and found to have the same characters of the of the group (see also Linnavuori 1959, Freytag and female genitalia noted above for the clade found by Ma 1988, Nielson 1996), it would not conform to the Zahniser and Dietrich (unpublished data). most recent and more restrictive deÞnition of Ham- In addition to the morphological characters cited ilton (2000). This deÞnition was based on the ocelli above, the limited ecological data for the Faltala group being closer to the eyes than to the facial sutures. also support placement in the grass-feeding clade de- Although this feature is consistently observed in Heca- scribed above. All records of hosts or habitat for spe- lini sensu Hamilton, it also occurs in other Deltoceph- cies of the Faltala group indicate that they feed on alinae, e.g., Balclutha Kirkaldy, Cerrillus Oman, Drak- grasses. DeLong and Thambimuttu (1973) reported ensbergena Linnavuori, and Ball. that they “occur upon grasses and are found in mead- Contrary to Linnavuori (1975b), Hamilton (2000) also ows where sheep or cattle are grazing.” Of the species considered Hecalini to be closely related to Eupelici- described in this study, the two from Peru were swept nae (Dorycephalini) based on the following synapo- or vacuumed from grasses, and the Brazilian species morphies: male apodemes of second sternite as long as was collected in yellow pan traps in the Serra da Mesa tergal apodemes, yet narrow; “anal tube” withdrawn region, characterized by cerrado/cerradinho vegeta- into the pygofers; male pygofers with broad ventral tion, which is typiÞed by grasses, herbs, bushes, and lobes that meet or cross at lower edges; ovipositor small trees. curiously shaped, being straight or concave ventrally In conclusion, based on the above evidence, the and hump-backed near midlength (Readio 1922, Plate Faltala group taxa, together with Hecullus and Tenu- 29, Figs. 4Ð8). cephalus, are at present a nondeÞned assemblage that Based on the absence of the above ocellicharacter, could be placed as a subgroup in Hecalini sensu lato, Hamilton (2000) removed Hecullus Oman from Heca- having many of the characters noted above for this July 2004 ZAHNISER AND WEBB:THE Faltala OMAN LEAFHOPPER GROUP 669

Figs. 6–13. K. junina n. sp. (6) Connective and styli, ventral view. (7) Valve and plate, ventral view. (8) Aedeagus, lateral view. (9) Female seventh sternite. (10) Second valvulae. (11) Detail of Þrst valvula sculpturing pattern. (12) Apex of Þrst valvula. (13) First valvula, lateral view. group, but lacking the distinct hump-backed ovipos- considered adult characters in hemimetabolous , itor, ocelli closer to the eyes than the facial sutures, and that their absence in the adult is a kind of neoteny. and other characters found in Hecalini sensu stricto This might be determined genetically or in combination (Hamilton 2000). Alternatively, these taxa could form with environmental factors (e.g., temperature) affecting a separate lineage within the grass-feeding clade noted hormonal levels in the . Members of the Faltala above. However, for the present, these taxa are re- group corroborate the correlation of brachyptery with tained in Athysanini pending further studies. reduced or absent ocelli, because its members charac- teristically have reduced ocelli. Brachyptery Key to the Faltala Group Genera With respect to brachyptery in the grass-specializ- ing clade described above, subbrachypterous or sub- 1. Ocellinear to eyes, separated by Ϸ3ϫ their own macropterous species are prevalent in some groups diameter from eyes ...... Faltala (e.g., Hecalini), but fully brachypterus species (e.g., Ocellidistantfrom eyes, 1⁄3 to 1⁄2 distance from some Dorycephalini) are less common. However, in eye to crown apex ...... 2 the Faltala group (all genera) and in Acrolithus and 2. Frontoclypeus glabrous or with indistinct mi- Hecullus (females only), fully brachypterous species crosculpturing ...... Clorindaia have been reported. An interesting character related Frontoclypeus distinctly shagreened .....3 to brachyptery is the reduction or absence of ocelli 3. Male pygofer with several macrosetae; base of (Kalmus 1945). Evans (1968) reported this phenom- aedeagus broad in lateral view and subrectan- enon in ulopine and cephaleline leafhoppers and dis- gular in shape, sometimes incised anteriorly cussed the signiÞcance of high altitude and low tem- (K. mella)...... Kramerana peratures in the production of brachypters in Male pygofer without macrosetae. Base of polymorphic species. Southwood (1961) suggested that aedeagus not broad in lateral view, sickle- the characters of fully developed wings and ocelli can be shaped or falcate ...... 4 670 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 4

Figs. 14–18. K. adusta n. sp. (14) Male, dorsal habitus. (15) Male, lateral habitus. (16) Connective and style, ventral view. (17) Male, face. (18) Aedeagus, lateral view.

4. Crown shorter than wide or subequal in length sinuate. Brachypterous; venation reticulated or some- and width; discal region indistinctly microscu- times obscure. Color stramineous to dark brown; gen- lptured. Male subgenital plates with a projec- eral color pattern varying in intensity between and tion or knob in center of plate on dorsal within species. Dorsal side of abdomen with six ivory side...... Virganana longitudinal stripes outlined with dark brown; the Crown much longer than wide; discal region median pair sometimes conßuent, especially caudad shagreened and rugose anteriorly. Male sub- on each abdominal segment; mottled with fuscous and genital plates without dorsal projection .... dark brown between ivory stripes; ivory on lateral ...... Aequicephalus margin of abdomen; stripes less apparent and general The specimens examined in this study are housed in coloration lighter in some species and/or specimens. the following institutions: Museo de Historia Natural, Chaetotaxy normal for deltocephalines (see Rakitov Universidad Nacional Mayor de San Marcos, Lima, 1998, Dietrich and Rakitov 2002); protrochanter with Peru (MHNSM); Museu de Zoologia, University of stout apical seta in anteroventral (AV) position; pro- Sao Paulo, Brazil (MZSP); Illinois Natural History femur row AV with 7Ð12 setae, longer than most del- Survey, Champaign, IL (INHS); Ohio State University tocephalines; profemur intercallary row with 5Ð8 long Insect Collection, Columbus, OH (OSU); The Natural Þne setae; protibia formula 4 ϩ 4; hind femur knee History Museum, London, Unite Kingdom (BMNH); formula 2 ϩ 2 ϩ 1 (except K. adusta n. sp., 2 ϩ 2 ϩ 0). Museu Nacional do Rio de Janeiro, Universidade Fed- Male pygofer quadrate, with several macrosetae eral do Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ). (Ϸ5Ð9) near caudal margin; caudal margin with nu- merous very short Þne setae. Valve large, triangular. Plates triangular, apices rounded, usually several mac- Additional Characters for Kramerana DeLong and rosetae medially and/or laterally on basal half. Base of Thambimuttu aedeagus broad, subrectangular in shape; shaft much In addition to the characters cited by DeLong and narrower (except in K. linnavuorii); apex biÞd. Con- Thambimuttu (1973), the following characters occur nective gracile. Styles broadly bilobed basally, preapi- in Kramerana. Crown gradually angled to anterior cal lobe small, distinct; apophysis Þnger-like. margin (Fig. 3) or upturned near apex (K. adusta, n. Apodemes of aedeagus well developed; bulbous near sp., Fig. 15). Clypellus parallel sided or widening phragma, extending caudad and turning dorsad; at- slightly apically; apical margin slightly notched or tenuate at apex. July 2004 ZAHNISER AND WEBB:THE Faltala OMAN LEAFHOPPER GROUP 671

Figs. 19–27. C. brasileira n. sp. (19) Male, dorsal habitus. (20) Male, lateral habitus. (21) Male, face. (22) Nymph, dorsal habitus. (23) Aedeagus and right apodeme, anterior view. (24) Aedeagus, lateral view. (25) Male pygofer, lateral view. (26) Connective and styli, ventral view. (27) Valve and plates, ventral view.

Kramerana junina Zahniser, n. sp. nal pit, but becoming obscure near lateral margin; dark (Figs. 1Ð13) brown band sometimes underlined with transverse Diagnosis. This species is similar to K. hypera fuscous band (sometimes obscure), underlined by Blocker and Fang, but can be distinguished by its thin transverse dark brown line; gena ivory on ventral smaller size and by the relative length of the connec- half. Lorum mottled with dark brown and fuscous. tive, which is as long as the style in the new species, Clypellus dark brown with ivory or fuscous laterally but Ϸthree-fourths/four-Þfths the length of the style and dorsally. Crown (Fig. 1) with median thick, but in K. hypera. Also, the apophysis of style is elongate in ill-deÞned dark brown or black longitudinal band, K. hypera, but not in K. junina; the aedeagus of the new with pair of small ivory spots at caudal margin. Re- species does not have serrations on the caudoventral mainder of vertex mottled with fuscous or dark brown, margin and is completely ßat along the ventral margin. except for pair of longitudinal ivory stripes outlined Male. Length 3.0Ð3.3 mm; vertex length 0.53Ð0.54 with dark brown running from caudal corners of mm; width of pronotum 0.92Ð0.98 mm. crown to near anterior margin. Pronotum and scute- Color pattern: Frontoclypeus (Fig. 5) dark brown llum mottled with dark brown and fuscous, and with or black with thin fuscous transverse bands; transverse ivory stripes loosely following from ivory spots and ivory band at anterior margin between eyes. Gena stripes on crown. Forewing reticulated, with various with dark brown band below eye beginning in anten- dark brown, fuscous, and ivory markings vaguely co- 672 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 4

Figs. 28–32. C. brasileira n. sp. (28) Second valvulae. (29) Female, seventh sternite. (30) Apex of Þrst valvula. (31) First valvula, lateral view. (32) Third valvula, lateral view. inciding with stripes of abdomen. Abdominal stripes Þne setae, as in male. Anal tube mostly retracted into and coloration distinct (Figs. 1 and 2). Venter mostly pygofer; segment X sclerotized only as a pair of ven- dark brown. Legs dark brown with various fuscous and trolateral bands and a distal sclerotized ring. First ivory markings. Genital segment with varying degrees valvulae as in Fig. 13; dorsal sculpturing submarginal of pigmentation, usually with median dark brown spot and maculose, the maculae well separated (Fig. 11); or stripe on dorsal half with broad transverse stripe; with a well-delimited apicoventral sculptured area, valve and plates fuscous, sometimes with dark brown. one-fourth/one-Þfth the length of the valvulae Chaetotaxy with hind femur knee formula: 2 ϩ 2 ϩ 1. (Fig. 12); Þrst valvifer subquadrate, but anterior mar- Male pygofer with strongly sclerotized hook at cau- gin arched (Fig. 13). Second valvulae (Fig. 10) slightly doventral margin, directed ventrally. Valve (Fig. 7) humpbacked medially; ventral margin relatively length about half width. Connective (Fig. 6) long; straight; without teeth dorsally. Third valvulae with anterior arms with small lateral lobes. Style (Fig. 6) as Ϸ15 macrosetae near ventral and caudal margins; long as connective. Aedeagus (Fig. 8) recurved; ven- similar in shape to that of Clorindaia brasileira n. sp. tral margin ßat. Anal tube retracted into pygofer; seg- (Fig. 32). ment X membranous dorsally, with two longitudinal Nymph. Similar in color pattern (Fig. 4) to adult, sclerotized bands ventrally. but often pale; median ivory stripes completely con- Female. Length 3.7Ð4.1 mm; vertex length 0.63Ð0.67 ßuent, extending onto crown and partly taking place mm; pronotum width 1.11Ð1.21 mm. of dark brown median stripe seen in adult. Abdomen Color pattern (Fig. 2) same as the male, but fre- with only four macrosetae on VIII tergite: two dorso- quently lighter, making some of the more distinctive laterally and one each at caudolateral margins. Pygofer markings (e.g., the longitudinal ivory bands on the with several macrosetae. Otherwise, normal for heca- vertex) less apparent. lines (Dmitriev 2002). Bases of the Þrst valvulae retracted into abdomen, Type Material. HOLOTYPE, (: PERU, Junõ´n, 42 km reaching the middle of the sixth abdominal segment. NE La Oroya, 4,000 m, 11Њ24Ј18ЉS75Њ50Ј31ЉW, Seventh sternite as in Fig. 9. Pygofer with Ϸ10Ð15 17-X-2002, R.A. Rakitov, vacuum, 02-15-5. PARA- macrosetae irregularly arranged along the ventral and TYPES: 36 ((,17&&, and 26 nymphs of various caudal margins. Caudal margin also with very small stages, same collection data; 10 ((,4&& same data,

Figs. 33 and 34. Hecullus sp. (33) Second valvulae, lateral view. (34) First valvula, lateral view. July 2004 ZAHNISER AND WEBB:THE Faltala OMAN LEAFHOPPER GROUP 673 except the collecting code, 02-15-3; 2 ((,2&&, same 23-X-2002, C. H. Dietrich, vacuum, 02-35-1. PARA- locality data, C. Dietrich and R. Rakitov, sweeping, TYPE: 1(, same data. These were taken in a grazed 02-15-2; “sweep and vacuum grasses and shrubs.” pasture containing bunchgrass. Holotype deposited at Holotype and 10 ((,5&&, and 5 nymph paratypes MHNSM; paratype deposited at INHS. deposited at MHNSM; 5((,3&&, and 3 nymph Etymology. The speciÞc name “adusta” is the fem- paratypes deposited at OSU; 25 ((,9&&, and 12 inine of the Latin adjective adustus, meaning tanned nymph paratypes deposited at INHS; 3((,3&&, and or brown. This refers to the golden brown or tanned 3 nymph paratypes deposited at BMNH; 5 ((,3&&, color of the crown and pronotum. and 3 nymph paratypes deposited at MNRJ. Additional Material Examined. Numerous other Clorindaia brasileira Zahniser, n. sp. nymphs were taken at the type locality; numerous ((, (Figs. 19Ð32) &&, and nymphs taken at PERU, Junõ´n, 3 km W Curi- pata, 3800 m, 11Њ36Ј36ЉS75Њ57Ј25ЉW, 17-X-2002; and at Diagnosis. Distinguished from other Clorindaia spp. PERU, Junõ´n, 20 km NE La Oroya, 3600 m, 11Њ30Ј9ЉS by the process of the pygofer (Fig. 25) and the color 75Њ56Ј28Љ, 17-X-2002. pattern of the crown (Fig. 19). Etymology. This species is named for the Peruvian Male. Length 4.8Ð5.4 mm; vertex 0.67Ð0.73 mm; department in which it was collected, Junõ´n. pronotum width 1.47Ð1.57 mm. Color pattern: Frontoclypeus (Fig. 21) variable in degree of pigmentation; alternating dark brown or Kramerana adusta Zahniser, n. sp. black and fuscous bands running transversely. Ante- (Figs. 14Ð18) rior margin of head with a transverse ivory band run- Diagnosis. Distinguished from other Kramerana ning from lateral frontal suture to near eye. Dark spp. by the strongly upturned anterior margin of the brown or black band beginning in antennal pit running head (Fig. 13), its generally dark color pattern (Figs. transversely beneath eye to margin of gena, and ap- 14 and 16), the hind femur knee formula 2 ϩ 2 ϩ 0, and parently extending onto venter and widening poste- the form of the aedeagus (Fig. 18). riorly (Fig. 20). Remainder of gena, lorum, and cly- Male. Length 2.9 mm; vertex 0.47Ð0.5 mm; prono- pellus mostly ivory to fuscous, dark brown or black tum width 0.93Ð0.97 mm. at sutures and especially at apex of clypellus. Median Color pattern: Frontoclypeus (Fig. 17) dark brown markings of vertex outlining ivory area taking form or black with transverse fuscous or ivory stripes. An- of a human standing, legs spread, and arms reaching terior margin of the head with pair of ivory bands outward and upward; body of human mostly ivory bordered by dark brown running from lateral frontal or ochraceous; outlined with dark brown. Rest of suture to eye. Gena dark brown below eye, extending vertex ochraceous, an irregular longitudinal ivory to margin; fuscous or ivory laterad of lorum. Lorum band outlined with dark brown running next to each entirely fuscous in type specimen, but fuscous at lat- eye. Pronotum with six irregular longitudinal ivory eral margin and dark brown at median margin in para- bands outlined with dark brown; margins ivory. type. Clypellus variable in pigmentation. Crown (Fig. Scutellum roughly following pattern of pronotum. 14) golden brown, with dark brown longitudinally Wings reticulated; the veins ivory, with dark brown or along midline. Pronotum mostly light golden brown, fuscous between veins. Color pattern of abdomen as with scattered fuscous or dark brown; dark brown in K. junina n. sp., but ochraceous between dark brown especially concentrated near caudal margin. Scutel- bordered ivory bands. Pygofer ochraceous with pair of lum with dark brown medially, light brown laterad of brown spots caudodorsally (Fig. 19). dark brown, and ivory at lateral corners. Forewing Chaetotaxy of normal deltocephaline type. Protro- venation reticulated and somewhat obscure, with scat- chanter with stout apical seta in AV position. Profemur tered dark brown, fuscous, and ivory; thick stripe of row AV with Ϸ10Ð12 setae; intercallary row with 4Ð7 ivory running along costal margin. Abdomen with lon- long Þne setae; protibia formula 4 ϩ 4; hind tibia knee gitudinal stripes slightly more fuscous than ivory. formula 2 ϩ 2 ϩ 1. Chaetotaxy with hind femur knee formula: 2 ϩ 2 ϩ 0. Pygofer (Fig. 25) quadrate; Ϸ4 macrosetae near Pygofer with 3Ð4 short macrosetae; caudoventral dorsocaudal margin; well-developed hook at ventro- margin rounded without a hook or process. Plates with caudal margin. Valve (Fig. 27) triangular; less than half few small macrosetae near lateral margin; with nu- as long as wide. Plates (Fig. 27) triangular with Ϸ6Ð7 merous short Þne hairs near lateral margin, and most macrosetae scattered near middle; some very small developed (longer) at apex of the plate. Anterior arms Þne setae near lateral margins, but nearly absent at of connective (Fig. 16) relatively long and divergent. apex. Connective (Fig. 26) a little shorter than style; Aedeagus (Fig. 18) strongly recurved; shaft incised on anterior arms with lateral lobes; stem wide near base caudal surface along apical one-third, margins serrate; of arms, constricted medially, and widening distally. incised only apically on anterior surface. Basal and Style (Fig. 26) broad basally; preapical angle small; apical parts apodemes forming separate processes, apophysis Þnger-like with a rippled texture. Aedeagus connected by membranous tissue. Segment X not scle- (Fig. 24) very broad basally, especially dorsoventrally; rotized dorsally; sclerotized weakly lateroventrally. shaft about as long as base, slightly recurved, apex Type Material. HOLOTYPE, (: PERU, Pasco, 8 km divided with small lateral triangular processes (Fig. E Carhuamayo, 4,000 m, 10Њ53Ј7ЉS75Њ56Ј44ЉW, 23). Apodemes (Fig. 23) and anal tube similar to those 674 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 97, no. 4 of Kramerana. Segment X sclerotized as a transverse Dietrich, C. H., and R. A. Rakitov. 2002. Some remarkable band dorsally and as a pair of longitudinal bands ven- new deltocephaline leafhoppers (Hemiptera: Cicadelli- trolaterally. dae: Deltocephalinae) from the Amazonian rainforest Female. Length 6.0Ð6.1 mm; vertex length 0.78Ð canopy. JNY Entomol. Soc. 110: 1Ð48. 0.94 mm; pronotum width 1.69Ð1.75 mm. Dmitriev, D. A. 2002. General morphology of leafhopper nymphs of the subfamily Deltocephalinae (Hemiptera: Color pattern same as male, but frequently paler. Ϸ Cicadellidae). Acta Entomol. Slovenica 10: 65Ð82. Pygofer with 10 macrosetae ventrally and with 4 Evans, J. W. 1968. Some relict New Guinea leafhoppers and macrosetae near the caudodorsal margin. Seventh their signiÞcance in relation to the comparative morphol- sternite as in Fig. 29. First, second, and third valvulae ogy of the head and prothorax of the Homoptera- (ϭgonoplac) as in Figs. 31, 28, and 32, respectively. Auchenorrhycha (Homoptera: Cicadellidae: Ulopinae). First valvulae apicomedial sculptured area (Fig. 30) Pac. Insects 10: 215Ð229. thinner dorsoventrally than in K. junina n. sp. First Freytag, P. H., and N. Ma. 1988. An unusual new genus valvifer (Fig. 31) anterior margin straight. and species of hecaline leafhopper from Venezuela Nymph. Color pattern as in Fig. 18. With only two (Homoptera: Cicadellidae). Entomol. News 99: 153Ð156. Hamilton, K.G.A. 1975. Review of the tribal classiÞcation of macrosetae on abdominal tergum, positioned at cau- the leafhopper subfamily Aphrodinae (Deltocephalinae dolateral corners of tergite VIII. Several macrosetae of authors) of the Holarctic region (Rhynchota: Ho- on pygofer. moptera: Cicadellidae). Can. Entomol. 107: 477Ð498. Type Material. HOLOTYPE, (:14Њ17.2ЈS48Њ55.5ЈW, Hamilton, K.G.A. 2000. Five genera of New-World “Shovel- BRASIL, Goia´s: Uruac¸u, Serra da Mesa Survey, 28-V- Headed” and “Spoon-Bill” leafhoppers (Hemiptera: Ci- 1996, yellow pans, cerrado-cerradinho, Johnson, cadellidae: Dorycephalini and Hecalini). Can. Entomol. Sharkov, Ejchel. PARATYPES: 3 &&, 3 nymphs, same 132: 429Ð503. locality, 24-V-1996; 1 (,1&, same locality, 25-V-1996; 2 Kalmus, H. 1945. Correlations between ßight and vision, ((, same locality, 26-V-1996; 2 ((, 2 nymphs, same and particularly between wings and ocelli, in insects. (( & Proc. R. Entomol. Soc. Lond. A: Gen. Entomol. (http:// locality, 27-V-1996; 2 ,1 , same locality, 28-V-1996; csssrrr.entnem.uß.edu/ϳpmc/journals/all-journals.htm). 1 (,1&, same locality, 29-V-1996. Holotype, and 4 ((, && (( Knight, W. J., and M. D. Webb. 1993. The phylogenetic 3 , and 2 nymph paratypes deposited at MZSP; 2 , relationships between virus vector and other genera of 1 &, and 1 nymph paratypes deposited at INHS; 1(,1&, macrosteline leafhoppers, including descriptions of new and 1 nymph paratype deposited at OSU; 1 (,1&, and taxa (Homoptera: Cicadellidae: Deltocephalinae). Syst. 1 nymph paratype deposited at BMNH. Entomol. 18: 11Ð55. Etymology. The species name highlights the Þrst Linnavuori, R. 1959. Revision of the Neotropical Delto- Þnding of a species of the Faltala group from Brazil. cephalinae and some related subfamilies (Homoptera). Ann. Zool. Soc. Zool. Bot. Fennicae ÔVanamoÕ 20: 1Ð370. Linnavuori, R. 1975a. Studies on Neotropical Deltocephali- nae (Homoptera: Cicadellidae). Notulae Entomol. 45: Acknowledgments 49Ð52. We thank Norman Johnson, Luciana Musetti, and Peter Linnavuori, R. 1975b. Revision of the Cicadellidae (Ho- Kovarik of OSU for the loan of specimens and for the op- moptera) of the Ethiopian region. III. Deltocephalinae, portunity to examine material from their survey of the Serra Hecalini. Acta Zool. Fennica 143: 1Ð37. da Mesa of Brazil. Thanks also to Zachary Falin of the Snow Linnavuori, R., and D. M. DeLong. 1977. The leafhoppers Entomology Collection at the Kansas University Museum of (Homoptera: Cicadellidae) known from Chile. Brenesia Natural History for the loan of material relevant to this study. 12-. 13: 163Ð267. A preliminary version of this manuscript beneÞted greatly Nielson, M. W. 1996. A new genus, Hecalocorica, and a new from the comments of C. H. Dietrich and Dmitri Dmitriev, species of hecaline leafhopper from Costa Rica (Ho- both of the INHS. An NSF PEET (Program for Enhancing moptera: Cicadellidae). Entomol. News 107(3): 125Ð127. Expertise in ) grant (DEB 9978026) and a National Oman, P. W. 1938. A generic revision of American Bytho- Science Foundation grant (DEB 0089671) to C. H. Dietrich scopinae and South American Jassinae. Univ. Kansas Sci. are gratefully acknowledged for supporting this research. Bull. 37: 343Ð420. Rakitov, R. A. 1998. On differentiation of cicadellid leg chaetotaxy (Homoptera, , Membra- References Cited coidea). Russian Entomol. J. 6: 7Ð27. Readio, P. A. 1922. Ovipositors of Cicadellidae (Ho- Blocker, H. D., and Q. Fang. 1992. A synopsis of the Faltala moptera). Kansas Univ. Sci. Bull. 14: 215Ð298. group (Homoptera: Cicadellidae) with descriptions of Southwood, T.R.E. 1961. A hormonal theory of the mechanism new taxa. J. Kansas Entomol. Soc. 65: 341Ð346. of wing polymorphism in Heteroptera. Proc. R. Entomol. DeLong, D. M., and C. C. Thambimuttu. 1973. Three Soc. Lond. A: Gen. Entomol. (http://csssrrr.entnem.u- closely related new genera and Þve new species of short- ß.edu/ϳpmc/journals/all-journals.htm). winged Chilean leafhoppers (Homoptera: Cicadellidae). Fla. Entomol. 56: 165Ð172. Received 8 October 2003; accepted 24 March 2004.