Some Unusual Neotropical Neocoelidiinae with a Redefinition of the Subfamily (Hemiptera: Membracoidea: Cicadellidae)

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1 SYSTEMATICS Some Unusual Neotropical Neocoelidiinae with a Redefinition of the Subfamily (Hemiptera: Membracoidea: Cicadellidae) C. H. DIETRICH Center for Biodiversity, Illinois Natural History Survey, 607 E. Peabody Drive, Champaign, IL Ann. Entomol. Soc. Am. 96(6): 700Ð715 (2003) ABSTRACT Diagnostic characters for the New World leafhopper subfamily Neocoelidiinae are reviewed. Krocodona Kramer (with new synonym Krocobella Kramer), Krocozzota Kramer, and the fossil genus Krocarites Dietrich and Vega, previously placed in Nirvaninae based on their depressed, strongly produced heads, are transferred to Neocoelidiinae (new placements) based on characters of the head, wing venation, leg chaetotaxy, and male genitalia. The related neocoelidiine leafhopper genera Retrolidia, new genus, based on three new species, and Krocolidia, new genus, based on two new species, all from South America; four new South American species of Krocodona; and a new species of Krocozzota (including the Þrst known male of the genus) are described and illustrated. Keys to Krocodona and Retrolidia species and a checklist of neocoelidiine genera are provided. KEY WORDS leafhopper, classiþcation, nomenclature, morphology, identiþcation THE LEAFHOPPER FAMILY Cicadellidae comprises 36 currently recognized subfamilies (Oman et al. 1990, Godoy and Webb 1994, Dietrich and Rakitov 2002, Dietrich and Dmitriev 2003). Many of these subfamilies are poorly characterized morphologically, and phylogenetic analyses of morphological (Hamilton 1983, Dietrich 1999) and molecular data (Dietrich et al. 2001) indicate that some, as traditionally deþned (Oman et al. 1990), are para- or polyphyletic. One such group is Neocoelidiinae. This subfamily comprises 18 genera and 130 described species known only from the New World. The vast majority of species are restricted to the tropics, but the ranges of a few extend into temperate North America. In the tropics, neocoelidiines have most often been collected at lights or by fogging rain forest canopies, and their host associations remain undocumented. Nearctic species have been recorded from various shrubs and trees, especially Pinus, Arctostaphylos, and Artemisia, but a few apparently feed on herbs (DeLong 1953). Linnavuori (1959) suggested that the group is related to Deltocephalinae, based on the bifurcate anterior tentorium branches. Recent molecular phylogenetic analyses (Dietrich et al. 2001) provided weak support for this relationship, but a preliminary morphology-based cladistic analysis (Dietrich 1999) suggested that the group belongs to a lineage comprising Coelidiinae, Nirvaninae, and Typhlocybinae. Kramer (1964a, 1967) revised the group and provided a key to genera, characterizing them mainly based on differences in the male genitalia, some of which appear to be arbitrary (e.g., symmetrical versus asymmetrical aedeagus). He deþned Neocoelidiinae based on the presence of a distinct antennal ledge, the exceptionally long antennae, and the obscure forewing venation. Evans (1947) had previously noted that Neocoelidiinae usually have forewing crossvein r-m 1 absent. Other possibly synapomorphic characters of Neocoelidiinae include the presence of a gibbosity or tubercle on the clypellus in many species, the fusion of hindwing veins R 4 5 and M 1 2, the macrosetal formula of the hind femur, the fusion of the male valve (sternum IX) to the subgenital plate, and presence of a ventral preapical spine or process on the male pygofer. Godoy and Webb (1994) noted that some of these characters are also present in certain genera placed by Kramer (1964b) in Nirvaninae. The molecular phylogeny of Dietrich et al. (2001)) grouped the four included representatives of Neocoelidiinae (Biza sp., Chinaia sp., and two Neocoelidia spp.) together with high (97Ð100%) bootstrap support, but did not include any New World Nirvaninae. Thus, the phylogenetic status of Neocoelidiinae has not yet been adequately tested. Recent sampling, particularly by fogging Amazonian rain forest canopies in Brazil, Ecuador, and Peru (Erwin 1983, 1989, Lucky et al. 2002), has yielded numerous new neocoelidiine taxa, as well as the previously unknown males or females of some genera. Many of these species are not easily referable to previously described genera. This undoubtedly reßects the extremely limited material upon which Kramer (1964a, 1967) based his revisionary studies of the subfamily. Comparative morphological study of the Neocoelidiinae and the New World Nirvaninae, based largely on this new material, indicates that the classi- Þcations of both subfamilies need to be revised. In this paper, I review the morphological characters deþning Neocoelidiinae, transfer some genera previously placed in Nirvaninae to Neocoelidiinae, and describe /03/0700Ð0715$04.00/ Entomological Society of America

2 November 2003 DIETRICH: UNUSUAL NEOCOELIDIINAE 701 Table 1. Checklist and distribution of the genera of Neocoelidiinae Biza Walker 1858b:253 (type species: B. crocea Walker 1858b). Brazil, Costa Rica, Ecuador, Panama, Peru. Chinaia Bruner and Metcalf 1934a:120 (type species: C. bella Bruner and Metcalf). Brazil, Colombia, Costa Rica, Ecuador, Guatemala, Guyana, Mexico, Panama, Peru. Chinchota Kramer 1967b:41 (type species: C. styx Kramer). Colombia. Cocoelidia DeLong 1953a:126 (type species: Neocoelidiana antlera DeLong). Mexico Coelana DeLong 1953a:128 (type species: Neocoelidia modesta Baker). Argentina, Bolivia, Brazil. Coelella DeLong 1953a:125 (type species: Neocoelidia distincta Oman 1931a:67). Mexico, U.S.A. Coelidiana Oman 1938b:397 (type species: Neocoelidia rubrolineata Baker 1898h). Brazil, Costa Rica, Guatemala, Guyana, Mexico, Panama, Peru. Coelindroma Kramer 1967b:43 (type species: C. fungosa Kramer). Peru. Deltocoelidia Kramer 1961a:238 (type species: D. maldonadoi Kramer). Venezuela. Krocarites Dietrich and Vega 1995:266 (type species: K. reflexa Dietrich and Vega), new placement. Hispaniola (Dominican amber fossil). Krocodona Kramer 1964b:114 (type species: K. sauridion Kramer), new placement. Brazil, Costa Rica, Ecuador, Honduras, Peru. Krocobella Kramer 1964b:118 (type species: K. colotes Kramer), new placement, new synonym. Krocolidia, new genus (type species: K.rufilinea, n. sp.). Brazil, Ecuador, Peru. Krocozzota Kramer 1964b:115 (type species: K. languria Kramer), new placement. Colombia, Panama. Megacoelidia Kramer and Linnavuori 1959a:55 (type species: M. splendida). Bolivia, Brazil. Nelidina DeLong 1953a:129 (type species: Coelidiana defila DeLong). Colombia, Peru. Neocoelidia Gillette and Baker 1895a:103 (type species: N. tumidifrons Gillette and Baker). Colombia, El Salvador, Guatemala, Honduras, Mexico, U.S.A. Neocoelidiana DeLong 1953a:125 (type species: Neocoelidia obscura Baker 1898h). Mexico, U.S.A. Retrolidia, new genus (type species: R. bimaculata, n. sp.). Ecuador, Peru. Salvina Melichar 1926a:344 (type species: Tettigonia dosisignata Fowler 1900d). Panama. Tichocoelidia Kramer 1962a:104 (type species: T. clarkei Kramer). Colombia. Tozzita Kramer 1964d:267 (type species: T. ips Kramer). Bolivia, Brazil. Xenocoelidia Kramer 1959a:30 (type species: X. youngi Kramer). Brazil, Colombia, Peru. Xiquilliba Kramer 1964d:268 (type species: X bellator Kramer). Brazil. Nomenclatural changes are indicated in bold. Note: Oman et al. (1990) erroneously listed Bolotheta Kramer 1963 as belonging to Neocoelidiinae; the genus was correctly placed by Kramer (1963) in Deltocephalinae. or redescribe some unusual genera and species. The classiþcation of Neotropical Nirvaninae will be revised in a separate paper (unpublished data). Materials and Methods Morphological techniques and terminology follow Oman (1949) and Kramer (1950), except that groups of macrosetae on the legs are named using the system of Rakitov (1998). Setal rows on the femur and tibia are referred to according to their position, assuming that the leg is fully extended perpendicular to the sagittal plane of the body, as follows: AD anterodorsal; AM anteromedial; AV anteroventral; PD posterodorsal; and PV posteroventral. Rakitov recognized two additional rows between AD and AV on the front femur: the intercalary and the more dorsal anteromedial (AM) row. Individual setae are numbered sequentially, beginning with the most distal. Numerical formulae are used to indicate the numbers of setae on the pro- and mesothoracic tibiae and the metathoracic femur. For tibiae, the formula given is AD PD; for the femur, the formula follows standard practice in which, for example, means that a distal pair, a penultimate pair, and a single antepenultimate seta are present. Body length is measured from the apex of the head to the apex of the forewing in repose. Digital photographs were taken using a Kodak DC-120 or Q-Imaging Micropublisher digital camera mounted on a stereo or compound microscope. Specimens examined are deposited in the following institutions: The Natural History Museum, London (BMNH), Escuela Politécnica Nacional, Quito, Ecuador (EPNQ, specimens currently held in trust at United States National Museum, WA [USNM]), Humboldt Institute, Villa Leyva, Colombia (HIC); Illinois Natural History Survey (INHS); Museum National dõhistoire Naturelle, Paris (MNHN); USNM. Specimens of some previously described species were not available for study; thus, it was necessary in some cases to refer to Þgures in earlier publications in the descriptions and keys below. Nomenclatural changes proposed herein are summarized in Table 1, which also lists all currently recognized genera of Neocoelidiinae and their type species. Results Subfamily Neocoelidiinae Oman 1943 Revised diagnosis. Neocoelidiinae differ from leafhoppers of other subfamilies in having the following combination of characters: ocelli on or near crown margin near eyes; antennae longer than half body length; forewing crossveins r-m 1 and m-cu 2 absent; hind wing with R 4 5 and M 1 2 conßuent (except Chinaia Bruner and Metcalf); hind femur macrosetal formula with penultimate pair well separated; male valve fused to plates, but articulated to pygofer; female second valvulae slender and elongate with toothed portion less than half total length. Revised description. Medium-sized to large, usually pale yellow, green, or orange leafhoppers, often with black markings dorsally. Crown usually well developed and depressed, margins usually raised and shelflike mesad of eyes, expanded laterad just anterad of eyes, anterior margin usually carinate; ocelli on or

3 702 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 6 Fig Neocoelidiinae. 1Ð4, Krocodona, dorsal habitus: 1, K. bipendula;2,k. circumflexa;3,k. janauaca;4,k. triloba. 5Ð6, Krocozzota magdalensis, dorsal and lateral habitus. 7Ð8, Retrolidia bimaculata, dorsal and lateral habitus. 9 and 10, R. nigricephala: 9, anterior dorsum; 10, head, ventral view. 11, Krocolidia obscura, anterior dorsum; 12 and 13, K. rufilinea, dorsal and lateral habitus. Scale bar, 1 mm. slightly above or below anterior margin of crown, closer to eyes than to midline; lateral frontal sutures obsolete, or present and parallel sided, or divergent dorsally; upper portion of frontoclypeus often with median longitudinal carina; antennal ledge well developed and oblique, or obsolete, ßagellum longer than half body length; clypellus parallel sided or distally expanded, even with or not attaining ventral margin of gena, often with medial gibbosity or tubercle; lorum small, not extended to genal margin; gena broadly rounded, only partially concealing procoxale ( proepisternum ), without erect seta ventrad of

4 November 2003 DIETRICH: UNUSUAL NEOCOELIDIINAE 703 Fig and 15, Retrolidia nigricephala, forewing and hind wing; 16, R. serrata, forewing; 17, Krocolidia sp. (Rio Tambopata, Peru). antennal pit; maxillary sensillum poorly delimited, closely adjacent to lorum near midlength; procoxale large, ßaplike; frontoclypeus convex, not extended onto crown; rostrum short, tapered, not surpassing prothoracic trochanters. Pronotum with maximum width exceeding that of head; anterior margin weakly produced; posterior margin broadly and angulately emarginate; lateral margins carinate (except Chinaia), divergent posteriorly, carina even with or below posterior corner of eye. Forewing crossveins r-m 1 and m-cu 2 absent (inner and middle anteapical cells open, Fig. 17); Cu connected to submarginal vein well distad of claval apex; anal veins free, without crossveins; appendix narrow, restricted to anal margin. Hind wing (Fig. 15) with R 4 5 and M 1 2 conßuent (except Chinaia), m-cu short. Front femur (Figs. 18Ð20) slender, AM 1 well developed, AV with 1Ð3 small preapical setae, intercalary row beginning at approximately midlength with widely spaced setae subequal or decreasing in size toward apex, PV absent; tibia slender, more or less cylindrical with variable numbers of dorsal macrosetae, row AV well developed, PV reduced; accessory row between AV and AD absent (except Chinaia). Middle trochanter with a few Þne ventral setae; femur broader than that of prothorax, macrosetal rows reduced or absent. Hind femur (Figs. 21Ð23) with setal formula usually and penultimate pair usually widely separated; tibial row PD with numerous close set macrosetae of approximately equal length, AD with 1Ð3 short setae between consecutive macrosetae; tarsomere I with dorsoapical pair of macrosetae and two longitudinal plantar rows, pecten with four or more platellae. Male pygofer usually with preapical ventral tooth or process (Fig. 24), without macrosetae; tenth segment long and well sclerotized, semicylindrical; valve not fused to pygofer, fused to subgenital plates; plates (Fig. 25) usually partially to completely fused medially, usually large and boatlike, with or without macrosetae; connective Y shaped, without median anterior lobe (except Chinaia and Salvina Melichar); style with apodeme much shorter than apophysis. Female pygofer with longitudinal row of macrosetae ventrolaterally; ovipositor extended beyond pygofer apex; dorsal sculpturing of Þrst valvula (Figs. 67Ð 70) strigate to reticulate, marginal or submarginal; second valvulae (Figs. 71Ð74) with weakly sclerotized area on dorsal margin just basad of toothed distal blades, with single distinct duct channel ventrally, dorsal teeth numerous, close set, quadrate and usually serrate; third valvulae with small setae ventroapically. Notes. As indicated in the above desciption, Chinaia is unusual in several respects, but its placement in Neocoelidiinae is supported by the partial or complete fusion of the valve and subgenital plates in the male, and by molecular evidence (Dietrich et al. 2001). The preceding redeþnition of Neocoelidiinae encompasses all of the genera placed in that group by Kramer (1964a, 1967) as well as the following genera, previously included in Nirvaninae: Krocobella Kramer, Krocodona Kramer, and Krocozzota Kramer (1964b); and Krocarites Dietrich and Vega (1995, described from Dominican amber). These genera have the head strongly produced with the antennal ledges reduced, and thus disagree with KramerÕs concept of Neocoelidiinae. Nevertheless, they exhibit several possibly synapomorphic traits characteristic of Neocoelidiinae that are absent or rare in other leafhopper subfamilies. These include: male valve (sternum VIII) fused to plates, but articulated to pygofer; pygofer with posteroventral tooth or spine; and hind wing with R 4 5 and M 1 2 conßuent. Other features present in these genera that support their placement in Neocoelidiinae (and exclusion from Nirvaninae) include: hind femoral macrosetal formula 2 2 1, with the penultimate pair widely separated; clypellus expanded distally and usually with medial gibbosity or tubercle; front femur without an enlarged subbasal intercalary seta and PV 1 absent. Each of these genera was described based on a single specimen. Additional specimens of the fossil

5 704 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 6 Fig Ð20, Front leg, anterior view: 18, Krocodona bipendula; 19, Retrolidia serrata; 20, Krocolidia sp., female from Rio Tambopata, Peru. 21Ð23, hind leg, anterior view: 21, K. bipendula; 22, Krocolidia sp. (Rio Tambopata, Peru); 23, R. serrata. AV anteroventral seta; IC intercalary row. genus Krocarites are unknown, but recent sampling in South America has yielded several additional specimens of KramerÕs genera. Based on this material, the characters given in KramerÕs (1964b) key do not consistently distinguish Krocobella from Krocodona. The holotype of Krocobella colotes Kramer was not available for study, but KramerÕs (1964b) description and illustrations indicate that the forewing venation is identical with that of Krocodona spp., and the male genitalia, particularly the shape of the style, seem

6 November 2003 DIETRICH: UNUSUAL NEOCOELIDIINAE 705 Fig Krocodona, male genitalia. 24Ð28, K. bipendula: 24 and 25, genital capsule, lateral and ventral views; 26 and 27, genitalia, lateral and ventral views; 28, aedeagal shaft apex, posterior view. 29Ð32, K circumflexa: 29, genital capsule, lateral view; 30 and 31, genitalia, lateral and ventral views; 32, aedeagus, dorsal view. 33Ð35, K. triloba: 33, genital capsule, lateral view; 34 and 35, genitalia, lateral and ventral views.

7 706 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 6 Fig Ð39, Krocodona janauaca: 36, genital capsule, lateral view; 37 and 38, genitalia, lateral and ventral views; 39, aedeagal apex, posterior view. 40Ð42, Krocozzota magdalensis: 40, genital capsule, lateral view; 41 and 42, genitalia, lateral and ventral views (aedeagus rotated 90 with respect to connective and styles). sufþciently similar to justify treating the two genera as synonyms. Revised diagnoses for Krocodona and the related genus Krocozzota are provided below. Two new neocoelidiine genera are also described in this work because, based on the previous concept of Neocoelidiinae (Kramer 1964a), they would have been excluded from this subfamily and placed in Nirvaninae. Krocodona Kramer 1964b: 114 Krocobella Kramer 1964b: 118, new synonym. Redescription. Small (5.8Ð6.5 mm), tan neocoelidiines mottled with brown, black, and red. Head (Figs. 1-4) strongly produced, 3 longer than basal width between eyes; in lateral view slightly declivous with apex obliquely truncate; crown only weakly elevated mesad of eyes, uniformly shagreen, margin carinate, in dorsal view gradually tapered anterad, apex angulate, arcuate in anterior view; coronal suture extended to crown apex, not carinate; ocelli well developed, on crown near lateral margin well anterad of eyes, in submarginal groove that extends nearly to crown apex; lateral frontal suture extended anterodorsad from antennal pit, not reaching crown margin; antennal ledge weak, not ßattened, ßagellum ca. half body length; frontoclypeus parallel-sided ventrad of antennal pits, with median keel dorsoapically; lorum slightly wider than clypellus; clypellus with apex as wide as base, margins parallel sided or slightly constricted medially, not extended to genal margin, with weak median gibbosity; genal margin broadly rounded. Pronotum in lateral view with dorsum nearly horizontal, continuing contour of crown; surface transversely rugose; lateral carina ventrad of posterior corner of eye. Exposed part of mesonotum and scutellum as wide as long. Forewing venation obscure, except

8 November 2003 DIETRICH: UNUSUAL NEOCOELIDIINAE 707 Fig Retrolidia, male genitalia. 43Ð46, R. bimaculata: 43 and 44, genital capsule, lateral and ventral views; 45 and 46, genitalia, lateral and ventral views. 47, R. nigricephala, genital capsule, lateral view; 48, R. serrata, genital capsule, lateral view; 49 and 50, R. nigricephala, genitalia, lateral and ventral views; 51 and 52, R. serrata, genitalia, lateral and ventral views.

9 708 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 6 Fig Krocolidia, male genitalia. 53Ð56, K. rufilinea: 53 and 54, genital capsule, lateral and ventral views; 55 and 56, genitalia, lateral and ventral views. 57Ð60, K. obscura: 57 and 58, genital capsule (genitalia not removed), lateral and ventral views; 59 and 60, genitalia, lateral and ventral views. near apex; R 1 and R 2 3 strongly reßexed; outer anteapical cell absent; inner apical cell wide, apex oblique; second apical cell widened distally; third apical cell small, petiolate. Hind wing with R 4 5 and M 1 2 conßuent. Front femur (Fig. 18) intercalary row with six setae, AV 1 well basad of apex, tibia dorsal setae 1

10 November 2003 DIETRICH: UNUSUAL NEOCOELIDIINAE 709 Fig Female seventh sternites: 61, Krocodona bipendula; 62, Krocozzota magdalensis; 63, Retrolidia bimaculata; 64, R. nigricephala; 65, R. serrata; 66, Krocolidia sp. (Rio Tambopata, Peru). 1; middle femur shorter and broader than that of prothorax, tibia with dorsal macrosetae 1 1, AV with few scattered setae, PV absent; hind femur (Fig. 21) with penultimate pair of macrosetae widely spaced, tibia AV arising one-thirds distance from base, tarsomere I pecten with Þve platellae. Male pygofer (Fig. 24) with a few small scattered setae posterodorsally, ventral margin with conical projection preapically, posterior lobe with or without processes; segment X large and well sclerotized; plates (Fig. 25) large, fused, except at apices, widest near midlength, without macrosetae; connective (Fig. 27) large, Y shaped; style apodeme (Fig. 26) foot shaped in lateral view, apophysis robust, with prominent setigerous preapical lobe extended nearly to apex of slender distal hook; aedeagus with (Fig. 26) or without (Fig. 30) large depressed posteriorly directed ventral process; gonopore-bearing shaft slender, recurved, with paired apical processes; gonopore apical. Female sternum VII (Fig. 61) produced, concealing base of ovipositor; Þrst valvulae (Fig. 67) slender, dorsal sculpturing submarginal, reticulate dorsally, becoming strigate more ventrad, ventral apical sculpturing reticulate; second valvulae (Fig. 71) slender, elongate, distal blades slightly expanded, dorsal teeth somewhat rounded, serrate; third valvulae with ca. seven scattered small ventroapical setae. Fifth instar nymph (based on Krocodona sp. prob. triloba, n. sp., Fig. 75) shiny brown, without distinct markings, apices of head and abdomen darker and sutures pale; legs pale yellow. Head structure similar to that of adult; coronal suture extended full length of crown; transverse suture near crown midlength extended ventrad onto face, curving posterad to antennal pit and continuing as clypeogenal suture; small pale impression anteromesad of eye; antennal ßagellum subequal to body length; lorum small, not extended to genal margin. Wing pad without apical setae. Chaetotaxy of legs similar to that of adult, except row PV of hind tibia with setae elongate, length 2 maximum width of tibia. Abdominal terga IV-VIII each with pale posterolateral seta; tergum VIII also with dark seta ventrad of pale seta. Tergum IX with basal section subequal in length to that of tergum VIII and pair of apically rounded tectiform distal lobes; basal section of tergum with two large pale setae on either side dorsolaterally, a dark seta ventrad of these, and a ventrolateral brush of dark setae more distad; distal lobes each bearing three evenly spaced preapical setae and a distal brush of ca. nine setae, all dark. Notes. The hind wing venation and morphology of the male genitalia; particularly the presence of a ventral tooth on the male pygofer and the subgenital plates, which are fused to the valve and to each other; leaves little doubt that the genus is properly placed in Neocoelidiinae. The genus was described based on a unique female specimen of K. sauridion Kramer from Honduras. Males of four additional species have been discovered in canopy fogging samples from Brazil and Ecuador in the USNM. The differences among these species in their male genitalia would cause them to key out to three different genera in KramerÕs (1964a) key to neocoelidiine genera. Krocodona bipendula, n. sp. and K. triloba, n. sp. (described below) would key out to Neocoelidia. Krocodona janauaca shares at least one key character with Cocoelidia DeLong (forked ventral hook on anal tube), and the remaining Krocodona species key to Nelidina DeLong. That such variation occurs within a single genus strongly indicates that a comprehensive reevaluation of the genus-level classiþcation of Neocoelidiinae is needed. In addition to the specimens listed under each species treatment below, three additional females from the USNM collection were examined with the following data: Costa Rica, Heredia, La Selva Field Station near Puerto Viejo, 21Ð28 March 1988; Brazil, Amazonas, Paraná do Xiboreninho, 7 August 1979; Peru, Madre de Dios, Rio Tambopata Res., 30 km (air) SW Puerto Maldonado, 10 November Based on differences in coloration, these specimens apparently represent additional undescribed species, but because males are lacking, it seems inadvisable to describe them here. Key to Species of Krocodona 1 Color pattern of crown consisting of a series of transverse bands, each broken to some extent medially (Figs. 1 and 4) Crown either unmarked or with color pattern consisting of oblique or angulate markings (Figs. 2 and 3) Transverse bands of crown broadly connected to each other laterally (Kramer 1964b: Fig. 2); range, Central America... sauridion Kramer 2 Transverse bands of crown mostly separate laterally, South America (Figs. 2 and 3)...3

11 710 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 6 Fig Ovipositors. 67Ð70, Þrst valvulae: 67, Krocodona bipendula; 68, Krocozzota magdalensis; 69, Retrolidia bimaculata; 70, Krocolidia sp. (Rio Tambopata, Peru). 71Ð74, second valvulae: 71, K. bipendula; 72, K. magdalensis; 73, R. bimaculata; 74, Krocolidia sp. (Rio Tambopata, Peru).

12 November 2003 DIETRICH: UNUSUAL NEOCOELIDIINAE 711 Fig Fifth instar nymphs, dorsal habitus: 75, Krocodona sp. prob. triloba; 76, R. bimaculata; 77, Krocolidia sp. (Onkonegare, Ecuador). 3 Ventral process of aedeagus expanded with pair of lateral preapical lobes, margin serrate (Fig. 35)... triloba, n. sp. 3 Ventral process of aedeagus evenly tapered, without lobes, margin entire (Fig. 27) bipendula, n. sp. 4 Male anal tube (segment X) with pair of large ventrolateral processes (Fig. 36) janauaca, n. sp. 4 Male anal tube without processes Aedeagal shaft in lateral view elongate, forming nearly complete circle (Fig. 30), apex with pair of lateral processes (Fig. 32) circumflexa, n. sp. 5 Aedeagal shaft in lateral view short, not forming circle, apex without processes (Kramer 1964b: Fig. 15)... colotes (Kramer) K. bipendula, n.sp. (Figs. 1, 24Ð28, 61, 67, 71) Description. Length male 5.8, female 6.5 mm. Color pattern of crown similar to that of K. sauridion, consisting of Þve irregular transverse brown bands preapically, each broken to some extent medially (Fig. 1). Male pygofer (Fig. 24) with small acute emargination posterodorsally; subgenital plate apex unmodiþed (Fig. 25); aedeagus (Fig. 26) with ventral process longer than shaft, evenly tapered in ventral view; apex of shaft strongly depressed with pair of short broad triangular processes (Fig. 28); female sternum VII with median trapezoidal posterior process (Fig. 61). Material examined. Holotype male, Ecuador, Orellana, Tiputini Biodiversity Station, near Yasuni National Park, 220Ð250 m, 5 February 1999, T. L. Erwin et al., lot 2096, transect T/10, fogging terra Þrme forest (EPNQ). One male paratype, Ecuador, Orellana, Transect Ent. 1 km S Onkonegare Camp, Reserva Etnica Waorani, S, W, 10 October 1994, T. L. Erwin et al., fogging terra Þrme forest, lot 931 (USNM). A female specimen from Rio Tambopata Reserve, Madre de Dios, Peru (USNM), is tentatively assigned to this species based on the similar color pattern of the crown and forewing. K. triloba, n. sp. (Figs. 4 and 33Ð35) Description. Length male 5.8 mm. Coloration (Fig. 4) and male genitalia similar to those of K. bipendula, but with male pygofer margin entire posterodorsally (Fig. 33); aedeagus with ventral process expanded and serrate preapically with lateral preapical lobes (Fig. 35), and apex of shaft with slightly asymmetrical pair of slender laterally directed processes (Fig. 34). Material examined. Holotype male, Ecuador, Orellana, Transect Ent., 1 km S Onkonegare Camp, Reserva Etnica Waorani, S, W, 23 June 1996, T. L. Erwin et al., fogging, terra Þrme forest, lot 1610 (EPNQ). Two Þfth instar nymphs from the same locality (lot 1183, 4 October 1995; lot 1563, 22 June 1996 [USNM]) are tentatively identiþed as this species. Krocodona circumflexa, n. sp. (Figs. 2, 18, 21, 29Ð32) Description. Length male 6.3 mm. Color pattern of crown consisting of an indistinct transverse brown band between eyes and an extensive brown area medially outlining a pair of pale sigmoid lines (Fig. 2). Posterior lobe of pygofer with pair of short posterodorsally directed processes, dorsal process slightly longer with expanded serrate apex, ventral process slightly tapered and truncate (Fig. 29); subgenital plate apex with folded medial lobe; aedeagus without ventral process, shaft in lateral view forming nearly complete circle with apex extended anterad of dorsal apodeme (Fig. 30), apical half strongly depressed and straplike, apex with pair of short slender tapered processes (Fig. 32), each with preapical dorsal tooth (Fig. 30). Material examined. Holotype male, Ecuador, Orellana, Transect Ent., 1 km S Onkone Gare Camp, Reserva Etnica Waorani, S, W, 4 October 1996, T. L. Erwin et al., fogging, terra Þrme forest, lot 1751 (EPNQ). One male paratype, Ecuador, Orellana, Tiputini Biodiversity Station near Yasuni National Park, 220Ð250 m, S, W, 8 February 1999, T. L. Erwin et al., lot 2022, transect T/3, fogging terra Þrme forest (USNM).

13 712 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 6 K. janauaca, n. sp. (Figs. 3 and 36Ð39) Description. Length male 5.8 mm. Color pattern of crown consisting of paired angulate and oblique brown lines (Fig. 3). Male pygofer with posterior lobe bearing a short, acute, posteriorly directed spine (Fig. 36); tenth segment with pair of large ventral processes arising near midlength, extended ventrally, and then curved posterolaterad, apex of each process compressed and expanded with irregular marginal teeth (Fig. 36); subgenital plate apex unmodiþed; aedeagus without ventral process, shaft straight, tapered, depressed, with pair of short lateral preapical processes (Fig. 39). Material examined. Holotype male, Brazil, Amazonas, Rio Janauaca, 40 km sw Manaus, 10 March 1979, S W; Mongomery, Erwin, Sucharov, Schimmel, Krischik, Date, Bacon, Collectors; white water innundation forest canopy fogged with pyrethrum, sample 58 (USNM). Krocozzota Kramer 1964b: 115 Diagnosis. Structurally similar to Krocodona, but differing as follows: head shorter and more strongly tapered in dorsal view, pair of bold dark lateral stripes extended from the crown onto the forewings (Fig. 5); ovipositor with a few preapical teeth of second valvulae more prominent than others (Fig. 72). Notes. Kramer described this genus based on the female holotype of K. languria Kramer from Panama. Malaise trap samples from Colombia yielded two males and a female of a closely related species, described below, that differs in the shape of the female seventh sternite. Krocozzota magdalenensis, n.sp. (Figs. 5, 6, 40Ð42, 62, 68, 71) Description. Length male 4.8Ð5.0, female 5.9 mm. Coloration similar to that of K. languria, except forewing with dark macula completely enclosing small pale area in inner anteapical cell and oblique macula extended from inner anteapical cell to costal margin (cf. Fig. 6 with Kramer 1964b, Fig. 9). Male pygofer (Fig. 40) with ventral preapical tooth well developed, plates in ventral view strongly constricted preapically, fused to apical one-fourths, with a few scattered subapical setae; connective (Fig. 41) short, Y shaped; style apodeme short, apophysis long with prominent setigerous preapical lobe and strongly recurved apical hook; aedeagus (Figs. 41 and 42) slender, shaft sinuate, apex recurved with pair of short slender recurved lateroapical processes. Female sternum VII (Fig. 62) broad and weakly bilobed apically. Material examined. Holotype male, Colombia, Magdalena, PNN Santa Marta El Ramo, N73 39 W, 2500 m, Malaise trap, 5/25/00Ð6/9/00, I. Uribe, M. 198; 1 male and one female paratype, same data (HIC). Retrolidia, New Genus Type species: R. bimaculata, n. sp., here designated. Diagnosis. This genus differs from other Neocoelidiinae in having the crown parabolically produced and distinctly concave, the penultimate pair of macrosetae on the hind femur close set, and the forewing with a false eyespot near the apex. Description. Medium-sized neocoelidiines (6.7Ð9.0 mm). Ground color stramineous; head (Fig. 8) with large black spot apically ventrad of crown margin; crown heavily marked with black; broad black marking extended longitudinally from anterior margin of pronotum covering much of clavus, roundly expanded onto corium near midlength of forewing and tapering toward apex, expanded area with orange crescentshaped mark (Fig. 7); black false eyespot outlined with orange present in distal portion of middle anteapical cell (Fig. 14); three divergent brown lines extended from middle of corium just basad of eyespot to costal margin; overall color pattern in lateral view giving the appearance of an insect facing toward the caudal end of the leafhopper (Fig. 8). Head (Figs. 7Ð10) produced, in lateral view with dorsum slightly declivous and apex obliquely truncate; crown 2 longer than its basal width between eyes, margin carinate anterad of ocelli, apex in dorsal view narrowly parabolic, surface not elevated mesad of eyes, concave, texture rugulose with sparse, minute, pale setae; coronal suture extended to apex of crown; ocelli well developed, just below crown margin above anterior edge of antennal pit; lateral frontal sutures indistinct, divergent, not extended to ocelli; antennal ledge weak, not ßattened; ßagellum ca. half body length; frontoclypeus parallel sided ventrad of antennal pits, with strong median anterodorsal keel; lorum narrower than clypellus; clypellus parallel sided, weakly gibbous medially, apex truncate, not extended as far as genal margin. Pronotum in lateral view (Fig. 8) with dorsal margin weakly declivous, continuing contour of crown; surface transversely striate and minutely punctate; lateral carina even with posterior corner of eye. Exposed part of mesonotum and scutellum approximately as wide as long. Forewing venation (Fig. 14) obscure, except near apex; R 1 well basad of origin of R 4 5 ; outer anteapical cell small and triangular; middle anteapical cell expanded and bulbous distally; second and third apical cells short, quadrate; inner apical cell long, apex oblique; m-cu 3 connected well basad of M fork. Hind wing (Fig. 16) with R 4 5 and M 1 2 conßuent. Front femur (Fig. 19) with AV 1 small, only slighly offset and poorly differentiated from intercalary row, intercalary row with 9Ð10 widely spaced setae approximately subequal in size; tibia with dorsal macrosetae 1 1; middle femur shorter and 2 broader than front femur; hind femur (Fig. 23) with penultimate macrosetal pair close set, but widely divergent; tibia with row AV originating near base; tarsomere I pecten with four platellae. Male pygofer (Fig. 43) with short posterodorsal lobe and long, well-sclerotized curved distal process; anal tube without processes; plates (Fig. 44) free, except near base, with preapical longitudinal row of submesal macrosetae terminating in dense apical mac-

14 November 2003 DIETRICH: UNUSUAL NEOCOELIDIINAE 713 rosetal group; connective (Fig. 46) Y-shaped style with (Fig. 46) or without (Fig. 52) digitiform ventrolateral process near midlength of apophysis, apex curved ventrad, with two prominent preapical setae, apodeme with quadrate dorsolateral lobe; aedeagus (Fig. 45) slightly asymmetrical, with paired processes. Female pygofer with sparse macrosetae distally; Þrst valvulae (Fig. 69) with dorsal margin humped medially, dorsal sculpturing strigate/concatenate, extended to margin only in distal one-third; second valvulae (Fig. 73) with area of dorsal fusion shorter than distal blade, blade broad with numerous close-set square-tipped serrate teeth dorsally, apex blunt, minutely serrate ventrally; third valvula with distal row of ca. seven setae. Fifth instar nymph (based on R. bimaculata, n. sp., Fig. 76) pale yellow with symmetrical brown markings dorsally, glabrous. Head structure similar to that of adult; crown depressed, midline weakly carinate, marginal carina well developed; face with prominent median longitudinal carina, apex subquadrate in proþle; antennal ßagellum slightly longer than body, not including pygofer process. Forewing pads without apical setae. Leg chaetotaxy resembling that of adult, except mesothoracic tibia with scattered Þne elongate setae dorsally. Abdominal terga III-VIII each with pair of large pale dorsolateral setae, those of tergum IV laterad of others; terga VII and VIII also with pair of pale spatulate setae ventrolaterally. Tergum IX with four elongate pale setae in irregular vertical row laterally and 5Ð 6 shorter darker spatulate setae ventrolaterally near apex; distal lobes greatly elongated (broken in all three specimens examined), subsegmented, bearing scattered setae, and ßexibly articulated to pygofer. Notes. Although Retrolidia may fall within KramerÕs (1964b) concept of Nirvaninae, given its strongly produced head, the genus is clearly referable to Neocoelidiinae based on the same criteria used to place Krocodona and Krocozzota within this subfamily (see above). The genus is unusual among neocoelidiines in having the antennal ledges greatly reduced, the surface of the crown even with the dorsum of the compound eyes, the penultimate pair of hind femoral macrosetae relatively close set, the male pygofer with an elongate posterior process (cf. Coelidiana ancora Kramer), and the subgenital plates with numerous macrosetae distally. The genus is described based on three species known only from the Amazonian rain forest canopy. The genus name is feminine and combines the preþx retro-, meaning backward, and -lidia from the type genus of Neocoelidiinae. Key to Species of Retrolidia 1 Crown with two black maculae (Fig. 7) bimaculata, n. sp. 1 Crown entirely black medially (Fig. 9) Forewing clavus with oblique preapical orange vitta broadly connected to commissural margin (Fig. 14); male pygofer process terminating in long dorsal and short ventral spine (Fig. 47).... nigricephala, n. sp. 2 Forewing clavus with oblique preapical orange vitta not or only slightly connected to commissural margin (Fig. 16); male pygofer process terminating in ventrally curved hook (Fig. 48)... serrata, n. sp. R. bimaculata, n.sp. (Figs. 7, 8, 45, 46, 63, 69, 73, 81) Description. Length male 6.7Ð7.1, female 7.4Ð7.6 mm. Crown (Fig. 7) with pair of broad transverse black maculae, one behind the other; dorsum of thorax dark orange-brown medially, contrasting with black sublateral stripe; clavus with oblique preapical orange vitta broadly connected to commissural margin (Fig. 7). Male pygofer (Fig. 43) with distal lobe broadly rounded; process cylindrical, curved mesad, then strongly arched dorsally and hooked ventrolaterad preapically; slender spine of variable length arising just basad of arch apex and extended ventrolaterad, apex of spine even with or shorter than base and apex of process. Style (Fig. 46) with broad mesal preapical lobe, apex weakly bidentate. Aedeagus (Fig. 45) with shaft broad, compressed, curved posterodorsad, then posterad; apex with pair of sinuately curved processes extended posteroventrally, each process with dorsal subbasal posterodorsally curved spine and 1Ð3 ventral subbasal teeth; another pair of processes arising between Þrst pair, extended dorsally and then bent abruptly anterad, irregularly tapered apically, apex arcuate in lateral view, forcipate in dorsal view, with minute ventral teeth; gonopore between bases of processes. Female sternum VII (Fig. 63) with posterior margin roundly produced with small medial notch, concealing base of ovipositor. Material examined. Holotype male, Ecuador, Orellana, Transect Ent. 1 km S Onkonegare Camp, Reserva Etnica Waorani, S, W, 4 October 1994, T. L. Erwin et al., fogging terra Þrme forest, lot 869 (EPNQ). One paratype female, same data. One paratype female and three Þfth instar nymphs, same locality, lot 1420, 4 February 1996 (USNM); paratype male, Ecuador, Orellana, Tiputini Biodiversity Station near Yasuni National Park, 220Ð250 m, S, W, 26 October 1998, T. L. Erwin et al., lot 1958 (INHS). Retrolidia nigricephala, n. sp. (Figs. 9, 10, 14, 15, 23, 47, 49, 50, 64) Description. Length male 7.8, female 8.9 mm. Crown (Fig. 9) black, except for narrow pale marginal band; dorsum of thorax uniformly black medially; forewing clavus with oblique preapical orange vitta broadly connected to commissural margin (Fig. 14). Male pygofer (Fig. 47) with posterodorsal lobe narrow; process compressed, curved ventrad then posterad, weakly sclerotized ventral ßange terminating in angulate process preapically, apex with long dorsal

15 714 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 6 and short ventral spine, each spine weakly recurved. Style (Fig. 50) without mesal preapical lobe, apophysis tapered and pointed distally. Aedeagus (Fig. 49) with pair of long slender ventral processes arising from base, each with preapical dorsal lobe and sinuate apex; shaft slender, weakly curved posterodorsad, apex with pair of small compressed, rounded lateral lobes and short depressed anterodorsal process; gonopore between lobes. Female sternum VII (Fig. 64) posterior margin with trapezoidal median lobe and pair of shorter, rounded lateral lobes. Material examined. Holotype male, Ecuador, Orellana, Tiputini Biodiversity Station near Yasuni National Park, 220Ð250 m, S, W, 26 October 1998, T. L. Erwin et al., lot 1988, transect T/9, fogging terra Þrme forest (EPNQ); two paratype females, same locality, 5 February 1999, lot 2086, same transect (USNM, INHS). Retrolidia serrata, n.sp. (Figs. 16, 19, 48, 51, 52, 65) Description. Length male 7.6, female 8.2 mm. Coloration similar to that of R. nigricephala, except oblique orange vitta near apex of clavus not extended to commissural margin (Fig. 16). Male pygofer (Fig. 48) with well-developed, weakly sclerotized posteroventral lobe; posterodorsal process compressed, irregularly serrate dorsally, gradually curved posterodorsad with prominent ventral preapical lobe and apical hook. Style (Fig. 52) without preapical lobes, apophysis tapered and pointed distally, with two long ventral preapical setae. Aedeagus (Fig. 51) with preatrium elongate; shaft tubular basally, recurved, depressed distally with pair of branched preapical lateral processes extended toward apex, apex bifurcate with distal lobes serrate, gonopore at base of bifurcation. Female sternum VII (Fig. 65) with truncate median lobe shorter than paired lateral lobes. Material examined. Holotype male, Ecuador, Orellana, Transect Ent. 1 km S Onkonegare Camp, Reserva Etnica Waorani, S, W, 22 June 1996, T. L. Erwin et al., fogging terra Þrme forest, lot 1572 (EPNQ); paratype female, same locality, 9 February 1999, lot 956 (USNM). Krocolidia, New Genus Type species K. rufilinea, n. sp., here designated. Diagnosis. This genus resembles Retrolidia in having the head strongly produced, but is easily distinguished by the distinct median longitudinal carina on the crown and the absence of mimetic color patterns. Description. Structurally similar to Retrolidia, but differing as follows. Color pale stramineous with or without red median dorsal longitudinal stripe. Head (Fig. 11) 2.5 longer than basal width between eyes, crown with distinct median longitudinal carina, margins carinate and ßangelike anterad of ocelli, apex in dorsal view evenly rounded, surface between median and lateral carina concave, uniformly shagreen, without distinct setae; gena not completely concealing proepisternum in anterior view; antennal ledge well developed, oblique. Pronotum with lateral carina below posterior corner of eye. Forewing (Fig. 17) with outer anteapical cell large, crossveins nearly even with r-m; middle anteapical cell weakly expanded distally, second and third apical cells relatively long; inner apical cell with apex tapered; m-cu 3 connected to M 3 4. Front femur (Fig. 20) slightly shorter than middle femur, hind femur (Fig. 22) with penultimate pair of macrosetae widely separated, tibial row AV originating approximately half distance from base. Male pygofer (Fig. 53) with large posterodorsal process arising as sclerotized fold along dorsal margin, and short recurved ventral tooth; plates (Fig. 54) with a few sparsely distributed macrosetae preapically, fused to just beyond midlength, in ventral view with broad lateral lobe near midlength; connective Y shaped; style (Fig. 56) apophysis elongate, with broad low setigerous lobe, apex weakly hooked; aedeagus (Fig. 55) with pair of basal processes, shaft short, compressed, without apical paired processes. Female sternum VII (Fig. 66) broadly rounded posteriorly; Þrst valvula (Fig. 70) similar to that of Retrolidia, except median dorsal hump less pronounced; second valvula (Fig. 74) slender, area of dorsal fusion longer than distal blade, toothed distal blade with prominent basal and preapical tooth in addition to numerous smaller teeth; third valvula with 2Ð3 small preapical ventral setae. Fifth instar nymph (based on unidentiþed species from Ecuador, Fig. 77 [USNM]) pale, unmarked, except pair of small black spots, one anterad of other, on lateral margin of abdominal tergum VI; glabrous. Head structure similar to that of adult, antennal ßagellum exceeding length of body. Leg chaetotaxy resembling that of adult, except setae of hind tibial row PV long and Þne, length more than twice maximum width of tibia. Abdominal terga IV-VIII each with enlarged seta posterolaterally; tergum IX slightly longer than tergum VIII, with two subbasal setae and three more distal setae. Notes. Females probably representing additional, undescribed species of this genus were examined from the following localities: Ecuador, Orellana, Tiputini Biodiversity Station (lot 2090, 5 February 1999, USNM); Peru, Madre de Dios, Rio Tambopata Reserve (12 November 1983, USNM); Brazil, Para, Santo Antonio do Taua (29 January and 15Ð23 October 1979, MNHN). Two nymphs of an undescribed species were examined from Ecuador, Orellana, Reserva Etnica Waorani, 1 km S Onkonegare Camp (lot 1739, 3 October 1996, USNM). The genus name is feminine and is a combination of the preþx Kroco- from the three neocoelidiine genera described by Kramer and originally placed in Nirvaninae, with the sufþx -lidia, from the type genus of Neocoelidiinae. Krocolidia is described based on two species. K. rufilinea, n.sp. (Figs. 12 and 13, 53Ð56) Description. Length male 5.9 mm. Dorsum (Fig. 12) with broad bright reddish orange median longitudinal

16 November 2003 DIETRICH: UNUSUAL NEOCOELIDIINAE 715 stripe extended from apex of crown to apex of clavus. Male pygofer (Fig. 53) with posterodorsal process extended well beyond posteroventral tooth; plates (Fig. 54) narrowly rounded apically, without dorsal macrotricha; aedeagus (Fig. 55) with basal processes short, evenly tapered, extended dorsolaterad; shaft with numerous small lateral preapical spines, extended posterad of basal processes. Material examined. Holotype male, Brazil, Para, Jacareacanga, I-1970, F. Barbosa, B.M ; one male paratype, same data, except VI-1970 (both BMNH). Krocolidia obscura, n.sp. (Figs. 11 and 57Ð60) Description. Length male 6.5 mm. Dorsum (Fig. 11) with obscure orange median longitudinal stripe and pair of orange lateral stripes restricted to crown. Pygofer (Fig. 57) with posterodorsal process not surpassing ventral tooth, toothlike apical process present just dorsad of ventral tooth; plates (Fig. 58) broadly rounded apically, with dense longitudinal band of sublateral macrotrichia on dorsal surface in distal half; aedeagus (Fig. 59) with basal processes arising ventrally and extended posterolaterad, each process slender, tapered, with mesial preapical tooth just distad of midlength (Fig. 60); shaft without lateral spines, extended anterodorsad of basal processes. Material examined. Holotype male, Brazil, Para, Jacareacanga, I-1970, F. Barbosa, B.M ; one female paratype, same data, except IX-1970 (BMNH). Acknowledgments I am indebted to Terry Erwin and Warren Steiner for access to the Smithsonian canopy fogging samples from western Amazonia; to Paul Freytag and Michael Sharkey (University of Kentucky, Lexington) for access to their Malaise trap samples from Colombia; and to Dmitry Dmitriev, Roman Rakitov, Daniela Takiya, Mick Webb, Jamie Zahniser, and two anonymous referees for comments on various drafts of the manuscript. This work was supported in part by Grants DEB and DEB from the National Science Foundation. References Cited DeLong, D. M A synopsis of the tribe Neocoelidinii [sic] in the Americas (Homoptera-Cicadellidae). Lloydia 16: 93Ð131. Dietrich, C. H The role of grasslands in the diversi- Þcation of leafhoppers (Homoptera: Cicadellidae): a phylogenetic perspective, pp. 44Ð48. In C. Warwick (ed.), Proceedings, Fifteenth North American Prairie Conference, 23Ð26 October 1996, St. Charles, IL. Natural Areas Association, Bend, OR. Dietrich, C. H., and D. A. Dmitriev Reassessment of the leafhopper tribes Koebeliini Baker and Grypotini Haupt (Hemiptera: Cicadellidae). Ann. Entomol. Soc. Am. 96: 766Ð775. Dietrich, C. H., and R. A. Rakitov Some remarkable new deltocephaline leafhoppers (Hemiptera: Cicadellidae: Deltocephalinae) from the Amazonian rainforest canopy. J. NY Entomol. Soc. 110: 1Ð48. Dietrich, C. H., and F. E. Vega Leafhoppers (Homoptera: Cicadellidae) from Dominican amber. Ann. Entomol. Soc. Am. 88: 263Ð270. Dietrich, C. H., R. A. Rakitov, J. L. Holmes, and W. C. Black IV Phylogeny of the major lineages of Membracoidea (Insecta: Hemiptera: Cicadomorpha) based on 28S rdna sequences. Mol. Phyl. Evol. 18: 293Ð305. Erwin, T. L Tropical forest canopies: the last biotic frontier. Bull. Entomol. Soc. Am. 29: 14Ð19. Erwin, T. L Canopy arthropod biodiversity: a chronology of sampling techniques and results. Rev. Peru. Entomol. 32: 71Ð77. Evans, J. W A natural classiþcation of leaf-hoppers (Homoptera, Jassoidea). Part 3. Jassidae. Trans. R. Entomol. Soc. London 98: 105Ð271. Godoy, C., and M. D. Webb Recognition of a new subfamily of Cicadellidae from Costa Rica based on a phenetic analysis with similar taxa (Hemiptera Homoptera Auchenorrhyncha). Trop. Zool. 7: 131Ð144. Hamilton, K.G.A ClassiÞcation, morphology and phylogeny of the family Cicadellidae (Rhynchota, Homoptera), pp. 15Ð37. In W. J. Knight, N. C. Pant, T. S. Robertson, and M. R. Wilson (eds.), Proceedings, First International Workshop on Biotaxonomy, ClassiÞcation and Biology of Leafhoppers and Planthoppers of Economic Importance, 4Ð7 October 1982, London, United Kingdom. Commonwealth Institute of Entomology, London, England. Kramer, J. P. 1964a. A generic revision of the leafhopper subfamily Neocoelidiinae (Homoptera: Cicadellidae). Proc. US Nat. Mus. 115: 259Ð287. Kramer, J. P. 1964b. A review of the Neotropical Nirvaninae (Homoptera: Cicadellidae). Entomol. News 75: 113Ð128. Kramer, J. P New Neotropical Neocoelidiinae with keys to the species of Coelidiana, Xenocoelidia, and Nelidina. Proc. Entomol. Soc. Wash. 69: 31Ð46. Kramer, S The morphology and phylogeny of Auchenorrhynchous Homoptera (Insecta). Illinois Biol. Monogr. 20: 1Ð111. Linnavuori, R Revision of the Neotropical Deltocephalinae and some related subfamilies (Homoptera). Ann. Soc. Zool. Bot. Fenn. Vanamo 20: 1Ð370. Lucky, A., T. L. Erwin, and J. D. Witman Temporal and spatial diversity and distribution of arboreal Carabidae (Coleoptera) in a western Amazonian rain forest. Biotropica 34: 376Ð386. Oman, P. W The Nearctic leafhoppers: a generic classiþcation and check list. Mem. Entomol. Soc. Wash. 3: 1Ð253. Oman, P. W., W. J. Knight, and M. W. Nielson Leafhoppers (Cicadellidae): a bibliography, generic checklist, and index to the world literature 1956Ð1985. C.A.B. International Institute of Entomology, Wallingford, United Kingdom. Rakitov, R. A On differentiation of cicadellid leg chaetotaxy (Homoptera: Auchenorrhyncha: Membracoidea). Russian Entomol. J. 6: 7Ð27. Received for publication 25 March 2003; accepted 11 July 2003.