Proceedings of ^^\the United States IK-S^^National Museum ^'^^^SMITHSONIAN INSTITUTION ? WASHINGTON, D.C.
Volume 115 1964 Number 3484
A GENERIC REVISION OF THE LEAFHOPPER SUBFAMILYNEOCOELIDIINAE(HOMOPTERA: CICADELLIDAE)
By James P. Kramer'
The leafhopper subfamily Neocoelidiinae was erected by Oman(1943) in order to accommodate the two North American genera,Neocoelidia and Paracoelidea, which had been placed in the Jassinae(Deltocephalinae of modern workers) for many years. Evans (1947)added the Neotropical genera Biza, Chinaia, Coelidiana, and Salvinain his generic checklist of the subfamily. DeLong (1953) partiallyrevised the group, adding new genera and subgenera as well as manynew species. He included only one of the genera added by Evans andattempted to place the Neotropical Neocoelidiinae described by Fowlerand Osborn with reference to the literature alone. More recently newgenera and species have been added by Kramer (1959, 1961, and 1962)and by Kramer and Linnavuori (1959). Although our knowledge ofNeotropical fauna is still quite meager, it seems appropriate to synthe-size the information we do have at present in order that future workersmay have a foundation upon which to build.
I Entomology Research Division, Agricultural Research Service, U.S. Department of Agriculture.259
260 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115Subfamily NeocoelidiinaeDescription.?Small to large leafhoppers (3-13 mm.). Clypelluseither approximately parallel-sided or widened distally. Lateralfrontal sutures extended near or to ocelli, which are on or near anteriormargin of crown or less often on face. Ocellocular areas developed asdistinct ledges above antennal pits. Antennae long to very long, fre-quently exceeding entire length of body including forewings at rest.Face and crown either separated by carina or with carina lacking;carina, when present, usually limited to distance between ocelli. Headin dorsal view angled or rounded apically, often strongly producedbeyond eyes. Pronotum short, much wider than long, with lateralmargins carinated. Scutellum large and well developed. Venationof forewings usually obscure except apically and with either two orthree preapical cells. Male genital structures variously modified butstyles usually simple. Ground color various shades of white, yellow,or orange. There are often contrasting markings of various hues onthe head, thorax, and forewings.Diagnosis.?The most useful set of characters for recognition of theNeocoelidiinae consists of the distinct ledge above each antennal base,the exceptionally long antennae (particularly as found in many of theNeotropical genera), and the venation of the forewings, which isusually highly obscure except apically.Discussion.?As far as it is known, the subfamily is restricted tothe Americas, with good representation in both the temperate andtropical areas. The available host and food plant records seem toindicate that most of the species occur on trees or shrubs. Accordingto DeLong (1953), various Nearctic species have been taken on the fol-lowing genera of plants: Pinus, Acacia, Rhus, Arctostaphylos, andSphaeralcea. Except for one species of Chinaia known to breed onavocado, the plant relationships of the Neotropical species areunknown.The key to genera which follows is based almost entu-ely upon char-acters of the male genitalia. Although most of these structures areconcealed within the genital capsule and require dissection and clear-ing in order to be observed, they provide the most concrete evidencefor an arrangement of genera within the subfamily. Emphasis on themale genitalia for the delimiting of genera is not without precedent.Oman (1949) used this feature extensively in arranging the NorthAmerican Deltocephalinae, as did Young (1952) in revising the NewWorld Typhlocybinae. Both men provided generic definitions withinthe respective subfamilies that were more definite and better delimitedthan any prior to their work. It is hoped that this study will servea similar purpose.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?^KRAMER 261Key to Genera of Neocoelidiinae
1. Aedeagus consisting of two shafts, one above other, dorsal shaft bearinggonoduct (figs. 2, 8, 17) Neocoelidia Gillette and BakerAedeagus consisting of single shaft, which may be simple or elaborated withprocesses (figs. 42, 99) 22. Face and crown separated by carina, which is as long as distance betweenocelli 5Face and crown not separated by carina (i.e., carina absent) 33. Ocelli located on face below anterior margin of crown; clypellus distinctlyexpanded distally; connective cruciform . . Chinaia Bruner and MetcalfOcelli located on anterior margin of crown; clypellus not distinctly ex-panded distally; connective Y-shaped or nearly so 44. Aedeagus without apical modifications but with long recurved lateral,paired processes; male plates fused for entire length.Deltocoelidia KramerAedeagus with apical modifications but without lateral processes; maleplates not fused for entire length Xenocoelidia Kramer5. Exceedingly large and robust forms, males over 12 mm. in length; head in-cluding eyes less than two-thirds pronotal width.Megacoelidia Kramer and LinnavuoriSmall to moderate-sized species, males less than 10 mm. in length; head in-cluding eyes always more than two-thirds pronotal width 66. Ventral margin of male pygofer with distinct apical or preapical hook orpygofer terminating with pointed process (figs. 75, 81) 8Ventral margin of male pygofer without hooks and pygofer not terminatingwith pointed process 77. Aedeagus strongly asymmetrical; pygofer with dorsal process; styles ofmoderate length, not wrinkled, well sclerotized (figs. 49, 45, 47).Tozzita, new genusAedeagus symmetrical; pygofer without dorsal process; styles long, wrinkled,and weakly sclerotized (figs. 58, 55, 60) Xiqilliba, new genus8. Male pygofer inflated, laterally with distinct dorsoventral suture, hook onventral margin large and heavy (fig. 50) Coelana DeLongMale pygofer not inflated, laterally without distinct dorsoventral suture,hook on ventral margin small, slender, or absent 99. Aedeagus asymmetrical and deeply cleft dorsoventrally ; pygofer withcrossed internal processes at apex Tichocoelidia KramerAedeagus symmetrical but at times twisted, never deeply cleft; crossedprocesses of pygofer, if present, external 1010. Apex of pygofer in lateral view appearing cleft due to long mesally curveddorsal hook and weakly sclerotized ventral extension; stylar apex indorsal view curving laterally and caudally (figs. 62, 65) . Salvina MelicharApex of pygofer in lateral view and stylar apex in dorsal view not as above . 1
1
11. Anal tube of male with forked ventral hook; ventral cover of genital capsulesharply narrowed on distal half appearing as two extended "fingers"(figs. 67, 70) Cocoelidia DeLongAnal tube of male with single simple or pair of ventral hooks or none; ventralcover of genital capsule not as above 1212. With small but distinct dark-brown or black spot at apex of crown. ... 15Without spot at apex of crown 13
262 PROCEEDINGS OF THE NATIONAL MUSEUM vol. us
13. Cercopidlike species (fig. 114); forewing venation distinct; aedeagus simpleor with long lateral processes Biza WalkerNot cercopidlike species; forewing venation obscure; aedeagus simple orwith apical processes 1414. Aedeagus slender, with paired apical processes; male plates with long mac-rosetae (figs. 80, 78) Nelidina DeLongAedeagus not particularly slender, without apical processes; male plateswithout macrosetae (figs. 95, 98) Coelidiana Oman15. All veins of forewings brown and distinct; male plates with macrosetae(figs. 88, 89) Coelella DeLongMost veins of forewings yellowish and highly obscure; male plates withoutmacrosetae (figs. 81, 85) Neocoelidiana DeLongNeocoelidia Gillette and BakerFigures 1-35Neocoelidia Gillette and Baker 1895, p. 103. Type of genus Neocoelidia tumidi-frons Gillette and Baker by original designation.Paracoelidea Baker 1898, p. 292. Type of genus Paracoelidea tuherculata Bakerby original designation. New synonymy.Stenocoelidia DeLong 1953, p. 104. Type of genus Stenocoelidia virgata DeLongby original designation. New synonymy.Stenocoelidia subgenus Eurycoelidia DeLong 1953, p. 112. Type of subgenusNeocoelidia pulchella Ball by original designation. New synonymy.Description.?Carina separating face and crown distinct, weaklydeveloped, or absent. Shape of crown in dorsal view highly variable,extremes with anterior margin broadly rounded to sharply angular,nearly always longer at middle than next to eye. OceUi occur onanterior margin of crown. Head including eyes usually narrowerthan pronotum, which is more or less indented on posterior margin.Antennae from about half to three-fourths total length of bodyincluding forewings. Venation of forewings highly obscure, exceptapically, in most species.Ground color ivory white to yellow. Often with spot at apex ofcrown and pair of dark spots on scutellum. Many with dorsal stripefrom apex of crown to distal portion of forewings. Species rarelyimmaculate.Male genitaha as discussed below.Discussion.?The reduction of Paracoelidea, Stenocoelidia, andEurycoelidia to synonymy under Neocoelidia is based upon the consist-ently uniform pattern of the male genitalia and the lack of sharply de-fined limits in gross characters between the groups as previouslydefined.Attention is called to the following drawings for the pm'poses ofthe discussion: N. tumidifrons (figs. 1-6), A^. tuherculata (figs. 7-13),N. virgata (figs. 14-16), and A^. pulchella (figs. 17-24). The malegenitalia of these various type-species show that in all cases theventer of the capsule is covered by a single plate presumably formed
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 263by a complete fusion of the normally paired plates. Other mutuallyshared characters are the comparatively simple pygofers distinguishedby a single ventral hook or tooth and the pair of slender simple proc-esses at the base of the anal tube. In all of them the aedeagus consistsof two shafts which are arranged one above the other with the dorsalshaft bearing the gonoduct. The connective is uniformly Y-shapedand the styles are basically alike.DeLong's separation of Stenocoelidia and Eurycoelidia from Neo-coelidia on the basis of elongate and slender forms versus short androbust forms holds for some of the species, but others cannot beseparated on this basis because of their intermediate condition.Paracoelidea, although striking because of the tuberculate clypellus,is considered here as a group within Neocoelidia because of thesimilarities in male genitalia.The genus Neocoelidia is primarily North and Central Americanbut a few species range into northern South America. Knull (1942)has reviewed the North American species, providing a key basedlargely on color and size. DeLong (1953) described many newspecies primarily from Mexico and Central America; and althoughhe provided no key, his illustrations of the pygofers and aedeagi arehighly adequate for species recognition.Three species of Neocoelidia, the types of which have not beenstudied since their original description, are discussed here.Neocoelidia fuscodorsata (Fowler), new combinationFigures 25-28Tettigonia fuscodorsata Fowler 1900, p. 269, tab. 18, fig. 6.Stenocoelidia fuscodorsata (Fowler), DeLong 1953, p. 121.Stenocoelidia clara DeLong 1953, p. 104, new synonymy.Fowler described T. fuscodorsata from a long series of specimenscollected at several localities in Mexico and Guatemala. DeLongassumed that Fowler's series included more than one species, whichis probably true, but he studied no type material. Through thecooperation of Dr. W. E. China and the British Museum (NaturalHistory), three of Fowler's syntypes, one male and two females,were made available for study. The male with data " Teapa, Tabasco,Mexico" is hereby designated as the lectotype. A comparison ofthis lectotype and the type of S. clara DeLong show that these areconspecific. The colored illustration of T. fuscodorsata whichappeared with Fowler's description is not diagnostic for the species.The general coloration is as figured, with the following modification:distinct black spot at apex of crown and near each basal angle ofscutellum; apical spot fused with dorsal stripe whereas spots on scutel-lum free. The male genitaha are shown in figures 25-28.
264 PROCEEDINGS OF THE NATIONAL MUSEUM vol. iisNeocoelidia verecunda (Fowler), new combinationTettigonia verecunda Fowler 1900, p. 269, tab. 18, fig. 7.Stenocoelidia verecunda (Fowler), DeLong 1953, p. 122.Unfortunately this Guatemalan species is knoAvn from femalesonly, and its generic placement is open to question. I have studiedFowler's two syntypes in the British Museum and two additionalspecimens in the collection of the U.S. National Museum. In noneof the specimens are the longitudinal red markings on the crown,pronotum, and forewings quite as distinct as illustrated with Fowler'soriginal description. Neocoelidia crenulata OsbornFigures 29-35Neocoelidia crenulata Osborn 1923, p. 79.This species was described from a single male from Minca, Colombia.The type was made available through the courtesy of Dr. G. E. Wal-lace and the Carnegie Museum. In general it is marked like N.fuscodorsata except that there are no spots on the scutellum and thelongitudinal dorsal stripe is crenulate on its margins. The genitaliaof the type are illustrated in figures 29-35.Chinaia Bruner and MetcalfFigures 37, 113Chinaia Bruner and Metcalf 1934, p. 120. Type of genus Chinaia bella Brunerand Metcalf by original designation.Description.?Without carina separating face and crown. Cly-pellus expanded distally. Shape of crown in dorsal view broadlyrounded, wider than long. Ocelli on face distinctly below anteriormargin of crown. Head including e^^es narrower than pronotum,which is not indented on posterior margin. Antenna very long,exceeding total length of body including forewings. Venation offorewings obscure except at apex.Ground color yellowish to pale orange, with orange to bright red-orange markings on pronotum and extensively on forewings. Dark-brown markings often found on clavus and apical portion of forewingsas well.Male genitalia: Valve obscure. Plates deep and scoop-shaped.Pygofer either with dorsal processes or an elongation of terminus.Anal tube simple. Connective cruciform and not fused with aedeagus.Aedeagus moderately straight or recurved with or without lateralflaps.Discussion.?This is a Neotropical genus; most of the describedspecies occur in Central America, but a few are South American.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 265Chinaia was recently revised by Kramer (1959). A habitus drawingof a typical Chinaia can be seen in figure 113.Deltocoelidia KramerDeltocoelidia Kramer 1961, p. 238. Type of genus Deltocoelidia maldonadoiIvramer by original designation.Description.?Without carina separating face and crown. Shapeof crown in dorsal view bluntly angular. Ocelli located on anteriormargin of crown. Head including eyes about as wide as pronotum,which is broadly indented on posterior margin. Antennae about aslong as total length of body including forewings. Venation of fore-wings obscure except apically.Ground color stramineous with contrasting markings on crown,pronotum, and forewings in form of stripes and spots.Male genitalia: Male plates solidly fused and appear as single plateon venter of genital capsule. Both pygofer and anal tube simple.Connective Y-shaped and poorly sclerotized. Aedeagus slender,recurved, and with paired lateral processes.Discussion.?The single leafhopper at present referable to thisgenus is the type-species, Deltocoelidia maldonadoi Kramer, knownonly from Venezuela. Illustrations of the male genitaha appearedwith the original description.Xenocoelidia KramerFigures 38-44Xenocoelidia Eo-amer 1959, p. 30. Type of genus Xenocoelidia youngi Kramerby original designation.Description.?Without carina separating face and crown. Shapeof crown in dorsal view rounded or subangular. Ocelli located onanterior margin of crown. Head including eyes either as wide aspronotum or slightly narrower. Posterior margin of pronotumbroadly and very shallowly indented. Antennae as long as totallength of body including forewings. Venation of forewings obscureexcept at tip.Ground color ivory to yellowish with or without contrasting mark-ings on crown, pronotum, and forewings.Male genitalia: Valve lacldng. Male plates fused at least basally,at times for most of length. Pygofer with or without ventral toothbut always with terminus slightly thickened and bearing setae. Analtube simple. Connective Y-shaped and closely associated with aedea-gus but joint between them flexible. Style long with mesal curvature.Aedeagus slender and broadly U-shaped in lateral view with apicalelaborations.
266 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115Discussion.?This genus was originally described to receive thespecies X. youngi Kramer and X. colombiana Kramer. Osborn (1923,p. 77) described Neocoelidia inflata, which has been found to be con-generic with the two originally included species. Through thecooperation of Dr. G. E. Wallace of the Carnegie Museum, the types ofOsborn's species were made available for study. The generic transferis made at this tune: Xenocoelidia inflata (Osborn), new combination.Key to Species of Xenocoelidia1. Coloration almost uniform ivory white except for tiny fuscous spot at tip ofeach clavus and yellow hyaline apical portion of each forewing; apex ofaedeagus in lateral view with sharp tooth and broad acute bladelike dorsalexpansion (Colombia) X. youngi KramerColoration involving distinct markings on head and pronotum; apex ofaedeagus in lateral view appearing as two sharp points 22. Head with two pairs of orange spots, one marginal between ocelh and onediscal on crown. Pronotum with two orange-margined black spots nearanterior margin. Each lateral margin of scutellum with black spot.Orange triangular spot at scutellar apex. Forewings unmarked except forvery slight enbrowning along commissural margin. Aedeagal apex inposterior view with one pair of long antlerlike processes (Brazil) (genitaliaillustrated in figs. 39-44) X. inflata (Osborn)Head with pale-yellow band below and a bright-orange band above anteriormargin. Pronotum with lateral margins and an irregular U-shaped anteriorcentral marking bright orange. Scutellum unmarked. Forewings markedas follows: claval suture with a pale yellow band becoming obscure dis-tally; clavus with dusky orange stripe running along scutellum and com-missural margin; a brown spot flanks stripe laterally before apex of eachclavus. Aedeagal apex in posterior view with two pairs of comparativelyshort processes (Colombia) X. colombiana KramerMegacoelidia Kramer and LinnavuoriMegacoelidia Kramer and Linnavuori 1959, p. 55. Type of genus Megacoelidiasplendida Kramer and Linnavuori by original designation.Description.?With distinct carina separating face and crown.Shape of crown in dorsal view subquadrate, wider than long, scarcelyangular apically, distinctly concave, and with lateral and posteriormargins carinate. Ocelli on anterior margin of crown. Head includ-ing eyes comparatively small, less than two-thirds as wide as pro-notum. Posterior margin of pronotum broadly but slightly indented.Antennae longer than entire length of body including forewings.Venation of forewings obscure except apically.Ground color rich deep orange with or without black markings onlegs, pronotum, and forewings.Male genitalia : Valve lacking. Male plates fused basally. Pygofervariously modified distally with elongations or processes. Anal tubeand styles simple. Connective Y-shaped and clearly articulatedwith aedeagus. Aedeagus stout with apical portion recurved.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 267Discussion.?Megacoelidia contains only two known South Amer-ican species, both of which appeared with the original generic descrip-tion. The genus contains the largest members of the subfamily.Key to Species of MegacoelidiaDorsal markings consisting of single narrow black border on posterior margin ofpronotum and on apex of each forewing. Male plates in ventral view pointedapically and aedeagus with long paired lateral processes (Brazil).M. splendida Kramer and LinnavuoriDorsal markings limited to single narrow black border on apex of each forewing.Male plates in ventral view rounded apically and aedeagus with short pairedlateral processes (Bolivia) M. aurantia Kramer and LinnavuoriTozzita^ new genusFigures 45-49Type of genus Tozzita ips, new species.Description.?With slender but distinct carina separating face andcrown. Crown in dorsal view much longer than wide, bluntly angularapically, and strongly produced beyond eyes. Ocelli near anteriormargin of crown but posterior to carina. Head including eyes dis-tinctly narrower than pronotum. Posterior margin of pronotumbroadly indented. Antennae as long as body including forewings atrest. Venation of forewings highly obscure except apically.Ground color stramineous to light brown without definite markingsexcept for black spot at apex of crown. Forewings stramineoushyahne.Male genitalia.?Valve lacking. Male plates fused for nearlyentire length. Pygofer modified only dorsally with processes or exten-sions. Anal tube with heavily sclerotized portions. Connective moreor less V-shaped and articulated with strongly asymmetrical aedeagus.Tozzita ips, new speciesFigures 45-49Length.?Male 6.75 mm.Coloration.?Stramineous with indefinite brownish areas on headand thorax. Only distinct marking is black apical spot on crown.Forewings stramineous hyaline.Male genitalia.?Capsule in ventral view with slight apical notchon ventral cover which is exceeded by the pygofer (fig. 48). Capsulein lateral view with few tiny spines on ventral margin of pygofer,dorsum with blunt, sclerotized process; anal tube with two heavilysclerotized plates, posterior one notched dorsally (fig. 45). Aedeagusin ventral aspect slender and strongly asymmetrical (fig. 49), withapex forked and gonopore opening on narrow mesal extension (fig. 46).681-546?63 2
268 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115Aedeagus slender in lateral view with dorsal hump on basal half andwith apical mesal extension slender and recurved (fig. 47).HoLOTYPE.?Male, Riberalta, Bolivia, W. M. Mann, January,1921-22, Mulford Biological Expedition. USNM type 65826.Female unknown. Xiqilliba, new genusFigure 55-60Type of genus Xiqilliba hellator, new species.Description.?With distinct carina separating face and crown.Crown in dorsal view wider than long and bluntly angular apically.Ocelli near anterior margin of crown but sHghtly posterior to carina.Head including eyes narrower than pronotum. Posterior margin ofpronotum mesally indented. Antennae as long as body includingforewings. Venation of forewings comparatively distinct.Ground color yellow marked with brown or black on head,pronotum, and forewings.Male genitalia.?Valve lacking. Male plates fused basally forabout half length. Pygofer simple. Anal tube with long paired proc-esses extending anteriorly into genital chamber. Styles very long,wrinkled, slender, and poorly sclerotized. Connective modifiedY-shaped. Aedeagus slender and simple.Xiqilliba bellator, new speciesFigures 55-60Length.?Male 6.5 mm.Coloration.?Grossly appearing as yellow leafhopper with brownstripe extending from anterior margin of crown across pronotum andscutellum on to forewings, where abruptly widens near midpoint ofeach clavus extending laterally to costal margin of each forewing,thus covering entire distal portion of forewings.Ground color yellow. Black spot at apex of crown below carina.Crown mesally brown but color vaguely delimited. Pronotum mesallybrown with color widest posteriorly. Scutellum with four very darkstripes: two wide lateral and two narrow double-toothed centralstripes. Forewings brown except for large yellow patches in anteriorcostal area.Male genitalia.?As defined generically with additional charac-ters as follows : long paired processes of anal tube moderately slenderbut irregular in lateral view (fig. 55), sharply pointed and partiallycrossed anteriorly and double-pronged basally where joining anal tubein ventral view (fig. 56). Partiallj^ crossed anterior portions ofprocesses form base on which aedeagus rests. Aedeagus uniformlyslender, upturned distally, with apex projecting shghtly caudally (fig. 58)
.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 2G9HoLOTYPE.?Male, and one paratype male, Itaituba, Brazil, noother data. USNM type 65827. Female unknown.Coelana DeLong, new statusFiGUBES 50-54, 108-109Coelidiana subgenus Coelana DeLong 1953, p. 128. Type of subgenus Neocoelidiamodesta Baker by original designation.Description.?With carina separating face and crown. Crown indorsal view broadly angular at apex and slightly wider than long.Ocelli on anterior margin of crown. Head including eyes distinctlynarrower than pronotum. Posterior margin of pronotum broadly andsharply indented. Antennae about half as long as body includingforewings. Venation of forewings obscure except apically.Ground color stramineous with small black spot at coronal apex.Forewings stramineous hyaline.Male genitalia: Valve lacking. Male plates fused basally. Py-gofer greatly inflated, clearly exceeding length of plates, and ventralmargin with large and heavy hook. Anal tube with ventral process.Connective approximately Y-shaped and articulated with simpleaedeagus.Discussion.?Ooelana includes two species, G. modesta (Baker)(figs. 50-54) and C. drakei new species, from South America. Bothare known from Bolivia, but G. modesta is recorded also from northernArgentina and southern Brazil.Key to Species of CoelanaMALES ONLYLength 7 mm. or more; pygofer in lateral view broadly rounded apically and withinner process entire distally (fig. 50); aedeagus transverse (fig. 54).C. modesta (Baker)Length 6.5 mm. or less; pygofer in lateral view narrowed apically and with innerprocess dentate distally (fig. 108); aedeagus short-coupled (fig. 109).C, drakei, new speciesCoelana drakei, new speciesFigures 108-109Length.?Male 6.3 mm.Coloration.?Uniformly stramineous with only distinct markingconsisting of black spot at coronal apex.Male genitalia.?Capsule in lateral view with pygofer narrowedapically and inner processes double-toothed ventrally at apex, only oneprocess visible in drawing (fig. lOS). Aedeagus in lateral view withshaft sharply upturned (fig. 109).
270 PROCEEDINGS OF THE NATIONAL MUSEUM vol. ii5HoLOTYPE.?Male, Bolivia, no other data. USNM type 66368Female unknown.Discussion.?This species is very close to C. modesta but is sepa-rated easily from it by the characters in the above key. The species isnamed for Dr. Carl John Drake, from whose collection the specimenwas obtained. Tichocoelidia KramerTichocoelidia Kramer 1962, p. 104. Type of genus Tichocoelidia clarkei Kramerby original designation.Description.?With carina separating face and crown, lateral andposterior coronal margins carinate. Concave crown in dorsal viewangled apically, approximately pentagonal in shape and well pro-duced beyond eyes with ocelli on anterior margin. Head includingeyes distinctly narrower than pronotum. Antennae about three-fourths as long as body including forewings. Posterior margin ofpronotum mesally indented. Venation of forewings comparativelydistinct.Ground color stramineous to light brown with few additional mark-ings, most of which are inconspicuous. Forewings stramineoushyaline.Male genitalia: Valve lacking. Male plates short and fused ex-cept apically. Pygofers with ventral processes and paired internal proc-esses at apex. Anal tube with pair of ventral hooks. Connectivemodified Y-shaped and firmly fastened to slender, cleft, asjrmmetricalaedeagus by flexible joint.Discussion.?Tichocoelidia clarkei Kramer, a Colombian species, isthe lone representative of the genus. The genital structures werefully illustrated with the original description cited above.Salvina MelicharFigures 61-65Salvina Melichar 1926, p. 344. Type of genus Tettigonia dorsisignata Fowler bysubsequent designation of China 1938, p. 184.Description.?With carina separating face and crown. Crown indorsal view wider than long, sharply rounded apically, and producedbeyond eyes. Ocelli near anterior margin of crown but posterior tocarina. Head including eyes slightly narrower than pronotum. An-tennae nearly as long as body including forewings. Posterior marginof pronotum broadly and sharply indented. Venation of forewingsobscure except apically.Ground color yellow to orange with contrasting markings, espe-cially on forewings, of black and brighter hues.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 271Male genitalia: Valve lacking. Male plates separated only api-caliy. Pygofer in lateral view with ventral tooth and appearing cleftat apex due to long mesally curved dorsal hook and weakly sclerotizedventral extension. Anal tube simple but long. Stylar apex in dorsalview curving laterally and caudally. Connective heavy and Y-shaped. Aedeagus simple, compressed and upturned at apex.Discussion.?The type species was illustrated in color whenoriginally described by Fowler (1900, p. 282, tab. 19, fig. 6). Thisillustration is very good for showing the general markings of theleafhopper; however, in the specunen at hand the ground color is abrighter yellow-orange and the stripe on the commissural claval areais of a red-wine shade. This species is Ivnown from three syntypes, allof which are in the British Museum. Through the kindness of Dr.W. E. China, one male specimen was made available for this study.This male with data "Panama, Volcan de Chiriqui 4000 to 6000 feet,Champion" is hereby designated as the lectotype. The male genitaliaof Salvina dorsisignata (Fowler), the only included species, are illus-trated in figures 61-65.Cocoelidia DeLong, new statusFigures 66-74Neocoelidiana subgenus Cocoelidia DeLong 1953, p. 126. Type of subgenusNeocoelidiana antlera DeLong by original designation.Desckiption.?With carina separating face and crown. Crown indorsal view wider than long, bluntly angular apically, and producedbeyond eyes. Ocelli on anterior margin of crown. Head includingeyes narrower than pronotum. Antennae only about half as long asbody including forewings. Posterior margin of pronotum broadlyindented. Venation of forewings obscure except at apex.Ground color sordid yellow to light brown with weakly contrastingyellowish markings on head and dorsum of thorax. Apex of crownwith black spot. Forewings brown hyaline.Male genitalia: Valve lacking. Ventral cover of genital capsulesharply narrowed on distal half appearing as two extended "fingers."Pygofer in lateral view with ventral tooth and dorsal distally hookedprocess. Anal tube with short, forked ventral process. ConnectiveY-shaped with stalk bifurcate to receive aedeagus. Style simple.Aedeagus slender, recurved distally with pair of preapical processes.Discussion.?The Mexican leafhopper, Cocoelidia antlera (De-Long), is the only included species. The male genitalia are illustratedin figures 66-74.
272 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115Nelidina DeLong, new statusFigures 75-80, 110-112Coelidia7ia subgenus Nelidina DeLong 1953, p. 129. Type of subgenus Coelidianadefila DeLong by original designation.Description.-?With carina separating face and crown. Crown indorsal view wider than long, sharply rounded apically, and producedbeyond eyes. Ocelli on anterior margin of crown. Head includingeyes narrower than pronotum. Length of antennae at least half aslong as body. Posterior margin of pronotmn approxmiately straightacross or slightly indented. Venation of forewings most distinctapically.Ground color various shades of yellow without strongly contrastingmarkings.Male genitalia: Valve lacking. Male plates fused basally andwith long apical macrosetae. Pygofer in lateral view with a dorsalprocess or terminal hook. Anal tube simple. Connective verybroadly Y-shaped with stalk bifurcate to receive aedeagus. Stylarapex weakly hooked in lateral view. Aedeagus slender with pairedapical processes.Discussion.?Nelidina includes two species, A^. defila (DeLong)(figs. 75-80) and A^. taeniola, new species, from South America.The type of the genus, A^. defila, is known only from Peru, and A^.taeniola is recorded only from Colombia.Key to Species of NelidinaMALES ONLYLength 7 mm.; aedeagal processes uniformly slender and not twisted (figs. 79, 80).N. defila (DeLong)Length 4.5 mm.; aedeagal processes ribbon-like and twisted (figs. 111,112),N. taeniola, new speciesNelidina taeniola, new speciesFigures 110-112Length.?Male 4.5 mm.Coloration.?Uniformly yellowish or yellowish-green withoutdistinct markings. Exceedingly faint dark longitudinal striping onforewings.Male genitalia: Posterior margin of pygofer beset with numerousfine setae and with small sharp hook or tooth ventrally (fig. 110).Rest of capsule like A^. defila. Aedeagus transverse with long, twisted,ribbon-Uke, pahed apical processes (figs. Ill, 112).HoLOTYPE.?Male, Chic6, Colombia, elevation 2,900 meters,January 2, 1959, R. F. Ruppel. USNM type no. 34882. Paratype,male with same data.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 273Neocoelidiana DeLongFigures 81-86Neocoelidiana DeLong 1953, p. 122. Type of genus Neocoelidia ohscura Bakerby original designation.Description.?With carina separating face and crown. Crown indorsal view wider than long, very broadly or bluntly angled apically,and produced beyond eyes. Head including eyes narrower thanpronotum. Ocelli on anterior margin of crown. Antennae from halfto three-fourths as long as body including forewings. Posteriormargin of pronotum broadly and often sharply indented. Venationof forewings highly obscure except apically.Ground color stramineous to yellow, often with tliree inconspiciouslongitudinal stripes of slightly darker shade on crown and pronotum.Apex of crown with black spot. Forewings yellowish hyaline, attimes with few additional brown marldngs or tinges, with veinsconcolorous.Male genitalia: Valve lacldng. Male plates fused basally formore than half length, without short macrosetae apically. Pygofer inlateral view with dorsal process and either ventral process or ventralhook. Anal tube with single ventral hook. Connective broadlyY-shaped. Apex of style strongly hooked in lateral view. Aedeagusin lateral view approximately S-shaped, often somewhat twisted, andwith some sort of apical modifications.Discussion.?The genus as here defined contains seven species fromwestern United States and Mexico, all of which were treated byDeLong (1953). The male genital structures of the type-species,Neocoelidiana ohscura (Baker), are shown in figm-es 81-86.Coelella DeLong, new statusFigures 87-93Neocoelidiana subgenus Coelella DeLong 1953, p. 125. Type of subgenusNeocoelidia distincta Oman by original designation.Description.?With carina separating face and crown. Crownin dorsal view wider than long, bluntly angled apically, and producedbeyond eyes. Head including eyes narrower than pronotum. Ocellion anterior margin of crown. Antennae from half to three-fourthsas long as body including forewings. Posterior margin of pronotumbroadly and sharply indented. Venation of forewing highly distinct.Ground color stramineous to yellow, usually with three more or lesswell-defined longitudinal stripes of slightly darker hue on crown andpronotum. Apex of crown with black spot. Forewings hyaline withveins uniformly brown.
274 PROCEEDINGS OF THE NATIONAL MUSEUM vol. usMale genitalia. Valve lacking. Male plates fused for basal two-thirds, with short macrosetae apically. Pygofer with both dorsalprocess and ventral hook. Anal tube simple. Connective broadlyY-shaped. Apex of style hooked in lateral view. Aedeagus in lateralview with shaft turned either dorsally or ventrally in apical portion.Aedeagal apex simple or elaborated with short paired processes.Discussion.?The genus contains only two species. The type-species, Coelella distincta (Oman), occurs in the southwestern UnitedStates, while the second species, C. venosa (DeLong) 1953, p. 126, isMexican. The male genital structures of the type-species are illus-trated in figures 87-93. Bisa WalkerFigures 36, 114Biza Walker 1858, p. 253. Type of genus Biza crocea Walker by original designa-tion.Description.?With carina separating face and crown. Crown indorsal view subquadrate, wider than long, produced beyond eyes,rounded apically, and carinated laterally and posteriorly. Ocellion anterior margin of crown. Antennae at least half as long as bodyincluding forewings. Head including eyes narrower than pronotum.Posterior margin of pronotum broadly but rather shallowly indented.Forewings broad with venation distinct.Ground color yellow to orange with extensive brown or fuscousmarkings on the forewings.Male genitalia: Valve lacking. Male plates fused only basally.Pygofer in lateral view with only ventral hook or tooth, no dorsalprocesses, apex acute or rounded. Anal tube and styles simple. Con-nective Y-shaped and clearly articulated with aedeagus. Aedeagussimple, upturned apically, with or without lateral processes.Discussion.?As can be seen by reference to the habitus drawing(fig. 114), the general cercopid-like appearance is striking. Theincluded species are known from Central and South America. Thegenus Biza was recently revised by Kramer (1962).Coelidiana OmanFigures 94-107Coelidiana Oman 1938, p. 397. Type of genus Neocoelidia rubrolineata Bakerby original designation.Acocoelidia DeLong 1953, p. 130. Type of genus Acocoelidia unipunda DeLongby original designation. New synonomy.Description.?With carina separating face and crown. Crown indorsal view variable with length and width subequal or length exceed-ing width, angled apically, and produced beyond eyes. Ocelli onanterior margin of crown. Head including eyes narrower than
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 275pi'onotum. Antennae from half to nearly equal length of body includ-ing forewings. Posterior margin of pronotum broadly and usuallysharply indented. Venation of forewings highly obscure exceptapically.Ground color stramineous to yellow. Head, pronotum, and scutel-lum immaculate or with distinct or inconspicuous yellow-to-redmarkings in form of longitudinal and/or lateral stripes. Apex ofcrown without black spot. Forewings varying from concolorous tomoderately heavily marked with dark brown or black.Male genitaha: Valve lacking. Male plates fused basally andoften for nearly entire length. Pygofer in lateral view quite variable,no true dorsal process, but either terminating with spine of variablelength and development or simple; ventral margin with hook or hooks,long apical spine or simple. Anal tube with paired ventral hooks ornone. Connective Y-shaped. Style in lateral view strongly hookedapically. Aedeagus simple, slender, or moderately stout, and up-turned at apex.Discussion.?The male genitalia of Coelidiana rubrolineata (Baker)and Coelidiana unipuncta (DeLong) are illustrated in figure 99 andfigures 100-107. Members of Coelidiana, as here defined, rangefrom southern Mexico to Brazil. DeLong (1953) treated sevenmembers of this group. C. undata (Linnavuori) was transferred tothis genus by Kramer (1959).Two species of Coelidiana, the types of which have not been studiedsince their original description, are discussed below.Coelidiana coronata (Ball), new combinationNeocoelidia coronata Ball 1916, p. 208.This rather long-crowned species described by Ball was based upona unique female from Guatemala. Its generic placement wUl not becertain until males are available for study. The red markings ofthe dorsum are quite similar to some other Coelidiana.Coelidiana croceata (Osborn), new combinationFigures 94-98Neocoelidia croceata Osborn 1923, p. 78.This Brazilian species is very close to the type-species, C. rubro-lineata, also from Brazil. The only characters that will successfullyallow differentiation are found in the aedeagus. In C. croceata theaedeagus is crenulated ventrally and narrowed apically (fig. 95),whereas in C. rubrolineata the aedeagus is smooth ventrally andbroad apically (fig. 99). The drawings of C. croceata male genitalia(figs. 94-98) are based upon the allotype, which is in the CarnegieMuseum, Pittsburgh, Pennsylvania.
276 PROCEEDINGS OF THE NATIONAL MUSEUM vol. lisLiterature CitedBaker, C. F.1898. Notes on Jassini with some new species. Canadian Ent., vol. 30,pp. 289-292.Ball, E. D.1916. Some new species of Athysanus and related genera (Homoptera).Ent. News, vol. 27, pp. 204-208.Bruner, S. C, and Metcalf, Z. P.1934. A new Bythoscopidae from Costa Rica. Bull. Brooklyn Ent. Soc,vol. 29, pp. 120-124.China, W. E.1938. Melichar's "Monographie der Cicadellinen." Ann. Mag. Nat. Hist.,ser. 11, vol. 2 pp. 182-185.DeLong, D. M.1953. A synopsis of the tribe Neocoelidinii in the Americas (Homoptera-Cicadellidae). Lloydia, vol. 16, no. 2, pp. 93-131.Evans, J. W.1947. A natural classification of leaf-hoppers (Jassoidea, Homoptera), Part3: Jassidae. Trans. Roy. Ent. Soc. London, vol. 98, no. 6, pp.105-271.Fowler, W. W.1900. In Godman and Salvin, Biologia Centrali-Americana, vol. 49 (Hemip-tera-Homoptera, vol. 2), pt. 1, xii+ 339 pp., 21 pis.Gillette, C. P., and Baker, C. F.1895. A preliminary list of the Hemiptera of Colorado. Bull. ColoradoAgric. Exper. Sta., vol. 31, pp. 1-137.Knull, D. J.1942. The genus Neocoelidia in the United States (Homoptera: Cicadel-lidae). American Midi. Nat., vol. 28, no. 3, pp. 680-692.Kraaier, J. P.1959. An elucidation of the Neotropical genus Chinaia with a key to malesand a new allied genus (Homoptera: Cicadellidae: Neocoelidiinae).Proc. Biol. Soc. Washington, vol. 72, pp. 23-32.1961. New Venezuelan leafhoppers of the subfamilies Xestocephalinaeand Neocoehdiinae (Homoptera: CicadeUidae) . Proc. Biol. Soc.Wasliington, vol. 74, pp. 235-240.1962. A synopsis of Biza and a new allied genus (Homoptera: CicadeUidae:Neocoelidiinae). Proc. Biol. Soc. Washington, vol. 75, pp. 101-106.Kramer, J. P., and Linnavuori, R.1959. A new genus and two new species of leafhoppers from South America(Homoptera: Cicadellidae: Neocoehdiinae). Proc. Biol. Soc.Washington, vol. 72, pp. 55-58.Melichar, L.1926. Monographie der CicadeUinen, III. Ann. Mus. Nat. Hungarici,vol. 23, pp. 273-394.Oman, P. W.1938. A generic revision of American Bythoscopinae and South AmericanJassinae. Univ. Kansas Sci. Bull., vol. 24, pp. 343-420.1943. A generic revision of the Nearctic CicadeUidae. Summary of doctoralthesis, George Washington University.1949. The Nearctic leafhoppers (Homoptera: CicadeUidae): A genericclassification and check list. Ent. Soc. Washington Memoirs,no. 3, pp. 1-253.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 277OSBOBN, H.1923. Neotropical Homoptera of the Carnegie Museum: Part 2, Recordsand descriptions of five new genera and sixty-five new species ofthe subfamily Jassinae. Ann. Carnegie Mus., vol. 15, pp. 27-79.Walker, F.1858. List of the specimens of Homopterous insects in the collection of theBritish Museum, suj^pl., pp. 1-307.Young, D. A.1952. Reclassification of Western Hemisphere Typhlocybinae (HomopteraCicadellidae) . Univ. Kansas Sci. Bull., vol. 35, no. 1, pp. 3-217.
278 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115
Figures 1-13. Neocoelidia tumtdifrons Gillette and Baker: 1, lateral view of genital capsule;2, lateral view of aedeagus; 3, ventral aspect of stylar apex; 4, ventral cover of genitalcapsule; 5, ventral view of connective and style; 6, lateral aspect of stylar apex. Neo-coelidia tuherculata (Baker) : 7, lateral view of genital capsule; 8, lateral view of aedeagus;9, ventral cover of genital capsule; 10, ventral view of connective; 11, ventral aspectof stylar apex; 12, dorsal view of apical portion of lower aedeagal shaft; 13, lateralview of style.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 279
Figures 14-24.?Neocoelidia virgata (DeLong): 14, lateral view of genital capsule; 15,lateral view of aedeagus; 16, ventral cover of genital capsule. Neocoelidia pulchellaBall: 17, lateral view of aedeagus; 18, lateral view of genital capsule; 19, lateral view ofstyle; 20, dorsal aspect of stylar apex; 21, lateral aspect of stylar apex; 22, ventral coverof genital capsule; 23, dorsal view of apical portion of lower aedeagal shaft; 24, dorsalview of connective.
craspaFigures 25-36?Neocoelidia fuscodorsata (Fowler): 25, lateral view of genital capsule; 26,lateral view of aedeagus; 27, ventral cover of genital capsule; 28, dorsal view of styleand lateral view of stylar apex. Neocoelidia crenulata Osborn : 29, lateral view of genitalcapsule; 30, ventral cover of genital capsule; 31, lateral view of aedeagus; 32, dorsalview of apical portion of lower aedeagal shaft; 33, lateral aspect of stylar apex; 34,ventral view of connective and style; 35, dorsal view of stylar apex. Biza craspaKramer; 36, ventral cover of genital capsule.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 281
Figures SJ^i.?Chinaia bella Bruner and Metcalf: 37, ventral cover of genital capsule.Xenocoelidia youngi Kramer: 38, ventral cover of genital capsule. Xenocoelidia inflata(Osborn): 39, ventral cover of genital capsule; 40, lateral view of genital capsule; 41,ventral view of connective and style; 42, lateral view of aedeagus; 43, posterior view ofaedeagal apex; 44, lateral view of stylar apex.
282 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115
Figures 45-54.
?
Tozzita ips, new species: 45, lateral view of genital capsule; 46, dorsalview of aedeagal apex; 47, lateral view of connective, style, and aedeagas; 48, ventralcover of genital capsule with pygofers showing; 49, ventral view of aedeagus. Coelanamodesia (Baker): 50, lateral view of genital capsule; 51, lateral view of stylar apex;52, ventral cover of genital capsule with pygofers and anal tube showing; 53, ventralview of connective and style; 54, lateral view of aedeagus.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 283
Figures 55-65.
?
Xiqilliba bellator, new species: 55, lateral view of genital capsule; 56,ventral cover of genital capsule showing processes of anal tube; 57, lateral view ofaedeagal apex; 58, lateral view of aedeagus; 59, lateral view of stylar apex; 60, dorsalview of connective, style, and aedeagus. Salvina dorsisignata (Fowler): 61, ventralcover of genital capsule; 62, lateral view of genital capsule; 63, dorsal view of hooksat apex of pygofer; 64, lateral view of aedeagus; 65, dorsal view of connective and style.
284 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 115
Figures 66-80.
?
Cocoelidia antlera (DeLong): 66, lateral view of genital capsule; 67,posterior view of processes on anal tube; 68, dorsal view of apex of pygofer; 69, ventralview of connective with basal portion of aedeagus; 70, ventral cover of genita capsule;71, lateral view of aedeagal apex; 72, lateral view of aedeagus; 73, posterior view ofaedeagal apex; 74, lateral view of style. Nelidina defila (DeLong): 75, lateral viewof genital capsule; 76, ventral view of stylar apex; 77, ventral view of connective andstyle, also with lateral view of stylar apex; 78, ventral cover of genital capsule withpygofer showing; 79, ventral view of aedeagal apex; 80, lateral view of aedeagus.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 285
Figures 81-93.
?
Neocoelidiana obscura (Baker): 81, lateral view of genital capsule; 82,lateral view of aedeagus; 83, posterior view of aedeagus; 84, posterior view of aedeagalapex; 85, ventral cover of genital capsule; 86, lateral view of style. Coelella distincta(Oman): 87, lateral view of style; 88, lateral view of genital capsule; 89, ventral coverof genital capsule; 90, ventral view of aedeagus; 91, posterioventral view of con-nective; 92, lateral view of aedeagus; 93, lateral view of aedeagus, a variant.
286 PROCEEDINGS OF THE NATIONAL MUSEUM vol. u5
Figures 94-107.
?
Coelidiana croceata (Osborn): 94, lateral view of genital capsule; 95>lateral view of aedeagus; 96, lateral view of stylar apex; 97, ventral view of stylarapex; 98, ventral cover of genital capsule. Coelidiana ruhrolineata (Baker): 99,lateral view of aedeagus. Coelidiana unipuncta (DeLong) : 100, lateral view of genitalcapsule; 101, ventral view of connective; 102, ventral view of abdominal base showingapodemes; 103, ventral cover of genital capsule; 104, lateral view of style; 105, lateralview of aedeagus; 106, ventral view of stylar apex; 107, lateral view of stylar apex.
LEAFHOPPER SUBFAMILY NEOCOELIDIINAE?KRAMER 287
Chinaia lepida Biza craspa114Figures 108-1 14.?Co^/anA drakei, new species: 108, lateral view of genital capsule; 109,lateral view of aedeagus. Nelidina taeniola, new species : 1 10, lateral view of pygofer andanal tube; 111, lateral view of aedeagus; 112, ventral view of aedeagal apex. Chinaialepida Kramer: 113, habitus in dorsal view. Biza craspa Kramer: 114, habitus indorsal view.
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