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Two new species of Hanshumba from Southeastern Brazil and a key to males of the genus (Insecta: Hemiptera: Cicadellidae: Cicadellini)

ABSTRACT

The genus Hanshumba Young, 1977 is recorded from Southeastern and Southern Brazil (Atlantic Forest) and currently includes only three species. Here we describe and illustrate, based mainly on features of the male terminalia, two new species from State of Espírito Santo, Municipality of Santa Teresa: H. setifera sp. nov. and H. teresa sp. nov. The former can be distinguished by the male pygofer and anal tube with large processes bearing numerous setae and the aedeagus with pair of dentiform processes on median portion, whereas the latter has three pairs of longitudinal flanges on the aedeagal shaft. A key to males of the genus is added and its taxonomic status is briefly discussed.

Keywords
Cicadellinae; Leafhopper; Morphology; Sharpshooter; Taxonomy

Introduction

The genus HanshumbaYoung, 1977Young, D.A., 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. Bull. N. Carol. Agric. Exp. Stn 239, vi+1135 pp. is recorded from Southeastern and Southern Brazil (Atlantic Forest) and currently comprises only three species of small and similarly colored sharpshooters (Young, 1977Young, D.A., 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. Bull. N. Carol. Agric. Exp. Stn 239, vi+1135 pp.; Mauro-Barr and Carvalho, 2008Mauro-Barr, T.T., Carvalho, R.A., 2008. Two new species of the Neotropical genus Hanshumba Young from Brazil (Hemiptera: Cicadellidae: Cicadellini). Stud. Neotr. Fau. Envir. 43, 227-235.; Takiya et al., 2018Takiya, D.M., Cavichioli, R.R., Mejdalani, G., Felix, M., Gonçalves, C.C., Camisão, B., Barbosa, J.F., 2018. Cicadellidae in Catálogo Taxonômico da Fauna do Brasil. PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/ (accessed 21.06.18).
http://fauna.jbrj.gov.br/fauna/faunadobr...
): H. brasura Young (type-species), H. similis Mauro-Barr & Carvalho, and H. cavichiolii Mauro-Barr & Carvalho. Young (1977, p. 82)Young, D.A., 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. Bull. N. Carol. Agric. Exp. Stn 239, vi+1135 pp. included Hanshumba in his Dilobopterus generic group. Among the 27 genera assigned to this diverse group, Hanshumba appears to be most similar to Exogonia Melichar, 1926 (Mauro-Barr et al., 2009Mauro-Barr, T.T., Carvalho, R.A., Felix, M., Mejdalani, G., 2009. Taxonomy of the Neotropical leafhopper genus Exogonia with description of seven new Brazilian species (Hemiptera: Cicadellidae: Cicadellini). Zootaxa 2046, 43-68.) and Eldarbala Young, 1977 (Young, 1977Young, D.A., 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. Bull. N. Carol. Agric. Exp. Stn 239, vi+1135 pp.) based on features of the external morphology, color markings, and male terminalia. Hanshumba can be distinguished from those two genera, as well as from the remaining genera of the Neotropical Cicadellini, by the following combination of features: (1) total body length 5-7 mm; (2) dorsum yellow to green mottled with many irregular dark brown vermiculations and spots; (3) head well produced anteriorly and deltoid, without apical thornlike projection; (4) male pygofer well or strongly produced posteriorly, without processes or with peculiar inner process bearing setae, without area of flexion in the middle portion; (5) subgenital plate triangular, gradually tapered toward apex, not extending as far posteriorly as pygofer apex; (6) aedeagus expanded apically; (7) paraphyses present, symmetrical, with one or two pairs of rami.

In this paper, two new species of Hanshumba from Southeastern Brazil (State of Espírito Santo, Municipality of Santa Teresa) are described and illustrated. A new key to males of Hanshumba, based on features of the aedeagus and paraphyses, is provided.

Material and methods

Photographs of the body in dorsal view and of the male terminalia were prepared with CombineZP (free software developed by Alan Hadley), using a digital camera attached to a Zeiss Stemi 2000-C stereomicroscope. Dissected male terminalia parts were stored in small glass vials with glycerin and attached to the pins, below the specimens, as suggested by Young and Beirne (1958)Young, D.A., Beirne, B.P., 1958. A taxonomic revision of the leafhopper genus Flexamia and a new related genus (Homoptera: Cicadellidae). Tech. Bull. U. S. Dep. Agric. 1173, 1-53.. Descriptive terminology adopted herein followed mainly Young (1968Young, D.A., 1968. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 1. Proconiini. Bull. U. S. Nat. Mus. 261, 1-287., 1977Young, D.A., 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. Bull. N. Carol. Agric. Exp. Stn 239, vi+1135 pp., 1986)Young, D.A., 1986. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 3. Old World Cicadellini. Bull. N. Carol. Agric. Exp. Stn 281, v+639 pp., except for the facial areas of the head (Hamilton, 1981Hamilton, K.G.A., 1981. Morphology and evolution of the rhynchotan head (Insecta: Hemiptera, Homoptera). Can. Entomol. 113, 953-974.; Mejdalani, 1993Mejdalani, G., 1993. Morfologia da cabeça de Versigonalia ruficauda (Walker, 1851), com notas sobre a terminologia (Homoptera, Cicadellidae, Cicadellinae). Rev. Bras. Entomol. 37, 279-288., 1998Mejdalani, G., 1998. Morfologia externa dos Cicadellinae (Homoptera, Cicadellidae): comparação entre Versigonalia ruficauda (Walker) (Cicadellini) e Tretogonia cribrata Melichar (Proconiini), com notas sobre outras espécies e análise da terminologia. Rev. Bras. Zool. 15, 451-544.). Studied specimens are deposited at Museu Nacional (MNRJ) - Universidade Federal do Rio de Janeiro (Rio de Janeiro). Label data were quoted exactly with a reversed virgule (\) separating lines on labels.

Results

Hanshumba setifera sp. nov.

(Figs. 1-5)

Figs. 1-5
Hanshumba setifera sp. nov., male. 1, body, dorsal view, length 5.6 mm (antennae and legs not depicted). Terminalia: 2, pygofer lobes, lateral view; 3, subgenital plates, connective, styles, and paraphyses, dorsal view; 4, aedeagus and anal tube, lateroventral view; 5, aedeagus and anal tube, ventral view. APR, anterior ramus of paraphyses; EJB, ejaculatory bulb; LPR, lateral process of aedeagus; PPR, posterior ramus of paraphyses; SPR, setose process.

Diagnosis. Hanshumba setifera sp. nov. can be readily distinguished from the other four known species of the genus by the following combination of features: (1) apical portions of male pygofer (Fig. 2) and segment X (anal tube) (Figs. 4, 5) with conspicuous process bearing numerous setae; (2) style (Fig. 3) with apex obliquely truncate, foot-shaped; (3) aedeagal shaft (Figs. 4, 5) with distal half curved dorsally and with pair of dentiform processes on median portion; (4) paraphyses (Fig. 3) with two pairs of rami, anterior one directed anterad, thick, and obtuse apically, whereas posterior one directed posterad, slender, and acute apically.

Length of male holotype 5.6 mm; male paratype 5.1 mm.

Holotype description

Head (Fig. 1), in dorsal view, well produced anteriorly, deltoid, obtuse apically; median length of crown approximately 7/10 of interocular width and 4/10 of transocular width. Surface of crown convex, without sculpturing or setae, minutely punctate behind apex; frontogenal suture extending onto crown to near ocellus. Ocelli located on imaginary line between anterior eye angles, slightly closer to anterior eye angle than to median line of crown on right side and approximately equidistant on left side. Antennal ledge, in dorsal view, slightly protuberant; in lateral view, with anterior margin slightly oblique and convex. Frons with median portion slightly convex, muscle impressions inconspicuous; epistomal suture obscure medially; clypeus not protuberant, its contour continuing profile of frons.

Thorax (Fig. 1) with pronotal width slightly greater than transocular width of head; pronotum with lateral margins slightly convergent anteriorly, posterior margin slightly emarginate; dorsolateral carena declivous anterad, indistinct at area adjacent to eye; disk transversely rugulose. Mesonotum with scutellum not transversely striate. Forewing with membrane not sharply delimited, including at least apical cells, much of anteapical cells, and costal apical cell; with three closed anteapical cells, their bases proximal to claval apex; with four apical cells, base of fourth more proximal than base of third. Hind wing with vein R2 + 3 incomplete. Hind leg with femoral setal formula 2:1:1; length of first tarsomere approximately equal to combined length of two more distal tarsomeres, with two parallel rows of small setae on plantar surface.

Color (Fig. 1). Ground color of dorsum yellow to green mottled with many irregular dark brown vermiculations and spots. Anterior margin of crown and mesonotum with larger spots. Membrane of forewing translucent, with green spots on anteapical cells. Face, lateral and ventral portions of thorax, and legs mostly yellow; superior portions of antennal ledge and lateral lobe of pronotum with brown spot.

Male terminalia with pygofer (Fig. 2), in lateral view, strongly produced posteriorly; distal half distinctly narrower than basal half; posterior margin convex; posteroventral portion with conspicuous inner process bearing numerous setae, additional processes absent; macrosetae distributed on posterior third and forming ventral group on median third. Subgenital plate (Fig. 3), in ventral view, subtriangular, with basal fourth expanded and then narrowing gradually toward apex; surface with mostly small macrosetae along outer margin, microsetae also present; plates not fused to each other at base; in lateral view, not extending as far posteriorly as pygofer apex. Connective (Fig. 3), in dorsal view, Y-shaped; stalk broad and with median keel. Style (Fig. 3), in dorsal view, extending posteriorly beyond apex of connective; with distinct outer preapical lobe; outer preapical area bearing setae; apex obliquely truncate, foot-shaped. Aedeagus (Figs. 4, 5) symmetrical; shaft, in lateral view, with distal half curved dorsally; gonoduct distinct, gonopore located ventroapically; in ventral view, shaft with pair of small dentiform processes on median portion. Paraphyses (Fig. 3), in dorsal view, with stalk broad, articulated with apex of connective; with two pairs of rami: anterior one directed anterad, thick, obtuse apically and posterior one directed posterad, slender, acute apically. Anal tube (Figs. 4, 5), in lateral view, with segment X bearing conspicuous inner process covered by numerous setae (similar to that of pygofer).

Female unknown.

Material examined. Male holotype: "BR/ES [State of Espírito Santo], Santa Teresa \ Estação Biológica Santa \ Lucia 9-12/XI/2007 \ S. Buys Col." (MNRJ). Paratype, one male: same data as the holotype except "16/XII/2007" (MNRJ).

Known distribution. Atlantic Forest, Southeastern Brazil, State of Espírito Santo (new genus record).

Etymology. The new species name, setifera, refers to the conspicuous setose processes of the male pygofer and anal tube (Figs. 2, 4, 5).

Hanshumba teresa sp. nov.

(Figs. 6-12)

Diagnosis. This new species can be recognized by the following combination of features: (1) apical portion of male pygofer (Figs. 7, 7a) with small inner process bearing apical setae and segment X (anal tube) (Figs. 9, 10) without such process; (2) style (Fig. 8) with apex transversely truncate, not foot-shaped; (3) aedeagal shaft (Figs. 9-12) with three pairs of longitudinal flanges, two of them elongate and forming spiniform processes at apex; (4) paraphyses (Fig. 8) with two pairs of rami, anterior one directed anterad, elongate, slender, and with subacute apex, whereas posterior one directed posterad, fused to each other for most of their length, and with obtuse apex.

Figs. 6-10
Hanshumba teresa sp. nov., male. 6, body, dorsal view, length 5.9 mm (antennae and legs not depicted). Terminalia: 7, pygofer lobe, lateral view (inner setose process illustrated in Fig. 7a); 8, subgenital plates, connective, styles, and paraphyses, dorsal view; 9, aedeagus and anal tube, lateral view; 10, aedeagus and anal tube, ventral view (red arrows indicate the aedeagal shaft). APR, anterior ramus of paraphyses; EJB, ejaculatory bulb; PPR, posterior ramus of paraphyses; SPR, setose process.

Length of male holotype 5.9 mm; male paratype 5.8 mm.

Holotype description

Head (Fig. 6), in dorsal view, well produced anteriorly, deltoid, subacute apically; median length of crown approximately 7/10 of interocular width and 4/10 of transocular width. Surface of crown slightly depressed between apex and ocelli, without sculpturing or setae, minutely punctate behind apex; frontogenal suture extending onto crown to near ocellus. Ocelli located on imaginary line between anterior eye angles, each slightly closer to adjacent anterior eye angle than to median line of crown. Antennal ledge, in dorsal view, slightly protuberant; in lateral view, with anterior margin vertical, concave superiorly and convex inferiorly. Frons with median portion flattened, muscle impressions inconspicuous; epistomal suture obscure medially; clypeus not protuberant, its contour continuing profile of frons.

Thorax (Fig. 6) with pronotal width approximately equal to transocular width of head; pronotum with lateral margins slightly convergent anteriorly, posterior margin emarginate; dorsolateral carena declivous anterad, indistinct at area adjacent to eye; disk transversely rugulose. Mesonotum with scutellum not transversely striate. Forewing with membrane not sharply delimited, including at least apical cells, much of anteapical cells, and costal apical cell; with three closed anteapical cells, their bases proximal to claval apex; with four apical cells, base of fourth more proximal than base of third. Hind wing with vein R2 + 3 incomplete. Hind leg with femoral setal formula 2:1:1; length of first tarsomere approximately equal to combined length of two more distal tarsomeres, with two parallel rows of small setae on plantar surface.

Color (Fig. 6). Ground color of dorsum pale yellow to green mottled with many irregular brown vermiculations and spots. Anterior margin of crown and mesonotum with larger spots. Membrane of forewing translucent, with green spots on anteapical cells. Face, lateral and ventral portions of thorax, and legs mostly yellow; superior portions of antennal ledge and lateral lobe of pronotum with brown spot.

Male terminalia with pygofer (Figs. 7, 7a), in lateral view, strongly produced posteriorly; distal half narrower than basal half; posterior margin subtruncate; posteroventral portion with small inner process bearing apical setae, additional processes absent; macrosetae distributed on posterior third and extending ventroanteriorly onto median third. Subgenital plate (Fig. 8), in ventral view, subtriangular, with basal fourth distinctly expanded and then narrowing gradually toward apex; surface with mostly small macrosetae along outer margin, microsetae also present; plates not fused to each other at base; in lateral view, not extending as far posteriorly as pygofer apex. Connective (Fig. 8), in dorsal view, Y-shaped; stalk with high median keel. Style (Fig. 8), in dorsal view, extending posteriorly beyond apex of connective; with distinct outer preapical lobe; outer preapical area bearing setae; apex transversely truncate, not foot-shaped. Aedeagus (Figs. 9-12) symmetrical; shaft, in lateral view, distinctly expanded apically; in dorsolateral view, with three pairs of longitudinal flanges (ventral one small, median and dorsal ones elongate and forming spiniform processes at apex), borders of flanges minutely serrate; apical portion of shaft membranous; gonoduct distinct, gonopore located ventroapically. Paraphyses (Fig. 8), in dorsal view, with stalk gradually narrowing posterad, articulated with apex of connective; with two pairs of rami: anterior one directed anterad, elongate, slender, with subacute apex and posterior one directed posterad, fused to each other for most of their length, with obtuse apex bearing small keel. Anal tube (Figs. 9-12), in lateral view, with segment X not bearing inner process.

Figs. 11-12
Hanshumba teresa sp. nov., aedeagus and anal tube. 11, lateral view. 12, ventral view. DFL, dorsal flange; MFL, median flange; VFL, ventral flange.

Female unknown.

Material examined. Male holotype: "BR/ES [State of Espírito Santo], Santa Teresa\Estação Biológica Santa\Lucia 9-12/XI/2007\S. Buys Col." (MNRJ). Paratype, one male: same data as the holotype (MNRJ).

Known distribution. Atlantic Forest, Southeastern Brazil, State of Espírito Santo (new genus record).

Etymology. The new species name, teresa (name in apposition), refers to the Municipality of Santa Teresa in the State of Espírito Santo.

Key to males of Hanshumba Young, 1977

1. Paraphyses with two pairs of rami, one directed anterad and another posterad (Figs. 3, 8)........... 2

1ʹ. Paraphyses with single pair of rami directed posterad……………………………………........... 3

2. Aedeagus, in ventral view, with pair of small dentiform processes and without longitudinal flanges (Fig. 5)................................................................ H. setifera sp. nov. [State of Espírito Santo]

2ʹ. Aedeagus, in ventral view, without small dentiform processes and with three pairs of longitudinal flanges (Figs. 10, 12)................................... H. teresa sp. nov. [State of Espírito Santo]

3. Paraphyses rami, in dorsal view, with conspicuous spiniform process directed posterad (Mauro-Barr and Carvalho, 2008Mauro-Barr, T.T., Carvalho, R.A., 2008. Two new species of the Neotropical genus Hanshumba Young from Brazil (Hemiptera: Cicadellidae: Cicadellini). Stud. Neotr. Fau. Envir. 43, 227-235., p. 229, fig. 7)....... H. similisMauro-Barr & Carvalho, 2008Mauro-Barr, T.T., Carvalho, R.A., 2008. Two new species of the Neotropical genus Hanshumba Young from Brazil (Hemiptera: Cicadellidae: Cicadellini). Stud. Neotr. Fau. Envir. 43, 227-235. [State of Rio de Janeiro]

3ʹ. Paraphyses rami, in dorsal view, without such process.................................................................. 4

4. Aedeagal shaft, in lateral view, with constriction delimiting lobe at apical third (Mauro-Barr and Carvalho, 2008Mauro-Barr, T.T., Carvalho, R.A., 2008. Two new species of the Neotropical genus Hanshumba Young from Brazil (Hemiptera: Cicadellidae: Cicadellini). Stud. Neotr. Fau. Envir. 43, 227-235., p. 232, fig. 17)......... H. cavichioliiMauro-Barr & Carvalho, 2008Mauro-Barr, T.T., Carvalho, R.A., 2008. Two new species of the Neotropical genus Hanshumba Young from Brazil (Hemiptera: Cicadellidae: Cicadellini). Stud. Neotr. Fau. Envir. 43, 227-235. [State of Paraná]

4ʹ. Aedeagal shaft, in lateral view, gradually expanded posterad, without distinct constriction (Young, 1977Young, D.A., 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. Bull. N. Carol. Agric. Exp. Stn 239, vi+1135 pp., p. 260, fig. 211f)............................ H. brasuraYoung, 1977Young, D.A., 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella. Bull. N. Carol. Agric. Exp. Stn 239, vi+1135 pp. [State of Santa Catarina]

Discussion

The genus Hanshumba shares peculiar features of the male terminalia with Exogonia and, interestingly, with Ochrostacta Stål, 1869, which is currently assigned to the Proconiini (Young, 1968Young, D.A., 1968. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 1. Proconiini. Bull. U. S. Nat. Mus. 261, 1-287.; McKamey, 2007McKamey, S.H., 2007. Taxonomic catalogue of the leafhoppers (Membracoidea). Part 1. Cicadellinae. Mem. Amer. Entomol. Inst. 78, 1-394.). These features are as follows (Carvalho, 2010Carvalho, R.A., 2010. Análise filogenética dos gêneros neotropicais Exogonia Melichar, Hanshumba Young e Ochrostacta Stål (Insecta: Hemiptera: Cicadellidae). Unpublished D.Sc. dissertation, Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro, xxi+84 pp.): (1) male pygofer and anal tube (segment X) with setose processes (Figs. 2, 4, 5, 7a); (2) aedeagus distinctly expanded apically (Figs. 4, 9, 11); and (3) paraphyses with two pairs of rami (Figs. 3, 8). Preliminary morphological phylogenetic analyses carried out by Carvalho (2010)Carvalho, R.A., 2010. Análise filogenética dos gêneros neotropicais Exogonia Melichar, Hanshumba Young e Ochrostacta Stål (Insecta: Hemiptera: Cicadellidae). Unpublished D.Sc. dissertation, Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro, xxi+84 pp. suggested that the three genera should be synonymized, being Ochrostacta the oldest name. Indeed, Takiya's (2007)Takiya, D.M., 2007. Systematic studies on the leafhopper subfamily Cicadellinae (Hemiptera: Cicadellidae). Unpublished Ph.D. dissertation, University of Illinois, Urbana-Champaign, xvii+166 pp. comprehensive morphological and molecular phylogenetic analyses of the Cicadellinae indicated that Ochrostacta should be transferred to the Cicadellini. Nevertheless, this bold hypothesis of synonymization of these three genera has yet to be carefully evaluated by means of analyses employing both molecular and morphological data. Due to its inflated head, which is quite distinct from those of Hanshumba and Exogonia, Ochrostacta is among the most peculiar genera of the Cicadellinae (Young, 1968Young, D.A., 1968. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 1. Proconiini. Bull. U. S. Nat. Mus. 261, 1-287.; see images in Wilson et al., 2009Wilson, M.R., Turner, J., McKamey, S.H., 2009. Sharpshooter leafhoppers of the world (Hemiptera: Cicadellidae subfamily Cicadellinae), http://naturalhistory.museumwales.ac.uk/sharpshooters/home.php (accessed 23.06.18).
http://naturalhistory.museumwales.ac.uk/...
).

Carvalho's (2010)Carvalho, R.A., 2010. Análise filogenética dos gêneros neotropicais Exogonia Melichar, Hanshumba Young e Ochrostacta Stål (Insecta: Hemiptera: Cicadellidae). Unpublished D.Sc. dissertation, Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro, xxi+84 pp. study suggested a monophyletic group including the previously known Hanshumba species (H. brasura, H. cavichiolii, and H. similis). This clade had the herein described H. setifera sp. nov. as its sister group. The clade comprising these four species was supported by three unambiguous synapomorphies related to the characteristic yellow to green dorsum mottled with many irregular dark brown vermiculations and spots (Fig. 1). This color pattern is also found in H. teresa sp. nov. (Fig. 6), which is thus a putative member of the aforementioned monophyletic group. Due to the very similar color pattern of known Hanshumba species, it is often difficult to associate males with females. The discovery of our two new species shows once again how the insect fauna from the Brazilian Atlantic forest is still poorly known.

Acknowledgements

The manuscript benefited from the useful comments of Victor Quintas (MNRJ) and two anonymous reviewers. Gabriel Mejdalani and Rodney R. Cavichioli are research productivity fellows from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, processes 303627/2014-0 and 305484/2014-1, respectively). Dr. João Roberto Spotti Lopes (Escola Superior de Agricultura "Luiz de Queiroz" - ESALQ) is the advisor of Joyce Froza and his laboratory provided financial support for her visits to the Museu Nacional. Dr. Roberto Antonio Zucchi and Dr. Sinval Silveira Neto (ESALQ) kindly provided access to laboratory facilities. The first author has a stipend from Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) in connection with her D.Sc. studies. The specimens described here were collected by Dr. Sandor Buys.

References

  • Carvalho, R.A., 2010. Análise filogenética dos gêneros neotropicais Exogonia Melichar, Hanshumba Young e Ochrostacta Stål (Insecta: Hemiptera: Cicadellidae). Unpublished D.Sc. dissertation, Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro, xxi+84 pp.
  • Hamilton, K.G.A., 1981. Morphology and evolution of the rhynchotan head (Insecta: Hemiptera, Homoptera). Can. Entomol. 113, 953-974.
  • Mauro-Barr, T.T., Carvalho, R.A., 2008. Two new species of the Neotropical genus Hanshumba Young from Brazil (Hemiptera: Cicadellidae: Cicadellini). Stud. Neotr. Fau. Envir. 43, 227-235.
  • Mauro-Barr, T.T., Carvalho, R.A., Felix, M., Mejdalani, G., 2009. Taxonomy of the Neotropical leafhopper genus Exogonia with description of seven new Brazilian species (Hemiptera: Cicadellidae: Cicadellini). Zootaxa 2046, 43-68.
  • McKamey, S.H., 2007. Taxonomic catalogue of the leafhoppers (Membracoidea). Part 1. Cicadellinae. Mem. Amer. Entomol. Inst. 78, 1-394.
  • Mejdalani, G., 1993. Morfologia da cabeça de Versigonalia ruficauda (Walker, 1851), com notas sobre a terminologia (Homoptera, Cicadellidae, Cicadellinae). Rev. Bras. Entomol. 37, 279-288.
  • Mejdalani, G., 1998. Morfologia externa dos Cicadellinae (Homoptera, Cicadellidae): comparação entre Versigonalia ruficauda (Walker) (Cicadellini) e Tretogonia cribrata Melichar (Proconiini), com notas sobre outras espécies e análise da terminologia. Rev. Bras. Zool. 15, 451-544.
  • Takiya, D.M., 2007. Systematic studies on the leafhopper subfamily Cicadellinae (Hemiptera: Cicadellidae). Unpublished Ph.D. dissertation, University of Illinois, Urbana-Champaign, xvii+166 pp.
  • Takiya, D.M., Cavichioli, R.R., Mejdalani, G., Felix, M., Gonçalves, C.C., Camisão, B., Barbosa, J.F., 2018. Cicadellidae in Catálogo Taxonômico da Fauna do Brasil. PNUD, http://fauna.jbrj.gov.br/fauna/faunadobrasil/ (accessed 21.06.18).
    » http://fauna.jbrj.gov.br/fauna/faunadobrasil/
  • Wilson, M.R., Turner, J., McKamey, S.H., 2009. Sharpshooter leafhoppers of the world (Hemiptera: Cicadellidae subfamily Cicadellinae), http://naturalhistory.museumwales.ac.uk/sharpshooters/home.php (accessed 23.06.18).
    » http://naturalhistory.museumwales.ac.uk/sharpshooters/home.php
  • Young, D.A., 1968. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 1. Proconiini. Bull. U. S. Nat. Mus. 261, 1-287.
  • Young, D.A., 1977. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New World Cicadellini and the genus Cicadella Bull. N. Carol. Agric. Exp. Stn 239, vi+1135 pp.
  • Young, D.A., 1986. Taxonomic study of the Cicadellinae (Homoptera: Cicadellidae). Part 3. Old World Cicadellini. Bull. N. Carol. Agric. Exp. Stn 281, v+639 pp.
  • Young, D.A., Beirne, B.P., 1958. A taxonomic revision of the leafhopper genus Flexamia and a new related genus (Homoptera: Cicadellidae). Tech. Bull. U. S. Dep. Agric. 1173, 1-53.

Edited by

Associate Editor: Daniela Takiya

Publication Dates

  • Publication in this collection
    Oct-Dec 2018

History

  • Received
    24 July 2018
  • Accepted
    29 Aug 2018
  • Published
    10 Sept 2018
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