Research Article |
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Corresponding author: Luísa Alasmar ( lualasmar@ufpr.br ) Academic editor: Lara-Sophie Dey
© 2025 Luísa Alasmar, Alexandre Cruz Domahovski, Rodney Ramiro Cavichioli.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
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In a morphological-based analysis using implied weighting for 23 taxa and 50 characters, we provided a phylogenetic study for Ciminius. The genus was recovered as monophyletic with a high support. We also provided a taxonomic revision, with description of three new species, increasing the diversity of Ciminius from seven to ten species: the Neotropical C. albolineatus, C. callosa, C. platensis, C. yana, C. autumnalis sp. nov., C. dissidens sp. nov., C. sesamum sp. nov., and the Nearctic C. hartii, C. sidanus, and C. taosus. A new monotypic genus, Arcanus gen. nov., recovered as sister to Ciminius is described. Tylozygus Fieber, the new genus, and Ciminius were recovered forming a clade here nominated as Articulate Stem Clade, due to its unique connective morphology amongst all Cicadellini. The immature stages of Ciminius sp. are described and a key to males, an occurrence map, and notes of parasitism and submacroptery are provided. We discussed the Ciminius relationships according to our analysis, the genus distribution, and the taxonomic problems about C. callosa.
Auchenorrhyncha, Cicadellini, Neotropical region, Nearctic region, new taxa, sharpshooters
The genus Ciminius was erected by Metcalf and Bruner in 1936 to accommodate the species Tettigonia hartii Ball, 1901 recorded from Cuba and United States. Afterward,
Throughout history, the species Tettigonia hartii has had its name misspelled in some publications, including the author himself, in the original description of C. sidanus and C. taosus work (
Ciminius is composed of robust rather small sharpshooters, with coloration varying from pale yellow to black, and is abundant in grassland fields. According to
Cicadellini representatives are considered as potential vectors of Xylella fastidiosa (
Despite the abundance of Ciminius specimens in grasses (Poaceae), the last taxonomic study was realized in 1977. The strong similarity between Ciminius species and the occurrence of intraspecific variations, both in male genitalia and overall coloration aspects, indicates the need for a detailed study and revision of the genus. In this present work, we proposed the genus revision, and a phylogenetic analysis based on morphology. We also described three new Brazilian species and the immature stages of Ciminius sp., as long as redescriptions and illustrations of the valid species. A new genus Arcanus gen. nov. is described, based on two specimens from Southern Brazil. A brief discussion involving C. callosa and their similarities with C. platensis, C. albolineatus, and C. yana is provided. In addition, notes of parasitism and submacroptery are reported, and an occurrence map and a key to males of Ciminius are provided.
In this study, we examined 624 pinned adult specimens of Ciminius from the
UColeção Entomológica Pe. Jesus Santiago Moure, Universidade Federal do Paraná, Curitiba, Brazil (
All the described Ciminius and three additional new species from Brazil were included in the phylogenetic analysis, except for C. callosa, known only by two females (holotype and paratype) and an undissected male (allotype) from Bahia State, Brazil (Table
Specimens included in the phylogenetic analysis. Number of analyzed specimens, sex, and type material are provided for each taxon. PT = paratype, HT = holotype, * = specimens studied throughout photography.
| Taxon | Specimens | Type | ||
| ♂ | ♀ | ♂ | ♀ | |
| Tapajosa ocellata (Osborn) | 1 | 1 | — | — |
| Cicadella viridis Linneaus | 1 | 1 | — | — |
| Chlorogonalia coeruleovittata (Signoret) | 1 | 1 | — | — |
| Plesiommata corniculata Young | 1 | 1 | — | — |
| Plesiommata mollicella Fowler | 1 | 1 | — | — |
| Rotigonalia larissae Cavichioli | — | — | 1 (PT) | 1 (PT) |
| Rotigonalia olivacea Cavichioli | — | — | 1 (PT) | 1 (PT) |
| Segonalia machadoi Cavichioli and Takyia | — | — | 1 (PT) | 1 (PT) |
| Segonalia steinbachi Young | 1 | 1 | — | — |
| Syncharina argentina (Berg) | 1 | 1 | — | — |
| Syncharina punctatissima (Signoret) | 1 | 1 | — | — |
| Tylozygus fasciatus (Walker) | 1 | 1 | — | — |
| Tylozygus geometricus (Signoret) | 1 | 1 | — | — |
| Arcanus gen. nov. | — | — | 1 (HT), 1 (PT) | — |
| Ciminius albolineatus (Taschenberg) | 1 | 1 | — | — |
| Ciminius platensis (Berg) | 1 | 1 | — | — |
| Ciminius yana Young | 1 | 1 | — | 1(HT)* |
| Ciminius autumnalis sp. nov. | — | — | 1 (HT), 11 (PT) | 5 (PT) |
| Ciminius dissidens sp. nov. | — | — | 1 (HT), 44 (PT) | 51 (PT) |
| Ciminius sesamum sp. nov. | — | — | 1 (HT), 50 (PT) | 15 (PT) |
| Ciminius hartii (Ball) | 3 | 3 | — | 1(HT)* |
| Ciminius taosus (Ball) | 1 | 1 | — | 1(HT)* |
| Ciminius sidanus (Ball) | 3 | 3 | — | 1(HT)* |
For the taxa descriptions and redescriptions, the terminology adopted follows mainly
For genitalia dissections, the technique follows
The characters coding was based on direct observations of adult specimens of males and females, coded as binary or multistate, following the homology proposition as suggested by
All characters were treated as nonadditive (
The occurrence map was performed with the software QGis v. 3.34.9 “Prizren” (https://qgis.org/) with a World shapefile, provided by World Food Programme, (https://public.opendatasoft.com/explore/dataset/world-administrative-boundaries), cropped in the study area. The quoted data were based on
Tettigonia hartii Ball, 1901: 6, by original designation.
Small sharpshooters, from 3.2 to 5.7 mm in length. Overall coloration pale-yellow (Figs
Ciminius albolineatus, male. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Coloration
: Overall coloration (Figs
Ciminius albolineatus, female. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G median portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
Ciminius hartii, male. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Ciminius hartii, female. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G median portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
Ciminius platensis, male. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Ciminius platensis, female. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G median portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
Ciminius specimens have resemblances in the male genitalia to Arcanus gen. nov. and Tylozygus, especially in the morphology of the connective, but Ciminius can be promptly differentiated from them and from all the Cicadellini by the forewing presenting only two anteapical cells, with the outer one opened basally, and with a plexus of veins anteapically. Amongst all the New World Cicadellini, only Ciminius and Hadria Metcalf and Bruner, 1936 presents two anteapical cells, but in the latter, both are opened basally. In addition, Hadria specimens present a bright colorful aspect and their distribution is restricted to Cuba, Haiti and Dominican Republic (
The distribution is based on
C. albolineatus (Taschenberg, 1884). Argentina, Brazil (Goiás [new record], Mato Grosso [new record], Paraná [new record], Rondônia [new record], Roraima [new record], São Paulo), Bolívia, Cuba, El Salvador, French Guiana, Isle of Pines, Panama, Venezuela.
C. autumnalis sp. nov. Brazil (Paraná).
C. callosa (Osborn, 1926). Bolivia.
C. dissidens sp. nov. Brazil (Paraná).
C. hartii (Ball, 1901). Cuba, Mexico, United States (Florida, Georgia, Illinois, Indiana, Kansas, Louisiana, Mississippi, Missouri, New Mexico, Ohio, Oklahoma, Pennsylvania, Tennessee, Texas, Virginia).
C. platensis (Berg, 1879). Argentina, Brazil (Bahia [new record], Ceará [new record], Espírito Santo [new record], Mato Grosso do Sul [new record], Minas Gerais [new record], Paraná [new record], Pernambuco [new record], Piauí [new record], Rio de Janeiro, Rio Grande do Sul [new record], São Paulo), Paraguay, Peru, Venezuela.
C. sesamum sp. nov. Brazil (Paraná).
C. sidanus (Ball, 1936). Mexico, United States (Arizona).
C. taosus (Ball, 1936). United States (New Mexico).
C. yana Young, 1977. Argentina, Brazil (Mato Grosso do Sul, Paraná [new record], Rondônia, São Paulo [new record]).
Tettigonia albolineata Taschenberg, 1884: 446.
Ciminius albolineatus Young, 1977: 593.
Body pale yellow (Figs
Measurements
: Total length: males (n = 20): 3.6–4.4 mm, females (n = 20): 4.0–4.6 mm. — Crown: (Figs
BRAZIL – Mato Grosso • 3♂♂, 1♀; Chapada; Nov. 1963; M. Alvarenga leg.; DZUP • 1♂, 1♀; Chapada dos Guimarães; 19 Nov. 1983; Exc. Dep. Zool. UFPR leg.; Polonoroeste; DZUP • 1♀; Cáceres; 3 Apr. 1985; C. Elias leg.; Polonoroeste; DZUP • 1♀; Diamantino; 16 Feb. 1965; S. Laroca leg.; DZUP • 16♂♂, 12♀♀; Nova Ubiratã, E.S.E.C. Rio Ronuro; 13.1122°S, 54.4436°W; 330 m; light trap; 11–16 Jun. 2017; R.R. Cavichioli & A.C. Domahovski leg.; DZUP • 1♀; same collection data as for preceding; areia rio; DZUP • 15♂♂, 22♀♀; Novo Mundo, Parque Estadual do Cristalino; 09.4517°S, 55.8396°W; 240 m; light trap; 21–25 Jun. 2017; R.R. Cavichioli & A.C. Domahovski leg.; DZUP • 1♀; same collection data as for preceding; sweep; DZUP • 1♀; 11 km south of Rio Verde; 5 Nov. 2003; Mielke & Casagrande leg.; DZUP – Roraima • 1♂; Tepequém, Amajari; 820 m; 14–15 Jul. 2009; O.M. Mielke & M.M. Casagrande leg.; DZUP. – Rondônia • 8♂♂, 6♀♀; Guaporé; 12°16’05”S, 60°42’30”W; light trap; 23 Apr. 2006; J.A. Rafael & F.F. Xavier F. leg.; DZUP. – Paraná • 1♂, 1♀; Foz do Iguaçu; 11 Dec. 1966; Exc. Dept. Zool. leg.; DZUP. • 1♂; Foz do Iguaçu; 3 Dec. 1966; Exc. Dept. Zool. leg.; DZUP • 2♂♂; Tibagi, Parque Estadual do Guartelá; 1000 m; 24°33’47”S, 50°15’26”W; sweep; 21–24 Feb. 2017; A.C. Domahovski leg.; DZUP • 6♂♂; Tibagi, Parque Estadual do Guartelá; 980 m; 24°33’42”S, 50°13’31”W; 11–16 Jan. 2024; A. Paladini, L. Alasmar & A.C. Domahovski leg.; DZUP • 5♂; Curitiba, Centro Politécnico, UFPR; 20–25 Jan. 2022; sweep; A.C. Domahovski & L. Alasmar leg.; DZUP • 1♂, 2♀♀; Curitiba, Centro Politécnico, UFPR; 25°26’49”S, 49°13’54”W; 925 m; 22–25 Nov. 2022; A.C. Domahovski leg.; DZUP • 3♂♂; Curitiba, Centro Politécnico, UFPR; Apr. 2023; L. Alasmar leg.; DZUP • 15♂♂, 6♀♀; Ponta Grossa, Parque Estadual de Vila Velha; 25.247579°S, 49.992188°W; 930m; sweep; 14 Dec. 2022; A.P. Pinto, A.C. Domahovski, L. Alasmar, J. Ehlert & L.P. Aguiar leg.; DZUP. São Paulo • 1♂, 1♀; São José do Barreiro, Parque Nacional da Serra da Bocaina, near lodging; 22.733757°S, 44.616643°W; 1522 m; sweep; 9–11 Jan. 2023; A.P. Pinto, A.C. Domahovski, J. Ehlert & L.P. Aguiar leg.; DZUP.
Specimens of C. albolineatus and C. platensis are the most abundant species amongst all studied specimens, and have external resemblances to each other, but they can be differentiated by the male genitalia characters presented in the former as: aedeagus with ventral margin lacking processes and paraphysis curved ventrally, with ventral margin bearing a preapical constriction. The specimens of C. albolineatus collected in “Centro Politécnico, UFPR, Curitiba, Paraná” and “Parque Estadual Vila Velha, Ponta Grossa, Paraná” presented a great number of males and a few females with black coloration. Even when specimens are black, they present the forewings with veins paler, unlike the other black specimens of Ciminius, like C. hartii, C. dissidens sp. nov.. males, and C. sesamum sp. nov. which present the veins mostly concolor with the membrane. Until the present moment, this species presents the only occurrence of black females for the genus.
Cicadella callosa Osborn, 1926: 179.
Ciminius callosa Young, 1977: 593.
Overall coloration (Fig.
Coloration
: Overall coloration (Fig.
Holotype : BOLIVIA • ♀; Puerto Suarez; 150 m; J. Steinbach leg.; CMNH-IZ 724,504; CMNZ.
Cicadella callosa was described based on a female holotype analyzed herein, and a female paratype, both from Bolivia. In the original description, a male allotype was designed, from Bom Fin, probably misspelled from the municipality of Bonfim, in Salvador, Bahia State, Brazil.
Tettigonia hartii Ball, 1901: 61.
Ciminius hartii Metcalf and Bruner, 1936: 944.
Coloration of males (Fig.
Measurements
: Total length: males (n = 3) 3.9–4.1 mm; females (n = 3) 5.1–5.3 mm. — Crown: (Figs
Lectotype (Fig.
UNITED STATES OF AMERICA – Florida • 1♂, 1♀; Hudson; 13 Jul. 1939; Oman leg.; USNM • 1♂; Dunellon 7 Oct. 1938; Oman leg.; USNM • 1♂; New Port Ritchey; 7. Oct. 1938; Oman leg.; USNM • 1♀; Sof Picnic; 8 Oct. 1938; Oman leg.; USNM • 1♀; Sanford; 22 July. 1939; Oman leg.; USNM.
Ciminius hartii resembles externally to C. yana, C. sesamum sp. nov., and males of C. dissidens sp. nov.., but differs from them by the aedeagus not presenting any lobated process on dorsal margin nor serrated processes on ventral margin. Otherwise, the paraphysis is rectilinear, unlikely in C. yana and C. dissidens sp. nov.. Ciminius hartii is the most abundant species amongst Nearctic Ciminius reported in literature and in museum collections.
Acocephalus dubius Berg, 1879: 259.
Tettigonia dubia nomen nudum Berg, 1884: 162.
Tettigonia platensis Berg, 1884: 162.
Ciminius platensis Young, 1977: 593.
Coloration pale-yellow (Figs
Measurements
: Total length: males (n = 20) 4.0–4.6 mm, females (n = 20) 4.5–5.5 mm. — Crown: (Figs
BRAZIL – Rio Grande do Sul • 1♂; Areias Brancas, Torres; Fev.1983; K. Zanol leg.; DZUP • 3♀♀; Águas Belas, Viamão, 3–4 Mar.1980; K. Zanol leg.; DZUP • 1♂; Santa Maria, UFSM; 29°43’02.2”S, 53°43’07”W; 08 Fev. 2023; C.V. Rodrigues, F.D. Souza, G. Flores leg.; DZUP. – Paraná • 6♂, 6♀ Curitiba, Mar. 1983, Cavichioli leg.; DZUP • 2♂♂, 2♀♀; same collection data as for preceding; Sakakibara leg.; DZUP • 1♂, 7♀♀ same collection data as for preceding; A.M.S. e R.R.C. leg.; DZUP • 1♂, 1♀ same collection data as for preceding; Mar. 1982; Sakakibara leg.; DZUP • 1♂; Piraquara, Mananciais da Serra; 25°29’47”S, 48°58’54”W; 1021 m; 20 Feb. 2022; A.P. Pinto, J. Ehlert & L. Polizeli leg.; DZUP • 2♂♂, 2♀♀; Curitiba, Centro Politécnico, UFPR, grasslands fields near Biológicas Departament; 20–25 Jan. 2022; Sweep; A.C. Domahovski & L. Alasmar; DZUP • 2♂♂, 2♀♀; Diamante do Norte, E.E. Caiuá; 14 Dec. 2012; M. Savaris & S. Lampert leg.; Luiz de Queiroz trap; DZUP • 1♂, 7♀♀; Umuarama; Feb. 1986, A.F. Yamamoto leg.; DZUP • 1♂, 1♀; Porecatu; 20 Oct. 1970, Becker-Hatschback leg.; DZUP • 1♂, 9♀; Ponta Grossa, P.E. Vila Velha; 25.247579°S, 49.992188°W; 930 m; 14 Dec. 2022, A.P. Pinto, A.C. Domahovski, L. Alasmar, J. Ehlert & L.P. Aguiar leg.; Sweep; DZUP • 8♂♂, 1♀; São José dos Pinhais; 25°36’18’’S, 49°11’37’’W; 880 m; 06 May 2023; A.C. Domahovski leg.; sweep; DZUP • 3♂♂, 1♀; Tuneiras do Oeste, REBIO das Perobas; 01–04 APr. 2024; A.C. Domahovski & L. Alasmar; sweep; DZUP. – Espírito Santo • 1♂; Linhares, Res. Vale Rio Doce, main station; 19.0905°S, 40.0410°W; 56 m; 07 May 2007; J.A. Rafael & F.F. Xavier F. leg.; light; DZUP • 1♂, 3♀♀; Linhares; Sep. 1972; M. Alvarenga, leg.; DZUP. – Minas Gerais • 2♂♂, 4♀♀; Águas Vermelhas; Dec. 1983; M. Alvarenga leg.; DZUP • 2♂♂, 12♀♀; São Gonçalo Rio Abaixo, Estação Ambiental Peti-Cemig; 15 Apr. 2013; 19°53’02”S, 43°22’21”W; A. Lima, A.F. Kumagai & P. Dias leg.; light; DZUP • 1♀; Lavras, Campus UFLA, 16–18 Sep. 2013, ~905 m, 21°13’50”S, 44°58’32”W, M.N. Morales leg.; Malaise; DZUP. – Mato Grosso do Sul • 3♂♂, 5♀♀; Porto Murtinho; 07 Dec. 2012; M. Savaris & S. Lampert leg.; light trap; DZUP. – Alagoas • 1♂ Jacaré dos Homens; 9.662725°S, 37.233245°W, Jun. 2024, S. Cajé & J. Duarte-De-Mélo leg; light; DZUP. – Bahia • 1♂, 11♀♀ Encruzilhada; Nov. 1974; 980 m; Alvarenga leg.; DZUP • 4♀♀; same collection data as for preceding; Dec. 1980; DZUP • 4♀♀; Senhor do Bonfim, Serra Santana; 10.23223°S, 40.1159°W; 520 m; 15 May 2007; J.A. Rafael & F.F. Xavier F., light; DZUP. – Ceará • 35♂♂, 27♀♀; Cratéus, RPPN Serra das Almas; 5.11°S, 40.87°W; 320 m; 18 Apr. 2014; Melo & Rosa; light trap; DZUP • 2♂♂, 1♀; same collection data as for preceding; 22 May 2014; DZUP • 3♂♂, 18♀♀; same collection data as for preceding; 4.8107°S, 38.9740°W; 220 m; 15 May 2014; DZUP • 2♂♂; Ubajara, PN de Ubajara, Park entry; 3°50’18,00”S, 40°53’54,00”W; 849 m; 23 Apr. 2012; Câmara, J.T. leg.; light trap; DZUP. – Paraíba • 4♂♂, 1♀; Mamanguape, Reserva Biológica Guaribas; 6.7421°S, 35.1426°W; 13–16 May 2023, A. Martins & G. Celante leg.; light trap; DZUP • 2♂♂, 5♀♀; same collection data as for preceding; sweep; DZUP • 1♂, 5♀♀; Pocinhos, Caatinga Km230, 7.1437°S, 36.1357°W; 15 May 2023; A.L. Martins leg.; sweep; DZUP. – Pernambuco • 3♂♂, 10♀; Afrânio; 08.3153°S, 41.0259°W; 550 m; 16 May 2007; J.A. Rafael & F.F. Xavier F. leg.; Manual; DZUP. – Piauí: 2♂♂, 9♀♀; Ubajara; 03°50’77”S, 40°53’53”W; 846 m; 22 Apr. 2012, R.R. Cavichioli leg.; light; DZUP • 1♀; Coronel José Dias, P.N. Serra da Capivara; 590 m; 08°44’3.89”S, 42°30’10.56”W; 08–12 Mar. 2016, R.M. Feitosa, G.P. Camacho & F.O. Marfins leg.; DZUP • 1♂; Piracuruca, PN de Sete Cidades, dormitory; 4°5’57,00”S, 41°42’34,00”W; 193 m; 18–19 Apr. 2012; Takiya, D.M. leg.; light; DZUP. – São Paulo • 5♂♂, 1♀; Marília; 22.18927°S, 49.92630°W; 18–20 Nov. 2022; L. Alasmar leg.; sweep; DZUP • 2♂♂ same collection data as for preceding; 22.19384°S, 49.92172°W; DZUP • 2♂♂; same collection data as for preceding; Condomínio Green Valley; 22.16342°S, 50.00271°W; 07 Jan. 2023; DZUP.
Ciminius platensis is a common species collected with sweeping net in open field grasses, and along with Ciminius autumnalis sp. nov. and C. sidanus, do not present male black specimens known. Ciminius platensis can be differentiated from the latter securely due to the aedeagus with few and large teeth ventrally and paraphysis rectilinear and short.
Cicadella sidana Ball, 1936: 21.
Ciminius sidanus DeLong and Knull, 1946a: 17.
Coloration pale-yellow (Figs
Ciminius sidanus, male. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Measurements
: Total length: males (n = 3) 3.8–4.0 mm; females (n = 3) 4.6–5.0 mm. — Head and thorax: Crown (Figs
Ciminius sidanus, female. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G median portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
Holotype
(Fig.
Other material. UNITED STATES OF AMERICA – Arizona • 2♂♂, 2♀♀; Altar Valley; 17 Oct. 1937; Oman leg.; USNM • 1♂, 1♀; Baboquivari; 8 Sep. 1935; E. D. Ball leg.; USNM.
Ciminius sidanus presents a unique coloration in Ciminius, which is responsible for its easy recognition. Otherwise, the male genitalia assemble with C. hartii differentiating in a few features, as the presence of rugosities basally on aedeagus ventral margin and aedeagus apex expanded, while in C. sidanus the rugosities are restricted to preapical portion and the aedeagus apex is not expanded in lateral view. Although the separation of these two species is possible using the coloration and male genitalia, the similarities between them are strong, which suggest the necessity of a future molecular analysis to confirm their status as separate species.
Cicadella taosa Ball, 1936: 20.
Ciminius sidanus DeLong & Knull, 1946a: 17.
Overall coloration (Figs
Ciminius taosus, male. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Measurements
: Total length: males (n = 1) 4.7 mm; females (n = 1) 5.7 mm. — Head and Thorax: Crown (Figs
Ciminius taosus, female. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G edian portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
Holotype
(Fig.
UNITED STATES OF AMERICA – New Mexico • 1♂; Belen; 20 Jul. 1936; R. H. Beamer leg; USNM • 1♀ same data collection as for preceding; D. R. Lindsay leg; USNM.
Ciminius taosus is the largest member of Ciminius. The males of C. taosus can be differentiated in coloration from the other black species by the forewings with yellow longitudinal veins, fading towards apex, and the crown and frons contrasting yellow and black marks. In dorsal view (Figs
Ciminius yana Young, 1977: 593.
Coloration pale-yellow (Fig.
Measurements
: Total length: males (n = 11) 3.2–3.7 mm, females (n = 6) 3.9–4.3 mm. — Head and Thorax: Crown (Figs
Ciminius yana, male A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Ciminius yana, female. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G median portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
Holotype
(Fig.
BRAZIL – Paraná • 1♂; Antonina, Res. [Reserva] Rio Cachoeira; 25.316°S, 48.696°W; 50 m; 23–27 Jan. 2017; A.C. Domahovski leg.; DZUP • 1♀ same data collection as for preceding; Entomologia UFPR leg.; suspended light trap; DZUP • 2♂♂; Tuneiras do Oeste, REBIO das Perobas; 01–04 Apr. 2024; A. C. Domahovski & L. Alasmar, leg.; sweep; DZUP. – Mato Grosso do Sul • 1♂ Barranco Branco; 22 Dec. 1925; Souto Maior leg.; DZUP. – São Paulo • 1♂; Marília; 22.19384°S, 49.92172°W; 20 Nov. 2022; L. Alasmar leg.; sweep; DZUP • 2♂♂, 2♀♀; same data collection as for preceding; 18–20 Nov. 2022; DZUP • 1♂; same data collection as for preceding; 22.18927°S, 49.92630°W; 26 Dec. 2022; DZUP • 6♂♂, 3♀♀; same data collection as for preceding; Condomínio Green Valley; 22.16342°S, 50.00271°W; 07 Jan. 2023; DZUP.
The black males of C. yana resemble Ciminius dissidens sp. nov.. and Ciminius sesamum sp. nov. but can be distinguished due to the distinct yellow markings on claval suture and basis of R+M and C veins. The pale-yellow to brown males are more similar to C. albolineatus and C. platensis. Furthermore, males of C. yana can be distinguished from other congeners by the conspicuous lobate process on the aedeagus dorsal margin. From all the studied material, a unique black specimen of C. yana presented forewings venation that was completely yellow.
Coloration pale-yellow (Figs
Ciminius autumnalis sp. nov., male holotype. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Measurements
: Total length: holotype male 4.0 mm; paratypes, males (n = 11) 4.0–4.3 mm; females (n = 5) 4.5–4.63 mm. — Head and Thorax: Crown (Figs
Ciminius autumnalis sp. nov., female paratype. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G median portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
Holotype : BRAZIL – Paraná • ♂; Ponta Grossa, Parque Estadual de Vila Velha; 11 Apr. 2015; A.C. Domahovski leg; sweep; DZUP. — Paratypes: BRAZIL – Paraná • 3♂♂, 1♀; same data collection as for holotype; DZUP • 2♂♂ same data collection as for holotype; 24 Mar. 2016; B. Rosa & C. Yamakawa leg.; Moericke; DZUP • 4♂♂, 1♀; Tibagi, P.E. Guartelá; 11–16 Jan. 2024; 24°33’42”S, 50°13’31”W; 980 m; A. Paladini, L. Alasmar & A. C. Domahovski leg.; DZUP • 1♂, 1♀; same data collection as for preceding; DZRJ • 1♂, 1♀; same data collection as for preceding MNRJ • 1♂, 1♀; same data collection as for preceding USNM.
The epithet “autumnalis” means “from autumn” in Latim, and refers to the specimens’ coloration, reminding autumn leaves. This is a noun in apposition.
Ciminius autumnalis sp. nov. resembles C. albolineatus and C. platensis due to their size and overall coloration. Ciminius autumnalis sp. nov. can be differentiated from the other Ciminius species by the following male genitalia aspects: the aedeagus dorsal margin distinctly rounded dorsally, ventral margin serrated medially and rectilinear subapically; and paraphysis rectilinae, directed dorsally, with a distinct preapical constriction ventrally. The specimens of C. autumnalis sp. nov. presents variations in the aedeagus. In seven specimens studied, there were variations in shaft roundness, which can be strongly or moderately rounded dorsally, and in basiventral margin, which can be rectilinear or with a reentrance. The most distinct patterns were provided in the illustration (Fig.
Coloration black (males) (Fig.
Ciminius dissidens sp. nov., male holotype. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Measurements
: Total length: holotype male 3.9 mm; paratypes, males (n = 19) 3.8–4.2 mm; females (n = 20) 4.6–5.2 mm. — Head and Thorax: Crown (Figs
Ciminius dissidens sp. nov., female paratype. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G median portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
The epithet “dissidens” means “different” in Latim. It refers to the reddish coloration of females that can also occur in males, and is different from any other color pattern known in Ciminius. This is a noun in apposition.
Holotype : BRAZIL – Paraná • ♂; Ponta Grossa, Parque Estadual de Vila Velha; 11 Apr. 2015; A.C. Domahovski leg.; sweep; DZUP. — Paratypes: BRAZIL – Paraná • 2♂♂, 4♀♀; same data collection as for holotype; DZUP; 1♂; Ponta Grossa, Parque Estadual Vila Velha; 25 Nov. 2011; 965 m; 25°13’49’’S, 49°59’65’’W; A.C. Domahovski leg.; sweep; DZUP • 1♂; same collection data as for preceding, 24 Mar. 2016; B. Rosa & C. Yamakawa; Moericke; DZUP • 1♂ same data as preceding; 21 Nov. 2015; DZUP • 9♂♂, 17♀♀ same data collection as for preceding; 25.247579°S, 49.992188°W; 930 m; 14 Dec. 2022; A. P. Pinto, A. C. \ Domahovski, L. Alasmar, \ J. Ehlert & L. P. Aguiar leg, sweep; DZUP • 10♂♂, 10♀♀; same collection data as for preceding; DZRJ • 10♂♂, 10♀♀; same collection data as for preceding; MNRJ • 5♂♂, 5♀♀; same collection data as for preceding; USNM • 5♂♂, 5♀♀; same collection data as for preceding; MZUSP.
The black males of Ciminius dissidens sp. nov. resemble C. hartii, C. yana, C. sesamum sp. nov. and the black male specimens of C. albolineatus, due to their coloration and size. Although Ciminius dissidens sp. nov. tends to present more distinct yellow areas on the pronotum and head, and reddish portions laterally, different from Ciminius sesamum sp. nov. and C. yana. In addition, the forewing veins are never yellow, differing from C. albolineatus. Ciminius dissidens sp. nov. males can be distinguished from the other Ciminius species by the distinct wide aspect of the aedeagus in lateral view, with serrated processes along the entire ventral margin. The females of C. dissidens sp. nov., stand out for their distinct reddish forewing coloration. In a total of 44 males of C. dissidens sp. nov. studied, 40 males were black and only four were reddish.
Coloration pale-yellow (Figs
Ciminius sesamum sp. nov., male holotype. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Measurements
: Total length: holotype male 3.6 mm; paratypes, males (n = 19) 3.4–3.8 mm; paratypes, females (n = 20) 3.7–4.1 mm. — Head and Thorax: Crown (Figs
Ciminius sesamum sp. nov., female paratype. A distal portion of abdomen, ventral view; B distal portion of abdomen, lateral view; C sternite VII, ventral view; D first valvifer and first valvula, lateral view; E apical portion of first valvula; F second valvula, lateral view; G median portion of second valvula; H apical portion of second valvula; I second valvifer and gonoplac, lateral view; J apical portion of gonoplac. Scale bars in mm.
Holotype: BRAZIL – Paraná • ♂; São José dos Pinhais; 25°36’18’’S, 49°11’37’’W; 880 m; 08–22 Apr. 2017; A.C. Domahovski leg.; sweep; DZUP. — Paratypes: BRAZIL – Paraná • 11♂♂, 1♀; same collection data as for holotype; DZUP • 3♂♂ same collection data as for holotype; MNRJ • 1♂, 1♀; same collection data as for holotype; 07–21 Jan. 2017; DZUP • 1♂; same collection data as for preceding; USNM • 1♂,1♀; same collection data as for preceding; 25–28 Feb. 2017; DZUP • 3♂♂; same collection data as for preceding; USNM • 1♂; same collection data as for preceding; 23 Mar. 2017; DZUP • 5♂♂, 2♀♀; same collection data as for preceding; 02–05 Mar. 2019; DZUP • 1♂, 2♀♀; same collection data as for preceding; MNRJ • 4♂♂, 1♀; same collection data as for preceding; DZRJ • 2♂♂; same collection data as for preceding; 01–31 Mar. 2021; DZUP • 1♂; same collection data as for preceding; 01–30 Nov. 2022; USNM • 1♀; same collection data as for preceding; DZUP • 10♂♂, 3♀♀; Curitiba, Centro Politécnico, UFPR, grasslands near to Biológicas department; 20–25 Jan. 2022; A.C. Domahovski & L. Alasmar leg.; sweep; DZUP • 2♂♂ same collection data as for preceding; MNRJ • 1♂; same collection data as for preceding; Dec. 2022; DZUP • 2♂♂, 1♀; same collection data as for preceding; DZRJ • 1♂, 2♀♀; same collection data as for preceding; USNM.
The word “sesamum” is from Latin and means “of sesame”, an allusion to the overall aspect of this species: small, robust, and blackened or yellowed, resembling sesame seeds. This is a noun in apposition.
Ciminius sesamum sp. nov. resemble externally to other black males of the genus: C. hartii, C. yana, C. albolineatus, and C. dissidens sp. nov., however, C. sesamum sp. nov. have a more homogeneous black aspect, with indistinct or faint yellow marks along the crown and pronotum. The only constant yellow mark is in the posterior half of the mesonotum. In the aedeagus, C. sesamum sp. nov. can be distinguished from C. yana due to absence of a lobate distinct process on dorsal margin, distinguished from C. hartii and C. albolineatus due to the aedeagus bearing the ventral margin serrated, and from C. dissidens sp. nov., due the aedeagus not strongly widened in lateral view. In the locality of Centro Politécnico, mostly females presented a submacroptery condition (Fig.
|
1 Aedeagus ventral margin without processes (Fig. |
2 |
|
1’ Aedeagus ventral margin with serrated processes (Fig. |
6 |
|
2 Aedeagus dorsal margin with a conspicuous lobate process preapically (Fig. |
C. yana Young, 1977 |
|
2’ Aedeagus dorsal margin without those processes (Fig. |
3 |
|
3 Paraphysis rectilinear (Fig. |
4 |
|
3’ Paraphysis curved (Fig. |
5 |
|
4 Aedeagus shaft expanded on dorsal and ventral margins, ventral margin with rugosities basally, apex expanded (Fig. |
C. hartii (Ball, 1901) |
|
4’ Aedeagus shaft expanded on dorsal and ventral margins, ventral margin without or with rugosities preapically, apex not expanded (Fig. |
C. sidanus (Ball, 1936) |
|
5 Aedeagus ventral margin excavated basally (Fig. |
C. albolineatus (Taschenberg, 1884) |
|
5’ Aedeagus ventral margin not excavated basally (Fig. |
C. taosus (Ball, 1936) |
|
6 Aedeagus with serrated processes along entire length of ventral margin (Fig. |
7 |
|
6’ Aedeagus with serrated processes restricted to apical third of ventral margin (Fig. |
8 |
|
7 Aedeagus, in lateral view, conspicuously broadened (Fig. |
C. dissidens sp. nov. |
|
7’ Aedeagus, in lateral view, slender, apical third expanded, in a hood-like structure (Fig. |
C. sesamum sp. nov. |
|
8 Aedeagus ventral margin apical third with two or three large serrated processes (Fig. |
C. platensis (Berg, 1879) |
| Aedeagus ventral margin apical third with numerous small serrated processes (Fig. |
C. autumnalis sp. nov. |
Coloration reddish yellow (Figs
Head (Fig.
Overall coloration (Figs
Head (Fig.
Overall pale-yellow coloration more homogeneous (Figs
Head (Fig.
Overall coloration (Figs
Head and thorax (Fig.
Overall coloration (Figs
Head (Fig.
Female (Fig.
Male (Fig.
Remarks. The first and second stages (Fig.
The third instar presents the macula and the paler areas on crown and pronotum very distinct (Fig.
The fourth and fifth stages are very similar in overall coloration (Fig.
All the immature specimens were collected with the sweep method and the resulting material was stored with the following precedence label: “Curitiba, Paraná, Brazil, UFPR – Centro Politécnico – Estacionamento, varredura, XII.2022”. The Ciminius nymphs present the particularity of showing, even at first instar, some of the maculae present in the adults, as the maculae on pronotum and mesonotum, allowing them to an easy recognition at genus level. As the nymph develops, it is possible to note the reddish coloration of abdomen turning into yellow, which in adults turns into an even more darkened coloration. Some specimens collected, especially in fourth and fifth instars, presented a darkened overall coloration, which could possibly mean to be a black adult. However, in the collection site, there are occurrence of two species, C. albolineatus and C. sesamum sp. nov., both presenting adults with black overall coloration and therefore was not possible to identify the nymphs species without usage of molecular methods or rearing nymphs, until the emergence of adults.
Arcanus academicus sp. nov.
Small sharpshooters, size about 5.5 mm with overall coloration (Fig.
Arcanus academicus gen. et sp. nov., male holotype. A habitus, dorsal view; B habitus, lateral view; C head, frontal view; D pygofer, valve and subgenital plate, lateral view; E valve and subgenital plates, ventral view; F connective and style, ventral view; G aedeagus, lateral view; H aedeagus, caudoventral view; I connective, stem and paraphysis, lateral view; J paraphysis, dorsal view. Scale bars in mm.
Head and Thorax
: Crown (Fig.
Unknown.
The genus name Arcanus (a neutral noun) is derived from Latin, meaning “mysterious” and “secret.” This name alludes to the difficulty of collecting specimens and their remarkable, unique morphological characteristics.
Arcanus gen. nov. shares similarities with Ciminius and Tylozygus, especially in the male terminalia, such as: (1) pygofer moderately produced, with macrosetae distributed along the entire disk, (2) subgenital plates triangular, short, not extending posteriorly to pygofer basal third (3) styles short with apex truncate (4) stem of connective occurring as a separated sclerite. Although, the new genus can be promptly differentiated from Ciminius and Tylozygus by the combination of characters below: (1) body somewhat flattened dorsoventrally; (2) head moderately produced, with anterior margin subtriangular; (3) forewings with three anteapical cells, median one opened basally; and (4) paraphysis large and well sclerotized. Until the present moment, Arcanus gen. nov. is known only for its type location.
Overall coloration (Fig.
Measurements
: Holotype male 5.28 mm; paratype male 5.65 mm. — Coloration: Crown (Fig.
Holotype : BRAZIL – Paraná • ♂; Curitiba, Centro Politécnico, Biológicas parking; 25°26’49”S, 49°13’54”W; 925 m; Dec. 2022; AC Domahovski & L. Alasmar leg.; DZUP. — Paratype: BRAZIL – Paraná • 1♂; same data as holotype; DZUP.
The epithet “academicus” is from Latin and means “academic”, an allusion to the type locality of the species, Universidade Federal do Paraná.
Figure
The data matrix (Supplementary material, Table S1) consisted of 23 taxa and 50 characters, being two of overall coloration, four from head, seven from thorax, 24 from male genitalia and 13 from female genitalia. From these characters, two were coded as multistate and 48 as binaries. In the characters listed below, we provided to each one the number of steps (L), consistency index (ci) and retention index (ri). The monophyly of Ciminius was strongly supported, using the implied weighting, with a default K = 3 (SR = 93). Our matrix produced only one tree for implied weighting, with 111 steps, consistence index = 50 and retention index = 79, that will be the hypothesis discussed (Fig.
1. Overall coloration: (0) green (Fig.
Characters map, states indicated at left bottom square. A–D habitus in dorsal view: A Tylozygus geometricus; B Ciminius platensis; C Ciminius hartii; D Arcanus academicus sp. et gen. nov. E–H Crown, pronotum and mesonotum, dorsal view: E Ciminius platensis; F Chlorogonalia coeruleovittata; G Plesiommata mollicella; H Ciminius albolineatus. I–L Head, frontal view: I Segonalia steinbachi; J Ciminius platensis; K Syncharina punctatissima; L Rotigonalia olivacea. M–P Crown and pronotum, dorsal view: M Ciminius hartii; N Syncharina punctatissima; O Rotigonalia olivacea; P Ciminius platensis. Scales in mm.
2. Overall coloration, polymorphism, presence: (0) absent (Figs
3. Crown, black rounded maculae medially, presence: (0) absent (Fig.
4. Crown, posterior margin, pair of dark spots behind ocelli, presence: (0) absent (Fig.
5. Frons, muscular impressions, coloration aspect: (0) same as frons (Fig.
6. Face, clypeus, shape in males: (0) flattened (Fig.
7. Pronotum, texture: (1) strigate (Fig.
8. Pronotum, coloration: (0) uniform (Fig.
9. Forewing, anteapical cells, number: (0) two (Fig.
Characters map, states indicated at left bottom square. Arrows indicating the character in specimen. A–J Habitus, lateral view: A Ciminius platensis; B Arcanus academicus sp. et gen. nov.; C, F Syncharina punctatissima; D, I Plesiommata mollicella; E, G Cicadella viridis; H, J Ciminius albolineatus; K detail of accessory veins. Scales in mm.
10. Forewing, inner anteapical cell, aspect of base: (0) closed (Fig.
11. Forewing, median anteapical cell, aspect of base: (0) closed (Fig.
12. Forewing, outer anteapical cell, aspect of base: (0) closed (Fig.
13. Forewing, R1 vein, plexus of anteapical veins, presence: (0) absent (Fig.
14. Abdomen, sternite II, inner apodemes, presence: (0) absent (Fig.
Male abdomen characters map, states indicated at left bottom square. Arrows indicating the character position in specimen. A–D Abdomen, basal portion in ventral view: A Segonalia steinbachi; B, D Tylozygus geometricus; C Ciminius hartii; E–J Pygofer in lateral view E Ciminius hartii; F Segonalia steinbachi; G Syncharina punctatissima; H Ciminius hartii; I Ciminius platensis; J Plesiommata mollicella; K, L Subgenital plates, ventral view: K Ciminius hartii; L Plesiommata mollicella.
15. Abdomen, sternite II, inner apodemes, aspect: (0) short (Fig.
16. Male pygofer, posterior margin, aspect: (0) rounded (Fig.
17. Male pygofer, macrosetae, distribution: (0) restricted to apical third (Fig.
18. Subgenital plates, length: (0) not attaining pygofer apex (Fig.
19. Subgenital plates, shape: (0) gradually tapering towards apex (Fig.
20. Connective, arms, aspect: (0) U-shaped (Fig.
Male abdomen characters map, states indicated at left bottom square. Arrows indicating the character position in specimen. A–D connective and styles, dorsal view. A Ciminius sidanus. B Chlorogonalia coeruleovittata. C Plesiommata mollicella. D Tylozygus geometricus. E, F connective, paraphysis and aedeagus, lateral view. E Rotigonalia olivacea. F Ciminius platensis. G, H connective and stem, dorsal view. G Rotigonalia olivacea. H Ciminius platensis. I, J internal structures of male genitália. I Plesiommata mollicella. J Ciminius platensis K–N paraphysis, lateral and dorsal view, respectively: K, L Ciminius sesamum sp. nov.; M, N Tylozygus geometricus.
21. Stem of connective, length: (0) shorter than style length (Fig.
22. Stem of connective, aspect: (0) not keeled (Fig.
23. Stem of connective, articulation with connective arms, presence: (0) absent (Fig.
24. Paraphysis or paraphyses, presence: (0) absent (Fig.
25. Paraphysis or paraphyses, number of rami: (0) one (Fig.
26. Paraphysis or paraphyses, arm-ramus/rami articulation, presence: (0) absent (Fig.
Paraphysis and anal tube characters map, states indicated at left bottom square. Arrows indicating the character position in specimen. A–H paraphysis, lateral view: A Rotigonalia olivacea; B Ciminius yana; C Ciminius sesamum sp. nov.; D Arcanus academicus sp. et gen. nov.; E Ciminius sidanus; F Ciminius autumnalis sp. nov.; G Ciminius hartii; H Ciminius dissidens sp. nov. I, J Paraphysis, dorsal view: I Ciminius platensis; J Ciminius hartii. K, L Aedeagus and anal tube, lateral view: K Tylozygus geometricus; L Ciminius sidanus.
27. Paraphysis or paraphyses, ramus/rami, aspect: (0) slender (Fig.
28. Paraphysis or paraphyses, ventral margin, preapical constriction, presence: (0) absent (Fig.
29. Paraphysis or paraphyses, ramus or rami, curvature: (0) rectilinear (Fig.
30. Paraphysis or paraphyses, ramus or rami in dorsal view, apical third, windings, presence: (0) absent (Fig.
31. Aedeagus – anal tube connection, lobated membrane, presence: (0) absent (Fig.
32. Aedeagus, basidorsal margin, aspect: (0) not expanded (Fig.
Aedeagus characters map, states indicated at left bottom square. Arrows indicating the character position in specimen: A–F aedeagus, lateral view; A Rotigonalia olivacea; B Ciminius sesamum sp. nov.; C Ciminius yana; D Ciminius dissidens sp. nov.; E Ciminius hartii; F Segonalia steinbachi. G–H Aedeagus apex, ventral view: G Segonalia steinbachi; H Rotigonalia olivacea. I–L Aedeagus, lateral view: I Ciminius sidanus; J Ciminius platensis; K Ciminius taosus; L Rotigonalia olivacea.
33. Aedeagus, ventral margin, serrated processes, presence: (0) absent (Fig.
34. Aedeagus, apical process/processes, presence: (0) absent (Fig.
35. Aedeagus, apical process/processes, number: (0) one (Fig.
36. Aedeagus, apical third, aspect: (0) continuing or slenderer than shaft contour (Fig.
37. Aedeagus, apex, shape: (0) rounded (Fig.
38. Female sternite VII, proportion: (0) as long as wide (Fig.
Female abdomen characters map, states indicated at left bottom square. Arrows indicating the character position in specimen: A, B pygofer, ventral view: A Ciminius platensis; B Rotigonalia olivacea. C, D Pygofer, lateral view: C Ciminius platensis; D Syncharina punctatissima. E–H First valvifer and valvula I basal portion, lateral view: E Chlorogonalia coeruleovittata F Ciminius yana; G Ciminius platensis; H Tylozygus geometricus. I–L Valvula II apex, lateral view: I Ciminius platensis; J Tylozygus geometricus; K Ciminius platensis; L Syncharina punctatissima. Scales in mm.
39. Female pygofer, apex, aspect: (0) rounded (Fig.
40. Valvifer I, shape: (0) suboval (Fig.
41. Valvula I of ovipositor, truncated process, pre basally, presence: (0) absent (Fig.
42. Valvula I of ovipositor, apical portion, aspect: (0) gradually narrowed towards apex (Fig.
43. Valvula I of ovipositor, dorsal margin sculpting, shape: (0) strigate (Fig.
44. Valvula II of ovipositor, expansion after basal curvature, aspect: (0) slight (Fig.
Female valvula II characters map, states indicated at left bottom square. Arrows indicating the character position in the specimen. A, B Valvula II basal portion, lateral view: A Rotigonalia olivacea; B Ciminius platensis. C, D Valvula II apical portion, lateral view: C Ciminius platensis; D Syncharina punctatissima; E–H Valvula II apex, lateral view: E Ciminius platensis; F Syncharina argentina; G Ciminius albolineatus; H Rotigonalia olivacea. I–N Valvula II median portion of blade, denticles details, lateral view: I Ciminius sesamum; J Rotigonalia olivacea; K Ciminius autumnalis sp. nov.; L Ciminius sesamum sp. nov.; M Ciminius hartii; N Chlorogonalia coeruleovittata.
45. Valvula II of ovipositor, blade apex, direction: (0) horizontal (Fig.
46. Valvula II of ovipositor, apex, aspect: (0) broad (Fig.
47. Valvula II of ovipositor, preapical prominence on ventral margin, presence: (0) absent (Fig.
48. Valvula II of ovipositor, primary teeth anterior margin, denticles, presence: (0) absent (Fig.
49. Valvula II of ovipositor, gap between primary teeth, presence: (0) absent (Fig.
50. Valvula II of ovipositor, primary teeth at median portion, shape: (0) subtriangular (Fig.
Figures
The species Ciminius hartii has its distribution in both Nearctic and Neotropical regions. Amongst the Nearctic species, Ciminius hartii is the most abundant, appearing in several studies, checklists, museum catalogs, and in online platforms. They have a spread distribution in the south region of the United States, while the other Nearctic species, C. sidanus and C. taosus occur only in some states, the latter being recorded only from Texas (Fig.
The two most abundant Neotropical species, based on the examined material, belong to C. albolineatus and C. platensis, showing a wide distribution in Brazil. According to our results, there is a lack of Ciminius occurrence in Brazil’s Northern region, which can be explained by the low amount of material samples from this area (Fig.
Figure
Nymphs and adults of Cicadellidae are reportedly attacked by Dryinidae, Encyrtidae (Hymenoptera), Pipunculidae (Diptera), Halictophagidae and Helenchidae (Strepsiptera) (
Figure
A total of 133 females of Ciminius sesamum sp. nov. were studied, of which 85 were submacropterous (~64%), being the first record to the genus. Interestingly, we found no male presenting this feature, or in any other species of Ciminius. Submacroptery seems to be a rare condition amongst Cicadellinae and was only reported in Oragua Young, 1977 and Teleogonia Melichar, 1925 for the Cicadellini tribe and in Splonia Signoret, 1851 for Proconiini. Submacroptery by definition is when the forewing is somewhat shortened, exposing genital capsule up to one or two segments, with appendix reduced or absent (
In our phylogenetic analysis, all genera compounding the outgroup were recovered with strong support: Syncharina (SR = 93), Plesiommata (SR = 99), Rotigonalia (SR = 76), Segonalia (SR = 98) and Tylozygus (SR = 97). Ciminius was recovered as monophyletic (SR = 94), supported by three unambiguous synapomorphies: the forewings with two anteapical cells (9-1) (Fig.
Our analysis recovered Arcanus academicus gen. et sp. nov. as sister of Ciminius, with a high support (SR = 84), with one synapomorphy: the crown bearing a pair of black spots behind the ocelli (4-1) (Figs
Tylozygus was recovered as sister of the clade Ciminius + Arcanus gen. nov., with moderate support (SR = 51), supported by three unambiguous synapomorphies: stem of connective keeled (22-1) (Fig.
The Ciminius internal relationships were mostly defined by few characters of the male genitalia, once all the species have external morphology strongly conserved, resulting in clades recovered with low support. The species of Ciminius can be divided into two groups, the paraphyletic group “Without Processes”, due to the sister group relationship of C. taosus to the other Ciminius, and also composed by the clade C. sidanus + (C. hartii + (C. albolineatus + C. yana)), which share the ventral margin of aedeagus smooth, and the monophyletic group “Serrated Processes”, due to the ventral margin of aedeagus with serrated processes, composed of Ciminius dissidens sp. nov. + (C. autumnalis sp. nov. + (C. platensis + C. sesamum sp. nov.)). The group “Without Processes” was recovered with very low support (SR = 6), sharing one homoplasy: the paraphysis in dorsal view with windings in apical portion (30-1) (Fig.
Ciminius is collected in open areas composed of grasslands with a predominance of native grasses (Poaceae) (Fig.
Within the family Cicadellidae, some aspects of sexual dimorphism are known, as size and brachyptery, the latter mostly associated with females, as reported herein. The sexual dimorphisms involving coloration are mostly known for Typhlocybinae and Eurymelinae subfamilies (
Ciminius callosa is a problematic species because the original description consists in coloration, external morphology and only external genitalia of male and female, with uninformative illustrations: the head, pronotum and mesonotum in dorsal view and the genital capsules in ventral views. This species was described based on two females, holotype (Fig.
Although Ciminius species can be readily differentiated through morphology, a future study based on molecular data may contribute new discoveries, especially about the inner relationships that were recovered here with low support. Due to the wide distribution of the genus, further studies using biogeographic methods should be made to understand the Ciminius origin, whether it is Nearctic or Neotropical.
Fundings: This work was supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) fellowships from the senior author (Processes 131208/2021-7 and 141245/2023-9). ACD is post-doctoral fellowship from Fundação Carlos Chagas Filho de Amparo à Pesquisa do Rio de Janeiro (FAPERJ, Proc. E-26/204.206/2021).
Author Contributions: Conceptualization: LA and RRC. Methodology: LA, ACD and RRC. Software: LA and ACD. Validation: LA, ACD and RRC. Formal analysis: LA and ACD. Investigation: LA and RRC. Resources: RRC. Data Curation: LA. Writing – Original draft: LA. Writing – Review and Editing: ACD and RRC. Visualization: LA. Supervision: ACD and RRC. Project administration: LA, ACD and RRC. Funding Acquisition: LA and RRC.
The authors have declared that no competing interests exist.
We are very thankful to Vanessa Verdecia (CMNH) that kindly provided us the image of Ciminius callosa type, and to James Zahnizer that not only provided us the types images of C. hartii, C. sidanus and C. taosus, but kindly loaned us representants of these species. We also thank Andressa Paladini (UFPR), Beatiz Camisão (UFRJ), and Jádila S. Prando (UFRJ) who reviewed the first version of this manuscript, suggesting improvements.
Table S1
Data type: .xlsx
Explanation notes: Data matrix for phylogenetic analysis.