Research Article |
Corresponding author: Gabriel Mejdalani ( mejdalani@mn.ufrj.br ) Academic editor: Bruno Clarkson
© 2023 Nathalia H. Pecly, Daniela M. Takiya, Rodney R. Cavichioli, Gabriel Mejdalani.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
|
The genus Dasmeusa is distributed in Northern and Northeastern Brazil and the Guianas. Until the present study, six species were recognized within Dasmeusa. The species of this genus are very similar in terms of color and external morphology, being distinguished mainly by the male terminalia. Here, we review and redescribe Dasmeusa and its species, describe four new species, and present the first phylogenetic analysis of the genus, including 40 morphological characters and 15 terminal taxa. Dasmeusa flavescens Metcalf and Erythrogonia bicolor Metcalf are considered junior synonyms of the type-species, Dasmeusa pauperata (Fabricius). Scanning electron microscopy was employed for a detailed study of the integument of the type-species, including sensilla, surface sculpturing, brochosomes, organ of Evans, and other structures. The phylogenetic analysis with equal weights resulted in nine most parsimonious trees. The implied weighting method resulted in two trees, both with the same ingroup topology as observed in one of the nine equal-weights trees. This preferred topology is as follows: ((D. basseti (D. mendica (D. rafaeli sp. nov., D. falcifera sp. nov.))) (D. isabellina (D. oriximina sp. nov. (D. pauperata (D. imperialis, D. dinizi sp. nov.))))). Dasmeusa was recovered as monophyletic in all trees, being supported by five apomorphic characters.
Cicadellinae, cladistics, morphology, Neotropical region, SEM, sensilla
Melichar produced an extensive monograph on the taxonomy of the Cicadellinae (“Monographie der Cicadellinen”), which was published, posthumously, between 1924 and 1951 (
Only in 1977, in the detailed revision of the New World Cicadellini published by Young, Dasmeusa was formally described, including characters of the external morphology, male and female terminalia, a key to species, and a new species (D. mendica Young, 1977). More than twenty years later,
In his revision of the Cicadellini,
Based on raw morphological similarities,
Although
In the present paper, we redescribe the genus Dasmeusa and describe four new species. All previously known species are redescribed, including new diagnostic morphological characters (with the exception of D. imperialis because it was recently described). Dasmeusa flavescens Metcalf, 1955 and Erythrogonia bicolor Metcalf, 1949 are herein considered junior synonyms of the type-species Dasmeusa pauperata (Fabricius, 1803). We also provide an identification key to males, a list of the valid species, and a map showing their distribution. Morphological data are employed to investigate the phylogenetic relationships among the species of Dasmeusa; our outgroups include six genera of the Paromenia group. Scanning electron microscopy (SEM) was employed for a detailed study of the integument of the type-species, including sensilla, surface sculpturing, brochosomes, organ of Evans, and various other structures.
The studied specimens belong to the following institutions: Coleção Entomológica Pe. Jesus Santiago Moure, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba (
The morphological terminology adopted here followed mainly
Ap = appendix; Br = brochosomes; Cl = clypeus; Cs = claval sulcus; Cx = coxa; DEN = denticle; DSA = dorsal sculptured area; DUC = duct; Fe = femur; Fl = flagellum; Fr = frons; Fw = forewing; Ge = gena; La = labium; Lb = labrum; Lg = laterotergite; Me = mesonotum; Mi = microtrichia; Ms = maxillary stylet; Mu = muscle impression; OE = organ of Evans; Om = ommatidium; Pe = pedicel; PPR = preapical prominence; RAM = ramus; Sb = sensilla basiconica; Sc = s. coeloconica; Scp = scape; Se = setae; Sp = s. placodea; Spi = spiracle; St = s. trichodea; TOO = tooth; Tr = trochanter; VID = ventral interlocking device; VLI = valvifer I; VSA = ventral sculptured area.
AM = Amazonas; AP = Amapá; BA = Bahia; RR = Roraima; SE = Sergipe.
The external morphology of D. pauperata was studied using scanning electron microscopy (SEM). Our descriptions of microstructures and sensilla followed mainly
The ingroup consisted of nine species of Dasmeusa. Due to difficulties of associating females to accurately identified males of five species of the genus, females were not included in the phylogenetic analysis, unfortunately. Our outgroup choice was based on the Paromenia generic group proposed by
Taxa included in the phylogenetic analysis of Dasmeusa and outgroups. The number of males examined, their collections, and geographical distribution are provided for each species.
Species | Specimens | Collection | Country (state, province, or department) |
Paromenia auroguttata | 1 |
|
Brazil (Rio de Janeiro) |
Onega bracteata | 1 |
|
Ecuador (Napo) |
Jozima leucopa | 2 |
|
Brazil (Amazonas) |
Sailerana solitaris | 2 |
|
Brazil (Amazonas) |
Tacora johanni | 2 |
|
Brazil (Roraima) |
Jeepiulus flavus | 3 |
|
Brazil (Mato Grosso) |
Dasmeusa basseti | 3 |
|
French Guiana (Laussat, Montagne des Chevaux), Guyana (Mabura Hill) |
Dasmeusa dinizi | 3 |
|
French Guiana (Laussat, Montagne des Chevaux) |
Dasmeusa falcifera | 3 |
|
French Guiana (Laussat, Montagne des Chevaux) |
Dasmeusa imperialis | 3 |
|
Brazil (Amazonas) |
Dasmeusa isabellina | 2 |
|
Brazil (Pará) |
Dasmeusa mendica | 3 |
|
French Guiana (Montagne des Chevaux, Réserve Naturelle de la Trinité) |
Dasmeusa oriximina | 1 |
|
Brazil (Pará) |
Dasmeusa pauperata | 3 |
|
Brazil (Amazonas, Bahia, Roraima) |
Dasmeusa rafaeli | 3 |
|
Brazil (Amazonas) |
Morphological characters of the head, thorax, and male terminalia were identified based on their topographical identity before proposing hypotheses of primary homology, i.e., character state identity (
The implicit enumeration algorithm of TNT was used for estimating most parsimonious trees (
Cicada pauperata Fabricius, by subsequent designation of
Specimens preserved in collections usually pale yellow; whitish-yellow to greenish-yellow in life; forewing with preapical area with irregular orange transverse band or with second apical cell with distinct red spot. Head moderately to strongly produced anteriorly; coronal suture distinct, elongate, extending anteriorly beyond interocellar line; frons, in lateral view, with inferior third slightly angulate. Pronotum with lateral margins convergent anteriorly; posterior margin rectilinear. Forewing subhyaline; apex slightly expanded and obliquely truncate; with four apical cells, base of fourth approximately aligned with base of third; costal apical cell broadened posteriorly. Male terminalia with pygofer bearing basiventral lobe; without processes; subgenital plate triangular, not fused basally to its counterpart; style without preapical lobe; aedeagus with shaft short, usually with single process, rarely with pair of processes; paraphyses present, biramous, with or without processes on stalk. Female terminalia with sternite VII well produced posteriorly; pygofer well produced posteriorly; valvula I abruptly narrowed apically, with ventroapical margin somewhat sinuous and apex acute; valvula II with dorsal margin convex, teeth non-contiguous, mostly subtriangular. See systematic notes after the generic description and section 4.3 of the discussion for additional information on the identification of the genus.
Length. ♂♂ 7.4–10.8 mm; ♀♀ 8.4–10.3 mm. Head: in dorsal view, moderately to strongly produced anteriorly; median length of crown varying from 5/10 to 9/10 of interocular width and from 4/10 to 6/10 of transocular width; anterior margin generally rounded; without carina at transition from crown to face; coronal suture distinct, elongate, extending anteriorly beyond interocellar line; frontogenal suture extending onto crown and usually attaining ocellus; ocelli large or of moderate size, located approximately on imaginary line between anterior eye angles, or slightly before or slightly behind this line, each ocellus approximately equidistant between median line of crown and adjacent eye angle; antennal ledge, in dorsal view, varying from not protuberant to slightly protuberant; in lateral view, with anterior margin oblique and convex. Frons, in anterior view, convex; median area mostly smooth; muscle impressions distinct; in lateral view, inferior third slightly angulate; epistomal suture incomplete medially; clypeus, in lateral view, convex, continuing inferior contour of frons. Thorax: pronotum, in dorsal view, with width slightly greater than or approximately equal to transocular width of head; lateral margins convergent anteriorly; posterior margin rectilinear; dorsolateral carina complete, rectilinear, declivent anteriorly; disk without pubescence or punctures. Mesonotum with scutellum not transversely striate and without punctures. Forewing with membrane indistinct; veins not elevated; apex slightly expanded and obliquely truncate; with four apical cells, base of fourth approximately aligned with base of third; with three closed anteapical cells, their bases located more proximally than claval apex; costal apical cell broadened posteriorly; without anteapical plexus of veins; texture subhyaline. Hind wing with vein R2+3 incomplete. Hind leg with femoral setal formula 2:1:1; first tarsomere longer than combined length of two more distal tarsomeres, with two longitudinal parallel rows of small setae on plantar surface. Coloration: head, pronotum, mesonotum, forewings, and legs of preserved specimens usually yellow (whitish-yellow to greenish-yellow in life); preapical area of each forewing with irregular orange transverse band or with distinct red spot on second apical cell. Male terminalia: pygofer, in lateral view, moderately to strongly produced posteriorly; with basiventral lobe; without processes; anteroventral margin with distinct group of microsetae. Subgenital plate, in ventral view, not fused basally to its counterpart; not extending as far posteriorly as pygofer apex. Connective, in dorsal view, usually T-shaped, rarely V-shaped; arms broad. Style, in dorsal view, without preapical lobe. Aedeagus, in lateral view, with shaft usually short and bearing single ventral process, more rarely with pair of processes or with apical digitiform projection. Paraphyses present, symmetrical or slightly asymmetrical, with or without processes on stalk. Female terminalia: sternite VII, in ventral view, well produced posteriorly; narrowing gradually towards apex. “Internal” sternite VIII, in dorsal view, usually without sclerotized areas. Pygofer, in lateral view, well produced posteriorly; posterior margin narrowly rounded to subacute; macrosetae distributed mostly on posterior half. Valvula I, in lateral view, abruptly narrowed apically, ventroapical margin somewhat sinuous, apex acute; dorsal sculptured area extending from basal portion to apex of blade, formed mostly by scale-like processes arranged in oblique lines (strigate); ventral sculptured area restricted to apical portion of blade, formed mostly by scale-like processes; base of valvula forming lobe directed anterad; ventral interlocking device located on basal third or basal half of blade. Valvula II, in lateral view, with dorsal margin convex; blade with about 45 to 60 non-contiguous, mostly subtriangular teeth; preapical prominence distinct; apex obtuse; denticles distributed on teeth and on dorsal and ventral apical portions of blade (ventral dentate apical portion longer than dorsal portion); valvula with ducts extending towards teeth and apical area. Gonoplac of the usual Cicadellinae type: in lateral view, with basal half narrow; apical half expanded, gradually narrowing towards apex; latter obtuse.
According to the results of our cladistic analysis, the genus Dasmeusa can be distinguished from other members of the Paromenia group, as well as from other sharpshooters, by a combination of the following synapomorphic traits: (1) posterior margin of pronotum rectilinear (Fig.
D. basseti Cavichioli & Chiamolera, 1999. French Guiana [new record] and Guyana.
D. dinizi Pecly, Takiya, Cavichioli & Mejdalani sp. nov. French Guiana.
D. falcifera Pecly, Takiya, Cavichioli & Mejdalani sp. nov. French Guiana.
D. imperialis Pecly, Takiya & Mejdalani, 2019. Brazil (Amazonas State).
D. isabellina Cavichioli & Chiamolera, 1999. Brazil (Pará State).
D. mendica Young, 1977. French Guiana and Guyana.
D. oriximina Pecly, Takiya, Cavichioli & Mejdalani sp. nov. Brazil (Pará State).
D. pauperata (Fabricius, 1803), type-species. Brazil (Roraima, Amazonas, Pará, Sergipe, and Bahia states), French Guiana, Guyana, and Suriname.
D. rafaeli Pecly, Takiya, Cavichioli & Mejdalani sp. nov. Brazil (Amazonas State) and Guyana.
1 | Forewing with orange tinge (Fig. |
D. isabellina Cavichioli & Chiamolera |
1’ | Without above combination of features | 2 |
2 | Forewing with small red spot near apex (Fig. |
3 |
2’ | Forewing without small red spot near apex; connective with stalk subequal or shorter than one arm width (Figs |
4 |
3 | Pygofer strongly produced posteriorly (Fig. |
D. falcifera sp. nov. |
3’ | Pygofer moderately produced posteriorly (Fig. |
D. rafaeli sp. nov. |
4 | Aedeagus with large ventral lobe and dorsoapical digitiform projection (Fig. |
D. pauperata (Fabricius) |
4’ | Terminalia not as above | 5 |
5 | Paraphyses without processes on stalk (Figs |
6 |
5’ | Paraphyses with processes on stalk (Figs |
7 |
6 | Aedeagus with pair of small, ventral preapical sclerotized dentiform processes (Fig. |
D. mendica Young |
6’ | Aedeagus with whole ventral margin produced into a single, strong, slightly bifid process directed ventrally (Fig. |
D. basseti Cavichioli & Chiamolera |
7 | Paraphyses with processes of stalk strongly developed (Fig. |
D. oriximina sp. nov. |
7’ | Paraphyses with processes of stalk small (Fig. |
8 |
8 | Paraphyses with single pair of processes on stalk, located dorsoapically, and rami slightly curved inwardly in dorsal view ( |
D. imperialis Pecly, Takiya & Mejdalani |
8’ | Paraphyses with two pairs of processes on stalk, one located dorsally at apical third and another ventrally at apex (Fig. |
D. dinizi sp. nov. |
♂ paratypes 8.8–9.2 mm (n = 2); ♀ paratype 9.6 mm; ♂♂ 8.8–9.2 mm (n = 3). Head (Figs
Dasmeusa basseti Cavichioli & Chiamolera, 1999. A–G Male. A head, pronotum, and mesonotum, dorsal view; B pygofer, lateral view; C subgenital plate, ventral view; D connective and style, dorsal view; E ejaculatory bulb and aedeagus, lateral view; F paraphyses, dorsal view; G paraphyses, lateral view; H–I female; H sternite VII, ventral view; I pygofer, lateral view.
Head, pronotum, and mesonotum light brown; ocelli orange. Forewing pale yellow, translucid, with veins light brown, preapical area with irregular, orange transverse band. Face, lateral and ventral portions of thorax, and legs mostly light brown.
Pygofer (Fig.
Terminalia with sternite VII (Fig.
French Guiana [new record] • 1 ♂: “FRENCH GUIANA: Laussat \ P3 \ 05°28′31.6″N – 053°35′07.3″W \ 30.IX.2010 \ Lamarre G. Leg”; “White sand forest \ Vitre trap (V 7)” (
♂ holotype 8.4 mm; ♂ paratypes 8.0–8.4 mm (n = 3). Male holotype. Head (Figs
Head, pronotum, mesonotum, and legs pale yellow. Forewing pale yellow, translucid, preapical area with irregular orange transverse band.
Pygofer (Fig.
The name of the new species, dinizi, refers to the biologist André Luis Diniz Ferreira, in recognition of his friendship to the first author and contribution as a skilled insect collector.
Laussat (French Guiana).
French Guiana • ♂ holotype: “French Guiana: Laussat \ P3 \ 05°28′31.6″N-053°35′07.3″W \ 12.ix.2010 \ Lamarre G. leg”; “White sand forest \ Light trap” (
Dasmeusa dinizi sp. nov. (Fig.
♂ holotype 7.4 mm; ♂ paratype 8.0 mm; ♀ paratypes 8.4–9.3 mm (n = 3). Male holotype. Head (Figs
Head, pronotum, and mesonotum pale yellow. Forewing pale yellow, second apical cell with distinct red spot.
Pygofer (Fig.
Sternite VII (Fig.
Dasmeusa falcifera sp. nov. A–G Male. A head, pronotum, and mesonotum, dorsal view; B pygofer, lateral view; C valve and subgenital plate, ventral view; D connective and style, dorsal view; E ejaculatory bulb and aedeagus, lateral view; F paraphyses, dorsal view; G paraphyses, lateral view; H–I female; H sternite VII, ventral view; I pygofer, lateral view.
The name of the new species, falcifera, refers to the falciform aedeagus (Fig.
Laussat (French Guiana).
French Guiana • ♂ holotype: “French Guiana: Laussat \ P3 \ 05°28′31.6″N - 053°35′07.3″W \ 12.ix.2010 \ Lamarre G. leg”; “White sand forest \ Light trap” (
As mentioned in the key, D. falcifera sp. nov. (Fig.
♂ paratype 8.4 mm; ♂ 7.8 mm. Head (Figs
Head, pronotum, and forewing mostly orange; mesonotum dull white. Frons mostly orange; clypeus pale yellow. Remainder of face (gena and lorum) and lateral and ventral portions of thorax pale yellow; legs mostly pale yellow to orange.
Pygofer (Fig.
Dasmeusa isabellina Cavichioli & Chiamolera, 1999, male. A Head, pronotum, and mesonotum, dorsal view (male holotype,
Brazil, Pará State • 1 ♂: “Belterra – PA [Pará State] \ Faz. [Fazenda] Treviso \ 01-10/XII/2018 \ Marcela Monné \ Pedro S. Dias” (
♂♂ 9.2–9.6 mm (n = 3). Head (Figs
Head, pronotum, and mesonotum pale yellow. Forewing translucent with transverse broad orange stripe across bases of apical cells.
Pygofer (Fig.
Dasmeusa mendica Young, 1977, male. A Head, pronotum, and mesonotum, dorsal view; B pygofer, lateral view; C subgenital plate, ventral view; D connective and style, dorsal view; E ejaculatory bulb and aedeagus, lateral view; F aedeagus, ventral view; G paraphyses, dorsal view; H paraphyses, lateral view.
French Guiana • 1 ♂: “FRENCH GUIANA: Montagne \ des Chevaux \ 4°44’56″N - 52°26’28″W, alt. 75 m \ 18.iii.2012 \ light trap (GEML) \ SEAG col.” (
According to
♂ holotype 8.0 mm. Male holotype. Head (Figs
Head, pronotum, mesonotum, and forewing pale yellow.
Pygofer (Fig.
Female unknown.
The name of the new species, oriximina, refers to the type locality (Oriximiná) in Pará State (Northern Brazil). It is a noun in apposition.
Brazil (Pará State).
Brazil, Pará State • ♂ holotype: “BR [Brazil]/ PA [Pará State] – Oriximiná \ Porto Trombetas \ 6-14.viii.2018 \ M.L. Soares & Y. \ Anthoinine \ Malaise [trap]” (
Dasmeusa oriximina sp. nov. (Fig.
Cicada pauperata Fabricius, 1803: 71.
Tettigonia lurida Signoret, 1853: 662.
Erythrogonia bicolor Metcalf, 1949: 260. Syn. nov.
Dasmeusa flavescens Metcalf, 1955: 264, new name for Tettigonia lurida Signoret, 1853: 662, which was preoccupied by T. lurida Germar, 1821: 70. Syn. nov.
Note:
♂♂ 9.5–10.8 mm (n = 5); ♀♀ 9.2–10.3 mm (n = 5). Head (Figs
Head, pronotum, and mesonotum mostly brownish-yellow. Forewing yellow, preapical area with irregular orange transverse band. Face, lateral and ventral portions of thorax, and legs pale yellow.
Pygofer (Fig.
Sternite VII (Fig.
Brazil, Amazonas State • 1 ♂: “BRASIL: AM [Amazonas State], 80km N [north of] \ Manaus, Reserva do \ PDBFF [Projeto Dinâmica Biológica de Fragmentos Florestais], Km 41 \ 02°24′S, 59°43′W \ 21-22.vii.2004” (
Dasmeusa pauperata (Fabricius, 1803), type-species, male. A Head, pronotum, and mesonotum, dorsal view; B forewing; C pygofer, lateral view; D subgenital plate, ventral view, with SEM of surface sculpturing (microtrichia); E connective and style, dorsal view; F aedeagus, lateral view; G paraphyses, dorsal view; H apical portion of paraphyses, posterior view; I paraphyses, lateral view.
Dasmeusa pauperata (Fabricius, 1803), type-species, female. A Sternite VII, ventral view; B pygofer, lateral view; C valvifer I and valvula I, lateral view; D dorsal sculptured area; E apical portion; F valvula II, lateral view (schematic, see associated photos [G, H, I] for details of teeth and other structures); G basal teeth; H median teeth; I apical portion. DEN = denticle; DSA = dorsal sculptured area; DUC = duct; PPR = preapical prominence; RAM = ramus; TOO = tooth; VID = ventral interlocking device; VLI = valvifer I; VSA = ventral sculptured area.
Dasmeusa pauperata can be readily distinguished from the remaining known species of the genus by the aedeagus with a large ventral lobe and a dorsoapical digitiform projection (Fig.
As mentioned above, Dasmeusa flavescens Metcalf, 1955 was proposed as a new name for Tettigonia lurida Signoret, 1853, which was preoccupied by T. lurida Germar, 1821.
Dasmeusa flavescens Metcalf, 1955, female lectotype. A Body, dorsal view; B body, lateral view (arrow indicates location of angle at inferior portion of frons); C labels; D face, anterior view; E apical portion of abdomen, ventrolateral view; F apical portion of abdomen, ventral view. The name flavescens was proposed by
Erythrogonia bicolor Metcalf, 1949, male holotype, which was treated by
♂ holotype 8.0 mm; ♂ paratypes 7.6–8.0 mm (n = 2). Male holotype. Head (Figs
Head, pronotum, mesonotum, and forewing pale yellow; second apical cell of forewing with distinct red spot.
Pygofer (Fig.
The name of the new species, rafaeli, is given in honor of Dr. José Albertino Rafael (Instituto Nacional de Pesquisas da Amazônia, Manaus) in recognition of his outstanding contribution to the knowledge of the Brazilian entomofauna.
Brazil (Amazonas State).
Brazil, Amazonas State • ♂ holotype: “BRASIL, AM [Amazonas State], Manaus, \ R. [Reserva] F. [Florestal] Ducke, AM-010 km 26 \ 02°25′49.5″S, 59°58′31.8″W \ Julho/Agosto 1979 \ Fumigação – Dossel \ T. E. Erwin”; “Brazil Canopy Fogging \ Project – July/August \ Dry Samples \ Trans. 10 (1 of 2)” (
Brazil, Amazonas State • 1 ♂: “Brasil AM [Amazonas State] CEPLAC \ Manaus – Rod. [Rodovia] AM010 \ Km 30: 5.X.1977 \ Col. I.S. Gorayeb” (
The color pattern of D. rafaeli sp. nov. (Fig.
Brazil: Amapá State [new record] • 1 ♀: “SERRA DO NAVIO \ Terr. [Território do] Amapá BRASIL \ 30–IX–1957 \ J. Lane leg.”; “Coleção \ J. Lane” (
Scanning electron microscopy (SEM) of the head of Dasmeusa pauperata (Fabricius, 1803). A Compound eye: ommatidia; B interommatidial sensillum trichodeum; C brochosomes on ommatidium; D antenna: scape, pedicel, and base of flagellum; E microtrichia sculpturing on antennal pedicel; F scale-like sculpturing on antennal flagellum; G surface of frons; H sculpturing on frons; I surface of gena covered by brochosomes; J sensillum placodeum on frons; K organ of Evans on maxillary plate; L organ of Evans at higher magnification, with adjacent sensillum trichodeum.
French Guiana: one specimen without abdomen: “FRENCH GUIANA: Laussat \ P8 \ 05°28′31.6″N – 053°35′07.3″W \ 18.x.2010 \ Lamarre G. leg”; “Flooded forest \ Vitre trap (V5)” (
Guyana: 1 ♀: “GUYANA: Essequibo, \ forest on plateau \ above Kaieteur Falls, \ approx. 400m. alt.”; “17 October 1991 \ J. H. Martin coll. \ B. M. 1991-182” (
The use of scanning electron microscopy (SEM) allowed us to study in detail the integument of Dasmeusa pauperata, the type-species of the genus. The following features of the integument were observed for the first time in this genus and are also poorly known in the Cicadellidae as a whole: microtrichia (Figs
Scanning electron microscopy (SEM) of the head of Dasmeusa pauperata (Fabricius, 1803). A Microtrichia at apex of clypeus; B apex of clypeus and labrum; C labrum and second article of labium; D third article of labium and apex of maxillary stylets; E apex of third article of labium and maxillary stylet; F sensilla coeloconica of third article of labium; G sensillum coeloconicum at higher magnification; H maxillary stylet and lateral projections of mandibular stylet (asterisks); I apex of maxillary stylets, with brochosomes.
The data matrix included 40 morphological characters, of which 14 are from the external morphology and coloration (head and thorax) and 26 from the male terminalia; these characters were coded for 15 terminal taxa (six of them outgroups) (Table S1). Thirty-one characters are binary and nine are multistate; only eight characters were non-informative for the parsimony analysis. The characters and their states are listed below (non-informative ones are indicated).
3.3.1.1. External morphology and coloration
1. Head, crown apex, carina (dorsal view): (0) absent (Fig.
2. Antennal ledge, degree of projection in relation to outline of crown (dorsal view): (0) not protuberant (Fig.
3. Ocellus, position in relation to imaginary line between anterior eye angles (dorsal view): (0) aligned with angle (Fig.
4. Head, extension of frontogenal suture on crown (dorsal view): (0) attaining ocellus (Fig.
5. Frons, form of superior third (lateral view): (0) rounded (Fig.
Scanning electron microscopy (SEM) of the thorax of Dasmeusa pauperata (Fabricius, 1803). A Mesonotum surface; B mesonotum surface at higher magnification, showing microtrichia; C higher magnification of mesonotal microtrichia; D surface of scutellum covered by brochosomes; E higher magnification of scutellar brochosomes; F base of forewing; G base of forewing at higher magnification, with microtrichia; H claval sulcus of forewing; I claval sulcus of forewing at higher magnification, with brochosomes; J appendix of forewing; K appendix of forewing at higher magnification, showing microtrichia; L same as preceding at still higher magnification.
6. Frons, form of inferior third (lateral view): (0) not angulate (
7. Frons, aspect of surface of superior portion (frontal view): (0) not depressed medially, (1) depressed medially. Non-informative.
8. Pronotum, form of posterior margin (dorsal view): (0) distinctly concave (
9. Forewing, form of apex (lateral view): (0) convex (
10. Forewing, coloration of apical portion, orange stripe: (0) absent (
11. Forewing, coloration of apical portion, red spot: (0) absent (
12. Forewing, form of costal apical cell: (0) not broadened posteriorly (
13. Forewing, position of base of fourth apical cell in relation to third apical cell: (0) proximal (
14. Forewing, aspect of surface: (0) mostly coriaceous (
3.3.1.2. Male terminalia
15. Pygofer, form of posterior margin (lateral view): (0) rounded (Fig.
16. Pygofer, degree of development (lateral view): (0) well produced posteriorly (Fig.
17. Pygofer, chaetotaxy, microsetae forming group on anteroventral portion: (0) absent, (1) present (Fig.
18. Pygofer, chaetotaxy, distribution of macrosetae: (0) mainly apical third (
19. Pygofer, posterodorsal margin, long and broad process with acute apex (lateral view): (0) present (
20. Pygofer, distinct lobe, basiventral portion (lateral view): (0) absent (
21. Subgenital plate, position of apex in relation to pygofer (lateral view): (0) extending as far posteriorly as pygofer, (1) not reaching pygofer apex. ci = 1.000, ri = 1.000.
Scanning electron microscopy (SEM) of the hind leg of Dasmeusa pauperata (Fabricius, 1803). A Surface of coxa, trochanter, and femur; B surface of coxa and femur at higher magnification, showing group of setae; C apex of femur showing macrosetal formula 2:1:1; D one seta of the macrosetal formula covered by brochosomes; E brochosomes on seta at higher magnification; F setae of tibia; G tarsus and pretarsus, ventral view; H detail of basal tarsomere; I articulation between median and distal tarsomere, showing microtrichia; J microtrichia of distal tarsomere; K apex of tarsus and pretarsus; L apex of tarsus and pretarsus at higher magnification.
22. Connective, shape (dorsal view): (0) Y-shaped (
23. Connective, length of stalk in relation to style (dorsal view): (0) not extending beyond style apex (Fig.
24. Connective, median keel of stalk (dorsal view): (0) present (Fig.
25. Style, form of apex (dorsal view): (0) acute (
26. Style, preapical lobe (dorsal view): (0) absent (Fig.
27. Paraphyses: (0) absent, (1) present (Fig.
28. Paraphyses, symmetry (dorsal view): (0) symmetrical (Fig.
29. Paraphyses, stalk length in relation to rami length (dorsal view): (0) shorter than (Fig.
30. Paraphyses, form of apex of each ramus (dorsal view): (0) acute (Fig.
31. Paraphyses, apical portion of stalk, pair of dorsal spiniform processes: (0) absent (Fig.
32. Paraphyses, apical portion of stalk, pair of ventral spiniform processes: (0) absent (Fig.
33. Paraphyses, orientation of rami (lateral view): (0) directed dorsally (Fig.
34. Paraphyses, position of apical portions of rami (dorsal view): (0) crossing each other (Fig.
35. Aedeagus, shaft aspect (lateral view): (0) elongate and slender (Fig.
36. Aedeagus, ventroapical portion of shaft, process (lateral view): (0) absent (Fig.
37. Aedeagus, ventroapical process, aspect of basidorsal surface (lateral view): (0) smooth, (1) irregularly serrate. ci =0.500, ri = 0.000.
38. Aedeagus, ventroapical process, aspect of ventral surface (lateral view): (0) smooth, (1) irregularly serrate. Non-informative.
Scanning electron microscopy (SEM) of the abdomen of Dasmeusa pauperata (Fabricius, 1803). A Surface of sternite V, showing rows of microtrichia; B sternite V, with sensillum coeloconicum; C sensillum coeloconicum at higher magnification; D sensillum trichodeum of sternite V; E sculpturing of sternal surface, including microtrichia, s. coeloconica, and s. trichodea; F surface of sternite V at higher magnification, showing microtrichia and s. coeloconicum; G s. coeloconicum; H s. coeloconicum at higher magnification; I s. coeloconicum of sternite V surrounded by microtrichia; J s. trichodeum from segment III; K laterotergite VIII showing spiracle; L spiracle at higher magnification.
One of the nine equally most parsimonious trees of the phylogenetic analysis of Dasmeusa (88 steps, CI = 0.580, RI = 0.602). Sailerana solitaris was employed for rooting this tree. This is same and only ingroup topology recovered by implied weighting (fit = 7.85000, k = 3). Accordingly, it is considered our preferred hypothesis. Character numbers are indicated above circles and character states below. White circles (○) indicate homoplastic transformations and black circles indicate (●) non-homoplastic transformations. Bootstrap support values (>50) and Bremer decay indexes, when applicable, are shown.
39. Aedeagus, ventroapical portion of shaft, pair of processes (lateral view): (0) absent (Fig.
40. Aedeagus, position of gonopore (lateral view): (0) apical (Fig.
The analysis with equal weights resulted in nine equally most parsimonious trees (L = 88, CI = 0.580, RI = 0.602). The strict consensus of these trees (Figure S1) is almost entirely polytomous within Dasmeusa, but D. rafaeli sp. nov. + D. falcifera sp. nov. and D. imperialis + D. dinizi sp. nov. consistently formed clades. All trees recovered Dasmeusa as monophyletic, although with relatively low support scores. The implied weighting analysis (k = 3) resulted in two trees (fit = 7.85000), both with the same topology for the ingroup. This ingroup topology, which was also found in one of the nine most parsimonious trees with equal weights, is considered the preferred hypothesis (Fig.
The monophyly of Dasmeusa was recovered in all nine most parsimonious trees (Figure S1) and in the implied weighting search (Fig.
The hypothesis of the sister group relationship between Dasmeusa and Jozima is based on two apomorphic features in the context of the Paromenia group: (1) short and compact aedeagal shaft (c. 35, s. 1, Fig.
The external morphology is apparently quite conservative within the genus Dasmeusa, with the exception of male terminalia characters (see taxonomic discussion below). Therefore, only a limited number of characters (40) could be considered in the present study for the phylogenetic analysis. However, we hope that our preliminary hypothesis of relationships within Dasmeusa (Fig.
Microtrichia (subcellular projections) were found in various parts of the body of D. pauperata, such as the antennal pedicel (Fig.
We observed brochosomes distributed close to microtrichia in various body regions (Fig.
The organ of Evans, sometimes referred to as the maxillary sensillum, is a peculiar structure located, in leafhoppers, on the maxillary plate, next to the lorum (Fig.
Insect sensilla are ectodermal organelles built up by a definite number of characteristic cells (
According to
Sensilla placodea were found on the frons of D. pauperata (Fig.
Sensilla trichodea (hair-like structures or setae) of distinct sizes, generally with a pointed apex (Fig.
With the addition of four new species and the treatment of D. flavescens as a junior synonym of D. pauperata, the genus Dasmeusa currently includes nine species. Our comparative morphological studies indicate that the following combination of easily observable features will most readily distinguish Dasmeusa (Figs
Although females of Dasmeusa are still poorly known, it appears that their terminalia structures are quite conservative, showing little significant interspecific variation. For the most part, we have found so far in these structures only subtle variations in the posterior margin of the sternite VII (Figs
Members of the subfamily Cicadellinae are exclusively xylem-feeders, being usually considered generalists (
The known distribution of Dasmeusa is shown in Fig.
The authors have declared that no competing interests exist.
Luiz A. A. Costa (
Table S1
Data type: .pdf
Explanation note: Data matrix for the phylogenetic analysis of Dasmeusa and outgroup taxa.
Figure S1
Data type: .pdf
Explanation note: Strict consensus of the nine equally most parsimonious trees of the phylogenetic analysis of Dasmeusa and outgroup taxa.