Corresponding author: Tatiana Petersen Ruschel (
Academic editors Christiane Weirauch, Martin Wiemers
The Linnean, Wallacean, and Darwinian shortfalls are knowledge gaps about species taxonomy, distribution, and evolution, respectively.
Ruschel TP, Bianchi FM, Campos LA, Carvalho GS (2023) Total evidence analysis elucidates the tangled systematic scenario within Fidicinini (Hemiptera, Auchenorrhyncha, Cicadidae). Arthropod Systematics & Phylogeny 81: 35–77.
The announcement of the current biological crisis warned biologists of the accelerated loss of biodiversity (
Cicadas are insects well known for their underground nymphal stage and the loud songs performed by the males to attract females to mating (
The uncus (10th abdominal segment) plays a primary role in cicada copula (
The timbal cover, a fold projecting forwards one each side of the second male urotergite (Pringle 1954;
Here we investigate the relationships of these new species of
Dry pinned specimens of both sexes were studied in a stereoscopic microscope. The male genitalia were extracted with forceps, heated in a potassium hydroxide aqueous solution (10% KOH), and posteriorly washed in water. The genitalia were conserved in micro vials filled with glycerine and attached to the specimen’s pin. The female reproductive system was not extracted due the distortion and decomposition observed in dried specimens (as reported by
Photographs of the major morphological structures were obtained with an AxioCam ERc 5s digital camera attached to a Stemi 2000 C P.06 stereoscopic microscope with Zen Lite 2011 software and with a Nikon AZ100M, followed by stacking with the Nikon NIS–Elements Ar Microscope Imaging Software. Vectorized drawings were made on the photographs. In addition, full-body images (dorsal view) were made with a digital camera. Terminologies adopted here follow
While studying specimens of
Other genera were chosen based on the current classification of the
The specimens examined in this study belong to the following collections:
The morphological characters were coded in Mesquite version 3.5.1 (
DNA was extracted from ethanol-preserved and pinned specimens. We preferably removed the right foreleg from the coxal cavity to access the indicated amount of muscular tissue. Genomic DNA was extracted using DNeasy Blood and Tissue kit (Qiagen, Valencia, CA, U.S.A.) according to the manufacturer’s instructions, eluting to a final volume of 100 μL when DNA was extracted from ethanol-preserved specimens or 50 μL from pinned specimens.
Two DNA markers were amplified, the nuclear elongation factor 1 alpha (
All PCR products were purified using Exonuclease I and shrimp alkaline phosphatase (Affymetrix, Inc. USB Products, Cleveland, OH, U.S.A.). Both DNA strands for all PCR products were sequenced by Macrogen, Inc. (Seoul, South Korea). Sequence chromatograms were visually inspected, verified, and manually edited using the Staden package (
Additional DNA sequences of
Two probabilistic methods were used to build the phylogenetic trees, Maximum-Likelihood (
Character states related to timbal cover and uncus shape (see results, characters 31–35, and 44) were optimized on the consensus resulting tree from the
The distribution map was made in Quantum Gis 3.16.4 (
Some taxonomic studies have identified species as
A total of 50 morphological characters of male adults were coded and included in a matrix (Table S2 and File S1), 42 out of these were characters of general morphology, and eight were genital.
Characters. Head and pronotum in dorsal view:
Types of uncus (character 44) in ventral and lateral views. The numbers indicate the character state. Scale bar: 1 mm.
The phylogenetic reconstructions under
Bayesian inference consensus tree based on the analysis of two molecular markers and 50 morphological characters for 61 species of
Besides
The optimization of the timbal cover characters (32–35) showed the convergence of the states by multiple transitions and reversions across the evolutionary scenario, lacking shared exclusive states in the recovered clades (Fig.
Characters optimization of timbal cover (31–35) and uncus shape (44) under parsimony based on Bayesian inference consensus tree. Capital letters in circles indicate the clades referred in discussion.
We georeferenced data for the 13 species of
Distribution map of
The Brazilian subregion is represented here by the Boreal Brazilian dominion (
The remaining ten species are recorded in the Chacoan subregion, represented here by the Chacoan dominion (
Three species occur exclusively in
The name refers to the long, acute median projection of sternite II, like a thorn. Latin, m.:
The new genus differs from any other
Head: including eyes, broader than the pronotum (except pronotal collar) and mesonotum; vertex short, wide and slender, the lateral ocelli widely separated, supra–antennal plates not prominent relative to the anterior margin of the head in dorsal view, distant to the eyes; apex of postclyepus convex, without a groove; postclypeus oval in ventral view, flat or slightly salient in lateral view; anteclypeus flat, with a concave basal area. Thorax: pronotal collar wide, not exceeding the lateral margin of eyes; paranota in ventro-posterior position relative to the eyes in lateral view, not exceeding the ventral limit of the eye; cruciform elevation with the central and lateral areas flat, and the arc of the posterior projections obtuse; basisternum 3 flat and prominent relative to the mesocoxae, with protuberances well developed and the posterior margin angled; three segmented tarsi; profemora armed with three spines: the proximal round, leaning forward at the apex, the median sharp and straight, and the distal shorter than the others; posterior tibiae with at least four spines: two dorsal, one of which is at the middle of the tibia and one of which is three-fifths the distance from the base to the apex of the tibia, and two ventral, one of which is three-quarters of the distance from the base to the apex of the tibia and one of which is subapical. Wings hyaline, veins not infuscated; forewings: vein RA divergent to Sc from the basal cell; basal vein of the second apical cell oblique (except in
French Guiana, Brazil and Peru.
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Timbal cover tumid (Fig. |
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– | Timbal cover flat (Fig. |
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Middle third of the anteromedial margin of the timbal cover convex (Fig. |
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– | Middle third of the anteromedial margin of the timbal cover concave |
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Labium reaching (Fig. |
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– | Labium not reaching or surpassing the middle of sternite II (Figs |
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Anterior margin of head slightly concave (Fig. |
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– | Anterior margin of head tightly concave (Fig. |
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Posterior margin of basisternum 3 straight (Fig. |
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– | Posterior margin of basisternum 3 angled (Figs |
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Posterior margin of basisternum 3 forming an obtuse angle (Fig. |
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– | Posterior margin of basisternum 3 in an acute angle (Fig. |
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Labium reaching or almost reaching the base of the basisternum (Fig. |
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– | Labium reaching the apex of the median projection of sternite II (Fig. |
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Tergites 2 to 7 with pilus setae in both lateral margins (Fig. |
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– | Tergites 2 to 7 without pilus setae in both lateral margins (Fig. |
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Apex of timbal cover almost reaching the lateral metascutellar plate (Fig. |
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– | Apex of timbal cover distant from the lateral metascutellar plate (Fig. |
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Anterior margin of head tightly concave (Fig. |
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– | Anterior margin of head slightly concave (Fig. |
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Timbal cover wide, almost covering the timbal cavity (Fig. |
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– | Timbal cover not covering the timbal cavity (Fig. |
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Tergites with pilus setae in both lateral margins of tergites 2 and 3 and in the anterior margin of tergite 6 (Fig. |
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– | Tergites without pilus setae in both lateral margins of tergites 2 and 3 and in the anterior margin of tergite 6 (Fig. |
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Bento Gonçalves, Rio Grande do Sul (Brazil).
The specific name refers to the distribution of species. The charruas were indigenous people who inhabited southern South America.
The species can be distinguished from all other species of
Body tawny with the head, thorax and abdomen marked in black.
Head (Fig.
N = 5 males and 5 females mean (range). Length of body: male 21.64 (20.39–23.09), female 21.44 (20.02–23.38); width of head including eyes: male 9.58 (9.07–10.13); female 9.46 (8.25–10.11); length of the head: male 2.42 (2.17–2.61), female 2.47 (2.26–2.67); width of pronotum including pronotal collar: male 9.38 (8.83–10.03), female 9.22 (8.04–9.80); length of pronotum including pronotal collar: male 3.36 (3.63–3.20), female 3.49 (3.40–3.56); width of mesonotum: male 7.93 (7.50–8.56), female 7.78 (6.41–8.46); length of mesonotum: male 6.01 (5.53–6.77); female 5.97 (4.89–6.65); length of forewing: male 28.72 (27.00–31.04), female 28.66 (25.72–30.39); width of forewing: male 9.82 (9.10–10.75), female 9.87 (9.56–10.55); length of hind wings: male 15.52 (14.38–16.63), female 15.55 (13.84–16.70).
Brazil (Rio Grande do Sul).
Pelotas, Rio Grande do Sul (Brazil).
The species is named in honor to the mother of the first author, Claudia Petersen Ruschel.
The species can be distinguished from all other species of
Body yellowish with the head, mesonotum and abdomen marked with black.
Head (Fig.
Holotype male. Length of body: 16.01; width of head including eyes: 6.90; length of the head: 1.60; width of pronotum including pronotal collar: 6.80; length of pronotum including pronotal collar: 2.20; width of mesonotum: male 5.70; length of mesonotum: 3.50; length of forewing: 19.90; width of forewing: male 6.5; length of hind wings: ale 11.20. Paratypes 8 males and 2 females, mean (range). Length of body: male 15.05 (14.40–15.80), female 15.00 (16.10–13.90); width of head including eyes: male 6.93 (6.70–7.00); female 7.25 (7.60–6.90); length of the head: male 1.60 (1.60–1.60), female 1.65 (1.60–1.70); width of pronotum including pronotal collar: male 6.73 (6.40–7.00), female 7.20 (7.50–6.90); length of pronotum including pronotal collar: male 2.38 (2.40–2.50), female 2.65 (2.70–2.60); width of mesonotum: male 5.58 (5.40–5.70), female 5.80 (5.70–5.90); length of mesonotum: male 3.68 (3.30–3.90); female 3.95 (4.10–3.80); length of forewing: male 19.50 (18.00–21.00), female 21.50 (21.80–21.20); width of forewing: male 6.80 (6.50–7.00), female 6.90 (7.00–6.80); length of hindwings: male 10.93 (10.20–11.80), female 11.55 (11.70–11.40).
Brazil (Rio Grande do Sul).
Espírito Santo, Brazil.
Holotype male, allotype female, and paratype male (
The species can be distinguished from all other species of
Body yellowish ventrally; head, pronotum, and mesonotum olive-green marked with black; abdomen dark castaneous with the tergites marked with black dorsally.
Head (Fig.
N = 5 males and 5 females, mean (range). Length of body: male 26.79 (24.64–28.24), female 23.56 (22.49–24.34); width of head including eyes: male 10.83 (9.87–11.51); female 10.60 (10.11–11.06); length of the head: male 2.71 (2.58–2.82), female 2.79 (2.45–3.58); width of pronotum including pronotal collar: male 10.48 (9.36–11.09), female 10.28 (9.68–10.79); length of pronotum including pronotal collar: male 4.14 (3.63–4.41), female 3.91 (3.71–4.16); width of mesonotum: male 8.99 (8.29–9.39), female 8.83 (8.18–9.18); length of mesonotum: male 7.25 (6.74–7.69); female 7.02 (6.45–7.34); length of forewing: male 34.45 (32.77–36.84), female 34.89 (33.84–35.57); width of forewing: male 11.46 (10.51–12.53), female 11.14 (10.68–11.57); length of hind wings: male 17.44 (16.31–18.36), female 17.39 (16.85–18.07).
2 females (
Brazil (Espírito Santo, Minas Gerais*, São Paulo*, Paraná*, Santa Catarina*, Rio Grande do Sul*).
Minas Gerais, Brazil.
The species can be distinguished from all other species of
Body dark castaneous, marked with black and tawny.
Head (Fig.
Unknown.
Syntype male (
Male (
Brazil (Goiás, Minas Gerais).
Itatiaia, Rio de Janeiro, Brazil.
This was the last species included in this study. Latin: faustus, lucky; psaltria, female harpist.
The species can be distinguished from all other species of
Body tawny with the head, thorax and abdomen marked with black, covered by golden setae.
Head (Fig.
Holotype male. Length of body: 26.28; width of head including eyes: 9.13; length of the head: 1.84; width of pronotum including pronotal collar: 8.54; length of pronotum including pronotal collar: 3.27; width of mesonotum: 7.37; length of mesonotum: 7.94; length of forewing: 40.67; width of forewing: 14.62; length of hind wings: 20.39.
Brazil (Rio de Janeiro).
Amazonas, Brazil.
The specific name refers to the distribution of the species. Iara is a mermaid from Brazilian folklore who lives in the Amazon River, who lures men singing with their enchanting music.
The species can be distinguished from all other species of
Body tawny with head, thorax and abdomen marked with black; pronotal collar and wing groove olive-green.
Head (Fig.
N = Holotype and paratypes (5 males and 2 females), mean (range). Length of body: male 19.22 (22.75–17.50), female 17.635 (17.64–17.63); width of head including eyes: male 8.698 (9.09–8.26); female 8.54 (8.64–8.44); length of the head: male 1.85 (1.62–1.97), female 1.575 (1.71–1.44); width of pronotum including pronotal collar: male 8.52 (8.90–7.82), female 8.28 (8.28–8.28); length of pronotum including pronotal collar: male 3.00 (2.72–3.26); female 2.835 (2.79–2.88); width of mesonotum: male 7.47 (6.80–7.88), female 7.065 (7.20–6.93); length of mesonotum: male 5.64 (5.94–5.10); female 5.49 (5.58–5.40); length of forewing: male 27.31 (28.80–25.38), female 26.385 (26.60–26.17); width of forewing: male 8.96 (9.73–8.00), female 9.095 (9.25–8.94); length of hindwings: male 12.98 (14.19–11.25), female 12.58 (12.90–12.26).
Brazil (Roraima, Amazonas, Rondônia).
Alto Paraíso de Goiás, Goiás, Brazil.
The specific name refers to the reddish color of species. Latin: ignis + arius, of fire.
The species can be distinguished from all other species of
Body castaneous marked with black and with pronotal collar and sternum tawny, operculum slightly reddish.
Head (Fig.
Holotype male. Length of body: 16.90; width of head including eyes: 7.62; length of the head: 1.89; width of pronotum including pronotal collar: 7.65; length of pronotum including pronotal collar: 2.61; width of mesonotum: 6.66; length of mesonotum: 4.95; length of forewing: 24.21; width of forewing: 8.43; length of hind wings: 11.82.
Brazil (Goiás).
French Guiana.
Holotype male, allotype female, paratypes 1 male and 1 female (
The species can be distinguished from all other species of
Body ochre marked with green and black.
Head: with a transverse wide black band extended over the vertex, covering the region of ocellus and reaches the apex of the postclypeus; base of eyes marked with black; anterior margin of head slightly convex; postclypeus unmarked, oval in ventral view, the apex not prominent in dorsal view relative to the supra-antennal plates; anteclypeus and carina tawny; lorum black; rostrum with a tawny mentum and labium; labium short, reaching the base of basisternum 3, black at the apex. Pronotum: paranota visible in dorsal view, wide, not reaching the eyes in dorsal view, and straight in lateral view. Mesonotum: submedian sigillae marked with black laterally; basisternum 3 with very developed protuberances relative to the median insertion, closely spaced forming an acute angle; posterior margin angled; cruciform elevation not covering tergite 1; apex of the posterior projections of the cruciform elevation obtuse; operculum obtuse; wings hyaline; forewings:, basal vein of the second apical cell oblique; hindwings: radius vein straight. Abdomen subcylindrical, the length is equivalent to the combined length of the head and thorax in dorsal view; timbal cover tumid, middle third of anteromedial margin concave; tergites 2 to 8 marked with black anteriorly; sternite VII sub-rectangular. Pygofer sub-cylindrical; the basal lobe long, reaching the uncal dorsal crest.
The female presents the same somatic characteristics as the male.
Holotype male and allotype female. Length of body: 24.3 and (missing); fore body length: 10.12 and 9.75; abdomen length: 11.5 and 10; width of head including eyes: 9 and 9.37; width of mesonotum: 7.5 and 7.5; wingspan: 63 and 64; total length including wings: 32 and 33; length of forewings: 28 and 28.5; width of forewings: 9 and 9.1 (
Photographs of original manuscript.
French Guiana.
Londrina, Paraná, Brazil.
The specific name refers to the olive green color of the species. Latin: oliva, olive.
The species can be distinguished from all other species of
Body green with head, thorax and abdomen marked with black.
Head (Fig.
Holotype male: Length of body: 19.25; width of head including eyes: 8.84; length of the head: 2.00; width of pronotum including pronotal collar: 8.70 length of pronotum including pronotal collar: 3.20; width of mesonotum: 7.44; length of mesonotum: 5.68; length of forewing: 28.28; width of forewing: 9.80; length of hind wings: 14.10. Paratypes (4 males): Length of body: 19.53 (20.78–18.73); width of head including eyes: 8.83 (9.12–8.56); length of the head: 1.76 (2.00–1.60); width of pronotum including pronotal collar: 8.62 (9.00–8.30); length of pronotum including pronotal collar: 3.18 (3.36–3.04); width of mesonotum: 7.26 (7.44–7.04); length of mesonotum: 5.70 (6.00–5.36); length of forewing: 26.27 (three paratypes are without the forewings); width of forewing: 9.43; length of hind wings: 14.92 (16.87–13.12).
Brazil (Paraná).
Magé, Rio de Janeiro, Brazil.
The specific name refers to the immortal bird of Greek mythology which obtains new life by arising from the ashes. The specimens designated as holotype and paratype were received by the first author before the Museu Nacional (Rio de Janeiro, Brazil) fire in 2018. The name is in honor of this important museum that we hope arises from the ashes like a
The species can be distinguished from all other species of
Body orange with head, thorax and abdomen marked with black.
Head (Fig.
Unknown.
Holotype male. Length of body: 22.60; width of head including eyes: 9.90; length of the head: 2.07; width of pronotum including pronotal collar: 9.72; length of pronotum including pronotal collar: 3.15; width of mesonotum: 8.28; length of mesonotum: 6.48; length of forewing: 29.62; width of forewing: 9.86; length of hind wings: 14.07. Paratype male. Length of body: 22.13; width of head including eyes: 10.41; length of the head: 2.25; width of pronotum including pronotal collar: 10.12; length of pronotum including pronotal collar: 3.37; width of mesonotum: 8.37; length of mesonotum: 6.61; length of forewing: 31.44; width of forewing: 11.04; length of hind wings: 15.15.
Brazil (Rio de Janeiro).
Serro, Minas Gerias, Brazil.
The species is named in honor to the father of the first author, Claudio Rubem Sassen Ruschel.
The species can be distinguished from all other species of
Body tawny with the head, thorax and abdomen marked with black.
Head (Fig.
Holotype male. Length of body: 17.06; width of head including eyes: 7.46; length of the head: 2.00; width of pronotum including pronotal collar: 7.76; length of pronotum including pronotal collar: 2.56; width of mesonotum: 6.72; length of mesonotum: 4.88; length of forewing: 23.35; width of forewing: 8.31; length of hind wings: 12.57. Paratypes (2 males). Length of body: 16.05 (15.36–16.75); width of head including eyes: 7.65, 7.71; length of the head: 1.60, 1.84; width of pronotum including pronotal collar: 7.655 (7.12–7.52); length of pronotum including pronotal collar: 3.23 (3.90–2.56); width of mesonotum: 6.00 (5.60–6.40); length of mesonotum: 4.36 (4.16–4.56); length of forewing: 22.55 (22.00–23.11); width of forewing: 7.39 (7.22–7.56); length of hind wings: 11.78 (11.10–12.46).
Brazil (Minas Gerais).
Cusco, Peru.
The specific name refers to the timbal cover tumid. Latin: tumidus, swollen.
The species can be distinguished from all other species of
Body green and castaneous marked with black.
Head (Fig.
Holotype male. Length of body: 22.30; width of head including eyes: 10.27; length of the head: 2.52; width of pronotum including pronotal collar: 9.84; length of pronotum including pronotal collar: 3.50; width of mesonotum: 8.40; length of mesonotum: 6.76; length of forewing: 32.85; width of forewing: 11.00; length of hind wings: 15.82. Paratypes (male and female). Length of body: 22.71, 22.29; width of head including eyes: 10.85, 10.55; length of the head: 2.81, 2.74; width of pronotum including pronotal collar: 10.37, 10.34; length of pronotum including pronotal collar: 3.69, 3.58; width of mesonotum: 8.79, 8.55; length of mesonotum: 6.90, 7.08; length of forewing: 32.86, 33.17; width of forewing: 11.35, 10.68; length of hind wings: 16.60, 16.96.
Peru.
Alto Caparaó, Minas Gerais, Brazil.
The specific name refers to the dorsal color of species (head, pronotum and mesonotum). Latin: viridis, green. Latin: nota, mark.
The species can be distinguished from all other species of
Head and thorax yellow greenish marked with black, and abdomen tayny marked with black.
Head (Fig.
Holotype male. Length of body: 20.41; width of head including eyes: 9.14; length of the head: 1.71; width of pronotum including pronotal collar: 8.91; length of pronotum including pronotal collar: 1.89; width of mesonotum: 7.65; length of mesonotum: 5.67; length of forewing: 28.88; width of forewing: 9.97; length of hind wings: 13.60.
Brazil (Minas Gerais).
For the first time, we presented a phylogenetic hypothesis of
The species currently classified in
Recently, twelve species were transferred from
The authorities of
The timbal cover surface (31) and its aperture (32) have at least two transitions within clade G, and the flat (311) and exposed (321) timbal cover are shared features between clades B and C. On the other hand, the same character states are spread out in the phylogeny. Thus, character states representing the putative diagnostic features of
Unlike the timbal cover, different works have proposed that the evolution of the uncus can be linked to phylogenetic relationships, at least at the genus and tribe level (
We recovered two clades with exclusive uncus states (type H in
We built the first phylogenetic hypothesis for
The high variability of the timbal cover shape and the polyphyly of the included
We are thankful to Conselho Nacional de Desenvolvimento Científico e Tecnológico – CNPq (L.A.C. 307204/2015–4) and to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – CAPES Finance Code 001, CAPES/CNPq PROTAX II [88882.156807/2017-01] (TPR). We would like to thank to the curators of the scientific collections for the loan of material, especially Dr Robin Thomson (UMSP), Dr Daniela Maeda Takiya (
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