Smilidarnis is fixed as the correct original spelling. Smilidarnis similifasciatus, sp. nov., from Bolivia and French Guiana, closely resembles S. fasciatus Andrade in being brightly colored but differs in the metathoracic tibial chaetotaxy, the male pygofer, first anal segment, aedeagus, and color pattern. In this new species, which is larger than S. fasciatus, females are larger than males. Photographs of the male and female adults and genitalia of the new species are provided. Bolivia is a new country record for the genus. A key to all species is provided.

Bolivia, French Guiana, Neotropical, spelling fixation

Membracid treehoppers are famous for their hyperdeveloped pronotum, which is usually expanded to cover much of the body or projected upwards and posteriorly over it. Many species are cryptically colored yet others exhibit the classic aposematic coloration, such as bright red, orange, or yellow, sometimes over a black ground color, advertising their presence to potential predators (Joron 2009).

The higher taxa of Membracidae are largely based on forewing venation, leg chaetotaxy, and coverage of scutellum and wings by the pronotum. Andrade (1989) established the genus Smilidarnis for two new species without placing the genus to higher taxon because they display features of both Smiliinae and Darninae. His diagnosis was expanded slightly by McKamey (2023) to accommodate the new variation found among three new species. Specifically, Andrade reported metathoracic tibial cucullate rows II and III double and, by implication, row I single, but S. duocornus McKamey has row I absent and S. robustus McKamey has rows I and III double and row II absent. In addition, two of the new species described by McKamey (2023) have suprahumeral horns. Smilidarnis is still unplaced within the family. Features indicating placement in Smiliinae include Andrade’s species having metathoracic tibial chaetotaxy matching that of Ceresini, and in S. concolor Andrade the forewing veins R and M are confluent near its apex, which is a diagnostic feature of Smiliinae (Deitz 1975). Smilidarnis also has features of Darninae: some Smilidarnis have the metathoracic tibia with setal row I or II absent (Hemikypthini and some Darnini lacking row I, some Hemikypthini lacking row II); and R₄₊₅ and M₁₊₂ are separate throughout (Fig. 2), which occurs in all Darninae and no Smiliinae. Consequently, leg chaetotaxy and forewing venation do not provide evidence resolving the relationship of Smilidarnis to other treehoppers. Based on morphology, the phylogenetic estimate by Dietrich et al. (2001) placed Smilidarnis as the sister-group to seven tribes of Smiliinae. A more recent, unpublished molecular phylogeny by Evangelista et al. (2017) placed Smilidarnis as the sister group of Ceresini (Smiliinae). No other phylogenetic estimates have included Smilidarnis.

Until now, S. fasciatus Andrade (1989) was the only brightly colored species in the genus. It is recorded only from French Guiana and Peru. Lapèze (2021) collected three specimens of S. fasciatus at lights in two localities in French Guiana. He noted that in contrast to most membracids, the female was smaller than the male. The female habitus photographs by Lapèze (2021, p. 118) match the line drawing by Andrade (1989; Figs 1, 2) in every respect. A newly discovered species closely allied to S. fasciatus, and as brightly colored, is represented by a series of six specimens of males and females, from Bolivia, which were also collected at lights. The series of the new species provides an opportunity to investigate male vs. female variation in size, morphology, and coloration. The color pattern in the photograph by Sakakibara of a specimen from French Guiana (Deitz and Wallace 2010; distribution by Dr. A. M. Sakakibara, pers. comm.) is consistent with that of the new species.

In quoting labels, quotation marks separate labels and a vertical line separates lines on a label.

Terminology for general morphology, forewing venation (except crossvein s), and leg chaetotaxy follows Deitz (1975) except the gonoplac for the female genitalia, which follows Mejdalani (1998).

The abdomen was detached, macerated in a 10% KOH solution at room temperature for 72 hr, bathed in water, then in acetic acid to stop the reaction. After dissection, structures were stored in a glass microvial containing glycerin and pinned beneath the specimen.

A Leica MZ12 stereomicroscope was used to examine structures. All measurements were taken directly from the metadata of images from all specimens.

Images were taken with a Canon 5Dsr camera with an adjustable 65mm lens. Photos were taken using Capture One Pro version 10.1.2, 64 bit, build 10.1.2.23 imaging software, aided by CamLift version 2.9.7.1. The specimen was lit using two adjustable Dynalite MH2050 RoadMax flash heads, each attached to a Manfrotto 244 arm. The light was diffused using a lampshade-style cone of translucent paper between the specimen and light sources. After individual focal planes were photographed, they were compiled into a single, composite image using Zerene Stacker - USDA SI-SEL Lab Bk imaging system, version 1.04, build T201706041920. Stacked images were enhanced and edited in Adobe Photoshop CSS Extended version 12.0. The scale bars were generated through Photoshop directly from the metadata of the photos.

All specimens are deposited in the U.S. National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM).

Smilidarnis Andrade (1989) is here fixed as the correct original spelling in accordance with ICZN Article 32.5 (International Commission on Zoological Nomenclature 1999); it was misspelled Smiliodarnis in the discussion. The Bolivian specimens of the new species represents a new country record for the genus.

Key to species

1	Pronotum with pair of stout suprahumeral spines	2
–	Pronotum without suprahumeral spines	3
2	Breadth across suprahumeral spines distinctly greater than breadth across posterior lateral spines (McKamey 2023; Fig. 2)	S. duocornus McKamey
–	Breadth across suprahumeral spines subequal, slightly less than breadth across posterior lateral spines (McKamey 2023, fig. 14)	S. robustus McKamey
3	Pronotum posteriorly with tips but not bases of lateral spines black; forewing with veins R and M distally fused then separated preapically	S. concolor Andrade
–	Pronotum posteriorly with bases of lateral spines black; forewing with veins R and M not fused at any point	4
4	Pronotum with central apical spine pale throughout; head vertex with ventrolateral margins and frontoclypeus forming evenly convex curve (McKamey 2023; Fig. 9)	S. erwini McKamey
–	Pronotum with central apical spine black in distal third; head vertex with ventrolateral margins straight and frontoclypeus forming an angle (Fig. 3)	5
5	Pronotum with lateral margin having an isolated yellow patch	S. fasciatus Andrade
–	Pronotum with most lateral yellow band bifurcate posteriorly, lower arm running along almost entirety of lateral margin (Figs 2, 5)	S. similifasciatus sp. nov.

 Smilidarnis similifasciatus sp. nov.

https://zoobank.org/FC4868AC-C340-4292-B7E5-778D81965EC2

Diagnosis

Pronotum without suprahumeral spines; brightly colored with orange, yellow, and black; most lateral yellow band bifurcate posteriorly, lower arm running along almost entirety of lateral margin.

Distribution

Bolivia (new country record for genus), French Guiana.

Description

Dimensions (mm). Pronotal length ♀10.3–10.8, ♂9.8–9.9; total length including wings in repose ♀12.1–12.6, ♂11.5; width between humeral angles ♀4.7–4.8, ♂4.5–4.7; head width including eyes ♀3.9–4.0, ♂3.7–3.9, head width excluding eyes ♀2.7–2.8, ♂2.5–2.7; head height ♀1.7–1.8, ♂1.7–1.8; distance between apices of posterolateral spine apices ♀2.6–2.9, ♂2.4–2.6. Head (Figs 3, 6). Vertex inclined forward, wider than tall, glabrous, vertical dark lines including ocelli depressed; dorsal margin weakly convex, lateral margins straight; ocelli circular, on imaginary middle line between eyes; distance from outer margin of ocellus to eye 1.13 × distance between inner margins of ocelli; frontoclypeal sutures prominent; frontoclypeus in anterior view ventrally convex, in line with vertex ventrolateral margins. Pronotum (Figs 1–6). Suprahumeral spines absent; weakly elevated immediately behind humeral angles; distally with 3 spines, all directed ventroposteriorly, lateral pair directed weakly laterally; lateral spines reaching to leveI above r-m crossvein, central spine surpassing forewing vein M₃₊₄. Legs. All femora lacking cucullate setae and spines; metathoracic tibia row I cucullate setae basally in single row, distally double, row II in single row, sparse and with minute basal hoods, row III cucullate setae in single row. Female terminalia. Sternite VII deeply emarginate medially; pygofer long; valvula I (Fig. 10) long, with one weak, wide dorsal process distally, apex rounded; valvula II (Figs 11, 12) in lateral view narrow and lanceolate with dorsal margin linear, lacking dorsal dentae or sinuations, apex subacute, ventrally striate; gonoplac (Fig. 13) long, distally wide, apex rounded, bearing macrosetae along entire ventral margin. Male terminalia (Figs 7–9). Abdominal segment X (1^st anal segment) with basaoventral lobe (Fig. 7); pygofer (Fig. 7) including lateral plate subquadrate, lateral plate with weak dorsal lobe, ovate, pilose throughout, subgenital plate fused basally, its sides subparallel, slightly narrowing distally, length in ventral view about 2.3 × width; style (Figs 8, 9) recurved with short acute apex bearing 4 setae preapically; aedeagus (Figs 8, 9) U-shaped in lateral view, shaft subparallel in anterior and lateral views, lacking dentae or other texture on distoanterior face but with minute dentae along lateral margins (Fig. 9); gonopore dorsal. Color. Male and female alike in color; pronotal color orange with 5 yellow longitudinal bands, medial and most lateral band nearly extending to large yellow spot in posterior fifth, most lateral longitudinal band bifurcate at mid length into 2 solid branches, 1 running along lateral margin, and black at bases and apices of posterior spines; head orange with 3 vertical yellow bands connecting to medial and closest yellow bands of pronotum; forewing anteriorly clear, posteriorly with amber tint.

Figures 1–6. 

Smilidarnis similifasciatus sp. nov. 1–3 male habitus in dorsal. lateral, and anterior views, respectively 4–6 female habitus in dorsal, lateral, and anterior views, respectively. Scale bar: 5 mm.

Figures 7–13. 

Smilidarnis similifasciatus sp. nov. 7 male pygofer, lateral view 8, 9 aedeagus and styles in dorsanterior and oblique view, respectively 10 female valvula I 11, 12 female valvula II 13 gonoplac.

Material examined

Smilidarnis similifasciatus, holotype • ♂ with labels “BOLIVIA, Dept. Santa Cruz | Prov. Andres Ibañez, Potreillos del Guenda, 17°40.27’ S, | 63°27.45’ W, 370 m, 5 Dec 2008 | T. J. Henry, at MV/black light” and red label “HOLOTYPE | Smilidarnis | similifasciatus | McKamey” (USNM). Paratypes • 4 ♀, 1 ♂, all same data as holotype but with blue paratype labels (USNM).

Etymology

The specific name is a masculine Latin adjective based on a combination of “similis” (similar to) and the allied species “fasciatus”.

Discussion

Smilidarnis fasciatus and S. similifasciatus key out in the same couplet because they are superficially similar. Nevertheless, they differ in several respects. Regarding coloration, in the new species all yellow stripes are longer in both males and females than those of S. fasciatus. Regarding metathoracic tibia, cucullate setal rows are double rows in S. fasciatus but single rows in the new species (row I distally double). Regarding terminalia, in the new species the pygofer and lateral plate combined is more quadrate than in S. fasciatus, the lateral plate itself is ovate (vs. subquadrate in S. fasciatus), segment X has a basoventral lobe present (vs. absent in S. fasciatus), and the aedeagal dentae are lateral (vs. on the face of the shaft in S. fasciatus).

The female habitus photographs by Lapèze (2021) match the illustration of S. fasciatus by Andrade (1989), while the photograph by Sakakibara (Deitz and Wallace 2010) corresponds to S. similifasciatus. Both species, therefore, are recorded from French Guiana. The S. similifasciatus male terminalia is most similar to that of S. erwini McKamey (2023) in that both have abdominal segment X bearing a basoventral lobe and the aedeagus lacking dentae or other texture on the dorsoanterior face of the shaft. The female genitalia of S. fasciatus has not been described so no comparison with that species is possible here, and only the second valvulae of S. concolor has been illustrated and described. The female genitalia of S. similifasciatus resembles those of S. duocornus, S. robustus and the second valvulae of S. concolor, in having the first valvulae long and narrow with poorly developed dentae distally, the second valvulae lanceolate, and the gonoplac distally wide with long setae ventrally.

Although the Bolivian males of the new species are slightly smaller than the lengths given by Andrade (1989) and Lapèze (2021) for S. fasciatus, in S. similifasciatus the four females are all larger than both males in every respect except head height, in contrast to Lapèze’s (2021) opposite finding about the relative lengths of genders of S. fasciatus in French Guiana. Also, the females of S. similifasciatus are longer than those of S. fasciatus as reported by Lapèze (2021).

I thank Alyssa Seemann (USDA Systematic Entomology Laboratory) for taking and processing photographs and compiling plates, and Lewis Deitz (emeritus, North Carolina State University), Olivia Evangelista (CSIRO, Australia), and an anonymous reviewer for comments on an earlier draft of the manuscript. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the USDA. The USDA is an equal opportunity provider and employer.