An Overlooked New Endemic Species of Renonus DeLong, 1959 (Hemiptera: Cicadellidae: Deltocephalinae: Athysanini) from the Seasonally Dry Forest of Western Mexico ^†

Abstract

The leafhopper genus Renonus is one of the rarest genera in the leafhopper tribe Athysanini. The Mexican endemic monotypic species, Renonus rubraviridis DeLong, is historically known from few localities, and since the original description, no additional data has been provided. During an ongoing survey conducted in western Mexico over the Seasonally Dry Tropical Forests, including the surroundings of the Estación de Biología Chamela (IB-UNAM), specimens of R. rubraviridis and others that do not morphologically fit with previously described species were collected. Herein, a new endemic species to Mexico, Renonus cuixmalensis sp. nov., is described and illustrated in detail. In addition, morphological notes of R. rubraviridis, the key to species, a map of distribution, a habitat description, and a discussion about the strong influence on distribution through the Seasonally Tropical Dry Forest are given.

1. Introduction

The prominent leafhopper family Cicadellidae (Hemiptera: Auchenorrhyncha: Cicadomorpha: Cicadoidea) comprises more than 23,000 valid species worldwide [1]. With up to 40 tribes and 991 genera, the subfamily Deltocephalinae is a vast group with exclusive herbivore habits and is highly specialized, wherein most tropical forests remain within those less documented [2,3,4,5,6], including the Neotropical realm.

The Seasonally Dry Forest of Mexico has a discontinuous distribution throughout the country and provides complex habitats supporting a diverse endemic leafhopper fauna strongly associated and not spread beyond forest boundaries [7,8]. Dry tropical forests around the globe, including the Mexican section, are among the most affected ecosystems in the world by constant pressures of industrialization, livestock, and deforestation [9,10]. According to Pinedo-Escatel et al. [2], Aguilar-Pérez et al. [11], and Pinedo-Escatel and Dietrich [12] the Seasonally Dry Forest of Mexico harbors 65% of species of the tribe Athysanini, a higher number than any other country in the Americas.

The Mexican endemic Athysanini leafhopper genus Renonus DeLong, 1959 was described with the inclusion of Renonus rubraviridis DeLong, 1959, based on two males collected in the State of Guerrero, Mexico; the species was included in the genus as a monotypic type. Most specimens were taken in the early or late 1930s over three collecting events: first in 1930 by Parra, second in 1938 by Dampf, and last in 1939 by DeLong. Two decades later, using such a small series of individuals, it was formally described [13]. The species was known solely from five localities in the Mexican states of Guerrero, Tamaulipas, and Tabasco. Since then, no additional data on whether the genus or species has been reported, even though long-term fieldwork was performed in the formal revision of Athysanini from Mexico [4], which did not provide information on this taxon besides the current historical knowledge.

The goal of this contribution is to summarize the current knowledge of the Mexican endemic leafhopper genus Renonus and to describe a new species, Renonus cuixmalensis Pinedo-Escatel sp. nov., which was found inhabiting the Biosphere Reserve of Chamela-Cuixmala and the surroundings of the Estación de Biología Chamela of the Universidad Nacional Autónoma de México (IBUNAM). After meticulous and detailed morphological examination of all available materials from recent fieldwork, including the type species of the genus, in accordance with the redefinition presented by Pinedo-Escatel et al. [4] for Athysanini to occur in Mexico, it was possible to allocate unique features to distinguish the genus Renonus and expand knowledge leading to the separate and discriminated morphological identification of the new species from its congener species. A first dichotomous key and a map of distribution for Renonus are given below.

2. Materials and Methods

2.1. Leafhopper Sampling

Sampling was performed contemplating trips in 2001, 2017, 2024, and 2025. The collecting methods used were (a) fogging, (b) swept net, and (c) light trap. Net diameter was 50 cm and 70 cm in depth, and light was running from 7 p.m. until 3 a.m., following the standard protocol of Aguilar-Pérez et al. [11]. The net per site comprises 800 sweeps in total, while the light trap used an 800-watt bulb running from 7 p.m. to 1 a.m., reflecting light to a blank sheet. Fog is based on a single application at 4 a.m. to one targeted tree. Specimens captured using both methodologies were placed in a glass vial with 95% ethanol. Collecting sites lie between an altitude of 40 to 500 masl. covered by deciduous vegetation, which broadly spans other regions of Western Jalisco and most areas of the municipality of La Huerta, where specimens were taken. Collecting sites lie between an altitude of 40 to 500 masl. covered by deciduous vegetation (see below in habitat), which broadly spans to other regions of Western Jalisco and most areas of the municipality of La Huerta. Collecting events vary; one location with multiple points was run in the perimeter of the Estación de Biologiá Chamela, UNAM, and a few others beyond the limits. A swept net was focused on climbing trees to collect individuals near the canopy, while light trapping was always positioned in proximity to the tallest trees with a clear window without vegetation from the ground facing canopies.

2.2. Morphological Terminology and Criteria

Overall terminology herein follows Dietrich [14], wing venation follows the system proposed by Anufriev and Emeljanov [15], and leg chaetotaxy follows Rakitov [16]. Nomenclatural changes and valid names followed Dmitriev et al. [1], Oman et al. [17], Zanol [18], and Linnavuori [19]. Description of the body color is based on dry pinned specimens.

2.3. Procedures of Specimen’s Preparation

Abdomens of all specimens were processed into a morphological study by using external and internal features following the conceptualization based on Pinedo-Escatel et al. [4]. Male abdomens were removed carefully from each individual pinned, then submitted to a standard leafhopper clearing protocol with the following modifications: (i) abdomens immersed overnight in a cold 15% KOH solution, (ii) rinsed five times with distilled water, (iii) soaked with clove essence, and (iv) rinsed one time with ethanol at 100% to remove any remaining KOH. Terminalia was retained in a glass microvial with pure glycerin and pinned beneath its respective specimen. Labels are literally quoted exactly as written. Total length was measured from the anterior margin of the crown to the tips of the forewings in the resting position.

2.4. Imaging, Edits, and Vectorization

Specimens’ photographs of overall external habitus (dorsal, ventral, and anterior views) and internal (male and female genitalia) genitalia were taken at the Laboratorio Nacional de Biodiversidad (LANABIO of the Instituto de Biología, Universidad Nacional Autónoma de México) using a stereomicroscope ZEISS Axio Zoom V 16 multifocal photography equipped with an AxioCam MRc5 camera and the ZEN 2012 program (blue edition). Measurements were extracted using an electronic vernier with the ZEN 2012 program. Equipment automatically takes about 50 series of multifocal images to generate a single image; then, the color and extra edition are performed in Photoshop^® version 24.1, respecting the color pattern based on pinned organisms.

The illustrations were generated using a stereomicroscope with a lucid camera attached, drawings were vectorized and edited using Photoshop Illustrator^®. A map of distribution in Mexico was projected using QGIS^® version 3.14, taking available georeferenced data known from all specimen labels. Each checklist entry provides information in the order as follows: taxon name, author and year of description, synonyms and original combination, citation, and distribution.

2.5. Entomological Repositories

The voucher specimens studied are housed at the following entomological collections:

CNIN	Colección Nacional de Insectos, Instituto de Biología, Universidad Nacional Autónoma de México, Mexico City, Mexico.
USNM	National Museum of Natural History, Washington DC, USA.
AMNH	American Museum of Natural History, New York, USA.
FSCA	Florida State Collection of Arthropods, Gainesville, Florida, USA.
CAS	California Academy of Sciences, San Francisco, California, USA.
NHM	National History Museum, London, UK.
OSUC	Ohio State C.A. Triplehorn Insect Collection, Ohio, USA.

The above acronyms used refer to material studied and deposited for each taxon, including the new species.

3. Results

Systematic Entomology

Family Cicadellidae Latreille, 1825

Subfamily Deltocephalinae Dallas, 1870

Tribe Athysanini Van Duzee, 1892

Genus Renonus DeLong

  Renonus DeLong, 1959: 325.

Type species

Renonus rubraviridis DeLong, 1959.

Morphological remarks

Body length: males between 3.88 mm and 4.24 mm and females 4.24 mm (only known for R. cuixmalensis sp. nov.). Body pale with orange bands and stripes on crown and pronotum. Head: Slightly conical, somewhat produced and upturned; frontal sutures reaching ocellus centrally at bottom. Crown depressed and surface shagreened. Anteclypeus widening apically.

Pronotum: With lateral margin weakly carinate; without transverse striations. Forewing macropterous with three anteapical cells, inner anteapical cell open and outer anteapical cell truncate, several reflexed costal veins; A1 and A2 convergent medially; appendix restricted to anal margin. Profemur with only AM1 present at medial height; AD + PD seta 1 + 4. Metafemur setal formula 2 + 2 + 1. Segment X is membranous but laterally weakly sclerotized.

Pygofer: Incised near to base; macrosetae well differentiated into 3 or 4 rows; without processes. Subgenital plate: Subtriangular with uniseriate macrosetae laterally. Connective: Y-shaped, stem short without processes. Style: Bilobed at base, apophysis short with blunt apex. Aedeagus: curved dorsad with a pair of medial or preapical processes.

Species list of Renonus

Renonus cuixmalensis sp. nov.

    Distribution: Mexico

     Records: Biosphere Reserve of Chamela-Cuixmala; La Huerta (Jalisco).

     Repositories: Colección Nacional de Insectos (CNIN); National Museum of Natural History (USNM); American Museum of Natural History (AMNH); Florida State Collection of Arthropods (FSCA); California Academy of Sciences (CAS); National History Museum (NHM).

Renonus rubraviridis DeLong, 1959

    Distribution: Mexico

     Records: Iguala, Zincauro, and Santo Tomas (Guerrero); El Mante (Tamaulipas); Reforma (Tabasco).

     Repositories: Ohio State C.A. Triplehorn Insect Collection (OSUC); Colección Nacional de Insectos (CNIN).

Species description and new data

Renonus cuixmalensis sp. nov.

Figure 1, Figure 2, Figure 3, Figure 4 and Figure 5

Type locality

Biosphere Reserve of Chamela-Cuixmala (details below in material reviewed and habitat).

Description

Color: Body coloration pale with orange bands and stripes on the dorsum; wings brownish with hyaline veinlets.

Head: Crown pale with a broad transverse orange band (Figure 1A). Frontoclypeus mostly pale with a yellowish transverse band near the ecdysial line, surface weakly shagreened. The anteclypeus and lora have pale and light yellowish marks. Gena with the upper region near the eyes light yellowish and the remaining pale (Figure 2A).

Thorax: Pronotum with a broad transverse orange band with four longitudinal orange stripes interconnected near the anterior margin. Scutellum mostly orange with lateral pale marks (Figure 1A,B). Forewing brownish with translucent veinlets. Legs pale with yellowish dorsum and brownish setae. Venter yellowish, but dorsum of abdomen mostly light brownish with black specks distally (Figure 1A,B).

Figure 1. Male of Renonus cuixmalensis sp. nov. (A) Habitus, dorsal view. (B) Habitus, lateral view.

Figure 1. Male of Renonus cuixmalensis sp. nov. (A) Habitus, dorsal view. (B) Habitus, lateral view.

Male genitalia: Pygofer: 1.1× longer than high, posterior margin slightly pointed, without processes (Figure 3A), 3–4 stout elongate macrosetae near the second third of the pygofer distant from the posterior margin, minute fine setae on ventral near the ventral margin. Anal tube: As long as the pygofer with membranous apex, wider at base, laterally weakly sclerotized, with fine setae at apex (Figure 3B).

Valve: Wide at base, with anterior margin almost straight and posterior pointed. Subgenital plate: Triangular, mesal margin sclerotized, apex pointed, 14–18 uniseriate long fine setae (Figure 3C).

Style: Short, bilobed at base, medial lobe poorly developed, apophysis not elongated, blunt apex, without fine setae or tooth (Figure 3D). Connective: Y-shaped, stem short. Aedeagus: Wide, tubular, prominent in male capsule, curved dorsal, strongly sclerotized, apex truncate, a long pair of processes on posterior margin directed caudad with sharp tip (Figure 3E). Gonoduct tubular: Poorly defined, and gonopore ovoid in form, positioned near apical margin.

Female genitalia: Sternite VII slightly notched centrally.

Etymology

The species epithet is based on the Náhuatl name of the region, Cuixmala, meaning “place where the soul rests”, where specimens were taken inside the polygon of the Biosphere Reserve of Chamela-Cuixmala.

Distribution

Restricted to Mexico (Jalisco state), Figure 8.

Habitat

Seasonally Tropical Dry Forest, Figure 9.

Figure 2. Anterior view of Renonus spp. (A) R. cuixmalensis sp. nov. (B) R. rubraviridis DeLong.

Figure 2. Anterior view of Renonus spp. (A) R. cuixmalensis sp. nov. (B) R. rubraviridis DeLong.

Measurements, observed range of males (mm)

Total specimens = 12. Body: length, 3.88–3.90; width, 1.14–1.16. Head: width, 1.30–1.32; mid-length, 0.36–0.37; eye width 0.43–0.43; eye length 0.60–0.62; crown mid-width before eyes, 0.75–0.78; crown posterior width between eyes, 0.62–0.62; distance between ocelli, 0.64–0.65; frontoclypeus width, 0.66–0.67; frontoclypeus length, 0.66–0.67; anteclypeus width, 0.20–0.21; anteclypeus length, 0.31–0.31; lorum width, 0.16–0.16; lorum length, 0.24–0.24; gena width, 0.26–0.26; gena length, 0.50–0.50. Pronotum: width, 1.14–0.16; length, 0.55–0.56. Scutellum: width, 0.78–0.79; length, 0.52–0.53. Forewing: length, 3.01–3.05.

Measurements, observed range of female (mm)

Total specimens = 3. Body: length, 4.24–4.27; width, 1.32–1.34. Head: width, 1.38–1.39; mid-length, 0.38–0.39; eye width, 0.43–0.44; eye length, 0.67–0.67; crown mid-width before eyes, 0.78–0.79; crown posterior width between eyes, 0.66–0.68; distance between ocelli, 0.64–0.68; frontoclypeus width, 0.71–0.72; frontoclypeus length, 0.69–0.70; anteclypeus width, 0.21–0.22; anteclypeus length, 0.33–0.33; lorum width, 0.19–0.19; lorum length, 0.24–0.24; gena width, 0.29–0.29; gena length, 0.55–0.55. Pronotum: width, 1.21–1.23; length, 0.59–0.60. Scutellum: width, 0.81–0.83; length, 0.53–0.54. Forewing: length, 3.26–3.29.

Figure 3. Male genitalia of Renonus cuixmalensis sp. nov.; (A) Pygofer, lateral view. (B) Pygofer, dorsal view. (C) Subgenital plate and valve, ventral view. (D) Style, ventral view. (E) Aedeagus, lateral view.

Figure 3. Male genitalia of Renonus cuixmalensis sp. nov.; (A) Pygofer, lateral view. (B) Pygofer, dorsal view. (C) Subgenital plate and valve, ventral view. (D) Style, ventral view. (E) Aedeagus, lateral view.

Material examined

Holotype ♂ (CNIN)—MEXICO: Jalisco, La Huerta, Reserva de la Biosfera Chamela-Cuixmala, Estación de Biología Chamela UNAM, 19°29′56.9″ N 105°02′31.6″ W, 67 m, 4 Noviembre 2017, Torres-Moreno Col., Red Entomologica [MEXJAL09].

Figure 4. Female of Renonus cuixmalensis sp. nov. (A) Habitus, dorsal view. (B) Habitus, lateral view.

Figure 4. Female of Renonus cuixmalensis sp. nov. (A) Habitus, dorsal view. (B) Habitus, lateral view.

Figure 5. Renonus cuixmalensis sp. nov. at Estación Biológica Chamela, Instituto de Biología, UNAM. Photo courtesy of Emmanuel Rodríguez Rojas.

Figure 5. Renonus cuixmalensis sp. nov. at Estación Biológica Chamela, Instituto de Biología, UNAM. Photo courtesy of Emmanuel Rodríguez Rojas.

Paratypes 11♂♂ and 3♀♀; 2♂♂ and 2♀♀ (CNIN)—same data as holotype [MEXJAL09 and MEXJAL020]; 1♂ and 1♀ (CNIN), 1♂ (NHM)—MEXICO: Jalisco, near La Huerta, 480 m, 19°27′47″ N 104°39′13″ W, 15 Oct 2001, S. H. McKamey et al. Colls., fogging; 3♂♂ (CNIN).

1♂ (USNM), 1♂ (AMNH), 1♂ (FSCA), and 1♂ (CAS)—MEXICO: Jalisco, La Huerta, Reserva de la Biosfera Chamela-Cuixmala, Estación de Biología Chamela UNAM, 19°29′58.6″ N 105°02′18.0″ W, 39 m, 27 Julio 2024, Pinedo-Escatel J.A. Col., Red Entomologica [MEXJAL347].

Remarks

The new species is externally very similar to R. rubraviridis, but the disposition of aedeagal processes is a key distinguishable character between these species.

Renonus rubraviridis DeLong, 1959

Renonus rubraviridis DeLong, 1959: 326.

Figure 6 and Figure 7.

Remarks

Body coloration is very similar to R. cuixmalensis but slightly darker in color (Figure 1A,B and Figure 2B) and differs in the combination of features below.

Figure 6. Renonus rubraviridis DeLong. (A) Habitus, dorsal view. (B) Habitus, lateral view.

Figure 6. Renonus rubraviridis DeLong. (A) Habitus, dorsal view. (B) Habitus, lateral view.

Pygofer: 1.5× longer than high, posterior margin broadly rounded, without processes (Figure 7A), 3 stout macrosetae near the posterior margin, without minute fine setae on the ventral near the ventral margin. Anal tube: longer than pygofer and membranous, wider at mid-length, dense fine setae at apex (Figure 7A). Valve wide at base, with anterior margin straight and posterior rounded. Subgenital plate: slender and triangular, mesal margin weakly sclerotized, apex pointed, 15–17 uniseriate long fine setae.

Figure 7. Male genitalia of Renonus rubraviridis DeLong. (A) Pygofer, lateral view. (B) Style, ventral view. (C) Aedeagus, lateral view.

Figure 7. Male genitalia of Renonus rubraviridis DeLong. (A) Pygofer, lateral view. (B) Style, ventral view. (C) Aedeagus, lateral view.

Style: Moderately short, bilobed at base, medial lobe poorly developed, apophysis not elongated, apex digitate, with two fine setae on outer margin, without tooth (Figure 7B). Aedeagus: Wider at base, narrowing towards apex, not particularly prominent in male capsule, recurved, strongly sclerotized, apex sharply pointed, stout pair of processes arising on posterior margin well directed anterad (Figure 7C). Gonoduct: Wide, poorly sclerotized. Gonopore: Ovoid on subapical margin.

Measurements, male (mm)

Total specimens = 2. Body: length, 4.15–4.16; width, 1.17–1.18. Head: width, 1.33–1.34; mid-length, 0.35–0.36; eye width, 0.41–0.42; eye length, 0.60–0.61; crown mid-width before eyes, 0.79–0.80; crown posterior width between eyes, 0.70–0.71; distance between ocelli, 0.66–6.67; frontoclypeus width, 0.67–0.67; frontoclypeus length, 0.69–0.69; anteclypeus width, 0.19–0.19; anteclypeus length, 0.32–0.32; lorum width, 0.16–0.16; lorum length, 0.30–0.30; gena width, 0.26–0.26; gena length, 0.55–0.55. Pronotum: width, 1.19–1.21; length, 0.56–0.58. Scutellum: width, 0.87–0.90; length, 0.59–0.61. Forewing: length, 3.20–3.22.

Type of material examined

Holotype ♂ (OSUC)—MEXICO: Iguala, Guerrero, 11 September 1939, D. M. DeLong Coll.

Paratype ♂ (OSUC)—MEXICO: Iguala, Guerrero, 2 September 1930, J. Parra Coll.

Other material examined

1 ♂ (CNIN)—MEXICO: Guerrero, Zirándaro, Zirándaro de los Chávez, 18°28′8″ N 100°58′55″ W, 216 m, 15 Abril 2025, Vega-Ovando F.R. Col., Trampa Luz [MEXGUE401].

Distribution

Reported in the Mexican states of Tamaulipas, Tabasco, and Guerrero, Figure 8.

Figure 8. Distribution of Renonus over the Seasonally Tropical Dry Forest of Mexico.

Figure 8. Distribution of Renonus over the Seasonally Tropical Dry Forest of Mexico.

Key to male species of Renonus

1.

Posterior margin of pygofer pointed; style apex digitate; apex of aedeagus rounded; shaft with medial processes on posterior margin strongly directed anterad; ………………………………………………………………… Renonus rubraviridis DeLong, 1959

-

Posterior margin of pygofer truncate; style apex slightly pointed; apex of aedeagus acute; shaft with preapical processes on posterad margin directed caudad………………………………………………………………………… Renonus cuixmalensis sp. nov.

4. Discussion

The genus Renonus has a particular distribution in Mexico, where most records have been reported mainly from areas with almost complete coverage of Seasonally Tropical Dry Forest (Figure 9); for R. rubraviridis, 75% of records are known in this ecosystem. In the case of the new taxon, and despite being collected with several capture methods, it was also consistently found in this ecosystem (Figure 8) throughout several trips to Western Mexico. However, notably, the abundance gathered from collecting effort is low using a net and flight interception trap, in contrast to a light trap, which was the most successful method (70%), suggesting an arboreal preference as stated with data reported [2,3].

Figure 9. Vegetation surrounding the Biological Station of Chamela (IBUNAM) with Seasonally Tropical Dry Forest. Photo taken by Adrian Gómez-Jácome.

Figure 9. Vegetation surrounding the Biological Station of Chamela (IBUNAM) with Seasonally Tropical Dry Forest. Photo taken by Adrian Gómez-Jácome.

Renonus cuixmalensis sp. nov. is known from two zones that are relatively near to each other, and both lie in the Pacific Lowland province, based almost purely on the influence of Neotropical components; meanwhile, R. rubriviridis is known from five zones, of which two are from the Veracruzan province with strong elements of rainforest, and three are from the Balsas Basin province, a region known as one of the highest hotspots of data-based diversification for the tribe Athysanini in Mexico (Figure 10) and home to more than 25 genera, of which the external morphology is quite uniform, according to evidence provided by Pinedo-Escatel et al. [2,3,4].

Figure 10. Map of distribution of Renonus. Biogeographical provinces of Mexico shown in color.

Figure 10. Map of distribution of Renonus. Biogeographical provinces of Mexico shown in color.

Although several genera during the late 1940s to 1970s in Mexico were described and revised [20,21,22,23,24,25,26,27], the genus Renonus was largely omitted in most, and even those vast contributions performed on Neotropical fauna, e.g., Linnavuori [19] and Oman [27], did not provide informative data [28].

Although these two species of Renonus are very similar externally, slight differences persist in body size, and they cannot be differentiated using overall color pattern. To allocate species, we need to examine genitalia using (I) the shape of the pygofer, (II) the style, and (III) the orientation of the aedeagal processes. The morphology of the genus by itself can differentiate it easily from other similar Mexican genera (e.g., Sanuca by DeLong [29], Stoneana by DeLong [30], or Bandara by Ball [31]), which display similar but closely shared external and internal features and must be separated using (a) the presence of a depressed crown, (b) a unique orange color pattern, (c) the absence of pygofer appendages, and (d) a single pair of aedeagal processes, which might segregate it from even other southern Neotropical ones [4]. Further work is needed to uncover species relationships among several genera that do not spread beyond the boundaries of the Balsas Basin province.