taxonID	type	description	language	source
33469A2EFFF7FFF3FF4AF9FBFEF932ED.taxon	diagnosis	Diagnosis: Vertex, supra-antennal ledges, and frontoclypeus curving ventrally and posteriorly in lateral view, concave in frontal view. Pronotum with metopidium curved posteriorly, forming semicircular shape; dorsal margin highly sinuous, with depression in apical third that rises and descends again before apex; pronotum shorter than forewings, reaching to slightly above apex of fifth apical cell.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF7FFF3FF4AF9FBFEF932ED.taxon	description	Description: Color and surface: Overall coloration ferruginous, dark brown, or black with bluish-iridescent reflections. Body coarsely punctate, each puncture bearing a golden seta. Forewings punctate and sclerotized in basal cells C and R, extending to just before the mid-length of the wing, and also on basal third of clavus. Abdominal terga coarsely punctate; cleared abdomens exhibit granular texture, with each granule translucent and more visible on sternites. Head (Fig. 2 a, 2 c, 3 a, 3 c). Vertex, supra-antennal ledges, and frontoclypeus curving ventrally and posteriorly in lateral view, concave in frontal view. Frontoclypeus nearly as wide as vertex, forming a semicircular outline ventrally and pressed against anteclypeus. Vertex approximately 1.5 x wider than long. Thorax (Fig. 2, 3). Pronotum in lateral view with dorsal margin of metopidium curved posteriorly, forming a semicircular shape, followed by a highly sinuous dorsal margin; a marked depression present in apical third, which then rises and descends again before apex. Pronotum shorter than forewings, reaching to slightly above apex of fifth apical cell. Dorsomedian carina elevated; lateral or accessory carinae absent. In frontal view, humeral angles weakly projected; dorsal margin dome-like. Postocular lobes extending to near ventral margin of eyes. Forewing with R branching into R 1 and Rs near the base; two discoidal cells present; one s, one r-m, and two m-cu crossveins; s crossvein located distad to r-m (Fig. 2 b, 3 b). Hindwing with one r-m crossvein and one m-cu crossvein, M divided in M 1 + 2 and M 3 + 4 (Fig. 4 a). Metafemur with spine-like cucullate seta at apex. Pro- and mesotibiae dorsally flattened and slightly expanded, but not foliaceous. Metatibiae with posterodorsal row of cucullate seta bearing spine-like setae; anterodorsal row with approximately 4 – 5 spine-like cucullate setae; ventral row with small cucullate setae (Fig. 2 f). First metatarsomere with cucullate setae at apex of ventral surface. Abdomen (Fig. 4, 5, 6). Terga, pleurites, and sternites III – VII partially fused. Dorsum of terga III – VI each with a pair of small tuberosities. Sternum III with one pair of depressions and one central, keel-like elevation. Sternum VII slightly emarginate. Female genitalia. Pygofer with pointed lobule on posterodorsal margin, near anal tube. Gonoplac subrectangular, short (less than 3 × longer than wide), with setae and acanthae along entire surface. First valvulae robust, short, thick at base; dorsal margin tapering to rounded apex with apparent opening; ramus near dorsal margin; pores near base; ventral sculptured area with linear tegumentary processes on inner and ventral surfaces. Second valvulae short, robust, subovoid, in lateral view gradually expanded toward midlength, then tapering slightly toward apex; apex with dorsally directed hood-like structure (apical hood), laterally expanded and rounded in dorsal view; ramus near ventral margin at base, extending dorsally toward apex; valvulae forming rhomboidal aperture in ventral view (ventral aperture); pores restricted to apical hood and area surrounding ventral aperture.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF7FFF3FF4AF9FBFEF932ED.taxon	discussion	Remarks. Morphology: Paragara nigra and P. tholoidea share a very similar head and pronotal morphology. The distribution of maculae on the forewings differs only slightly in the intensity of pigmentation. One of the specimens collected and photographed in the field (Fig. 2) matches the coloration of the P. nigra holotype (Fig. 1 a – c), while a specimen examined from the ICN collection (Supplementary Fig. 1) corresponds to the coloration of the P. tholoidea type (Fig. 1 e – g). The overall coloration of P. nigra appears to be slightly darker than that of P. tholoidea, with darker and more pronounced wing maculae. Beyond this character, we found no other distinguishing features between the two species. In some cases, these differences in coloration could be attributed to factors such as collection method, preservation, degree of sclerotization, developmental stage, or even association with different mutualistic ant species. However, since the P. nigra specimens we collected in the field consistently share the same coloration and given the limited number of P. tholoidea specimens examined, we do not consider these species to be synonymous at this time. The abdominal terga with coarse punctation (Fig. 4 a – b) are similar to those of other membracids morphologically similar to Bolbonota, Tritropidia, some Leioscyta Fowler and Tropidoscyta Stål species, and Eunusa (Fig. 8 f). This coarse abdominal punctation is also present in other membracid groups, such as Hypsoprorini (see figures 29 – 31 in Creão-Duarte et al. 2017) and Boocerini (Wallace & Deitz 2004). The sternal and pleural regions lack coarse punctation but exhibit fine, translucent punctures and a high density of acanthae (Fig. 4 d). In addition, the pair of depressions on sternite IV (Fig. 4 d) is commonly found in other genera of Membracini. The female genitalia in Paragara are similar to Bolbonota, Tritropidia, and some Leioscyta and Tropidoscyta species, in which the genitalia are relatively short in proportion to abdominal length, which is often associated with oviposition on the surface instead of inside host tissue (as it is in Paragara). In Paragara, the gonoplac is no more than three times longer than high, unlike the condition found in most Membracidae, where it typically exceeds a ratio of 4: 1. The gonoplac almost completely covers the first and second valvulae, resembling the subgenital plate of males. As a result, distinguishing females from males is difficult in the field and even under laboratory conditions (Fig. 4 b, c, e, f). The second valvulae bear a dorsoapical structure resembling a hood that partially covers the remainder of the valvula (Fig. 5 d – g, 6 f – i )). Interestingly, this hood-like structure appears to have associated pores. This condition differs from most membracids, in which this area is exposed and typically bears small teeth on the dorsal margin. The distribution of pores in the second valvulae of this group is distinctive, being concentrated on the hood and the ventral surface, rather than distributed along the ramus or restricted to the dorso-apical region. Additionally, a prominent rhomboid-shaped opening is present ventrally, where the valvulae do not fully converge (Fig. 6 g). These features may be associated with both oviposition and the secretion of large amounts of wax over the eggs. The first valvula is also robust and short, with a rounded apex and an apparent opening (Fig. 5 b-c, 6 c – e). The ramus is positioned near the dorsal margin, and the pores are located close to the base. In ventral view, the basal half of the valvula appears somewhat globose, with linear tegumentary processes on the inner and ventral surfaces, a feature that differs markedly from other membracids for which the female genitalia have been described in detail (e. g. Creão-Duarte et al. 2017, Flórez-V 2019). Previous studies have noted shared morphological traits — such as a short ovipositor, sternum VII slightly emarginate and a slightly notched second sternite — among genera related to Bolbonota — and have even treated them collectively under the tribe Bolbonotini (Lin et al. 2004; Metcalf & Wade 1965). The latter authors formally resurrected Bolbonotini to include Bolbonota, Bolbonotodes Fowler, Erechtia Walker, Paragara, and Tritropidia. However, this classification has not been followed in subsequent studies, likely due to the lack of phylogenetic analyses including several key Membracinae genera such as Bolbonotodes, Eunusa, Havilandia Dietrich & McKamey, Lewdeitzia Dietrich & McKamey, Paragara, and Talipes Deitz (Deitz & Wallace 2010). Furthermore, many taxa currently placed in Membracini incertae sedis (Sakakibara 2012), as well as the genera Erechtia, Tropidoscyta, and Leioscyta, must be included in future phylogenetic analyses to robustly resolve relationships among Membracini, “ Bolbonotini ”, and Talipedini.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF7FFF3FF4AF9FBFEF932ED.taxon	biology_ecology	Biology Paragara is ant-mutualistic; the two species found in the field, P. nigra and P. chagrana sp. nov., were each observed in association with a single ant species: Dolichoderus bidens (Linnaeus, 1758) and D. septemspinosus Emery, 1894 (Dolichoderinae), respectively. In both cases, the general coloration of the treehoppers resembled, to some extent, that of their associated ants. Adults of both Paragara species were also found co-occurring with other membracids, sometimes forming mixed-species aggregations (Fig. 10 b), which may be advantageous to attracting ants. Females oviposit eggs superficially on plant tissue and cover them with a wax-like secretion (Fig. 10 a, 12 ab). The wax-covered egg mass forms an oval mound, with a striated pattern radiating outward from a central longitudinal line (Fig. 10 a). This structure closely resembles the wax patterns seen in egg masses of Leioscyta spiralis, Tritropidia spp. and cf. Tropidoscyta (Supplementary Fig. 2). It was also common (n = 9) to find clustered egg masses, with up to five laid in close proximity within a single area (Fig. 10 c, 11 c – d). It is unknown if these were oviposited from a single or multiple females. Egg masses of Paragara may be 2 – 3 times the length of the female’s body, and in several instances, females were found with greatly distended abdomens, markedly exceeding the length of the wings (Fig. 10 b, 10 d, 11 f). This capacity for abdominal expansion appears distinctive and has also been observed in females of Eunusa concolor Fonseca, which likewise produce disproportionately large egg masses relative to adult size. Thus, abdominal length relative to the pronotum and wings appears to be a highly variable character in these genera; in some females, the pronotum reaches the abdominal apex, while in others, the abdomen clearly surpasses this point. Females were observed depositing the wax-like substance, and some were found near their egg masses. However, no active defensive behavior was observed (e. g., sitting over the eggs, kicking, or producing vibratory buzzing). Instead, these treehoppers displayed a characteristic movement in which they raised and oscillated the hind legs back and forth (Fig. 10 b). This behavior has been described in other ant-associated membracids, such as Thuris, many tragopines (Smiliinae) and some biturritines (e. g. Biturritia Goding, Lophyraspis Stål and Mina Walker; Aetalionidae) and has been hypothesized to mimic ant antennal movements, potentially enhancing camouflage within ant aggregations (Quintero-M & Flórez-V 2025). Differences in coloration between species appear to correlate with the identity of their respective mutualistic ant partners. It is likely that ants exert strong selective pressure on the evolution of treehopper morphology and coloration. Ants play a fundamental role in the ecology of lowland membracids (Olmstead & Wood 1990) and are also among the dominant insect predators in lowland forest canopies (Camacho & Avilés 2019). By forming mutualistic associations with these dominant predators, treehoppers may reduce predation pressure (Wood 1984). In some cases, host plants could influence coloration differences. However, our observations of P. nigra show that coloration remains consistent despite the use of different host plants. The same pattern occurs in P. chagrana, which can occur on Vismia Vand. (Hypericaceae), Heliconia L. (Heliconiaceae), and several Melastomataceae, yet its coloration remains consistent. In many cases, particularly among species with high ant dependency and specialized ant mutualisms, treehoppers exhibit color patterns and morphological traits that resemble their associated ants. For example, certain species such as Paragara, Thuris, and some Lophyraspis have sinuously contoured dorsal profiles (Supplementary Fig. 3), which may mimic the narrow waist (petiole) between the mesosoma and gaster of ants. In terms of coloration, one of the most striking cases involves treehoppers associated with Cephalotes atratus, which typically exhibit black coloration and / or yellowish-brown maculae resembling the frontal lobes of the ants. Examples include Heteronotus sp. (Perger 2021), Anobilia nigra Tode, Todea peruviana (Funkhouser) and females of Phormophora maura (Fabricius), among others (Flórez-V pers. obs.). This was first reported by Richter (1945) after finding the association and morphological similarity between the gasters of Dolichoderus analis Emery (now as Dolichioderus quadridenticulatus (Roger, 1862 )) attending Tropidolomia involuta (Coquebert) (erroneously identified as Tragopa involuta; it corresponds to a species close to Anobilia guianae or Tragopa zebra) and of Cephalotes atratus (L.) attending Todea peruviana. This potential for morphological and chromatic convergence with ant partners is not limited to adults, but can also be observed in the nymphal stages of some species. For instance, the nymphs of Paragara chagrana sp. nov. and Thuris binodosus (Goding) share a remarkably similar texture, characterized by very short setae (Supplementary Fig. 3). Their coloration in field-collected specimens is also strikingly similar, both to each other and to their mutualistic ant, D. septemspinosus (Fig. 11 e – g). This resemblance may be related to that both treehopper species are mutualists of the same ant species. In addition, the nymphs of both P. chagrana and T. binodosus have long legs, particularly the hind legs, a trait common in several ant-associated treehopper species in which nymphs exhibit similar hind-leg movements as the adults (see above). These observations suggest that mimicry in some ant-associated treehoppers may be multimodal, involving not only visual resemblance (in shape and coloration), but also behavioral elements such as movement patterns. In some cases, such mimicry could enhance camouflage within the ant-attended aggregation, and reduce aggression by the tending ants. Although chemical mimicry has not yet been documented in Membracidae, similar strategies are well known in other mutualistic systems, such as lycaenid butterflies and their ant hosts (Pierce et al. 2002). This raises the possibility that multiple forms of mimicry — visual, behavioral, and perhaps chemical and through vibrational signals — may act in concert to reinforce and stabilize these strong and specialized mutualistic interactions.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF2FFF1FF4AF8FBFB9E35FC.taxon	description	(Fig. 1 e – h, Supplementary Fig. 1)	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF2FFF1FF4AF8FBFB9E35FC.taxon	diagnosis	Diagnosis: Overall coloration ferruginous with golden pubescence. Supraocular callosities pubescent. Forewing without, or with only a faint, basal and subapical transverse dark macula. In dorsal view, lateral margins of pronotum converge at approximately 40 ° to apex of posterior process.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF2FFF1FF4AF8FBFB9E35FC.taxon	materials_examined	Material examined and distribution (Fig. 14): Holotype female (photo): ECUADOR: Napo River: “ Paragargara \ tholo [letters difficult to transcribe] \ [female symbol] type ”, “ New Genus ”, “ Goding \ bequest \ 1934 ”, “ USNMENT \ 01113120 ”, “ Deitz Research \ 71 - 3639 abdomen missing ”. Other specimens examined: COLOMBIA: (1 female in ICN). Also, two records from iNaturalist in French Guiana from Elendil Cocchi show individuals that appear to be morphologically intermediate between P. tholoidea and P. nigra (https: // colombia. inaturalist. org / observations / 265033083 and https: // colombia. inaturalist. org / observations / 75457211). For the moment, we prefer to maintain it as P. tholoidea.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF2FFF1FF4AF8FBFB9E35FC.taxon	discussion	Remarks: As noted by Deitz (1975) and visible in the photographs of the P. tholoidea holotype, part of the posterior process of the pronotum, the left forewing, several legs, and the abdomen are missing from the specimen (Fig. 1 f).	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF0FFF1FF4AFD90F926327C.taxon	description	(Fig. 1 a – d, 2, 4, 5, 10)	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF0FFF1FF4AFD90F926327C.taxon	diagnosis	Diagnosis: Overall coloration dark brown with golden pubescence. Supraocular callosities pubescent. Forewing with a transverse dark brown macula crossing the apex of basal cells R and M, apex of second discoidal cell, and base of fifth apical cell, completely covering first discoidal cell; base of the wing with one distinct dark macula in basal M and Cu cells and covering almost completely clavus. In dorsal view, lateral margins of pronotum converge at approximately 40 ° to apex of posterior process.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF0FFF1FF4AFD90F926327C.taxon	description	Measurements: Female (CBUCES-F 17394) (mm): Body length: 3.5; body length (distended abdomen): 3.8; pronotum length: 2.8; pronotum height: 1.1; forewing length: 2.9; distance between humeral angles: 1.6; head width: 1.3; head height: 0.8.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF0FFF1FF4AFD90F926327C.taxon	biology_ecology	Biology: Paragara nigra was consistently found (n = 11) in association with D. bidens at both Leticia and Mitú (Fig. 10). Females of P. nigra were observed in proximity to Tropidolomia auriculata (Olivier) (Fig. 10 c), Neotynelia nigra (Funkhouser) (Fig. 10 b), and Anobilia aff. guianae (Haviland). These mixed-species aggregations were all associated with D. bidens. In some cases, egg masses of different species were found in close proximity, with females of T. auriculata and A. aff. guianae guarding their eggs while females of P. nigra were observed secreting wax over their own (Fig. 10 c). Despite forming such mixed-species aggregations, the number of P. nigra individuals was low; typically, only one or two adults were present, surrounded by numerous ants. Most of these mixed aggregations were found on Vismia, and on two occasions, on an unidentified vine.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF0FFF1FF4AFD90F926327C.taxon	materials_examined	Examined material and distribution (Fig. 14): Holotype female (photo) in USNM: PERU: “ Leonpampa \ Peru \ Dec. 1937 ”, “ WDFunkhouser \ Collection \ 1962 ”, “ HOLOTYPE \ Paragargara nigra \ W. D. Funkhouser ”. Other specimens examined: COLOMBIA: Amazonas: “ COLOMBIA. Amazonas, Leticia, antigua Estación Biológica El Zafire, 4.00286 ºS, 69.89558 ºW, 100 - 150 msnm, manual, IA 192, Jul. 27 / 2022, leg. M. Arcángel, C. Flórez-V, M. Idárraga, S. Quintero-M, CBUCES-F 16616 ” (1 female in CBUCES); other indivuals with the same data except by: “ IA 241, Ago. 3 / 2022, CBUCES-F 17382 ” (1 female in CBUCES), “ IA 245, Ago. 3 / 2022, CBUCES-F 17394 ” (1 female in CBUCES); Vaupés: “ COLOMBIA. Vaupés, Mitú, comunidad Cucura, 1.171402 ºN, 70.166708 ºW, Manual, IM 64, Feb. 20 / 2024, leg. Camilo Flórez-V, José Fernando Londoño, Santiago Quintero-M, CBUCES-F 20404 “ (1 female in CBUCES).	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF0FFF1FF4AFD90F926327C.taxon	discussion	Remarks: Funkhouser (1940) described and illustrated a short carina above humeral angles, but that was an error; it is not in the photo of the type.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFFEFFF9FF4AFF40FA0E371C.taxon	description	urn: lsid: zoobank. org: act: 91 AC 862 E- 7550 - 43 CD- 857 B-C 33664 FE 22 DE (Fig. 3, 6, 7, 11, 12)	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFFEFFF9FF4AFF40FA0E371C.taxon	diagnosis	Diagnosis: Head, pronotum, rest of thorax, coxae, femora, tibiae, and abdomen black with bluish iridescent reflections.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFFEFFF9FF4AFF40FA0E371C.taxon	description	Description: Holotype female. Color and surface: Head, pronotum, rest of thorax, coxae, femora, tibiae, and abdomen black with bluish iridescent reflections. Eyes gray; ocelli golden. Body coarsely punctate, each puncture bearing a golden seta. Supraocular callosities glabrous. Forewing punctate and sclerotized in basal cells C and R, extending to just before mid-length of wing, and also on basal third of clavus; these areas, along with basal cell M, basal half of Cu, and basal two-thirds of clavus, are black-pigmented; black macula at apex of clavus; broad black transverse macula spans first apical cell and adjacent apical limbus area, first discoidal cell, apex of basal cell R, second discoidal cell, and basal half of second, third, and fourth apical cells; it also completely covers fifth apical cell and part of adjoining apical limbus; membrane and veins between basal and transverse maculae white; apical portion of wing transparent and hyaline, with yellow veins. Joints, apices of tibiae, and tarsomeres golden. Head: Eyes subtriangular in frontal and frontolateral view. Thorax: Pronotum relatively robust, in lateral view with dorsal margin of metopidium curved posteriorly, forming a semicircular shape, followed by a highly sinuous dorsal margin; marked depression in apical third, which then rises and descends again before apex. Pronotum shorter than forewings, reaching a point above m-cu crossvein. Dorsomedial carina elevated; lateral or accessory carinae absent. In frontal view, humeral angles weakly projected; almost hexagonal. In dorsal view, lateral margins converge at approximately 50 ° at apex of posterior process. Postocular lobes extending to near ventral margin of eyes. Wings and legs as described for the genus. Abdomen: As described for the genus. Male unknown. Nymph (Fig. 7): Color and surface. Body oval and compact. Surface lacking chalazae and scoli; scattered very short setae present. Overall appearance dull; glossy black areas include supraocular callosities, forewing pads, legs, anal tube, and most of head except vertex area above ocelli, which bears a pair of oblique glossy lines. General coloration dark brown to black, with yellowish-brown on the coxae, trochanters, dorsal surface of the femora, margins of eyes, ocelli, and antennae. Head. Frontoclypeus, anteclypeus, and maxillary plates swollen (unlike adults); vertex slightly bilobed dorsally; rostrum reaching nearly to middle of abdomen. Thorax. Pronotum with rounded metopidium, followed by slightly sinuous dorsal margin; posterior process reaching fourth abdominal segment. Forewing pads with straight costal margin. Legs long relative to body size; tibiae distinctly extending beyond posterior margin of abdomen; hind tibia longer than pro- and mesotibiae. Abdomen. Abdomen length approximately half that of thorax. Abdominal terga each with one small mid-dorsal protuberance. Anal tube robust, cylindrical, lacking scoli, and nearly equal in length to preceding three abdominal segments. Measurements: Holotype female (mm): Body length: 3.3; pronotum length: 2.6; pronotum height: 1.1; forewing length: 2.7; distance between humeral angles: 1.9; head width: 1.5; head height: 1.0.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFFEFFF9FF4AFF40FA0E371C.taxon	biology_ecology	Biology: Paragara chagrana sp. nov. was consistently found (n = 16) in association with D. septemspinosus (Fig. 11, 12). This species was observed occasionally forming mixed-species aggregations with other membracid species, including Thuris binodosus (Fig. 12 f), Aphetea Fowler (Fig. 12 d), Anobilia aff. guianae (gray and black morph) (Fig. 12 d), and Neotynelia banderai Creão-Duarte & Sakakibara (Fig. 12 c). In some of these aggregations, egg masses of different species were found in close proximity. Paragara chagrana sp. nov. was collected from various host plants, including branches of several Melastomataceae species, leaves and stems of Vismia, and inflorescences of Heliconia. As with P. nigra, large aggregations of P. chagrana were not observed; typically, only 2 – 3 adults were present, and in just two cases were 4 – 5 nymphs found (on Heliconia and Vismia). However, these few individuals were consistently surrounded by numerous ants (typically 5 – 10 ants per treehopper). On one occasion, a parasitoid wasp (cf. Mymaridae) was observed near an egg mass while a female P. chagrana was depositing the wax-like substance over the eggs on the underside of an apical Vismia leaf. The female was surrounded by more than five ants, which moved rapidly over the leaf and nearby branch (Fig. 11 a), where other membracid species were also present. Despite walking close to or even over the wasp, the ants did not attack or chase it away (Fig. 11 b). The wasp approached the eggs and appeared undeterred by the presence of the ants or the wax-like secretion. Even after the female left the egg mass, the ants remained with the eggs. Although we were unable to determine exactly how long the ants stayed with the clutch, they remained for at least two days. These same plants also hosted other treehopper species, including females guarding eggs (Anobilia aff. guianae), solitary adults, and in some cases, aggregations of nymphs and adults. The tendency of multiple treehopper species to oviposit on the same plant may represent a strategy to retain ant attention on that plant, or at least on a specific area of it, thereby enhancing the protection of their offspring through collective association with ant defenders.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFFEFFF9FF4AFF40FA0E371C.taxon	materials_examined	Examined material and distribution (Fig. 14): Holotype female in CBUCES. COLOMBIA: Amazonas: Leticia: “ COLOMBIA. Amazonas, Leticia, antigua Estación Biológica El Zafire, 4.00286 ºS, 69.89558 ºW, 100 - 150 msnm, manual, IA 267, Ago. 4 / 2022, leg. M. Arcángel, C. Flórez-V, M. Idárraga, S. Quintero-M, CBUCES-F 17498 ” (1 female in CBUCES). Paratype females in CBUCES. Same data as holotype except by: “ IA 268, CBUCES-F 17500 ” (1 female in CBUCES-F), “ IA 292, Ago 6 / 2022, CBUCES-F 16660 ” (1 female in CBUCES), “ IA 34, Jul. 14 / 2022, CBUCESF 15180 ” (1 female in CBUCES). Other specimens examined: One nymph examined with the same holotype information.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFFEFFF9FF4AFF40FA0E371C.taxon	etymology	Etymology: The specific epithet chagrana refers to the “ chagra, ” traditional agroforestry plots maintained by Indigenous peoples in the Amazon, where crops such as cassava (Manihot esculenta), coca (Erythroxylum spp.), pineapple (Ananas comosus), plantains (Musa sp.) and many other edible plants are cultivated. Some of the specimens herein collected of this species were collected in or near these cultivated areas, highlighting its ecological and cultural association with this landscape.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFFEFFF9FF4AFF40FA0E371C.taxon	discussion	Remarks: The pronotum of P. chagrana sp. nov. is broader than that of P. nigra, with greater width-to-length and width-to-height ratios, giving it a more robust overall appearance (Fig. 3 a, 3 c). As a result, in dorsal view, the lateral margins of the pronotum converge at a less acute angle at the apex of the posterior process (approximately 50 °) (Fig. 3 e), compared to P. nigra (approximately 40 °) (Fig. 2 e). The wing coloration and pattern of maculae in P. chagrana are consistent among individuals observed in the field. No intermediate color forms were observed between P. chagrana and either P. tholoidea or P. nigra. Additionally, the glabrous supraocular callosities are unique to this species and show bluish iridescent reflections (Fig. 3 a), a trait also observed in the nymphs (Fig. 7 a-b). The female genitalia are very similar to those of P. nigra, with only minor differences in the degree of sclerotization. Nymphs of P. chagrana and T. binodosus are morphologically similar. As in the adults, the nymphs of T. binodosus have broadened and flattened maxillary and mandibular plates, while the frontoclypeus and anteclypeus are not as swollen. In addition, the nymphs of T. binodosus can be readily distinguished by a digitiform ventral extension on segment IX (Quintero-M and Flórez-V 2025; Supplementary Fig. 3 b). The type locality of this species is in the Amazon, near the tri-border region between Brazil, Colombia, and Peru (Fig. 14). It is therefore likely that the distribution of this new species extends into the neighboring territories of Brazil and Peru.	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF8FFE1FF4AFC48FE0E368C.taxon	description	(Fig. 8, 9, 13)	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF8FFE1FF4AFC48FE0E368C.taxon	discussion	Remarks. Morphology: Eunusa concolor is externally similar in overall morphology to Paragara, sharing features such as a head with the frontoclypeus pressed against the anteclypeus, a short pronotum lacking lateral carinae, a relatively sinuous dorsal margin, and a dark brown to brown body with golden pubescence. However, there are numerous morphological and ecological differences between the two genera. In E. concolor, the frontoclypeus and anteclypeus are proportionally smaller relative to head length, leaving the maxillary plates clearly visible in ventral view (Fig. 8 c). The supra-antennal margins are slightly elevated and do not form a continuous surface with the frontoclypeus, as they do in Paragara. The dorsal outline of the pronotum, while sinuous in lateral view, is considerably less contoured (Fig. 8 b) than in Paragara. The anterodorsal and posterodorsal rows of cucullate setae on the metatibia are spine-like and widely spaced (Fig. 8 b). In contrast, Paragara exhibits a posterodorsal row with more numerous and closely spaced spines, while the anterodorsal row bears only a few. The female genitalia of E. concolor are also markedly more elongate than those of Paragara, Bolbonota, and Tritropidia. This distinction was previously noted by McKamey (1992), who described the female genitalia of E. concolor as significantly longer than in these genera. The coloration of Eunusa concolor is variable; the maculae on the forewings may be more pronounced and extensive in some individuals than in others. Overall body coloration also tends to be lighter in younger, less sclerotized adults compared to older, more heavily sclerotized individuals. Variation is also observed in wing venation: some specimens have the M vein of the hind wing free (i. e., lacking a crossvein r-m), while in others it is not free (Fig. 8 g). The CuA vein is consistently free, with no m-cu crossveins present. Additionally, the M vein in E. concolor is unbranched (Fig. 8 g), whereas in Paragara it is bifurcated (Fig. 4 a). Eunusa concolor has a broader geographic distribution than Paragara, ranging from Central America and the Amazon Basin to the Guiana Shield and central Brazil (Mato Grosso) (Fonseca 1974; McKamey 1992). In contrast, Paragara is known only from the Amazon (Dmitriev et al. 2022), Guyana (vouchers in the United States National Museum of Natural History and the Illinois Natural History Survey), and French Guiana, with the latter based on iNaturalist records. The nymphs of Eunusa are characterized by a dense covering of peglike chalazae on the head, thoracic nota, and abdominal terga (McKamey 1992) (Fig. 9). The presence of a wax-like coating further distinguishes E. concolor nymphs from those of P. chagrana, which lack such a covering entirely, unlike most Membracini nymphs which are extensively covered with a wax-like coating (Lin 2006).	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF8FFE1FF4AFC48FE0E368C.taxon	biology_ecology	Biology Eunusa concolor is a mutualist of ants of the genus Azteca Forel. It inhabits protective shelters constructed by these ants, which can extend over large portions of the host plant. These structures have a dense, papery appearance and typically feature entrances and exits at the longitudinal ends (Fig. 13); in longer shelters, additional small openings may occur along the upper surface. These ant-built shelters housing E. concolor are usually located at plant nodes and axils, most frequently near the apices of the branches. Aggregations of E. concolor have been found on Vismia spp. (Hypericaceae), Bellucia Neck. and Miconia Ruiz & Pav. (Melastomataceae), typically in forestedge habitats. In Colombia, E. concolor has been observed across multiple regions, including the Magdalena Medio, the Amazonian foothills, and the Amazon Basin itself. In all cases, individuals were found inside protective shelters built by Azteca ants. This species inserts its eggs into plant tissue and subsequently covers them with a wax-like secretion (Fig. 13 a-c). This oviposition strategy differs from the more superficial egg-laying observed in Paragara, Bolbonota, Tritropidia and some Leioscyta and Tropidoscyta species (Supplementary Fig. 2). The behavior of piercing the plant to insert the eggs appears to be associated, at least in part, with the sword-like shape of the ovipositor. Females of Eunusa concolor remain positioned over their egg masses, although no other conspicuous defensive behaviors, such as kicking or lateral movements, were observed. Even when ant shelters are disturbed or opened, females may remain over the eggs (Fig. 13 a – c), although they also frequently flee. The degree of maternal investment in this species remains unclear, as the behavior of females inside the ant shelters cannot be easily observed. As in other membracids with close ant mutualisms, maternal care may involve maintaining the presence of ant partners until the nymphs hatch (Bristow 1983). However, it is also possible that this behavior serves to defend the eggs from natural enemies such as parasitoids or parasites that may not be detected or repelled by the ants. As they develop, the nymphs of Eunusa concolor gradually become covered in a white wax-like substance. This wax is lost during each molt and gradually reaccumulates over time. As in many treehopper species inhabiting humid tropical forests, E. concolor can be found in mixed-species aggregations, occasionally co-occurring with other species inside the same ant-built shelters. Females of Harmonides spp. were observed within shelters alongside E. concolor (Fig. 13 f), and individuals were also found in proximity to nymphs and adults of Stegaspis fronditia (Linnaeus) (Fig. 13 e). In addition, E. concolor was observed sharing shelters with lycaenid larvae (Fig. 13 g).	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
33469A2EFFF8FFE1FF4AFC48FE0E368C.taxon	materials_examined	Examined species: Lectotype in MZUSP: BRAZIL: “ Utiariti \ Rio Papagaio, Mt \ 3: XI: 1966 \ Lenko & Pereira \ 3916 ”, “ Cotypus ”, “ Eunusa \ concolor \ J. P. Fonseca det. [other letters difficult to read] ”, “ LECTOTYPE \ Eunusa \ concolor Fonseca \ desig. S. H. McKamey 1992 ”. Other examined specimens: COLOMBIA. Antioquia: Anorí: “ COLOMBIA. Anorí, vereda La Plancha, bosque de galeria cerca del ETCR Anorí, 7.127125 ° N, 75.220134 ° W, 1300 msnm, manual, Jul. 28 / 2018, leg. Camilo FlórezV (Expedición BIO Anorí), CBUCES-F 2119 ” (1 adult). Cocorná: “ COLOMBIA. Antioquia, Cocorná, vereda Los Mangos, cerca al PCH San Matías, 6.041479 ° N, 75.122633 ° W, 700 – 1100 msnm, manual, Dic. 4 / 2017, leg. C. Flórez-V, A. Ospina, CBUCES-F 13904 ” (3 adults, 1 nymph). Gómez Plata: “ COLOMBIA. Antioquia, Gómez Plata, vereda Vegas de la Clara, Finca Vegas de La Clara, 6 ° 34 ’ 53.37 ’’ N, 75 ° 11 ’ 43.28 ’’ W, 1080 msnm, manual, Dec. 13 – 14 / 2012, leg. Álvaro Cogollo, Camilo Flórez-V, Katherine Mejía, CBUCES-F 9891 ” (30 individuals); Remedios: COLOMBIA. Antioquia, Remedios, vereda La Cruz, finca La Brillantina, I 135, Jul. 1 / 2022, leg. C. Flórez-V, M. Idárraga, S. Quintero-Montoya, CBUCES-F 14350 (3 males, 3 females, 2 nymphs), sama data except by: “ I 27, Jun. 17 / 2022, CBUCES-F 14511 ” (1 nymph). Putumayo: Mocoa: “ COLOMBIA. Putumayo, Mocoa, vereda San Antonio, desde la vereda hasta la variante Sibundoy-Mocoa, 1.205502 ° N, 76.681895 ° W, 700 – 1000 msnm, manual, May. 5 / 2018, leg. S. Carvalho, C. Flórez-V, M. J. Sanín, CBUCES-F 9341 ” (6 adults in CBUCES). Risaralda: Pueblo Rico: “ COLOMBIA. Risaralda, Pueblo Rico, corregimiento Santa Cecilia, Área de Manejo Especial de Comunidades Negras Alto Amurrupá, 5.33526 ° N, 76.15723 ° W, 300 – 600 msnm, manual, Abr. 12 – 15 / 2017, leg. Blás Cardenás, Camilo Flórez-V, CBUCES-F 820 ” (1 adult), same data except by: “ CBUCES-F 821 ” (1 adult), “ Abr. 15 / 2017, CBUCES-F 418 ” (1 adult), “ Abr. 15 / 2017, CBUCES-F 560 ” (1 adult), “ Sep. 22 / 2016, leg. C. Bota, B. Cardenás, C. Flórez-V, CBUCES-F 1485 ” (1 adult), “ Sep. 25 / 2016, leg. C. Bota, B. Cardenás, C. Flórez-V, CBUCES-F 1064 ” (1 adult). Vaupés: “ COLOMBIA. Vaupés, Mitú, Reserva Chunduu, 1.238532 ºN, 70.214158 ºW, 189 – 212 msnm, manual, IM 7, Feb. 17 / 2024, leg. Camilo Flórez-V, Santiago Quintero-M, CBUCES-F 20350 ” (3 adults, 3 mutualistic ants in CBUCES), same data except by: “ IM 24, Feb. 18 / 2024, CBUCES-F 20367 ” (2 adults in CBUCES), “ IM 41, Feb. 19 / 2024, CBUCES-F 23984 ” (4 females in CBUCES), “ IM 114, Feb. 23 / 2024, CBUCESF 20452 ” (1 adult in CBUCES), “ IM 118, Feb. 23 / 2024, CBUCES-F 20456 ” (2 adults, 2 nymphs in CBUCES).	en	Flórez-V, Camilo, Urban, Julie M. (2026): New taxa, natural history and morphology of Paragara Goding and comparisons with Eunusa Fonseca (Hemiptera: Membracidae: Membracini). Zootaxa 5752 (1): 37-62, DOI: 10.11646/zootaxa.5752.1.2, URL: https://doi.org/10.11646/zootaxa.5752.1.2
