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The nymphs of Acutalis Fairmaire, Bordoniana Sakakibara, Thrasymedes Kirkaldy, and Micrutalis Fowler are described and illustrated (Bordoniana and Thrasymedes for the first time). The nymphs of all four genera are exceedingly cryptic. The nymphs of some species lack scoli on the head and pronotum but all have paired scoli on the meso- and metathoracic nota and abdominal segments III–IX. Some species also have lateral rows of enlarged chalazae on the abdomen, and even large scoli ventrolaterally—the latter condition is unique within Smiliinae. The eggs are deposited in stems (not in exposed masses) and nymphs are solitary and not ant-attended. The fifth instar nymphs of Micrutalini range in length from 3.0–3.5 mm, much smaller than the fifth instars of most other treehoppers.

Three new genera in Acutalini are described, two of which have two discoidal cells (R2+3 and M) in the forewing, as in Euritea Stål. Ceresinoidea zackigen. nov. et sp. nov., from Guatemala, differs from other acutalines in having a pair of suprahumeral spines and a stepwise convex pronotum in lateral view. Quinquespinosa septamaculagen. nov. et sp. nov., which is widely distributed in South America, differs in having a basal cell M and three posterior pronotal spines. Tectiforma guayasensisgen. nov. et sp. nov., from Ecuador, has the pronotum strongly tectiform throughout. A key to all genera of Acutalini is provided.

Andinodarnis atromeris sp. nov. urn:lsid:zoobank.org:act: 7A3F9F29-501E-4935-B93D-09960D69C292 (Figures 4–7) Diagnosis: Head and metopidium black; in dorsal view, black area V-shaped, extended diagonally from humeral angles to dorsomedial carina above semicircular lateral impression. Forewing veins black and membrane on anterior area of apex smoky hyaline. Description: Holotype male. Color: Head ochraceous black, with one yellow inverse T between ocelli (on epicraneal suture and dorsal margin of frontoclypeus), eyes silver, ocelli ochraceous yellow. Pronotum black from metopidium and diagonally from humeral angles toward dorsum over beginning of lateral impressions, and then bright yellow [in live specimens, this yellow area is green, see Fig. 1A–B]. Labium, thoracic pleurites and sternites, coxae and base of metathoracic femur black. Forewings smoky hyaline with black veins; one yellow band on sclerotized area of clavus; one darker macula between apex of M basal cell, first discoidal cell, second and third apical cell and adjacent area of apical limbus. Pro- and mesothoracic femora, tibiae and tarsomeres ochraceous brown; third tarsomeres with apex black. Abdomen dark brown (Figs. 4, 5). Abdomen: Subgenital plates triangular and slender, bilobed after half-length, bearing a quadrangular process at apex in ventral view, and triangular in lateral view. Styles hook-shaped, relatively slender, shank broader at middle. Aedeagus U-shaped, several rows of minute teeth along entire anterior surface of posterior arm (Fig. 6). Female. Same as male but larger. Abdomen: Gonoplac in lateral view with apex rounded, ventral outline widely arched, basal half distinctly narrower than apical half. First valvula blade-shaped, slender, dorsal outline widely arched upward; longitudinal dorsal axis on apical ¾ and apex ventrally ornate with oblique and lineal integumental processes, ramus extended up to apex, pores along ramus extension, ramus extended almost until apex, ventral sculptured area restricted to apex. Second valvula distinctly more sclerotized than first valvulae, dorsal outline widely arched upward, ramus extended almost until apex with pores along, canals at apical 1/3 extended toward rounded teeth which are only present on apical 1/3; two relatively more sclerotized teeth contiguous at base of apical 1/3, longitudinal ventral axis ornate basally with linear and oblique integumental processes, and apically with escamiform integumental processes (Fig. 7). Nymph: unknown. Measurements: Holotype male (mm): body length: 9.85; forewing length: 8.26; pronotal length: 8.12; pronotal height: 3.07; pronotal width: 3.78; head width: 3.38; vertex width: 2.29; vertex length: 1.36. Biology. Two females were recorded on Erato vulcanica (Asteraceae) (Fig. 1A) but only one was collected (designated here as the paratype). The holotype was collected on Acanthaceae (Fig. 1B). Additional field data is required to determine whether these are legitimate host plant associations. Examined material. Holotype male: COLOMBIA: Risaralda: Pueblo Rico: “COLOMBIA. Risaralda, Pueblo Rico, vereda Montebello, PNN Tatamá, subiendo hacia el cerro Montezuma, 5.23983°N, 76.08175°W, 2300 msnm, manual, en Acanthaceae, May. 25–26/2016, leg. C. Bota, C. Flórez-V, CBUCES-F 274” (CBUCES; collected on an undeterminied species of Acanthaceae) . Female paratype: COLOMBIA: Risaralda: Pueblo Rico: “ COLOMBIA. Risaralda, Pueblo Rico, vereda Montebello, PNN Tatamá, subiendo hacia el cerro Montezuma, 5.224971°N, 76.082570°W, 1800 msnm, manual, en Erato vulcanica (Asteraceae), Sep. 12–19 /2014, leg. A. Chinome, C. Bota, C. Flórez-V, M.A. Loaiza, CBUCES-F 253” (CBUCES; collected on Asteraceae: Erato vulcanica). Notes on type series. Holotype minuten-mounted, in good state of preservation, with dissected abdomen placed in vial with glycerine pinned underneath the specimen. Paratype equally well preserved, minute-mounted, with dissected abdomen placed in vial with glycerine pinned underneath the specimen. Distribution. COLOMBIA: Risaralda: Pueblo Rico (Vereda Montebello, PNN Tatamá— 1800–2350 masl) (Fig. 8). Etymology. The epithet specific ‘ atromeris ’ refer to the words ‘ atra ’, which means black, and ‘ meris, ’ which means portion, to denote the unique coloration of the head and pronotum on the anterior half. Remarks. This species is readily distinguished by its unique pronotal coloration, dark brown anteriorly and yellow beyond humeral angles (Fig. 4). The pronotal shape closely resembles A. pectoralis, except for a slightly lower dorsum and less pronounced suprahumeral horns. First and second valvulae exhibit some slight interspecific differences: in A. atromeris sp. nov. the prominent teeth of the second valvula are contiguous while in A. pectoralis they are separated by small teeth. This species has only been found in one site of the Cordillera Occidental in Colombia (Western Andean Mountain range), while A. pectoralis was collected in different locations across the Cordillera Central in Colombia, in Macizo Colombiano, and in southern Ecuador and Peru (Fig. 8)., Published as part of Flórez-V, Camilo & Evangelista, Olivia, 2022, Andinodarnis gen. nov., a new treehopper genus from the Andes (Hemiptera: Membracidae: Darninae), pp. 37-50 in Zootaxa 5200 (1) on pages 44-48, DOI: 10.11646/zootaxa.5200.1.3, http://zenodo.org/record/7251848

Andinodarnis gen. nov. urn:lsid:zoobank.org:act: 264E9AEB-7003-4D71-A3B7-C85CD1973882 Type-species: Tomogonia pectoralis Stål, 1869 Diagnosis: Head in anterior view subtriangular. Pronotum convex, finely punctate, smooth; suprahumeral horns acute, very close to humeral angles, slightly projected and curved backward in dorsal view; semicircular lateral impressions extending from behind humeral angles to apical third; posterior process extended slightly beyond M fork of forewing. Forewings entirely exposed, not concealed by pronotum. Pro- and mesothoracic femur bearing two rows of cucullate setae on ventral surface; metathoracic femur bearing one row of cucullate setae on ventral surface, fewer and comparatively smaller than other femora. Description: Color: Pale to ochraceous yellow, dark brown, or combination thereof; yellow patches bright green in live specimens. Labium, thoracic pleurites and sternites, coxae and base of metathoracic femur dark brown to black. Forewings amber to smoky hyaline with dark macula at apical third. Abdomen ochraceous, yellow to dark brown. Vestiture: Head and pronotum smooth, shiny, and finely punctate. Body glabrous, except thoracic pleurites and sternites, coxae, ventral surface of femora and tibiae covered in golden or silver pubescence. Head: Subtriangular; posterior margin of vertex, including supra-antennal ledges and frontoclypeus, forming slightly arcuate continuous outline; with feeble semicircular concavity between ocelli; lower half depressed, slightly directed posteriorly, frontoclypeus ovoid to subpentagonal with smooth round edges; dorsal margin arcuate; labium reaching mesothoracic coxae. Thorax: Pronotum convex, low, dorsally arched, moderately compressed laterally behind humeri; suprahumeral horns minute, conical, arising right above humeral angles, curved backward in dorsal view, slightly projected into acute tip; semicircular impressions, one at each side of posterior process, extending from behind humeral angles to apical third; dorsomedial carina inconspicuous anteriorly, gradually compressed and tectiform from mid-dorsum to pronotal apex; posterior process tapering into spine-like apex. Forewings entirely exposed, R initially divided into R 1+2+3 and R 4+5, with one s, one r-m, and two m-cu crossveins; crossvein s slightly distad of r-m; s and m-cu delimiting two discoidal cells. Hind wing with one r-m, and m-cu crossvein almost as long as M 3+4, and R 4+5 not fused with M 1+2. Pro- and mesothoracic femur bearing two rows of cucullate setae on ventral surface (Fig. 5A–D), metathoracic femur bearing one row of cucullate setae on ventral surface, each row with few spines (7 or less), much less acute and pronounced in metathoracic femur; well-defined rows of cucullate setae on metathoracic tibia (three single rows: antero-dorsal, postero-dorsal and ventral = I, II and III sensu Deitz, 1975) (Fig. 5E–G). Abdomen: Terga smooth, lacking fenestrae, tuberosities or dorsal scars (Fig. 3A, 7A). Male genitalia: Subgenital plates somewhat rectangular, elongate, weakly curved in lateral view. Styles slender, hook-shaped, broader at middle. Aedeagus U-shaped, bearing several rows of minute teeth on upper anterior surface of posterior arm. Female genitalia. Gonoplac round apically in lateral view, narrow basally and abruptly wider at apical half; ventral margin arched, sclerotized, covered in fine punctation and setae. First valvulae in lateral view weakly curved, apex triangular and acute; delicately ornate along dorsal margin, fine lines more or less perpendicular to longitudinal axis; ramus reaching apex of valvulae with pores along entire extension. Second valvulae blade shaped, distinctly more sclerotized than first valvulae, dorsally curved toward triangular apex, finely serrate; apical 1/5 covered in canals and pores; longitudinal ventral axis ornate basally with linear and oblique integumental processes, and apically with escamiform integumental processes; ramus extended almost to apex. Nymph: unknown. Distribution. COLOMBIA: Antioquia, Putumayo, Risaralda. ECUADOR: Pichincha (new record). PERU: Cuzco. Members of Andinodarnis gen. nov. are endemic to the Andean region. Recent specimens were collected in montane habitats ranging between 1800 and 3000 masl (Fig. 8). Etymology. The generic name is composed of two words: ‘ andinus ’, meaning ‘from the Andes’, and ‘ Darnis ’, a genus in the same tribe that is somewhat similar in appearance. The gender is feminine. Constituent species (2): Andinodarnis pectoralis Stål, 1869 Andinodarnis atromeris Flórez-V and Evangelista sp. nov. Remarks. Representatives of Andinodarnis gen. nov. are infrequently collected and scarce in entomological collections. Several collecting events targeting this group yielded only a few specimens (six new exemplars in four collecting sites). Current records indicate that A. pectoralis occurs at mid to high elevation along the Northern and Central Andes (1800–3000 masl), while A. atromeris sp. nov. is restricted to the western area of Colombian Andes (1800–2450 masl). The genus can be unambiguously ascribed to Darnini based on the femoral cucullate setae (Fig. 5). Tribal constituents in the clade with the same pronotal syndrome lack suprahumeral horns and have forewings partially to entirely concealed by the posterior process, in addition to more pronounced and numerous femoral cucullate setae. Andinodarnis gen. nov. is, therefore, the only genus in the raindrop group (‘dewdrop’ sensu Roy et al., 2007) with fully exposed wings and suprahumeral horns (Figs. 2, 4). These two traits are also observed in Sundarion, a genus in the tribe outside the raindrop group, which differs in having much larger, strongly projected suprahumeral horns, which are prismatic and often truncate apically. A phylogenetic framework is required to verify whether these topologically similar structures are homologous, considering stark differences in pronotal shape and vestiture. In Andinodarnis gen. nov., the suprahumeral horns are conical, minute, lacking a longitudinal carina, and the surface of the pronotum is ultra-smooth, shiny, and finely punctured. Additional distinguishing features of Andinodarnis gen. nov. include a triangular head, femora with few cucullate setae on the ventral surface (rows with seven spines or less), and a widely curved upward female second valvulae with one or two prominent teeth apically (Fig. 3, 7). Key to Andinodarnis species 1. Head, pronotum and legs pale to ochraceous yellow, forewings amber hyaline, highest point of dorsal outline past mid-dorsum (Fig. 2).......................................... A. pectoralis Stål comb. nov. (= Tomogonia pectoralis Stål, 1869) 1’. Head and metopidium dark brown, posterior process yellow from humeral angles to apex, forewings smoky hyaline, highest point of dorsal outline approximately at mid-dorsum (Fig. 4).............. A. atromeris Flórez-V and Evangelista sp. nov., Published as part of Flórez-V, Camilo & Evangelista, Olivia, 2022, Andinodarnis gen. nov., a new treehopper genus from the Andes (Hemiptera: Membracidae: Darninae), pp. 37-50 in Zootaxa 5200 (1) on pages 38-40, DOI: 10.11646/zootaxa.5200.1.3, http://zenodo.org/record/7251848, {"references":["Deitz, L. L. (1975) Classification of the higher categories of the New World treehoppers (Homoptera: Membracidae). North Carolina Agricultural Experiment Station, Technical Bulletin, 225, 1 - 177.","Roy, L., Guilbert, E. & Bourgoin, T. (2007) Phylogenetic patterns of mimicry strategies in Darnini (Hemiptera: Membracidae). Annales de La Societe Entomologique de France, 43 (3), 273 - 288. https: // doi. org / 10.1080 / 00379271.2007.10697523"]}

Flórez-V, Camilo, Evangelista, Olivia (2022): Andinodarnis gen. nov., a new treehopper genus from the Andes (Hemiptera: Membracidae: Darninae). Zootaxa 5200 (1): 37-50, DOI: https://doi.org/10.11646/zootaxa.5200.1.3

Oropedium tatamaense sp. nov. (Fig. 2–4) urn:lsid:zoobank.org:act: 30656033-7F76-4871-BA03-EB19BD541033 Diagnosis: Integument covered with dense golden pubescence; overall color black. Other diagnostic characters as for the genus. Description: Holotype male. Surface (Fig. 3): Head, pronotum, forewing (except by membranes), legs and ventral area of thoracic sclerites and abdomen with dense golden pubescence. Head and pronotum coarsely punctate; forewing sclerotized and punctate on basal 1/5; abdomen finely punctate. Color (Fig. 3): Overall color dark brown to black. Eyes reddish dark brown with scarlet borders, ocelli golden. Pronotum with a lighter spot on dorso-median carina on area above humeral angles. Forewing dark brown on sclerotized area and veins, membrane translucent and amber, one brown band (divided by dark brown veins) crossing basal 2/5 of basal cell R and Cu, base of basal cell M and at mid length of clavus; apex of clavus golden; apical limbus paler on area behind M 1+2 and M 3+4. External morphological characters as in the genus description. Abdomen: Male genitalia (Fig. 4). Pygofer with a transverse ridge from middle of posterior margin almost reaching dorsal outline. Subgenital plate divided almost from base, each lobe barely narrowed at apex; in ventral view, lateral margin slightly sinuous; in lateral view, apex barely bent upward. Lateral plate suboval, globular, dorsal margin broader than ventral margin, setae sparse. Style apex hooklike, slender throughout, with setae along external margin on apex. Aedeagus U-shaped, broad throughout, with minute teeth on anterior and anterolateral surfaces of the apex of the posterior arm, gonopore apical. Measurements (in mm): Holotype male. Body length: 5.05; forewing length: 4.65; pronotum height: 1.33; pronotum length: 4.00; width between humeral angles: 1.96; head width: 1.66; head length: 0.83. Material examined: Holotype male in CBUCES. “ COLOMBIA. Risaralda, Santuario, PNN \ Tatamá, páramo de Tatamá, Piedra Bomba, \ 5.120532°N, 76.057963°W, 3750 msnm, \ manual, en Ericaceae, \ Ene. 7/2018 \ leg. C. Flórez-V, A. Ospina, J. Ruiz \ CBUCES-F 11808”. Two male paratypes in ethanol 96% with the same information as the holotype except by the collection code: “CBUCES-F 7783” (in CBUCES). One male paratype is dissected, and the genitalia are preserved inside the abdomen in a microvial with glycerin. Distribution: COLOMBIA: Risaralda:Santuario(PNN Tatamá– Páramo de Tatamá, 5.120532°N, 76.057963°W, 3750 masl) (Fig. 1). Biology: Three males were found on different individuals of the same species of Ericaceae in the páramo (Fig. 2A–B). One early instar nymph, seemingly of a Membracinae, was also found on Ericaceae (Fig. 2C), but it could not be determined if belongs to O. tatamaense. Etymology: The specific epithet ‘ tatamaense’ refers to the Emberá word ‘Tatamá’ which means “The Grandfather of the Rivers.” It refers to the type locality where the genus was found in the Tatamá mountains. Remarks: This species was found during a short expedition to the Páramo of Tatamá and was the only treehopper species found above 3000 masl in this locality. The entomofauna of the Páramo of Tatamá is practically unknown and certainly many new species will be revealed by further exploration in this highly biodiverse area and other páramos in the Andes. On the other hand, Ericaceae has not been frequently recorded as host plant for treehoppers, even though it is a relatively common family in Alto-Andean forests and páramos. This family of plants is more common in temperate regions, and, in the tropical region, they are only found in mountains. I have found other treehopper species on this family of plants, including a possible new species of Metcalfiella in a páramo of the Cordillera Central in Colombia at 3250 masl. Oropedium tatamaense has irregular and superficial carinae on the apex of the pronotal posterior process. These carinae seem different from the latero-apical carinae described by Sakakibara (2012) for Talipedini, which have a well pronounced pair of carinae on the posterior third of the posterior process parallel to the lateral margins. Additionally, each specimen herein examined exhibits different venation on apical third at each forewing (left forewing different from right forewing in the same specimen), with different numbers of extra-crossveins. Despite this variation in number of crossveins, the branching pattern of vein R in R 1 +2+3 and R 4 +5 is consistent among the three specimens examined., Published as part of Flórez-V, Camilo, 2022, A new genus and a new species of treehopper (Hemiptera: Membracidae) from the páramo of Tatamá in Colombia, pp. 143-154 in Zootaxa 5195 (2) on pages 146-149, DOI: 10.11646/zootaxa.5195.2.3, http://zenodo.org/record/7184659

Flórez-V, Camilo (2022): A new genus and a new species of treehopper (Hemiptera: Membracidae) from the páramo of Tatamá in Colombia. Zootaxa 5195 (2): 143-154, DOI: https://doi.org/10.11646/zootaxa.5195.2.3

Key to tribes of Membracinae (including Oropedium gen. nov. in incertae sedis) 1. Forewing vein R initially divided into R1 and Rs (Fig. 5H, 5J, 6A, 6C)........................................... 2 1’. Forewing vein R initially divided into R1+2+3 and R4+5 (Fig. 3C, 5G, 5I, 6B)....................................... 3 2 (1). Pronotum with latero-apical carinae (Fig. 5J); metathoracic tibia club-shaped, dilated apically; metathoracic tarsus shorter than pro- and mesothoracic tarsi...................................................................... Talipedini 2’. Pronotum lacking latero-apical carinae (Fig. 5H); metathoracic tibia with anterolateral and posterolateral margins subparallel or slightly dilated apically (except clavate in Havilandia Dietrich and McKamey); pro-, meso- and metathoracic tarsi subequal in length................................................................................... Membracini 3 (1’). Metathoracic tibia clavate, metathoracic tarsus distinctly shorter than pro- and mesothoracic tarsi (Fig. 6B)..................................................................................................... Hoplophorionini 3’. Metathoracic tibia not clavate (Fig. 3C), or, if slightly expanded apically, then tarsus subequal in length to that of prothoracic leg................................................................................................. 4 4 (3’). Forewing with apical limbus extended over clavus (Fig. 5I); lateral margins of pronotum covering portions of apical limbus and veins of forewing (except Cladonota); abdominal terga coarsely pitted (Fig. 6E) (although finer in Cladonota; Fig. 6F).............................................................................................. Hypsoprorini 4’. Forewing with apical limbus not extended over clavus (Fig. 5G); lateral margins of pronotum covering at most part of apical limbus of forewing; abdominal terga finely punctuated (Fig. 4C)............................................... 5 5 (4’). Pronotum usually with anterior process, metopidium forming an acute angle with dorsal outline, pronotum without lateral carinae (Fig. 5G)............................................................................ Aconophorini 5’. Pronotum plateau-shaped, lacking anterior process, metopidium forming an angle of 90º with dorsal outline, lateral carina extended from above metopidium almost to humeral angles (Fig. 3B–C)............................................................................................. Oropedium tatamaense sp. nov. [Membracinae incertae sedis], Published as part of Flórez-V, Camilo, 2022, A new genus and a new species of treehopper (Hemiptera: Membracidae) from the páramo of Tatamá in Colombia, pp. 143-154 in Zootaxa 5195 (2) on pages 149-153, DOI: 10.11646/zootaxa.5195.2.3, http://zenodo.org/record/7184659

Oropedium gen. nov. urn:lsid:zoobank.org:act: 26B00DDC-7120-4773-95BF-14BFEA067C62 Type species: Oropedium tatamaense sp. nov. Diagnosis: Head octogonal, area around ocelli convex above, concave below; supra-antennal ledges and inferior margin of frontoclypeus foliaceous, slightly folded forward. Pronotum plateau-shaped anteriorly, metopidium outline in lateral view vertical, forming almost an angle of 90º, descending abruptly at posterior third to the acute posterior process apex; one dorsolateral carina on each side, extended from dorsomedial carina above metopidium to above humeral angles; humeral angles triangular, slightly laterally produced; lateral surface of pronotal posterior process undulate. Forewing vein R initially divided into R 1+2+3 and R 4+5. Pro- and mesothoracic tibia foliaceous. Metathoracic tibia with row I and II of cucullate setae enlarged as spines. Abdomen finely punctate. Description: Male. Head (Fig. 3): Octagonal in frontal view, area around ocelli convex above, concave below; supra-antennal ledges and inferior margin of frontoclypeus foliaceous, slightly folded forward. Thorax (Fig. 3): Pronotum plateau-shaped, metopidium forming an angle of 90º with dorsal outline, descending abruptly at posterior third to the apically acute posterior process; dorsomedial carina elevated and foliaceous; one dorsolateral carina, extended from dorsomedial carina above metopidium to above humeral angles; humeral angles triangular, slightly laterally produced; lateral surface of pronotal posterior process undulate, slightly depressed behind humeral angles, then inflated at anterior 1/3, depressed behind and then slightly inflated again at posterior 1/3 and then depressed again toward apex of posterior process. Forewing vein R initially divided into R 1+2+3 and R 4+5, 2 m-cu crossveins, 1 s crossvein, often with one or two additional r-m crossveins and/or irregular veins toward apex, clavus acuminate. Hindwing with 1 r-m and 1 m-cu crossvein, first and third apical cell slightly longer than second apical cell. Tibia foliaceous, tarsi subequal in length, metathoracic tibia with cucullate setal row III smaller than rows I and II, rows I and II enlarged as spines. Abdomen (Fig. 4): Male genitalia. Subgenital plate divided almost from base, each lobe barely narrowing at apex; lateral plate suboval, globular; style apex hooklike, slender throughout; aedeagus Ushaped, broad throughout, with minute teeth on anterior and anterolateral surfaces of the apex of the posterior arm. Female, egg masses, and nymphs unknown. Distribution: COLOMBIA: Risaralda: Santuario (páramo de Tatamá, 3750 masl) (Fig. 1). Species included: Oropedium tatamaense sp. nov. Etymology: The name ‘ Oropedium ’ refers to the Latin word ‘ oropedium ’, which means “plateau”. It denotes to the plateau-shape of the pronotum. The gender is neutral. Remarks: The new genus Oropedium belongs to the subfamily Membracinae based on the following combination of characters: pronotum well-developed, covering scutellum, wider at humeral angles than head width, forewing with two m-cu crossveins, metathoracic tibia with cucullate setal row III greatly reduced in comparison with rows I and II (Dietrich et al. 2001). Following Dietrich and McKamey (1995), Creão-Duarte and Sakakibara (1998) and Dietrich et al. (2001), this genus could be related to Aconophorini or Hypsoprorini, based on R vein divided in R 1+2+3 and R 4+5 on forewings, the non-clavate metathoracic tibia, and metathoracic tarsi subequal in length to pro- and mesothoracic tarsi. Despite overall body shape resemble species in Membracini (e.g. Leioscyta and Tropidoscyta) and Talipedini (e.g. Erechtia), Oropedium exhibits a unique combination of characters, and a comprehensive phylogenetic and taxonomic revision of Membracinae will be necessary to establish its correct position. Oropedium differs from the Membracinae tribes by the following characters: the new genus lacks a lateroapical carinae in the pronotum (unlike Talipedini; Sakakibara 2012); the lateral edges of the metathoracic tibia are parallel and tarsi are subequal in length, different from the metathoracic tibia clavate like Hoplophorionini (McKamey and Deitz 1996) or club shaped like Talipedini (Sakakibara 2012); the pronotum covers a small portion of the apical limbus of the forewing, the apical limbus does not extended over the clavus and the abdomen is finely punctate (unlike Hypsoprorini; Dietrich and McKamey 1995); the pronotum has a dorsolateral carina on each side, the metopidium outline in lateral view is vertical, forming almost an angle of 90º with dorsal outline (unlike Aconophorini; Dietrich and Deitz 1991); and the R vein of forewing is initially divides into R 1+2+3 and R 4+5 (unlike Membracini and Talipedini; Dietrich and McKamey 1995, Creão-Duarte and Sakakibara 1998, Dietrich et al. 2001, Sakakibara 2012). The forewing R vein divided into R 1+2+3 and R 4+5 is consistent among the three specimens examined in the new genus, and this branching pattern is like in Aconophorini, Hypsoprorini and Hoplophorionini (Fig. 5G, 5I, 6B), and different to Talipedini and Membracini where the R vein initially divided into R 1 and Rs (Fig. 5H, 5J, 6A). The branching pattern of vein R is apparently constant among tribes (Creão-Duarte and Sakakibara 1998). Phylogenetic analyses have shown Hypsoprorini as the sister group of the other Membracinae (Dietrich et al. 2001, 2017; Lin et al. 2004). Then, Aconophorini has been recovered as the sister group of a clade comprising Hoplophorionini +Tali pedini+ Membracini (Lin et al. 2004), while another phylogeny has recovered Hoplophorionini as the sister group of Aconophorini + Talipedini + Membracini (Dietrich et al. 2017). The relationships within this clade have not been resolved consistently, and Membracini has not been recovered as monophyletic in these analyses. Although this clade it is not fully resolved, it seems that bifurcation of R 1 and Rs is a derived character. The overall shape of the new genus resembles species in Leioscyta and some species of Membracinae in incertae sedis (see Sakakibara 2012 for a complete list of these species) which exhibit a lateral carinae on each side of the pronotum and lack accessory carinae (like Enchenopa -related groups, Fig. 5H) or suprahumeral carinae (as Talipedini, Fig. 5J). However, the branching pattern of vein R in these species is different from that in Oropedium, where the R vein initially divided into R 1 and Rs seems to be consistent among Leioscyta species (Fig. 6C) and Membracinae species in incertae sedis. This branching pattern along with other features such as the octagonal head, pronotum plateau-shaped, abdomen finely punctate distinguish the new genus from this group of species. This genus also differs in other external traits. The frontoclypeus is wider than long and subfoliaceous, different from Hypsoprorini (Fig. 5C) and many Membracini (e.g. Fig. 5B, 5E), while the supra-antennal ledges are slightly expanded over the frontoclypeus (Fig. 3A). The abdominal terga do not bear dorsal fenestrae or tuberosities as in some Aconophorini, Talipedini and Hoplophorionini, and they are finely punctate (Fig. 4C), contrary to the coarsely punctate pronotum in Hypsoprorini and Membracini species related to Bolbonota (Fig. 6D–F). Other features of the new genus resemble species in other groups. The overall shape of the head and pronotum of Oropedium resembles some Talipedini (e.g. Talipes appendiculatus and T. fenestratus, Fig. 5J). It has a lateral carina at each side as in Talipedini (Fig. 5J) and some Membracini (e.g., Enchenopa, Enchophyllum, Leioscyta, Tritropidia, Fig. 5H). The forewing has extra crossveins like some Hoplophorionini (Fig. 6B) and Hypsoprorini (Fig. 5I). The metathoracic tibia is subfoliaceous and tarsi are subequal in length like some Membracini (Fig. 6A), Aconophorini and Hypsoprorini (Fig. 5I). The single species of Oropedium has irregular and not well pronounced carinae in the posterior third of the posterior pronotal process (Fig. 3B–D). Using different light sources and views, it is possible to observe variation in the carinae among the three examined specimens. Members of Talipedini exhibit one pair of latero-apical carinae at the posterior third of posterior process (Sakakibara 2012). However, this carina in talipedines is well-marked, parallel to the lateral margins and merges with the median carina at the apex of the posterior process (Sakakibara 2012), while in Oropedium, this carina is irregular, barely elevated and not parallel to the lateral margins., Published as part of Flórez-V, Camilo, 2022, A new genus and a new species of treehopper (Hemiptera: Membracidae) from the páramo of Tatamá in Colombia, pp. 143-154 in Zootaxa 5195 (2) on pages 144-146, DOI: 10.11646/zootaxa.5195.2.3, http://zenodo.org/record/7184659, {"references":["Dietrich, C. H., McKamey, S. H. & Deitz, L. L. (2001) Morphology - based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea). Systematic Entomology, 26, 213 - 239. https: // doi. org / 10.1046 / j. 1365 - 3113.2001.00140. x","Dietrich, C. H. & McKamey, S. H. (1995) Two new Neotropical treehopper genera and investigation of the phylogeny of the subfamily Membracinae (Homoptera: Membracidae). Proceedings of the Entomological Society of Washington, 97, 1 - 16.","Creao-Duarte, A. J. & Sakakibara, A. M. (1998). Analise cladistica dos generos de Membracinae (Hemiptera, Auchenorrhyncha, Membracidae). Revista Brasileira de Zoologia, 15 (4), 823 - 846.","McKamey, S. H. & Deitz, L. L. (1996) Generic revision of the New World tribe Hoplophorionini (Hemiptera: Membracidae: Membracinae). Systematic Entomology, 21, 295 - 342. https: // doi. org / 10.1111 / j. 1365 - 3113.1996. tb 00602. x","Dietrich, C. H. & Deitz, L. L. (1991) Revision of the Neotropical treehopper tribe Aconophorini (Homoptera: Membracidae). North Carolina Agricultural research Service Technical Bulletin, 293, 1 - 134.","Lin, C - P., Danforth, B. N. & Wood, T. K. (2004) Molecular Phylogenetics and Evolution of Maternal Care in Membracine Treehoppers. Systematic Biology, 53 (3), 400 - 421. https: // doi. org / 10.1080 / 10635150490445869"]}

Machaerotypus stigmosus Li & Chen sp. nov. urn:lsid:zoobank.org:act: 60892F20-5E1B-4F61-BDCE-9B010ADAB636 Figs 1–4, 7, 8H Diagnosis Forewing dark brown with indistinct transverse white stripes (Figs 1E–F, 2A–B), half of anterior surface and margin densely clothed with coarse denticulation; aedeagal shaft with tiny dorsal hump at base (Fig. 3E), dorsal margin dentate in distal half; second valvulae with broad dorsal hump on basal half (Fig. 4E). Etymology The word ‘ stigmosus ’ is a Latinized adjective derived from the Greek word ‘stigma’ (spot), referring to the forewing with many spots. Material examined Holotype CHINA • ♂; Guizhou Province, Dafang County; 27°19′ N, 105°61′ E; 8 Mar. 2019; Xiu-Dong Huang leg.; collected with yellow pan traps; IEGU. Paratypes CHINA • 23 ♂♂, 5 ♀♀; same collection data as for holotype; IEGU. Description MEASUREMENTS. Male (n = 24) body length: 4.7–5.1 mm, forewing length: 3.9–4.0 mm, width between humeral angle apices: 1.5–2.1 mm, width between suprahumeral horn apices: 1.5–1.9 mm. Female (n = 5) body length: 5.2–5.3 mm, forewing length: 4.2–4.6 mm, width between humeral angle apices: 2.2–2.3 mm, width between suprahumeral horn apices: 1.6–1.8 mm. COLORATION. Brown. Male slightly darker than female overall. With the following parts black: head, pronotal callosity, femur and tarsal claws (female lighter), posterior pronotal process apical half except tip. Forewing veins brown with black on a few veins, pattern black, and brown brindle, with indistinct transverse white medial band; female with more extensive brown markings than male (Fig. 1). Abdomen black, with base of each segment narrowly yellow-white. HEAD AND THORAX. Vertex with dorsal margin trapezoidal, ocelli slightly closer to inner margin of eyes than to each other; frontoclypeus more than two-thirds length of vertex ventral margins, apex expanded (Fig. 1C–D), frontoclypeal lobes distinct, length more than half as long as frontoclypeus. Pronotum with punctures evenly distributed. Callosity nearly triangular, with sparse setae, surface smooth. Suprahumeral horns short. Posterior pronotal process ending at anal angle of forewing at rest, depressed at base, strongly expanded near middle in lateral view (Fig. 1E–F). Basal fourth of forewing punctate with opaque sclerotization, veins M and Cu fused on the first brown brindle area, veins M+Cu and R fused at center of opaque sclerotization (Fig. 2A–B). MALE GENITALIA. Pygofer with dorsal margin slightly convex in lateral view, sternite IX longer than wide in lateral view (Fig. 3A). Lateral plate upper half narrow and pointed, depressed slightly on bottom half, with setae on margin (Fig. 3G). Basal half of subgenital plate fused (Fig. 3D). Style clasp angled dorsally; distally recurved, distal half slender, dorsal surface with setae; connective n-shaped (Fig. 3A, F). Aedeagus in lateral view U-shaped, shaft with slight convexity near base, tapered distally with coarse denticles in distal half of dorsal margin; gonopore oblong, preapical on posterior surface (Fig. 3B, E). FEMALE GENITALIA. Sternite VII in ventral view with posterior margin deeply concave, both sides with setae (Fig. 4C). Pygofer longer than wide in lateral view with numerous setae; drop-shaped in ventral view (Fig. 4A–B). First valvulae gradually narrowed and tapered to acute apex (Fig. 4D). Second valvulae gradually widened from base to near midlength with slight dorsal hump near base, distal half of dorsal margin irregularly and finely dentate, apex bluntly pointed (Fig. 4E). Third valvulae (gonoplac) broadly rounded distally, ventral margin with numerous setae (Fig. 4F). Distribution China (Guizhou) (Fig. 7). Remarks This species is similar to M. sibiricus (Lethierry, 1876), but differs from the latter as follows: 1) forewing with many spots and brown wide stripe (hyaline in M. sibiricus); 2) posterior pronotal process depressed at base, strongly expanded near midlength (flat a base and center in M. sibiricus); 3) ocelli slightly closer to inner margin of eyes than to each other (equidistant in M. sibiricus)., Published as part of Yang, Lin, Long, Jian-Kun, Chang, Zhi-Min & Chen, Xiang-Sheng, 2022, Two new species of the treehopper genus Machaerotypus Uhler, 1896 from China (Hemiptera: Membracidae: Centrotinae), pp. 64-79 in European Journal of Taxonomy 826 (1) on pages 67-72, DOI: 10.5852/ejt.2022.826.1835, http://zenodo.org/record/6761661

Genus Machaerotypus Uhler, 1896 Figs 1–6, 8 Machaerotypus Uhler, 1896: 30. Machaerotypus – Goding 1931: 303. — Kato 1940: 147. — Funkhouser 1950: 207. — Metcalf & Wade 1965: 416. — Chou & Yuan 1981: 103. — McKamey 1998: 70. — Yuan & Chou 1988: 72; 2002: 254. Type species Machaerotypus sellatus Uhler, 1896 = M. sibirica (Lethierry, 1896). Diagnosis Body usually with bright red markings. Head with vertex slightly to strongly arcuate dorsally, ventral margins oblique. Ocelli ovoid, closer to inner margin of eyes than to each other. Suprahumeral horns rounded, each with crescent-shaped ridge, not extended laterad of humeral angles. Posterior pronotal process mostly concealing scutellum, depressed at base, flat or with slight bulge in center. Humeral angle triangular and stout. Callosity smooth or slightly bristly. Scutellum exposed on both sides and at apex. Forewing multi-striped or hyaline, base with opaque sclerotization. Metathoracic tibia with three longitudinal rows of cucullate setae, trochanter without teeth. Remarks Compared with Yuan & Chou’s (2002) record, pronotum characteristics of four species, M. subinermis, M. ishiharai, M. stigmosus Li & Chen sp.nov., and M. nodulus Li & Chen sp.nov., were supplemented, and the key to species of Machaerotypus was updated based on Yuan & Chou (2002). Host plant Unknown. Distribution China, Japan, Russia, Korea (Fig. 7). Checklist of species of Machaerotypus Uhler, 1896 M. arisanus (Kato, 1928); Japan, China (Taiwan). M. camelliae Chou & Yuan, 1981; China (Yunnan). M. ishiharai Kato, 1940; Japan (Honshu). M. mali Chou & Yuan, 1981; China (Shaanxi). M. nodulus Li & Chen sp. nov.; China (Guizhou). M. rubromarginatus Kato, 1940; China (Shaanxi, Liaoning). M. rubronigris Funkhouser, 1938; China (Zhejiang, Gansu). M. semirubronigris Yuan & Chou, 1992; China (Sichuan). M. sibiricus (Lethierry, 1876); Russia; Korea; Japan; China (Heilongjiang, Beijing, Shanxi, Shaanxi, Sichuan). M. stigmosus Li & Chen sp. nov.; China (Guizhou). M. subinermis Lindberg, 1927; Russia; Korea; China (Gansu, Heilongjiang). M. taibaiensis Yuan, 2002; China (Shaanxi). M. yananensis Chou & Yuan, 1981; China (Shaanxi). Key to species of Machaerotypus Uhler, 1896 (based on Chou & Yuan 2002) 1. Body or pronotum with obviously reddish markings....................................................................... 2 – Body dark or brown.......................................................................................................................... 8 2. Reddish markings on pronotum trident shaped (Fig. 8A, a)......... M. camelliae Chou & Yuan, 1981 – Reddish markings on pronotum non-trident shaped......................................................................... 3 3. Half of suprahumeral horn reddish (Fig. 8D, d)................ M. semirubronigris Yuan & Chou, 1992 – Suprahumeral horn entirely reddish.................................................................................................. 4 4. Posterior pronotal process with one reddish band (Fig. 8B, b).... M. rubronigris Funkhouser, 1938 – Reddish markings on posterior pronotal process more extensive, Y-shaped.................................... 5 5. Anterior parts of reddish Y-shaped pronotal marking enlarged (Fig. 8G, g)...................................................................................................................................................... M. mali Chou & Yuan, 1981 – Anterior parts of reddish Y-shaped marking narrow......................................................................... 6 6. Posterior pronotal process almost entirely reddish (Fig. 8C, c)..... M. rubromarginatus Kato, 1940 – Center of posterior pronotal process in dorsal view brown or black................................................ 7 7. Forewing with a hyaline transverse band near center............................... M. taibaiensis Yuan, 2002 – Forewing brown and slightly shallow edge................................ M. yananensis Chou & Yuan, 1981 8. Forewing veins with small nodules.................................................................................................. 9 – Forewing veins without nodules..................................................................................................... 10 9. Posterior pronotal process flat (Fig. 8J)............................................. M. subinermis Lindberg, 1927 – Posterior pronotal process with dorsal margin arched (Fig. 8M)...... M. nodulus Li & Chen sp. nov. 10. The apex of posterior pronotal process curved (Fig. 8L).......................... M. arisanus (Kato, 1928) – The apex of posterior pronotal process flat......................................................................................11 11. Posterior pronotal process with two depressions (Fig. 8I).......................... M. ishiharai Kato, 1940 – Posterior pronotal process with one depression.............................................................................. 12 12. Posterior pronotal process flat or slightly expanded (Fig. 8K)............ M. sibiricus (Lethierry, 1876) – Posterior pronotal process obviously expanded in lateral view (Fig. 8H).................................................................................................................................................. M. stigmosus Li & Chen sp. nov., Published as part of Yang, Lin, Long, Jian-Kun, Chang, Zhi-Min & Chen, Xiang-Sheng, 2022, Two new species of the treehopper genus Machaerotypus Uhler, 1896 from China (Hemiptera: Membracidae: Centrotinae), pp. 64-79 in European Journal of Taxonomy 826 (1) on pages 65-67, DOI: 10.5852/ejt.2022.826.1835, http://zenodo.org/record/6761661, {"references":["Uhler P. R. 1896. Summary of the Hemiptera of Japan, presented to the United States National Museum by professor Mitzukuri. Proceedings of the United States National Museum 1108: 255 - 297. https: // doi. org / 10.5479 / si. 00963801.1108.255","Goding F. W. 1931. Classification of the Old World Membracidae. Journal of the New York Entomological Society 39: 299 - 313.","Kato M. 1940. Notes on Membracidae from Easterm Asia. Entomological World 8: 1 - 21.","Funkhouser W. D. 1950. Homoptera. Fam. Membracidae. In: Wytsman P. (dir.) Genera Insectorum Fasc. 208. Luis Desmet-Verteneuil, Bruxelles. Available from https: // www. biodiversitylibrary. org / page / 18036561 [accessed 22 Nov. 2021].","Metcalf Z. P. & Wade V. 1965. General Catalogue of the Homoptera. A Supplement to Fascicle I - Membracidae of the General Catalogue of the Hemiptera. Membracoidea in two Sections. North Carolina State University, Raleigh. https: // doi. org / 10.5962 / bhl. title. 6822","Chou I. & Yuan F. 1981. A discussion on the genus Machaerotypus Uhler with descriptions of its Chinese species (Homoptera: Membracidae). Entomotaxonomia 3 (2): 103 - 110.","McKamey S. H. 1998. Taxonomic catalogue of the Membracoidea (exclusive of leafhoppers): Second supplement to fascicle 1 - Membracidae of the general catalogue of Hemiptera. Memoirs of the American Entomological Institute 60: 1 - 377.","Yuan F. & Chou I. 1988. Advances in the research of systematics of Membracoidea from China (Homoptera: Auchenorrhyncha). In: Vidano C. & Arzone, A. (eds) Proceedings of 6 th Auchenorrhyncha Meeting: 71 - 78. Consiglio Nazionale delle Ricerche, Turin.","Yuan F. & Chou I. 2002. Fauna Sinica. Insecta Vol. 28. Homoptera Membracoidea. Science Press, Beijing.","Kato M. 1928. Notes on Formosan Membracidae, with descriptions of one new genus and some new species. Insect World 32: 2 - 15.","Yuan F., Chou I. & Cui Z. X. 1992. Homoptera: Membracoidea. In: Chen S. - X. (ed.) Insects of the Hengduan Mountains Region. Vol. 1: 192 - 219. Science Press, Beijing.","Lindberg H. 1927. Zur Kenntnis der palaarktischen Cicadina IV. Notulae Entomologicae 7: 23 - 30."]}

Yang, Lin, Long, Jian-Kun, Chang, Zhi-Min, Chen, Xiang-Sheng (2022): Two new species of the treehopper genus Machaerotypus Uhler, 1896 from China (Hemiptera: Membracidae: Centrotinae). European Journal of Taxonomy 826 (1): 64-79, DOI: https://doi.org/10.5852/ejt.2022.826.1835, URL: http://dx.doi.org/10.5852/ejt.2022.826.1835

Machaerotypus nodulus Li & Chen sp. nov. urn:lsid:zoobank.org:act: 15335C7F-F247-48C1-B6EE-0B4E23E502A4 Figs 5–7, 8M Diagnosis Forewing membrane mostly hyaline with irregular transverse brown band near base and distinct nodules along veins (Fig. 5B, D). Aedeagal shaft abruptly narrowed at distal third in lateral view with small preapical hump (Fig. 6F). Lateral plate apex blunt (Fig. 6D). Etymology The species name ‘ nodulus ’ is derived from the Latin word ‘ nodus ’ (knot), referring to the nodules along the forewing veins. Material examined Holotype CHINA • ♂; Guizhou Province, Dafang Country; 27°19′ N, 105°61′ E; Mar. 2019; Xiu-dong Huang leg.; collected with yellow pan traps; IEGU. Description MEASUREMENTS. Male (n = 1) body length: 4.9 mm, forewing length: 4.1 mm, width between humeral angle apices: 1.9 mm, width between suprahumeral horn apices: 1.6 mm. COLORATION. Orange-brown marked with golden yellow except the head, ocelli, callosity, apical of humeral angle and posterior pronotal process, inner of legs black, surfaces of legs rusty red. Forewing and veins pale yellow with the following parts brown: nodulations, apical, anal angle, apical and onethird of apical, crescent-shaped stripe near basal fourth forewing. HEAD AND THORAX. Vertex nearly rectangular with dorsal and ventral margins slightly arcuate and oblique respectively, ocelli oblong, eyes oval, ocelli slightly closer to inner margin of eyes than to each other; frontoclypeus length more than half of vertex ventral margins, apical expanded slightly, frontoclypeal lobes distinct, length more than half of frontoclypeus, extended slightly across vertex ventral margins (Fig. 5C–D). Pronotum with big punctures on near middle carina, and small punctures evenly distributed. Callosity nearly oval, with small amounts of setae and puncture. Suprahumeral horn short and not exceeding humeral angle, nearly flat. Posterior pronotal process ending at Cu1, with triangular bulge on center and apex. Forewing with more than 15 nodules on veins evenly distributed. Basal one-fifth of forewing punctate with opaque sclerotization, veins M and Cu fused for nearly half length and divergent, veins M+Cu and R fused basally (Fig. 5D). Metathoracic tibia with three longitudinal rows of cucullate setae. MALE GENITALIA. Pygofer with dorsal margin concave slightly on the central in lateral view, sternite IX slightly curved, longer than wide in lateral view (Fig. 6A, C). Lateral plate apex blunt, depressed slightly on the central, with long setae on margin (Fig. 6D). Subgenital plates fused for more than half length (Fig. 6B). Style clasp angled dorsally; distally recurved, dorsal surface with setae; connective N-shaped (Fig. 6G). Aedeagus in lateral view nearly hook-shaped, abruptly narrowed at distal third and with slight preapical hump, narrow in anterior view, more than half of anterior surface and margin densely clothed with coarse denticulation (Fig. 6E–F). Distribution China (Guizhou) (Fig. 7). Remarks This species is similar to M. subinermis Lindberg, 1927, but differs from the latter in: 1) frontoclypeal lobes more than half as long as frontoclypeus (subequal in M. subinermis); 2) apex of frontoclypeus transverse (arcuate in M. subinermis); 3) posterior pronotal process arched (flat in M. subinermis)., Published as part of Yang, Lin, Long, Jian-Kun, Chang, Zhi-Min & Chen, Xiang-Sheng, 2022, Two new species of the treehopper genus Machaerotypus Uhler, 1896 from China (Hemiptera: Membracidae: Centrotinae), pp. 64-79 in European Journal of Taxonomy 826 (1) on pages 72-77, DOI: 10.5852/ejt.2022.826.1835, http://zenodo.org/record/6761661, {"references":["Uhler P. R. 1896. Summary of the Hemiptera of Japan, presented to the United States National Museum by professor Mitzukuri. Proceedings of the United States National Museum 1108: 255 - 297. https: // doi. org / 10.5479 / si. 00963801.1108.255","Kato M. 1928. Notes on Formosan Membracidae, with descriptions of one new genus and some new species. Insect World 32: 2 - 15.","Chou I. & Yuan F. 1981. A discussion on the genus Machaerotypus Uhler with descriptions of its Chinese species (Homoptera: Membracidae). Entomotaxonomia 3 (2): 103 - 110.","Kato M. 1940. Notes on Membracidae from Easterm Asia. Entomological World 8: 1 - 21.","Yuan F., Chou I. & Cui Z. X. 1992. Homoptera: Membracoidea. In: Chen S. - X. (ed.) Insects of the Hengduan Mountains Region. Vol. 1: 192 - 219. Science Press, Beijing.","Lindberg H. 1927. Zur Kenntnis der palaarktischen Cicadina IV. Notulae Entomologicae 7: 23 - 30.","Yuan F. & Chou I. 2002. Fauna Sinica. Insecta Vol. 28. Homoptera Membracoidea. Science Press, Beijing."]}

Treehoppers of the insect family Membracidae have evolved enlarged and elaborate pronotal structures, which is hypothesized to involve co-opted expression of genes that are shared with the wings. Here, we investigate the similarity between the pronotum and wings in relation to growth. Our study reveals that the ontogenetic allometry of the pronotum is similar to that of wings in Membracidae, but not the outgroup. Using transcriptomics, we identify genes related to translation and protein synthesis, which are mutually upregulated. These genes are implicated in the eIF2, eIF4/p70S6K and mTOR pathways, and have known roles in regulating cell growth and proliferation. We find that species-specific differential growth patterning of the pronotum begins as early as the third instar, which suggests that expression of appendage patterning genes occurs long before the metamorphic molt. We propose that a network related to growth and size determination is the more likely mechanism shared with wings. However, regulators upstream of the shared genes in pronotum and wings need to be elucidated to substantiate whether co-option has occurred. Finally, we believe it will be helpful to distinguish the mechanisms leading to pronotal size from those regulating pronotal shape as we make sense of this spectacular evolutionary innovation. UCR::Vicerrectoría de Docencia::Ciencias Básicas::Facultad de Ciencias::Escuela de Biología

Some insects possess complex three-dimensional (3D) structures that develop under the old cuticle prior to the last imaginal molt. Adult treehoppers (Insecta: Hemiptera: Auchenorrhyncha: Membracidae) have one such complex 3D structure, known as a helmet, on their dorsal side. The adult helmet likely forms inside the nymphal pronotum during the final instar nymphal stage.Previous morphological studies have reported that the adult helmet is a large, bi-layered, plywood-like structure, whereas the nymphal pronotum is a monolayer, sheath-like structure. The adult helmet is much larger than nymphal helmet. Thus, the emergence of the adult helmet involves two structural transitions: a transition from a monolayer, sheath-like pronotum to a bi-layer, plywood-like helmet, and a transition in size from small to large. However, when, how, and in what order these transitions occur within the nymphal cuticle is largely unknown.To determine how adult helmet development occurs under the nymphal cuticle, in the present study we describe the morphology of the final adult helmet and investigate developmental trajectories of the helmet during the final instar nymphal stage. We used micro-CT, scanning electron microscope and paraffin sections for morphological observations, and used Antianthe expansa as a model species.We found that the structural transition (from monolayer, sheath-like structure to bi-layer, roof-like structure) occurs through the formation of a “miniature” of the adult helmet during the middle stage of development and that subsequently, extensive folding and furrows form, which account for the increase in size. We suggest that the making of a “miniature” is the key developmental step for the formation of various 3D structures of treehopper helmets.

Stictocephala bisonia Kopp et Yonke, 1977 —Buffalo Treehopper (Fig. 7) Distribution in Turkey (Map 7): Ankara, Artvin, Balıkesir, Edirne, Kırklareli, Rize (Lodos & Kalkandelen 1981); Bolu, Bursa, Eskişehir, Giresun, İstanbul, Sakarya, Tokat, Zonguldak (this study). New records by citizen scientists in Turkey: 1. Bolu: Mudurnu, Karamurat, 10.X.2015 (Karacan 2015). 2. Bursa: Nilüfer, Ürünlü, 27.VI.2016 (Biçici 2016b). 3. Eskişehir, 23.VIII.2020 (GBIF.org 2021). 4. Giresun: Bulancak, Samugüney, 31.VIII.2017 (Karataş 2017). 5. İstanbul: Sarıyer, Bahçeköy, 4.IX.2019 (Güngör 2019a); 6. Ümraniye, 20.IX.2015 (Tosun 2016b). 7. Rize: Derepazarı, Tersane, 27.VII.2020: Ahmet Karataş. 8. Sakarya: Karasu, 08.IX.2015 (Eren G. 2015), ibid., 20.IX.2015 (Eren F. 2015), ibid., 07.VIII.2016 (Eren A. 2016). 9. Tokat (centre): Kömeç, 09.VIII.2017 (Uçar 2017). 10. Zonguldak: Alaplı, 09.IX.2014 (Yaşayacak 2015). Comments: S. bisonia is native to the central and eastern part of North America (Walczak et al. 2018). It was classified formerly as Ceresa bubalus (Fabricius, 1794) (Walczak et al. 2018). However, Andrade (1997) regarded it as a synonym of Stictocephala alta (Walker, 1851). This alien species was transported to eastern central Europe from North America and it was found for the first time in Serbia before 1912 (Walczak et al. 2018). It spread in most of the countries from Portugal to Azerbaijan in the Western Palearctic. As in North America, it is distributed just north of 50 o N latitude in Europe (Schedl 1991; Walczak et al. 2018). This insect was reported from Edirne in 1963 as Ceresa bubalus by Lodos & Kalkandelen (1981). Although it is the oldest introduced species among the eight bug species considered in the present study; the number of provinces in which it was reported is low, as there is not much work on it or alternatively, its spreading speed may be low in Turkey. It is mostly seen in Marmara and Eastern Black Sea provinces. There are 13 records from 11 localities (Map 7). Most of them were in autumn, with five records each in September and October (Graph 7). GRAPH 7. Seasonality of Stictocephala bisonia Kopp et Yonke, 1977, based on records from Turkey., Published as part of Çerçi, Bariş, Karataş, Ahmet & Karataş, Ayşegül, 2021, Insecta non gratae: New Distribution Records of Eight Alien Bug (Hemiptera) Species in Turkey with Contributions of Citizen Science, pp. 1-28 in Zootaxa 5057 (1) on pages 13-15, DOI: 10.11646/zootaxa.5057.1.1, http://zenodo.org/record/5585486, {"references":["Lodos, N. & Kalkandelen, A. (1981) Turkiye Auchenorrhyncha turlerinin yayilisi ve onemlerine ait notlarla on liste. VI.","Karacan, F. (2015) Dogal Hayat Observation Sharing Platform. Available from: http: // dogalhayat. org / property / stictocephalabisonia- 2 / (accessed 9 July 2021)","Bicici, B. (2016 b) Dogal Hayat Observation Sharing Platform. Available from: http: // dogalhayat. org / property / degisik- 4 / (accessed 9 July 2021)","Karatas, A. (2017) Dogal Hayat Observation Sharing Platform. Available from: http: // dogalhayat. org / property / bizim-koyden- 2017 - 39 - bogabasli-agaczipzipi / (accessed 9 July 2021)","Gungor, U. (2019 a) Dogal Hayat Observation Sharing Platform. Available from: http: // dogalhayat. org / property / membracidaestictocephala-bisonia / (accessed 9 July 2021)","Tosun, N. (2016 b) Dogal Hayat Observation Sharing Platform. Available from: http: // dogalhayat. org / property / sticocephalabisonia / (accessed 9 July 2021)","Ucar, S. (2017) Dogal Hayat Observation Sharing Platform. Available from: http: // dogalhayat. org / property / koluma-konantombalak / (accessed 9 July 2021)","Yasayacak, H. (2015) Dogal Hayat Observation Sharing Platform. Available from: http: // dogalhayat. org / property / stictocephalabisonia / (accessed 9 July 2021)","Walczak, M., Brozek, J., Junkiert, L., Kalandyk-Kolodziejczyk, M., Musik, K., Kaszyca, N., Lazuka, A. & Gierlasinski, G. (2018) Stictocephala bisonia Kopp et Yonke, 1977 (Hemiptera: Cicadomorpha, Membracidae) in Poland. Annals of the Upper Silesian Museum in Bytom Entomology, 27, 1 - 13 https: // doi. org / 10.5281 / zenodo. 2525549","Andrade, G. S. (1997) Stictocephala alta (Walker, 1851) sp. rev., comb. nov., the correct name for the \" buffalo treehopper \", with S. bisonia Kopp & Yonke, 1977 as a new synonym, and notes on Hadrophallus bubalus (Fabricius, 1794) comb. nov. (Homoptera, Membracidae). Transactions of the American Entomological Society, 123 (4), 289 - 295.","Schedl, W. (1991) Invasion der Amerikanischen Buffelzikade (Stictocephala bisonia Kopp und Yonke, 1977) nach Osterreich (Homoptera, Auchenorrhyncha, Membracidae). Anzeiger Fur Schadlingskunde Pflanzenschutz Umweltschutz, 64 (1), 9 - 13. https: // doi. org / 10.1007 / bf 01906190"]}

This study is prepared to provide an example for the utility of citizen science in faunistic research, which is becoming popular as a data source, especially for invasive alien species. With the globalization tendency of many living species by human impact, alien species rapidly spread far from their natural distribution range and become invasive in their newly settled ecosystems. Some of these invasive alien species become one of the most important problems for nature. This study focuses on distribution of following eight alien Hemiptera species that have previously been recorded from Turkey: Zelus renardii (Reduviidae), Corythucha arcuata and Corythucha ciliata (Tingidae), Leptoglossus occidentalis (Coreidae), Perillus bioculatus and Halyomorpha halys (Pentatomidae), Stictocephala bisonia (Membracidae) and Metcalfa pruinosa (Flatidae). New distributional and seasonality data for all these species are presented based on observations shared by citizen scientists in the websites Doğalhayat and iNaturalist. New provincial records for each species are given. As a result, it is apparent that citizen science is an important data source, supplies remarkable contribution for increasing the understanding on expansion of invasive alien species as well as providing clear picture on their distribution.

Eualthe punctum (Fairmaire, 1846) Hemiptycha punctum Fairmaire, 1846: 318 (Plate 6, figure 22); Funkhouser, 1927: 118; Metcalf & Wade, 1965: 678; McKamey, 1998: 476. Hyphinoe punctorum Buckton, 1903: 124 (Plate 28, figures 3, 3a), syn. nov. Remarks: Hyphinoe punctorum is proposed as a junior synonym of Eualthe punctum based on the original descriptions and illustrations. Fairmaire (1946) and Buckton (1903) characterized their respective species as having the following combination of features: an ochraceous-yellow pronotum with one pair of brown spot at each side; suprahumeral horns somewhat acute and divergent; and posterior process exceeding the apex of forewings. Buckton (1903) discussed a possible synonymy between these species, stating that Hyphinoe punctorum could be “a variety of Hemiptycha punctum ” (p. 124). Goding (1929) treated this species as Bubalopa based on his interpretation of the literature, and Funkhouser (1951) incorrectly treated H. punctorum as a synonym of B. furcata. This misidentification hindered the accurate determination of H. punctorum as well as Bubalopa and its constituent taxa., Published as part of Flórez-V, Camilo & Evangelista, Olivia, 2021, A revision of the treehopper genus Bubalopa Stål illuminates the systematics of Hyphinoini (Hemiptera: Auchenorrhyncha: Membracidae), pp. 529-551 in Zootaxa 5052 (4) on page 550, DOI: 10.11646/zootaxa.5052.4.4, http://zenodo.org/record/5577275, {"references":["Fairmaire, L. M. H. (1846) Revue de la tribu des Membracides. Annales de la Societe Entomologique de France, 4, 235 - 320.","Funkhouser, W. D. (1927) General Catalogue of the Hemiptera. Fascicle I: Membracidae. Smith College Press, Northampton, Massachusetts, 581 pp.","Metcalf, Z. P. & Wade, V. (1965) General Catalogue of the Homoptera. A supplement to fascicle I - Membracidae of the General Catalogue of Hemiptera. Membracoidea. In Two sections. Section I. Part 1. Membracidae, Centrotinae, Platybelinae, Hoplophorioninae, Darninae. North Carolina State University, Raleigh, North Carolina, 743 pp.","McKamey, S. H. (1998) Taxonomic catalogue of the Membracoidea (exclusive of leafhoppers): second supplement to fascicle 1 - Membracidae of the general catalogue of the Hemiptera. Memoirs of the American Entomological Institute, 60, 1 - 377.","Buckton, G. B. (1903) A Monograph of the Membracidae. Lovell Reeve, London, 296 pp.","Goding, F. W. (1929) The Membracidae of South America and The Antilles. IV. Subfamilies Hoplophorioninae, Darninae, Smiliinae, Tragopinae (Homoptera). Transactions of the American Entomological Society, 55 (3), 197 - 330.","Funkhouser, W. D. (1951) Homoptera Family Membracidae. Genera Insectorum, 208, 1 - 383."]}

Bubalopa iguaque Flórez-V and Evangelista sp. nov. (Figs. 9A–D, 10) Diagnosis: Overall coloration rusty brown, variegated with ochraceous yellow, body strongly pubescent. Suprahumeral horns subtriangular, robust, nearly horizontal, distinct round protrusion at base, orientation formula: 25º (frontal), 45º (dorsal); posterior process with marked dorsal hump, dorsal outline sinuous, descending in curve toward apex. Description: HOLOTYPE MALE. Color: Rusty brown; head, pronotum, femora and tibia variegated ochraceous yellow. Eyes silver with dark brown patches. Median carina, suprahumeral horns’ carina and lateral margins of pronotum slightly darker, spotted ochraceous yellow, more evidently on dorsal view. Forewings rusty brown, coriaceous area brownish yellow and opaque. Thoracic sternites and coxae dark brown, posterior margins yellow. Surface: Head, pronotum, legs and forewings with dense silvery pubescence; seta adjacent to each punctation in head and pronotum. Head: Subtriangular, supraantennal ledges with distinct indentation before middle, near eyes; frontoclypeus’ anterior 1/2 semicircular, posterior 1/2 triangular, apex slightly round. Thorax: Suprahumeral horns robust, subtriangular, nearly horizontal, 1.5–2× as long as wide, small budge on external surface at base of horns’ lateral carina, orientation formula: 25º (frontal), 45º (dorsal); suprahumeral horns’ carina distinctly marked, horn’s dorsal carina reaching median carina on anterior 1/3. Pronotal posterior process with dorsal hump past humeral angles, dorsal outline sinuous, descending in curve towards apex. Forewings distinctly reticulate with several supernumerary crossveins from apical 2/5. Abdomen: Terga III–VI with a pair of oval dorsal pits (scars), decreasing in size in posterior segments. Apodeme broad, finger-like, reaching posterior margin of segment IV, apex clavate in lateral view (Fig. 10C). Genitalia: Lateral plate fused with pygofer. Dorsal margin of subgenital plates sinuous, nearly undulate, as seen in lateral view. Parameres hook-shaped, relatively slender, shank broader at middle, bend and point triangular. Aedeagus U-shaped, posterior arm slender with teeth along anterior surface. Female and nymphs unknown. Measurements: Male (n=1, mm): Body length: 8.7; forewing length: 7.67; pronotal length: 7.88; pronotum height: 2.61; pronotal width: 2.93; head width: 2.91; vertex width: 2.08; vertex length: 1.54. Etymology: The epithet ‘iguaque’ (noun in apposition) refers to the name of the type-locality near the Iguaque lagoon, which was an important religious and cultural sanctuary for the indigenous people ‘Muiscas’, and they referred to this area as the “heart of the world”. Biology: The specimen was collected in a Malaise trap. Distribution: COLOMBIA: Boyacá: Villa de Leyva (SFF Iguaque – 5º25‘N 73º27‘W, 2855 masl) (Fig. 11). Coordinates on original label do not match with the locality SFF Iguaque. However, Mamarramas is a locality inside the SFF [Flora and Fauna Sanctuary] Iguaque. Examined Material: Holotype male from COLOMBIA: Boyacá: Villa de Leyva: “Col Boyacá SFF / Iguaque Cabaña Mamarramas / 5º25‘N 73º27‘W 2855m / Dic. 21 – Jan 07/2003 M. 1072/ R. Reina leg.” (ICN). Remarks: The holotype was collected at a higher elevation than other Bubalopa species (2800 masl). Bubalopa iguaque sp. nov. is unique in having a rusty brown, densely punctate and pubescent body, stout and coarsely margined suprahumeral horns, and pronotal posterior process with a dorsal hump. The abdominal pits are smaller than those in B. furcata, and an additional scar can be observed on tergum VI, although it is much less distinct, nearly vestigial. The male genitalia also exhibits unique features such as the shape of the parameres and the subgenital plate in lateral view. The abdominal apodeme is greatly enlarged, both in terms of width and length, and extends to the posterior margin of segment IV., Published as part of Flórez-V, Camilo & Evangelista, Olivia, 2021, A revision of the treehopper genus Bubalopa Stål illuminates the systematics of Hyphinoini (Hemiptera: Auchenorrhyncha: Membracidae), pp. 529-551 in Zootaxa 5052 (4) on pages 545-548, DOI: 10.11646/zootaxa.5052.4.4, http://zenodo.org/record/5577275

Bubalopa obscuricornis Stål, 1869 (Figs. 9E–H) Bubalopa obscuricornis Stål, 1869: 314; Funkhouser, 1927: 118; Metcalf & Wade, 1965: 702; McKamey, 1998: 476. Diagnosis: Overall color ochraceous yellow with brown spots and bands, legs brown. Suprahumeral horns direction formula of 35º (frontal), 25º (dorsal); dorsal carina of suprahumeral horns distinctly marked; dorsal contour widely arched from metopidium to its maximum elevation above humeral angles, then descending sinuously to apex of posterior process. Description: ADULT. Color: Overall color ochraceous yellow with brown spots and bands, punctation dark brown. Head ochraceous yellow spotted brown, ocelli yellow, frontoclypeus with two longitudinal brown bands sublaterally. Pronotum yellow ochraceous on metopidium, supra-ocullar callosities brown, suprahumeral horns and area between them brown, yellow on anterior and dorsal carina; impressions behind humeral angles ochraceousyellow, area above with a U-wide brown band extended until half of pronotum, then yellow ochraceous until apex of posterior process with a brown tip; lateral margins yellow anteriorly and brown posteriorly. Ventral area of head, thoracic sternites and coxae brown with ochraceous yellow margins. Forewings entirely brownish hyaline. Legs ochraceous yellow. Surface: Silver pubescence accompanied each punctation. Head: Subtriangular, supraantennal ledges sinuous, frontoclypeus ventral margin sinuous, apex acute. Thorax: Pronotum with suprahumeral horns 1.5X as long as wide in its base, suprahumeral horns direction formula 35º (frontal), 25º (dorsal); dorsal carina of suprahumeral horns distinctly marked; dorsal contour widely arched from metopidium to its maximum elevation above humeral angles, then descending sinuously to apex of posterior process. Male genitalia, female genitalia and nymph morphology unknown. Measurements: Female (n=1, mm): Body length: 10.09; forewing length: 8.94; pronotal length: 8.89; pronotum height: 2.08; pronotal width: 3.32; head width: 3.42; vertex width: 2.45; vertex length: 1.75. Biology: Unknown. Distribution: COLOMBIA (McKamey 1998), VENEZUELA (Fig. 11). Examined Material: VENEZUELA: Trujillo: “Venezuela, Trujillo, Boconó, 8/2/81, colector: Bordón”, “ Bubalopa obscuricornis Stål ” [A.M. Sakakibara det.] 1 female (DZUP). Remarks: This species can be easily distinguished from other Bubalopa by its unique color pattern, dark brown with nearly symmetrical, slightly irregular yellow patches, and robust suprahumeral horns, distinctly enlarged at the base. The posterior process is comparatively lower than other species, yielding a more slender and elongate body shape in lateral view. Although ‘Bogota’ was stated as the type-locality in the original description, the term was used in the 19 th century to refer to multiple sites in Colombia. The specimen examined in this study represents the first record of Bubalopa from Venezuela. The syntype (s) of B. obscuricornis could not be located, despite an extensive search of European repositories. Stål (1869) stated that the type material was deposited in the Museum of Stockholm [“Mus. Holm.”, NHRS]. The single specimen at NHRS marked as Bubalopa obscuricornis is clearly not part of Stål’s type collection (Gunvi Lindberg, pers. comm.). This specimen is also misidentified to species: it is, in fact, a representative of Eualthe laevigata (Fairmaire, 1846). In the absence of type specimens, we based our identification on the original description of B. obscuricornis, which independently matched a previous identification by A.M. Sakakibara. The specimen illustrated here (Figs. 9E–H) is a single female from the Bordón collection, currently on loan to DZUP and examined by Dr. Sakakibara., Published as part of Flórez-V, Camilo & Evangelista, Olivia, 2021, A revision of the treehopper genus Bubalopa Stål illuminates the systematics of Hyphinoini (Hemiptera: Auchenorrhyncha: Membracidae), pp. 529-551 in Zootaxa 5052 (4) on pages 548-550, DOI: 10.11646/zootaxa.5052.4.4, http://zenodo.org/record/5577275, {"references":["Funkhouser, W. D. (1927) General Catalogue of the Hemiptera. Fascicle I: Membracidae. Smith College Press, Northampton, Massachusetts, 581 pp.","Metcalf, Z. P. & Wade, V. (1965) General Catalogue of the Homoptera. A supplement to fascicle I - Membracidae of the General Catalogue of Hemiptera. Membracoidea. In Two sections. Section I. Part 1. Membracidae, Centrotinae, Platybelinae, Hoplophorioninae, Darninae. North Carolina State University, Raleigh, North Carolina, 743 pp.","McKamey, S. H. (1998) Taxonomic catalogue of the Membracoidea (exclusive of leafhoppers): second supplement to fascicle 1 - Membracidae of the general catalogue of the Hemiptera. Memoirs of the American Entomological Institute, 60, 1 - 377.","Fairmaire, L. M. H. (1846) Revue de la tribu des Membracides. Annales de la Societe Entomologique de France, 4, 235 - 320."]}

Hyphinoe Stål, 1867 Hyphinoe Stål 1867: 538 (type species: Hemiptycha cuneata Germar, 1835); Metcalf and Wade 1965: 691; McKamey 1998: 476. Ictaranthe Fowler 1895 b: 75 (type species: Ictaranthe latifrons Fowler, 1895). Tauriona Buckton 1903 a: 259 (type species: Tauriona obesum Buckton, 1903). Trapezoida Buckton 1905a: 335 (type species: Trapezoida hirsuta Buckton, 1905)., Published as part of Flórez-V, Camilo & Evangelista, Olivia, 2021, A revision of the treehopper genus Bubalopa Stål illuminates the systematics of Hyphinoini (Hemiptera: Auchenorrhyncha: Membracidae), pp. 529-551 in Zootaxa 5052 (4) on page 532, DOI: 10.11646/zootaxa.5052.4.4, http://zenodo.org/record/5577275, {"references":["Stal, C. (1867) Bidrag till Hemipterernas Systematik. Ofversigt af Kongliga Vetenskaps - Akademiens Forhandlingar, 7, 491 - 560.","Metcalf, Z. P. & Wade, V. (1965) General Catalogue of the Homoptera. A supplement to fascicle I - Membracidae of the General Catalogue of Hemiptera. Membracoidea. In Two sections. Section I. Part 1. Membracidae, Centrotinae, Platybelinae, Hoplophorioninae, Darninae. North Carolina State University, Raleigh, North Carolina, 743 pp.","McKamey, S. H. (1998) Taxonomic catalogue of the Membracoidea (exclusive of leafhoppers): second supplement to fascicle 1 - Membracidae of the general catalogue of the Hemiptera. Memoirs of the American Entomological Institute, 60, 1 - 377.","Buckton, G. B. (1903) A Monograph of the Membracidae. Lovell Reeve, London, 296 pp."]}

Key to Bubalopa species 1. Yellow, dark brown or combination thereof; dorsal outline of posterior pronotal process weakly to moderately arched past humeral angles (Figs. 4B, 4G, 5C, 5G, 9G); forewings not strongly reticulate (20 supernumerary veins distributed along apical 1/3).............. B. iguaque sp. nov. 2. Yellow with orange tinge and black punctations; suprahumeral horns slender, longer than wide basally, variable in length and orientation (Fig. 5); dorsal outline of posterior pronotal process feebly arched or gradually descending (Figs. 4B, 4G, 5C, 5G), lateral margins with weakly marked semi-circular impressions (Figs. 4F, 5B, 5F).................. B. furcata (Fairmaire) - Dark brown with reddish tinge and irregular yellow patches; suprahumeral horns robust and triangular, approximately as long as wide basally; dorsal outline of posterior pronotal process moderately arched, abruptly narrowed at posterior ½ (Fig. 9G); lateral margins with strong semi-circular depressions (Fig. 9F–G).............................. B. obscuricornis Stål, Published as part of Flórez-V, Camilo & Evangelista, Olivia, 2021, A revision of the treehopper genus Bubalopa Stål illuminates the systematics of Hyphinoini (Hemiptera: Auchenorrhyncha: Membracidae), pp. 529-551 in Zootaxa 5052 (4) on page 537, DOI: 10.11646/zootaxa.5052.4.4, http://zenodo.org/record/5577275

Bubalopa furcata (Fairmaire, 1846) (Figs. 3–8) Hemiptycha furcata Fairmaire, 1846: 314. Bubalopa furcata Stål, 1869c: 255; Funkhouser, 1927: 117; Metcalf and Wade, 1965: 701; McKamey, 1998: 476. Diagnosis: Overall color ochraceous yellow, punctation reddish black, antero-dorsal surface of suprahumeral horns dark brown with slight red tinge, tibia black, spotted yellow medially, anterior and posterior margins orange. Suprahumeral horn orientation: 35–50º (frontal view), 25–40º (dorsal view); dorsal outline slightly arched, descending in almost straight line from above humeral angles to apex of posterior process. Males: parameres Tshaped, subgenital plate abruptly narrow apically. Description: ADULTS. Color. Head and pronotum ochraceous yellow with reddish dark brown punctation. Eyes grey, ocelli yellow, surrounding cuticle forming a brown ring. Antero-dorsal surface of suprahumeral horns dark brown with slight red tinge. Forewings slightly to entirely opaque, amber to reddish dark brown with black punctation. Thoracic sternites, coxa, trochanter, femora and tarsomeres ochraceous yellow; dorsal surface of tibiae black spotted yellow, anterior and posterior margins orange, ventral surface ochraceous yellow. Head: Triangular, wider than long; suprantennal ledges discreetly sinuous, apex of frontoclypeus acute. Thorax: Suprahumeral horns 2–2.5x as long as wide basally, orientation: 35–50º (frontal), 25–40º (dorsal); dorsal carina of suprahumeral horns less sharp than others; dorsal contour weakly arched, highest above humeral angles, descending in almost straight line to apex of posterior process; semicircular impression at each side of median carina, approximately at middorsum. Abdomen: Terga III–V with a pair of oval dorsal pits (scars) (Figs. 7A, 7D, 8A, 8F). Male: Apodeme reaching posterior margin of sternite III. Lateral plate fused with pygofer. Subgenital plates wider at basal half, with slight dorsal hump, abruptly narrow in lateral view. Parameres broad, T-shaped, bifurcate distally into fingerlike lobes, weakly curved and round apically. Aedeagus U-shaped, several rows of minute teeth along entire length of anterior surface of posterior arm. Female: Gonoplac blade-shaped, covered in setae, distinctly more sclerotized along posteroventral margin. First valvulae blade-shaped, broad throughout; dorsal margin, from mid-length to apex, finely sculpted in short diagonal lines, gradually turning into well-marked striations, extended transversally across triangular and acute apex. Second valvulae blade-shaped, slightly broader at apex, two prominent teeth on apical third, quadrangular teeth on dorsal contour apically, located after apical prominent tooth, apical margin dorsally serrated, ramus extended up to apical 1/5. 4 TH /5 TH INSTAR NYMPH. Color : Mostly green in live specimens, brown in dry preserved specimens. Surface: vertex, pronotum and abdomen covered in minute, stout and subcylindrical setae, sparser on metathorax and mesonotum, abdominal segment I+II glabrous; elongate and slender setae covering ventral surface of body and legs, more densely distributed on ventral side of head. Tuberculate chalazae present on post-ocular lobes, suprahumeral buds, entire prothorax (except for pre-metopidium), anteroventral margin at base of forewing pads, mesonotal and abdominal scoli, ventrolateral margins of abdominal terga (Fig. 6E, ach), and anterodorsal and posterodorsal margin of tibia; chalazae and their setal portion varying in size, larger and prominent along the anterior surface of thoracic and abdominal scoli. Head: subtriangular; vertex bearing 10 scoli, arranged into 5 bilaterally symmetric pairs: 2 slender scoli on upper margin, equidistant from eyes and coronal suture, curved upwards (Fig. 6D, hs1); 2 stout scoli, each at external anterior margin, juxtaposed to eyes (Fig. 6D, hs3); 2 slender scoli (longest in vertex), each near external anterior margin, adjacent to latter pair, distinctly curved downwards (Fig. 6D, hs2); 2 smaller scoli, each adjacent to eyes, right below the latter pair (Fig. 6D, hs4); 2 small and stout scoli at inferior margin, right below eyes, slightly curved posteriorly (Fig. 6D, hs5). Thorax: prothorax well developed, tall and laterally flattened antero-dorsal projection, bearing 2 robust, conical scoli, curved forward apically (=postmetopidium scoli sensu McKamey et al. 2015) (Fig. 6F, pos); pair of minute scoli on metopidium (=premetopidium scoli sensu McKamey et al. 2015) (Fig. 6F, prs); suprahumeral buds flap-like, folded posteriorly onto sides of prothorax (Fig. 6C, shb); posterior process triangular, reaching anterior margin of metanotum; meso- and metanotum with 1 pair of small scoli each (Fig. 6F, mss, mts); ventral margin of forewing pads slightly emarginate anteriorly, sheltering metathoracic leg in repose, venation barely perceptible (tinged bright green in live specimens), pads reaching fourth abdominal segment; hindwing pads subtriangular, as long as forewing pads, anal margin mostly exposed. Abdomen: Abdominal terga III–V with pair of greatly enlarged dorsal scoli, each pair slightly arched in frontal and dorsal view in U-shaped outline (Fig. 6C, 6F, 6I, asIII-asV), terga VI–VIII with pair of small spinelike scoli (Fig. 6F, sps). Segment IX tubular, elongate (more than half the length of rest of abdomen), bearing one pair of apical spine-like scoli directed posteriorly (Fig. 6F, asIX) and slightly diagonally in V-shape. Measurements (in mm): Female (n=3, including lectotype)/male (n=1): Body length: 13.48/11.62; forewing length: 11.94/10.34; pronotal length: 11.91/9.99; pronotum height: 3.41/2.64; pronotal width: 4.50/3.46; head width: 4.08/3.28; vertex width: 2.81/2.19; vertex length: 2.17/1.72. Lectotype designation for Hemiptycha furcata Fairmaire, 1846: 314. In the original description of Hemiptycha furcata, Fairmaire (1846) specified Bogota as the type-locality, and the Signoret Collection as the repository of the examined material. When erecting Bubalopa and redescribing Bubalopa furcata, Stål (1869) also listed Bogota as the type-locality and mentioned that the specimen he examined had a mutilated head. The single specimen currently housed at the Naturhistorisches Museum Wien closely matches the description of Fairmaire and Stål both in terms of morphological features and its state of preservation. However, while the determination label of this specimen correctly states ‘ furcata ’, the locality label reads ‘ Bolivien’, contradicting the records from both of these authors. The Signoret collection was sold to the Naturhistorisches Museum Wien, Vienna, Austria (NMW) in 1890 after the death of Victor Signoret in 1889. After its acquisition, Signoret’s specimens were re-labelled by Anton Handlirsch (Rabitsch 2000). Each specimen received at least two labels, which were highly characteristic in terms of appearance and style. One label features a handwritten locality (usually one word, e.g., ‘ Bolivien ’) and a printed collection source (i.e., ‘Coll. Signoret’). The second label includes the species identification, containing a handwritten epithet (i.e. ‘ furcata ’), and the printed name of the identifier/author (i.e., ‘det. Signoret’) (Figure 4C). Fairmaire and Stål’s manuscripts, published between 1846 and 1869, indicate that they examined the type before it was transferred to NMW. The authors, therefore, correctly reported the locality from Signoret’s original labels before it was incorrectly transcribed by Handlirsch. Transcription errors, although rare, have been observed in other hemipteran specimens that Handlirsch curated (e.g., Reduviidae: Phymatinae; Rabitsch 2000). Despite this label inconsistency, there is strong indication that this exemplar is the syntype of Hemiptycha furcata, from Bogota, not Bolivia, which was examined first-hand by Stål. The term ‘Bogota’ was often used in the 19 th century to refer to multiple sites in Colombia, and therefore will not be included in the distribution of this species. Distribution: COLOMBIA: Antioquia: Medellín (Corregimiento de Santa Elena, 2500 masl); Puerto Berrío (Vereda Cristalina, 400 masl) (Fig. 11). Examined Material: Lectoype female (here designated): “ Bolivien \ Coll. Signoret.”, “ furcata \ det. Signoret.” [original type locality: ‘ Bogota’, incorrectly transcribed by A. Handlirsch] (NMW). COLOMBIA: Antioquia: Medellín: “ COLOMBIA. Antioquia, Medellín, Corregimiento Santa Elena, Reserva SMA San Pedro, 6.194048ºN, 75.499603ºW, 2600 msnm, Manual, en Senna pistaciifolia (Fabaceae), Jun. 2019, leg. Semillero de Entomología 2019-I (C. Flórez-V), CBUCES-F 9036 (1 male, 1 female, CBUCES); “ COLOMBIA. Antioquia, Medellín, Corregimiento Santa Elena, Reserva SMA San Pedro, 6.194048°N, 75.499603°W, 2600 msnm, Manual, en Senna pistaciifolia (Fabaceae), Jun. 2020, leg. C. Flórez-V, A. Ospina, CBUCES-F 8002 ”, “ CBUCES-F 8003 ”, “ CBUCES-F 8023 ” (1 male, 2 females, CBUCES); “ COLOMBIA. Antioquia, Medellín, Corregimiento Santa Elena, Reserva SMA San Pedro, 6.194048°N, 75.499603°W, 2600 msnm, Manual, en Senna pistaciifolia (Fabaceae), Sep. 5/2020, leg. C. Flórez-V, A. Ospina, CBUCES-F 8004 ” (1 nymph, CBUCES). Puerto Berrío: “ COant. Puerto Berrío Vda. / Cristalina R.U.N.A. 400–500m / Manual 22-sep-2005 / A. Acosta / CEUA 66707 ” (1 female, CEUA). Biology: Adults and immatures were observed feeding on branches of Senna pistaciifolia (Kunth) H.S. Irwin & Barneby (Fabaceae).Adults and nymphs are highly cryptic. The color, shape, and ultrastructure of scoli and chalazae help nymphs blend into the apical indumenta of the host plant. Nymphs closely resemble host plant stipules, where they usually feed. Adults were consistently observed on ligneous plant tissue feeding on subapical branches. Both nymphs and adults are solitary, not associated with ants or other hymenopterans, and were sparsely distributed in the surveyed transects. For example, only five adults and five nymphs were collected after thorough inspection of 50 host plants in that area, which suggests low population density. Other treehopper species were found to co-occur in the same host plant individual: Bordoniana nigricosta (Goding, 1926) (Smiliinae: Acutalini), an unidentified species of Membracinae, and several aggregations of Aetalion cf. nigromarginatum (Aetalionidae) including multiple adults and nymphs. Remarks: Representatives of B. furcata are easily distinguished from other Bubalopa by their overall color, ochre yellow and reddish brown punctation, acute frontoclypeus, slender suprahumeral horns, and a nearly diagonal dorsal outline. The supernumerary crossveins on the forewings vary intraspecifically and can be as numerous as in B. obscuricornis but still fewer than in B. iguaque. In males, suprahumeral horns are more horizontal (35º–40º, frontal view; n=2) than in females. A larger sampling of females indicates that there is considerable variation in horn orientation (35º–50º, frontal view; n=5), although four of the five examined females showed an angle from 40º–50º. While those differences are evident in frontal view, horns appear similar in lateral and dorsal view in both sexes for all exemplars. As observed in other species in the genus, B. furcata bears prominent oval pits on the dorsolateral area of abdominal terga III–V, which are thought to be scars resulting from nymphal scoli (Lencioni-Neto and Sakakibara 2013). The small semicircular impressions on the pronotal posterior process in adults, which appear to be topologically correlated with scoli in the nymphs, are also visible in A. robustus. These traits are not observed in other species of Bubalopa (see remarks after the redescription of the genus). Some structures of the male genitalia of B. furcata are highly characteristic, including the subgenital plate, with a large dorsolateral indentation, and the robust and T-shaped paramere. These two features are not observed in any other Membracidae. The internal lateral margin of abdominal pleurites, and the lateral and anterior margin of abdominal sternites are somewhat invaginated into the abdomen. The spiracles are located in the invaginations of the pleurite. The second valvulae of B. furcata (Figs. 8I–J) differs from the figure presented in Deitz (1975) for the same species (Fig. 24O, pp. 85). We believe that Deitz (1975) illustrated a specimen of Eualthe, which had been previously misidentified by Funkhouser as Bubalopa. This is the first species of Bubalopa for which nymphal morphology is described. The relevance of these characters to tribe and genus classification are discussed elsewhere in this paper. Nymphs were collected on the same date and same host plants as adults. The specimen from Puerto Berrío was incorrectly referred to in Flórez-V et al. (2015) as a representative of Sundarion. The authors commented that this exemplar did not exhibit femoral cucullate setae as in Sundarion but listed other characters as deep pronotal punctations or supernumerary forewing crossveins, which are characteristic of Bubalopa., Published as part of Flórez-V, Camilo & Evangelista, Olivia, 2021, A revision of the treehopper genus Bubalopa Stål illuminates the systematics of Hyphinoini (Hemiptera: Auchenorrhyncha: Membracidae), pp. 529-551 in Zootaxa 5052 (4) on pages 537-545, DOI: 10.11646/zootaxa.5052.4.4, http://zenodo.org/record/5577275, {"references":["Fairmaire, L. M. H. (1846) Revue de la tribu des Membracides. Annales de la Societe Entomologique de France, 4, 235 - 320.","Funkhouser, W. D. (1927) General Catalogue of the Hemiptera. Fascicle I: Membracidae. Smith College Press, Northampton, Massachusetts, 581 pp.","Metcalf, Z. P. & Wade, V. (1965) General Catalogue of the Homoptera. A supplement to fascicle I - Membracidae of the General Catalogue of Hemiptera. Membracoidea. In Two sections. Section I. Part 1. Membracidae, Centrotinae, Platybelinae, Hoplophorioninae, Darninae. North Carolina State University, Raleigh, North Carolina, 743 pp.","McKamey, S. H. (1998) Taxonomic catalogue of the Membracoidea (exclusive of leafhoppers): second supplement to fascicle 1 - Membracidae of the general catalogue of the Hemiptera. Memoirs of the American Entomological Institute, 60, 1 - 377.","McKamey, S. H., Wallner, A. M. & Porter, M. J. (2015) Immatures of the New World treehopper tribe Amastrini (Hemiptera, Membracidae, Smiliinae) with a key to genera. ZooKeys, 524, 65 - 87. https: // doi. org / 10.3897 / zookeys. 524.5951","Rabitsch, W. (2000) Type specimens of Phymatidae in the Natural History Museum Vienna (Insecta: Heteroptera). Kataloge der wissenschaftlichen Sammlungen des Naturhistorischen Museums in Wien, 15 (7), 1 - 21.","Lencioni-Neto, F., Sakakibara, A. M. (2013) Alcmeone robustus (Hemiptera: Membracidae: Darninae): Description of the lastinstar nymph and biological notes. Zoologia, 30 (5), 471 - 474. https: // doi. org / 10.1590 / S 1984 - 46702013000500001","Deitz, L. L. (1975) Classification of the higher categories of the New World treehoppers (Homoptera: Membracidae). North Carolina Agricultural Experiment Station, Technical Bulletin, 225, 1 - 177.","Florez-V, C., Wolff, M. & Cardona-Duque, J. (2015) Contribution to the taxonomy of the family Membracidae Rafinesque (Hemiptera: Auchenorrhyncha) in Colombia. Zootaxa, 3910 (1), 1 - 261. https: // doi. org / 10.11646 / zootaxa. 3910.1.1"]}