taxonID	type	description	language	source
5C16B55CFFA2882F46CDBFA7FA0EF930.taxon	type_taxon	Type species: Otiocerus rubescens Fowler 1900 by original designation	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA2882F46CDBFA7FA0EF930.taxon	diagnosis	Amended diagnosis. (modified from Fennah 1952). Elongate, fragile forms, ~ 5 – 9 mm in body length (incl. wings), often pale with varying degrees of reddish wash (wing veins often reddish) and posterior margin of mesonotum with row of 6 fuscous spots (sometimes obsolete). Head strongly compressed and projected in front of eyes (for a distance about equal to width of eyes in longest dimension); leading margin of head rounded in lateral view. Vertex about twice as long as broad at base; posterior margin excavate, lateral margins converging distally; disc strongly depressed, lateral carinae bearing pustules. Antennae elongate but not reaching anterior margin of head, scape wider than long, pedicle elongate, subantennal process absent. Pronotum short medially, strongly declinate in lateral view, in dorsal view anteriorly sharply convex, posteriorly deeply angularly concave; medially carinate. Mesonotum large, about as broad as long, weakly tricarinate; posterior margin of mesoscutellum transverse, ecarinate. Forewing elongate and broadly spatulate, widest about in distal three-fifths (composite vein Sc + R + M very short), fork of CuA about on level with fusion of Pcu and A 1 in clavus, clavus open, composite vein Pcu + A 1 joining with CuP and CuA before reaching wing margin. Apex of forewing broadly rounded, apical cells at least twice as long as broad. Male terminalia in lateral view with pygofer narrow, irregularly quadrate, anterior and caudal margins sinuate, medioventral process absent. Gonostyli elongate, exceeding anal tube, medially cupped, in ventral view medial margins sinuate (lacking median processes). Aedeagal shaft tubular, weakly upcurved bearing varied apical or subapical sclerotized processes and membranous retrorse endosoma bearing varied sclerotized processes. Anal segment short, apical angles shortly produced lateroventrad.	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA2882F46CDBFA7FA0EF930.taxon	discussion	Remarks. Among New World Otiocerini, Cobacella can be recognized by the superficial characteristics of being midsized and usually with a reddish wash, mesothorax bearing 6 fuscous spots along caudal margin (sometimes absent), head roundly produced (forward projecting, not upward), subantennal processes absent and forewings relatively broad. Fennah (1952) noted that in life they hold their wings slightly spread with the apices nearly touching the substrate (as opposed to upward and slightly spread in Anotia). The lack of a subantennal process distinguishes this genus from Apache Kirkaldy, 1901, Otiocerus Kirby, 1821, Shellenius Ball, 1928, and Kubilaya Koçak & Kemal, 2010 (see Fennah 1952, fig. 37 A and Fennah 1945, figs 39, 40 as Iquitosa Fennah, 1945) and Labicerus Erichson, 1848 (noted as present in the description Erichson, 1848). The rounded, forward-projecting head of Cobacella distinguishes it from Anotia, Sayiana Ball, 1928, Kubilaya and Heronax (Homometria), which are subconical and upward projecting, and from Platocerella Fennah 1952 and Platonax Metcalf, 1938 which are not as strongly projecting as Cobacella. Cobacella is an infrequently encountered genus and is poorly represented in institutional collections and on citizen science forums (4 putative records on iNaturalist). It appears that the species of Cobacella may possess sexual dimorphism (possibly differences in the projection of the head), intraspecific color variation, and may have some or all of the fuscous spots of the mesothorax obsolete. The lack of specimens in series and uncertainty in the association of males and females has made it difficult to assess intraspecific species variability. Cobacella rubescens (Fowler) was described (as Otiocerus (?) rubescens) based on 2 syntype specimens from Mexico (Figs 9 B, C, 10) Cobacella sexguttata was described based on a male holotype and female paratype from Trinidad. See ‘ Other material examined’ for specifics.	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA2882F46CDBFA7FA0EF930.taxon	biology_ecology	Plant associations. No plant associations are reported for Cobacella rubescens. The holotype of C. sexguttata was collected “ resting on leaves of wild tobacco, Acnistus arborescens ” (Solanaceae; Fennah 1952). The new species was collected in association with African oil palm, Elaeis guineensis Jacq. (Arecaceae).	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA2882F46CDBFA7FA0EF930.taxon	etymology	Etymology. The genus name is feminine in gender and appears to be formed from Cobax + - ella (diminutive suffix); Cobax is a derbid genus name, possibly originating from the Hungarian word ‘ kovács ’ (blacksmith) (Dmitriev 2002).	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	description	(Figures 2 – 6)	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	materials_examined	Type locality. Costa Rica, Puntarenas Province, Costa Rica.	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	diagnosis	Diagnosis. Description. Color. General body color yellow, darker in males, paler ventrad (Fig. 2). Head in lateral view dark orange, yellow around eyes and antennae, antennae concolorous with body. Mesonotum with six black spots, three each side of midline, anterior-most rounded adjacent to tegulae, second pair crescent-shaped at lateral extreme of scutum, third pair circular, near caudal extreme of lateral carinae (Fig. 2). Wings fuscous with red veins, darker in males (Fig. 2). Legs pale yellow, with variable reddish wash, especially proximad, male with reddist tarsi. Dorsum of abdomen brown. Structure. Body length (including wings) males: 7.0 mm (n = 1) and females: 7.8 mm (n = 2). Head. Strongly compressed, projecting beyond eyes for distance about equal to greatest width of eye, in lateral view, head rounded, vertex declinate smoothly rounded at fastigium. Vertex in dorsal view narrowly triangular, approximately twice as long as wide at base, disc strongly concave, posterior margin strongly concave, lateral margins converging anteriorly (nearly meeting at apex), strongly carinate and thickened bearing row of pits (Fig. 3 A), continued in reduced form on lateral margins of frons (Fig. 3 B). In frontal view, frons greatly compressed, lateral margins strongly carinate, closely approximate (Fig. 3 C). Antennae cylindrical and tubular, scape ring-like, pedicle approximately four times as long as wide (Fig. 3). Eyes roughly oval, emarginated above antennae, lateral ocelli faintly indicated anterior to antennae. Thorax. Prothorax steeply inclined in lateral view (Fig. 3 B), narrow in dorsal view, strongly convex at anterior margin, strongly concave at posterior margin, expanding slightly at lateral margins (Fig. 3 B); tricarinate, median carina strong, lateral carinae weak, arising from median carina near midlength, becoming obsolete laterad (Fig. 3 A, B). Mesonotum large, scutum convex in lateral view with concave inflection at leading margin of scutellum, in dorsal view posterior margin of mesoscutellum truncate; in dorsal view with tricarinate, median carina extending from anterior margin to posterior margin, inner lateral carinae sinuate, curved distad at anterior margin (Fig. 3 A), lateral margin of mesothorax carinate, arising near midpoint, forming posterior lip of the mesothorax (Fig. 3 B). Spinulation of hind tibiae 5 - 5 - 4. Forewings broad and spatulate, narrowest basally, widest near midlength, costal margin weakly convex, broadly rounded, trailing margin broadly convex except for concave expansion in jugal region. Clavus open, composite vein Pcu + A 1 projecting anteriorly to encompass CuP and branches of CuA before arched to wing margin past midlength; 22 vein branches reaching wing margin between ScP and A 1 + Pcu + Cu inclusive; marginal cells longer than wide. MP branching from ScP + R in base of wing just beyond basal cell, with fork of RP from Sc + RA shortly beyond fork of MP; ScP reaching wing margin just before wing midlength. Branching pattern RA 8 - branched, RP 2 - branched, MP 10 - branched, CuA 2 - branched (anastomosed to form closed procubital cell (Emeljanov 1996). Terminalia. Pygofer in lateral view narrow, irregular, strongly narrowed at dorsal margin, strongly sinuate on anterior margins, posterior margin bearing large rounded lobe in dorsal half, (Fig. 5 A); medioventral process absent (Fig. 5 B). Gonostyli in lateral view spatulate, medially cupped, irregularly sinuate on dorsal and ventral margins, inner dorsal margin with elongate process with sclerotized hooked apex (Figs 5 A – C); in ventral view, gonostyli narrow basally, expanding greatly at midpoint, truncate at apex with slight invagination near inner margin (Fig. 5 B). Aedeagal shaft tubular, simple, slightly upcurved, spinose flange on dorsal margin near midpoint (A 1) and process arising subapically on left, dorso-lateral margin (A 2), angled dorsad, curving slightly ventrad, nearly reaching apex of flagellum, reaching anterior margin of A 1 (Fig. 6), and serrulations on right lateral margin, from midlength to near apex (Fig. 6 A, 6 D). Flagellum bilaterally asymmetrical, two large processes on left lateral side along dorsal margin (F 1 & F 2), process F 1 on outer lateral margin, generally uniform in width, apex slightly constricted, inner and outer margins sinuate, apex curved slightly mesad; F 2 more constricted at apex and curved mesad, F 1 and F 2 curved ventrad, approximately equal length (Fig. 6 A & 6 B). Anal tube in lateral view slender, dorsal margin straight, ventral margin sinuate, apex angled ventrad, almost reaching apex of gonostyli (Fig. 6 A); in dorsal view, apex appears strongly forked and pointed (Fig. 6 B).	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	biology_ecology	Plant associations. African oil palm, Elaeis guineensis Jacq. (Arecaceae).	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	distribution	Distribution. Puntarenas Province, Costa Rica.	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	etymology	Etymology. The specific epithet is in reference to the town where specimens were collected “ La Palma ”, formed by the trucation of the town name as “ palm- “ with the suffex “ - ensis ”, meaning from a place or location. The specific name is intended as feminine to agree with the genus, although the spelling would be the same in either feminine or masculine form.	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	materials_examined	Material examined. Holotype male “ Costa Rica, Puntarenas Pr. / Finca La Palma / 6. VI. 2021, sweeping palms / Coll.: B. W. Bahder // Holotype / Cobacella palmensis ♂ / ” (FLREC); Paratypes same as holotype (2 females, FLREC). Sequence Data. For the barcoding region of COI, a 584 bp product was generated, for the 18 S gene, a 1,305 bp sequence was generated, and for the D 9 - D 10 region of the 28 S gene, a 776 bp sequence was generated. The phylogeny generated based on the available data and taxa showed low bootstrap support for most clades for COI, except for the genus Patara, which had moderate support (83) (Fig. 7 A). However, the phylogeny based on 18 S showed strong bootstrap support (97) for C. palmensis sp. n. resolving adjacent to Sayiana sayi (Fig. 7 B). The phylogeny generated based on 28 S showed moderate support (69) for placement of C. palmensis sp. n. adjacent to Anotia, however strong bootstrap support (100) for Anotia as a distinct clade from Cobacella (Fig. 7 C). Both the 18 S and 28 S derived phylogenies showed strong bootstrap support (100 and 99 respectively) for a clade comprised of Anotia, Cobacella, and Sayiana. The consensus tree based on all three loci also shows strong bootstrap support (87) for the placement of C. palmensis sp. n. adjacent to Sayiana sayi and also showed strong support (100) for the clade comprised of Anotia, Cobacella, and Sayiana (Fig. 7 D). Based on the pairwise comparison for the 18 S gene, the average intrageneric variability is 1.8 % (± 0.2) while the average intergeneric variability is 11.7 % (± 0.5) (Table 4). Cobacella palmensis sp. n. differs on average by 12.3 % (± 1.3) from all other genera included in this analysis with the lowest level of variation occurring when compared to Sayiana sayi where it differs by 5.3 % and the highest level of variance (excluding cenchreine outgroup) occurring with S. ballii and P. vanduzei, varying by 14.9 % (Table 4). Based on the pairwise comparison for 28 S, the average intrageneric variability is 3.9 % (± 0.1) whereas the average intergeneric variability is 16 % (± 0.7) (Table 5). Based on 28 S, C. palmensis sp. n. differs on average by 15.3 % (± 0.2) from the other genera analyzed, with the lowest level of variability observed with Sayiana sayi (7.7 %) and the highest level of variability observed with Mula resonans (23.2 %) (Table 5).	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	discussion	Remarks. The placement of C. palmensis sp. n. in Cobacella is supported by morphological characters observed and compared to both C. rubescens and C. sexguttata. However, a paucity of specimens from institutional collections, and limited observations on citizen science forums like iNaturalist, make it difficult to ascertain the extent of intraspecific variation or sexual dimorphism within species. In this manuscript we are annotating all the specimens or observations of which we are aware. Cobacella palmensis sp. n. differs from C. sexguttata in both the structure of aedeagus and wing coloration. Fennah (1952: 163, figs 34 H, 34 I) illustrated the aedeagus of C. sexguttata, which he described as “ … asymmetrical, sinuately tubular; flagellum with a single spinose process at its base, and a flattened plate acute at tip on left · on right two sclerties distally acute, supporting membranous portion ”. Fennah’s (1952) illustrations are imprecise, but it is evident that the aedeagus of C. sexguttata is more strongly asymmetrical, lacks the spinose flange on dorsal margin (found in C. palmensis sp. n., A 1 in Fig. 6) near midpoint, and has a different arragements of sclerotized spines of the flagellum. With respect to wing coloration, C. sexguttata has a more fuscous patch at the apex of the forewing with basal portion less fuscous while C. palmensis sp. n. has a uniform tint to the forewing. Concerning C. rubescens, it appears that the wing membrane is uniformly fuscous in C. palmensis sp. n. and more clearly hyaline in C. rubescens. In addition, C. palmensis sp. n. differs from C. rubescens in the size and shape of black spots on the mesothorax. Unfortunately, the male terminalia of C. rubescens has not been described and the syntypes are female. Aside from type material, the only other available specimen of Cobacella is a pallid female from La Selva research station in Costa Rica (Fig. 11) with the mesothoracic spots reduced to one on each side. This specimen likely represents an additional species. On iNaturalist there are four additional records of Cobacella. Two are from Puntarenas, Costa Rica (# 151547369, 146292818) that appear to represent C. palmensis sp. n., one from San José, Costa Rica (# 88345785) that may be C. rubescens, and one from S „ o Paulo State, Brazil (# 146995673) that we are reluctant to speculate on because it is geographically distant from the other observations.	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
5C16B55CFFA5882746CDBCD6FE55FE23.taxon	materials_examined	Other material examined. Cobacella rubescens (BMNH, female syntypes, Figs 9 B, C) Teapa, Mexico, H. H. Smith; (NHMW, female, Fig. 10) “ [Dominik] Bilimek / Mexico / 1871 / Orizaba // Juni [Duch for June] // rubescens / det. Fowler // Otiocerus? rubescens Fowler / Type ” (in Mus. Vind. Caes. = in Museo Vindobonense Caesarei = Imperial Museum Vienna). Cobacella sexguttata (BMNH, male); Amazonas, Fonteboa, Broomfield (det: Broomfield); and male holotype and female paratype (BMNH, in alcohol) “ Cobaccella / sexguttata / Fenn. Type / 1949 / Trinidad B. W. I. ” reported as La Reunion, Carapo, 14. ix. 1949, R. G. Fennah. Cobacella sp. (UDCC, female): Costa Rica, Heredia nr Puerto Viejo, La Selva Biol Sta. (at station) 179 ft, N 10 25 ’, W 84 00, 24 Feb. 2004, light, CR Bartlett, J. Cryan & J. Urban. iNaturalist records: Costa Rica (observations # 146292818, 151547369, 88345785); Brazil (S „ o Paulo) (observation # 146995673).	en	Zumbado Echavarria, Marco A., Barrantes Barrantes, Edwin A., Helmic, Ericka E., Bartlett, Charles R., Bahder, Brian W. (2023): A newspecies ofplanthopper in thegenus Cobacella (Hemiptera: Auchenorrhyncha Derbidae) from oil palm (Elaeis guineensis) in Costa Rica. Zootaxa 5351 (1): 107-121, DOI: 10.11646/zootaxa.5351.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5351.1.4
