A new species of planthopper in the genus Tico (Hemiptera: Auchenorrhyncha Derbidae) on palms from lowland tropical rainforest in Costa Rica Author Bahder, Brian W. University of Florida, Department of Entomology and Nematology-Fort Lauderdale Research and Education Center; 3205 College Ave., Davie, FL 33314 - 7719, USA. Author Zumbado Echavarria, Marco A. Universidad de Costa Rica-Sede San Ramón, Departamento de Ciencias Naturales, de la Iglesia el Tremedal 400 mts al Oeste carretera hacia San Pedro, San Ramón, Alajuela, Costa Rica. Author Barrantes Barrantes, Edwin A. Universidad de Costa Rica-Sede San Ramón, Departamento de Ciencias Naturales, de la Iglesia el Tremedal 400 mts al Oeste carretera hacia San Pedro, San Ramón, Alajuela, Costa Rica. Author Helmick, Ericka E. University of Florida, Department of Entomology and Nematology-Fort Lauderdale Research and Education Center; 3205 College Ave., Davie, FL 33314 - 7719, USA. Author Bartlett, Charles R. University of Delaware, Department of Entomology and Wildlife Ecology, 250 Townsend Hall, Newark, DE 19716 - 2160, USA. text Zootaxa 2022 2022-04-11 5125 3 283 294 journal article 56072 10.11646/zootaxa.5125.3.2 262e78ea-b4ce-45b0-bbf8-fc4ea6dac33d 1175-5326 6443733 F2940AF8-2CAE-4651-9100-A6C9F7430A9D Tico sierra Bahder & Bartlett sp. n. ( Figures 2–6 ) Type locality. Costa Rica , Heredia , La Selva Biological Station . Diagnosis. Distinguished from congeners by the red-tinted fuscous coloration over most of body and wings (but venter and legs pale). Pygofer with lateral margins of opening posteriorly expanded. Aedeagal shaft with dorsal process and pair of ventral serrated flanges. Gonostyli with sclerotized process on inner margin (near midlength) and pair of processes on dorsal margin (proximal process short and sclerotized, distal elongate, arched anteriorly). Anal tube with ventrodistal portion narrow and elongate (not broadened). Description. Color . General body brownish-orange (thorax) to fuscous (abdominal terga) with red tint ( Fig. 2 ); ventral portion of thorax (including legs) and abdomen ivory, clypeus fuscous dorsad, fading to testaceous yellow ventrad after midlength ( Fig. 3 ). Antennae whitish (tinged pink around flagellum). Nota of pro- and mesothorax (in dorsal view) orangish tan, margined with red; paranota (lateral view) paler ventrally. Forewings fuscous with red veins, more strongly tinted red on leading and apical margins, clear patches along wing margin at apex of RA, RP 2, and between MP 4 and CuA ( Fig. 4 ). FIGURE 2. Adult male habitus Tico sierra sp. n. ; A) body lateral view and B) body dorsal view, scale = 1 mm. Structure. Body length (without wings) 1.51–1.52 mm (males), 1.60–1.62 mm (females) ( Table 5 ). Head. In lateral view rounded (with slight hump marking transverse suture at fastigium) ( Fig. 3A ). In dorsal view, vertex trapezoidal, posterior margin truncately concave, lateral margins converging to weakly convex, transverse carina, vertex slightly more than 2X as wide on posterior margin as long at midlength; lateral carinae foliately keeled bearing single row of pits, disc strongly concave relative to keeled lateral carinae ( Fig. 3B ). In frontal view, transverse carina at fastigium distinct; lateral margins of frons foliately keeled, nearly parallel, converging slightly below eyes, expanding to frontoclypeal suture, a single row of pits running length of lateral margins, median carina absent ( Fig. 3C ). Frontoclypeal suture weakly arched (concave ventrally). Lateral ocelli distinct below eyes. Antennae short, posterior to eyes, scape very short, pedicel ovoid, bearing numerous sensillae and an elongate flagellum with bulbous base. Thorax. Pronotum (dorsal view, Fig. 3B ) at midline about ¾ length of vertex, anterior margin following contour of head, medially truncate behind vertex; posterior margin moderately concave; distinctly tricarinate (lateral carinae laterally arched, reaching hind margin or nearly so); in lateral view ( Fig. 3A ), posterior margin of pronotum raised, anteriorly declining; paradiscal regions broadly foliate, forming cup-like fossa surrounding antennae posteriorly ( Fig. 3A, a tribal feature), in frontal view, foliations broadly exceeding antennae, appearing rounded ( Fig. 3C ). Forewing ( Fig. 4 ) with apex of clavus near forewing midlength, fused vein Pcu+A1 reaching wing margin distantly proximad to CuP apex. Fork of R somewhat preceding claval apex, fork of CuA proximad of R fork (cell C5 longer than C1); composite vein M+R+ScP forming elongate stem from basal cell before branching of MP from ScP+R. Branching pattern: RA 1-branched, RP 2-branched (RP 1+2 and RP 3+4 ), MP 4-branched (a forked MP 1 plus MP 2 and MP 3+4 ; given in error as MP 1 , MP 2 , MP 3, and MP 4 in Bahder et al . 2021a fig. 6), CuA 2 branched (forming cells C5 and C5’ by icua crossvein and icu forming marginal C6 cell). TABLE 5. Biometric data for type material of Tico sierra sp. n.

Male	Female
Character	Range	Average ± SE	Range	Average ± SE
Body length, with wings	2.51–2.52	2.52±0.01	2.56–2.58	2.57±0.02
Body length, no wings	1.50–1.51	1.51±0.01	1.60–1.61	1.61±0.01
Forewing length	2.20–2.21	2.20±0.01	2.26–2.27	2.26±0.01
Vertex length	0.11–0.12	0.12±0.01	0.12–0.12	0.12±0.00
Vertex width, basal margin	0.22–0.22	0.22±0.00	0.22–0.22	0.22±0.00
Vertex width, distal margin	0.95–0.95	0.95±0.00	0.95–0.95	0.95±0.00
Pronotum length, midline	0.10–0.10	0.10±0.00	0.10–0.10	0.10±0.00
Mesonotum length, midline	0.41–0.41	0.41±0.00	0.41–0.42	0.42±0.01
Mesonotum width	0.53–0.54	0.53±0.01	0.54–0.54	0.54±0.00
Frons width, dorsal margin	0.95–0.95	0.95±0.00	0.95–0.95	0.95±0.00
Frons width, clypeal suture	0.17–0.17	0.17±0.17	0.17–0.17	0.17±0.00
Frons width, widest	0.17–0.17	0.17±0.17	0.17–0.17	0.17±0.00
Frons width, narrowest	0.09–0.09	0.09±0.00	0.09–0.09	0.09±0.00
Frons length, midline	0.34–0.34	0.34±0.00	0.35–0.35	0.35±0.00
Clypeus length	0.23–0.23	0.23±0.23	0.23–0.23	0.23±0.23

Terminalia. Pygofer in lateral view irregularly sinuate on anterior and posterior margins, wide dorsally, broadening near midlength, then narrowing to form large rounded projections on posterior margin (i.e., forming lateral projections of pygofer opening); a rounded tooth just on reclining face of posterior margin; lateral face of pygofer having appearance of a diagonal fold on medial portion (a similar structure evident in Tico emmettcarri , but less evident in T. pseudosororius , Bahder et al . 2021a , figs. 7A and 15A respectively), and appearing weakly sclerotized. In ventral view, medioventral process of pygofer opening absent (slightly invaginated at ventral midline, Fig. 5B ). Gonostyli in lateral view with angular projection at base, sinuate on ventral margin, distal end curving dorsad to semitruncate apex with slight invagination, dorsal margin with large, irregular process comprised of a smaller, sclerotized nodule at anterior end and longer, slender, pale process curved cephalad ( Fig. 6A ); in ventral view, expanded at base, constricting to narrowest point at 1/3 length, expanding to widest region at 2/3 length, narrowing to pointed apices curving mesad ( Fig. 5B ); inner face with sclerotized, crescent-shaped process near midlength ( Fig. 6B ). Aedeagus approximately bilaterally symmetrical, without processes along shaft (except subapically), shaft bearing rows of irregular serrations on weak flange ventrally and on middorsal projection ( Figs 7A, 7B ); apex with two pairs of elongate dorsal subapical processes, outer pair (A1 & A2) longer, twisted and robust, inner pair (A3 & A4) shorter, more slender, and not twisted; endosoma complex, with four pairs of processes, first (E1 & E2) dorsalmost, wide, flattened, auriform, and shortest of endosomal processes; second (E3 & E4) moderately long (exceeding A1 & A2) and angled dorsomediad, third (E5 & E6) about equal in length to E3 & E4, situated on lateral margin of endosoma, curved dorsoventrad; fourth (E7 & E8) longest, ventral-most, nearly reaching base of aedeagus ( Fig. 7 ). Anal tube, in lateral view, expanding distally, roughly triangular, elongate, reaching gonostyli apices, ventral margin weakly convex, distally projected, apex truncate ( Fig. 5A ), in dorsal view spatulate (expanded distally), apex truncate ( Fig. 5C ). Plant associations. Palm seedlings ( Arecaceae ); also swept off Heliconia L. ( Heliconiaceae ). FIGURE 3. Adult male Tico sierra sp. n. ; A) head, pronotum, and mesonotum right lateral view, B) head, pronotum, and mesonotum dorsal view, and C) head and paranotal fovea frontal view, scale = 1 mm. FIGURE 4. Male Tico sierra sp. n. wing venation; black = vein, italics = crossvein, green = cell. Distribution. Costa Rica ( Heredia ). Etymology. The specific name refers to the serrations on the aedeagus, a feature not observed in other Tico with the term ‘ sierra ’ Spanish for a saw. The specific name is intended as indeclinable. Material examined. Holotype male “ Costa Rica , Heredia / La Selva Biological Station / 15.V.2018 / Coll. : B.W. Bahder , sweeping palms / Holotype Tico sierra ♂ ” ( FLREC ); paratypes same as holotype ( 3 males , 2 females , FLREC and FSCA ). Sequence Data and Analysis. For the COI gene, a 632 bp product was generated (GenBank Accession No. OK575934 ) for Tico sierra sp. n. The phylogenetic analysis ( Fig. 8A ) demonstrated moderate bootstrap support (79) for the genus Tico with T. sierra sp. n. resolving adjacent to T. emmettcarri with that clade sister to T. pseudosororius . The mean variability within Tico , Agoo , and Omolicna is 13.7% (±0.3), 16.3% (±0.8), and 16.4% (±0.8), respectively while the mean variation among genera is 20% (±0.0). For the 18S gene, a 1,329 bp product was generated (GenBank Accession No. OK577947 ) for Tico sierra sp. n. The phylogenetic analysis ( Fig. 8B ) demonstrated strong bootstrap support (99) for the genus Tico with the arrangement of Tico species the same as for COI. FIGURE 5. Male Tico sierra sp. n. terminalia; A) left lateral view, B) ventral view, and C) dorsal view. FIGURE 6. Male Tico sierra sp. n. gonostylus; A) outer lateral view and B) inner margin showing sclerotized process. The consensus tree ( Fig. 8C ) supports placement of the novel taxon in the genus Tico . The mean variability within Tico , Agoo , and Omolicna is 1.2% (±0.2), 1.1% (±0.0), and 0.7% (±0.1), respectively while the mean variation among genera is 9.1% (±0.4). The combined tree supports Tico as monophyletic, with the arrangement of species the same as presented for the individual genes. FIGURE 7. Aedeagus of Tico sierra sp. n. : A) right lateral view, B) left lateral view, and C) dorsal view; A = aedeagal process, E = endosomal process. FIGURE 8. Maximum Likelihood phylogenetic tree based on 1,000 replicates: A) COI gene, B) 18S gene, and C) consensus of concatenated COI and 18S sequences. Remarks. Both molecular and morphological evidence supports the placement of the novel taxon in Tico , however, the coloration of this species sets it apart from its congeners. The three previously described species ( Tico emmettcarri , T. pseudosororius , and T. sororius ) are all pale with predominately clear wings bearing varied markings; whereas Tico sierra sp. n. is a dark species (except ventrally) with deeply red-infuscate forewings. In addition, the presence of serrations along the dorsal and ventral margin of the aedeagus is unique. Aside from color, the structure of the head, forewing venation, and male terminalia are all fundamentally comparable to the three previously described species. For both COI and 18S, the amount of variability that Tico sierra sp. n. exhibits relative to T. emmettcarri and T. pseudosororius is consistent with intrageneric variability assessed in this study for Agoo and Omolicna . Additionally, photographs of this species were taken from various species of Heliconia at the same location the type material was collected. The collection of T. emmettcarri and T. pseudosororius from a non-palm host and the association of Tico sierra sp. n. with both palms and non-palm monocots may indicate that members of Tico are not strict palm feeders.