Haplaxius pocococo Bahder & Bartlett, sp. n.

(Figures 2–7)

Type locality. 4.2 km SE of Santa Rosa de Pocosol (10˚ 35’ 27.14” N, 84˚ 30’ 28.83” W), Alajuela, Costa Rica.

Diagnosis. This species is recognized by a deep orange facial color traversed with a rough white arch, resulting in a bell-shaped orange patch along the frontoclypeal suture. Pygofer with triangular processes on the lateral margin of the opening and an ovoide medioventral process.Aedeagus with a large lateral process that is cupped and strongly curved inward, and a large swollen lobe on the opposite lateral margin of the aedeagus.

Description. Color. In life, adult males are generally pale green; in ethanol preserved specimens, green areas fade to stramineous yellow in males (Fig. 2A,B); females more strongly colored, reddish yellow (Fig. 2C,D). Males with vertex orangish anteriorly, paler posteriorly. Face orangish above frontoclypeal suture with rough white arc, resulting in bell-shaped orange patch below; clypeus and genae below antennae white; antennae orange. Pronotum anteriorly stramineous, posterior margin broadly white. Mesonotum orange-stramineous with irregular white patches along lateral carinae and posterior margin; mesonotum paler between lateral carinae forming weak median vitta; tegulae orangish, white patch on pleuron below tegulae; legs distally white. Wings clear with 2 diffuse dark patches on posterior margin, one near apex of clavus, one in cells formed by CuA (Fig. 4). Abdomen stramineous, dorsally darker. Females similar but more strongly patterned, red-orange.

Structure. Body length (including wings), males 2.58 mm, females 2.61 mm (Table 3). Head. Head with fastigium rounded in lateral view (Fig. 3A). Lateral carinae of vertex and frons keeled and foliaceous. Vertex with lateral margins subparallel, weakly converging anteriorly, posterior margin incised (Fig. 3B); posterior margin about level with posterior margin of compound eyes; disc concave. In frontal view, face ovoid, frons widest above frontoclypeal suture, strongly narrowing distally to fastigium, transverse carina present (Fig. 3C), median carina evident in ventral half of frons, obsolete dorsally, median ocellus present (Fig. 3C). Clypeus elongate-triangular. Antennae with scape very small, pedicel bulbous, about as wide as tall, bearing many irregularly placed sensory plaques. Lateral ocelli distinct, anterior to antennae, below compound eyes.

Thorax. Pronotum short, in dorsal view narrowest medially, wider laterally; posterior margin concave; in lateral view, paradiscal region widest near dorsal margin of tegulae, posterior margin concave, ventral margin extended to a rounded point. Pronotal carinae following posterior contour of eyes, turning posterior just above ventral margin of eye, terminating at ventro-lateral margin of prothorax (Figs. 3A, C). Mesonotum tricarinate, lateral carinae subparallel, curved laterally to reach posterior margin (Fig. 3B). Forewings transparent, with diffuse dark markings along trailing margin near apex of clavus and CuA (cell C5’, Fig. 4), with setal pits present along main veins. Claval apex near wing midlength, about parallel to ScP reaching wing margin; fusion of PCu and A1 about in basal third of wing, with PCu+A1 reaching wing margin before claval apex (i.e., clavus closed). Fork of CuA proximad of RA+ScP from RP fork, with both of these proximad of claval apex (Fig. 4); forewing branching pattern: RA 1-branched, RP 2-branched, MP 5-branched, CuA 2-branched with CuA connected with CuP by icu crossvein (Fig. 4).

Terminalia. Pygofer in lateral view broad, irregular in outline but approximately quadrate with ventral anterior process and large triangular median processes on lateral portion of pygofer opening (Fig. 5A). In ventral view, medioventral lobe elongate and ovoid, approximately 1.5x long as wide (Fig. 5B). Gonostyli in lateral view distally expanded and angled dorsad with rounded apex, strongly sinuate on dorsal margin with strong basal curve, resulting in constriction of gonostyli, appearing spear-like (Fig. 5A); in ventral view, appearing clubbed, inner margin strongly sinuate, outer margin strongly curved distad (Fig. 5B). Aedeagus asymmetrical and complex (Figs 6 & 7); aedeagal shaft with large, cupped process (near midlength) on right lateral margin (A1), curving anterio-dorsad (Figs 6 A-C & 7A-C), large, bulbous projection, angled dorsad, on left lateral side of shaft (A2) (Fig. 6 & 7); two elongate retrorse processes arising at dorsum of shaft apex, both strongly sinuate, angled anterio-dorsad, one (A3) more robust and angled to right lateral side, second (A4) slender angled to left lateral side (Figs. 6 & 7); flagellum with two large processes, one (F1) on right lateral side, nearly reaching A1, smaller process (F2) on left lateral side, surrounding A4. Anal tube in lateral view elongate, dorsal and ventral margins subparallel in basal 2/3, ventral margin projected in distal 1/3 (giving ventral margin a concave appearance), to elongate acute apex; ventral surface with large subapical median ventral process, irregularly triangular, angled ventrad (Fig. 5A); in dorsal view, lateral margins slightly sinuate, extending half beyond apex of pygofer lateral processes (Fig. 6C); anal column (paraproct, anal style) elongate, much longer than remainder of 11 th segment.

Plant associations. Coconut palm (Cocos nucifera), Arecaceae.

Distribution. Costa Rica (Alajuela)

Etymology. The specific name given is an amalgamation of the name of the nearest town, Pocosol and Spanish word for coconut—“coco”, the host of the new taxon, resulting in pocococo.

Material examined. Holotype male “ Costa Rica, Alajuela / Santa Rosa de Pocosol / 16.VI.2019 / Sweeping Cocos nucifera / Coll.: B.W.Bahder // Holotype / Haplaxius pocococo /” (FLREC); Paratypes, Santa Rosa de Pocosol [16.VI.2019] (1 male, 1 female — FSCA, 1female — FLREC).

Sequence Data. For Haplaxius pocococo sp. n. a 677 bp product was obtained for the COI gene (GenBank Accession No. MW 086873). On average H. pocococo sp. n. differed by 10.2% (SE±0.01) from other species of Haplaxius (with H. dougwalshi being most similar, 9.2% different) (Table 4) and the average variability among species of Haplaxius for COI was 13.49% (SE±0.01). Furthermore, Haplaxius pocococo sp. n. differed from Nymphomyndus, Oecleus, and Myxia by 16.9%, 16.8% and approximately 19.0%, respectively (Table 4). While Haplaxius pocococo sp. n. resolved within Haplaxius in the phylogenetic analysis for COI, there in general was weak support for Haplaxius (Fig. 8A). However, there was strong support (90) for Haplaxius pocococo sp. n. resolving next to H. dougwalshi (Fig. 8A).

A 1,396 bp product was obtained for the 18S gene (GenBank Accession No. MW 086509). On average Haplaxius pocococo sp. n. differed by an average of 0.48% (SE±0.02) from other species of Haplaxius (with H. dougwalshi being most similar, 0.1% different) (Table 5) and the average variability among species of Haplaxius for 18S was 0.6% (SE±0.00). Furthermore, Haplaxius pocococo sp. n. differed from Nymphomyndus, Oecleus, and Myxia by 2.3%, 1.6%, and approximately 2.4% respectively (Table 5). Based on the phylogenetic analysis for 18S, Haplaxius pocococo sp. n. resolved within the Haplaxius clade adjacent to H. dougwalshi with moderate support (87). However, there was strong support for Haplaxius (97) based solely on 18S (Fig. 8B). The consensus tree utilizing both COI and 18S data also showed Haplaxius pocococo sp. n. within Haplaxius with strong support in its placement adjacent to H. dougwalshi (Fig. 8C).

Remarks. Based on morphological characters and available molecular data Haplaxius pocococo sp. n. is placed in Haplaxius as currently comprised. Haplaxius dougwalshi is closely related to Haplaxius pocococo sp. n. based on both morphology (see below) and molecular data that suggest that Haplaxius pocococo sp. n. and H. dougwalshi form a clade within available Haplaxius species. There is also less variability in the sequence data between these two species compared to other Haplaxius for both loci. Furthermore, both species have a large, curved process near the midlength of the aedeagal shaft that is not present in the other species in this study. While other species of Haplaxius possess a process at the mid-point (viz. H. flocki (Kramer), H. viridis (Ball), H. ovatus (Ball) and H. simplicatus Caldwell), the strongly curved process is similar between H. dougwalshi and Haplaxius pocococo sp. n., and they have similar facial color patterns, lateral processes of the pygofer opening, and comparable gonostyli and medioventral process of the pygofer.