Bothriocera harthi Bahder & Bartlett, 2023

Bahder, Brian W., Myrie, Wayne, Helmick, Ericka E. & Bartlett, Charles R., 2023, A new species of planthopper in the genus Bothriocera (Hemiptera: Auchenorrhyncha: Cixiidae) from coconut palm (Cocos nucifera) in Jamaica, Zootaxa 5375 (1), pp. 111-127 : 115-124

Bothriocera harthi Bahder & Bartlett sp. n.

( Figures 2–6 )

Type Locality. Black Hill , Portland Parish, Jamaica.

Diagnosis. Medium-sized (for the genus) with strongly patterned forewings bearing a fuscous transverse band from stigma to apex of clavus and second highly irregular band along nodal line. Medioventral lobe of pygofer slightly wider than tall, rounded with slight point at apex. Phallus without processes along shaft, endosoma elongate, helically curved in sinistral direction bearing elongated basal projection (with 2 elongate subapical teeth) on left side and broad ‘fin-like’ projection on right side. Anal tube broadened subapically in left lateral view.

Description. Color. General body color brown ( Fig. 2 ), anterior portion of head darker, posterior region and genae paler, intercarinal regions of mesonotum slightly paler than lateral region, ventrally more pallid to testaceous on legs; forewings ( Fig. 4 ) with veins dark, cells translucent, bearing strong fuscous pattern: small dark patch around apex of Sc, broad, irregular transverse fuscous band from stigma to claval apex, very irregular transverse band at nodal level including apices of RA, RP and CA veins, small patch in cell C3aa.

Structure. Body length male (n = 2): 5.21 mm with wings; 3.28 mm without wings ( Table 3 ), female 5.25 mm with wings, 3.31 mm without.

Head. In dorsal view ( Fig. 3A ), vertex broadly pentagonal, approximately 2X wide as long at midpoint, subapical transverse carinaat fastigium distinct, jointing medially in broadly obtuse angle, connecting to median carina of frons, anterior margin concave (except projected at median carina), posterior margin of head broadly concave (nearly truncate medially). In frontal view ( Fig. 3B ) frons broadly and irregularly quadrate, laterally expanded at antennae (widest at midpoint of eye), median carina becoming obsolete ventrad; ventral margin of frons with broadly Ushaped ventral ridge. Median ocellus prominent. Frontoclypeal suture nearly transverse (slightly convex). In lateral view ( Fig. 3C ), head projected in front of eyes, margins irregularly sinuate, vertex declinate to level of antennae, lateral margins of frons forming reverse C-shaped concavity ahead of antennae, antennae subtended by genal carina from ventroanterior margin of eye to ventral margin of cup formed by lateral margins of frons. Clypeus, from anterior view triangular with incomplete median carina. Antennae short, scape very short and obscure, pedicle emarginate, appearing horseshoe-shaped in lateral view (bearing many sensory plaques) with central bristle-like flagellum with a bulbous base. Lateral ocelli prominent above antennae and anterior to dorsal margin of compound eyes. Compound eyes dorsoventrally elongate, anterior margin concave, posterior margin truncate.

Thorax. Pronotum very short medially, mostly hidden beneath posterior margin of head ( Fig. 3A ), deeply concave on posterior margin; paradiscal region foliate in front of tegulae, lateral margin, in lateral view broad, apex squared off well below lower level of compound eyes. Mesonotum elongated, much longer than wide (about 2.5x length of head at midline; Fig. 3A ), width approximately 2/3 length at midpoint, tricarinate with lateral carinae evident (subparallel, appearing to reach hind margin), median carina weak, in lateral view mesoscutum convex, inclected slightly upward at scutellum. Forewing relatively short and broadly spatulate ( Fig. 4 ), stigma distinct, claval apex before wing midlength, apex rounded, weakly projected near apex of RP 4; crossveins r-m and m-cu in proximal half; branching pattern; RA 2-branched, RP 3-branched, MP 5-branched, CuA 2-branched.

Terminalia. Pygofer in lateral view ( Fig. 5A ) irregularly triangular, shortest dorsally, expanded to midlength, narrowed to ventral margin (caudal margin appearing broadly rounded), dorsal margin rounded, posterior margin broadly rounded, anterior margin irregularly sinuate. In ventral view ( Fig. 5B ) medioventral process present, rounded, just wider than long bearing slight point at apex. Gonostyli in lateral view ( Fig. 5A ) narrowest basally, angled dorsad near midlength, then expanding to rounded apex, strongly concave along ventral margin in distal half; in ventral view ( Fig. 5B ), narrowest proximally, expanding distad, medial concavity near midlength point on inner lateral margin. Aedeagal shaft simple (without lateral or apical processes), cylindrical ( Figs. 6 & 7 ), approximately straight; endosoma elongate (longer than aedeagal shaft) and helically curved in sinistral direction distally narrowed to blunt, membranous apex; two primary processes arising subapically, the first (F1) arising on left lateral margin near endosoma base, comprised primarily of a process sclerotized along leading margin (otherwise membranous), curving ventrad on right lateral side with two sclerotized elongate teeth (F1a and F1b) arising subapically, F1a slightly longer and more slender than F1b; second process (F2) broadly falcate (“fin-like”), weakly sclerotized, with irregular serrulations along trailing margin; distal region of endosome bearing slender, elongated sclerotized internal rod (F3) extending nearly to apex. Anal segment in lateral view ( Fig. 5A ) elongated, distally downcurved; narrow basally, dorsal, and ventral margins subparallel, curved ventrad at 1/3 length, constricting slightly, then expanding near apex, generally rounded with slight point at apex; in dorsal view ( Fig. 5C ), broad, lateral margins sinuate, apex irregularly sinuate and slightly asymmetrical; paraproct conical, short and stout.

Plant associations. Coconut palm ( Cocos nucifera L. ).

Distribution. Black Hill, Saint Mary Parish, Jamaica.

Etymology. The specific name is given in reference to the field site where the holotype was collected (Hart Hill) that was amalgamated as “harthi”. The specific name is intended to be indeclinable.

Material examined. Holotype male “♂” ( FLREC) ; Paratypes 1 male, 2 females, same data as holotype ( FSCA).

Sequence Data. For Bothriocera harthi sp. n., a 642 bp product for COI, a 1,298 bp product for 18S, a 344 bp product for H3, and a 753 bp product for 28S were generated. In addition, products for COI, 18S, H3, and 28S were also generated for B. drakei ( Fig. 8 ), B. maculata ( Fig. 9 ), B. transversa ( Fig. 10 ), B. datuna ( Fig. 11 ), and B. basalis ( Fig. 12 ) with corresponding GenBank accession numbers presented in Table 2 . Based on the phylogenies, there is strong bootstrap support for all loci (89, 100, 95, and 100 for COI, 18S, H3, and 28S respectively) ( Fig. 13 ) and strong bootstrap support in the consensus tree (100) for Bothriocera being monophyletic ( Fig. 14 ). Furthermore, Bothriocera harthi sp. n. resolves within Bothriocera with strong bootstrap for COI (100, adjacent to B. drakei ),18S (86, adjacent to B. datuna , being 100% identical), H3 ((85, adjacent to B. datuna ). Finally, while the consensus tree generated shows strong bootstrap support (100) for the monophyly of Bothriocera (with Bothriocera harthi sp. n. resolving within the genus), the bootstrap support for relationshipsamong species available and loci analyzed was low (<52) ( Fig. 14 ).

TABLE 4. Pairwise comparison showing estimates of evolutionary divergence between sequences based on the COI gene for Bothriocera harthi sp. n. demonstrating intrageneric (orange) and intergeneric (blue) variability; the number of base differences per site from between sequences are shown. Standard error estimate(s) are shown above the diagonal and were obtained by a bootstrap procedure (1000 replicates).

TABLE 5. Pairwise comparison showing estimates of evolutionary divergence between sequences based on the 18S gene for Bothriocera harthi sp. n. demonstrating intrageneric (orange) and intergeneric (blue) variability; the number of base differences per site from between sequences are shown. Standard error estimate(s) are shown above the diagonal and were obtained by a bootstrap procedure (1000 replicates).

TABLE 6. Pairwise comparison showing estimates of evolutionary divergence between sequences based on the H3 gene for Bothriocera harthi sp. n. demonstrating intrageneric (orange) and intergeneric (blue) variability; the number of base differences per site from between sequences are shown. Standard error estimate(s) are shown above the diagonal and were obtained by a bootstrap procedure (1000 replicates).

Based on the pairwise comparison for COI nucleotide sequences, species in Bothriocera differed from each other by an average of 16.9% ( SE ±0.9) whereas variability among genera, on average, was 23.9% ( SE ±0.9). Bothriocera harthi sp. n. differed on average by 17.2% ( SE ±2.8) from other species within Bothriocera , and differed from genera in other higher taxa on average by 26.5% ( SE ±1.3) ( Table 4).

Based on the multiple pairwise comparisons for the loci analyzed, the highest levels of variance were observed for COI, followed by H3 with variances observed among 28S and 18S being significantly lower (with 28S being being slightly more variable than 18S). Regardless, Bothriocera harthi sp. n. varied from other congeners by levels that were within the observed range of intrageneric variability for all loci analyzed ( Table 7 ).

Remarks. Bothriocera harthi sp. n. is easily placed in the genus Bothriocera by head morphology (antennae in a cavity in front of compound eyes), the weakly overlapped wings (relative to Bothrioceretta ), and the short medioventral lobe of the pygofer. This placement is also supported by molecular data (COI, 18S, H3, and 28S genes). In general, Bothriocera harthi sp. n. superficially resembles the other Bothriocera in the pattern of the forewings. The forewing pattern may (as Caldwell 1943 asserted) be useful for species diagnosis, but better documentation of variation in color pattern of the known species is needed before relying on this feature.

Bothriocera harthi sp. n. appears most like Bothriocera datuna Kramer (see Fig. 11 and Kramer 1983, figs. 39–41). Aside from geography, Bothriocera harthi sp. n. differs most readily from B. datuna in that the latter species has less extensive wing markings the medioventral lobe of the pygofer bears an apical projection; also, and differences in the shape of the gonostyli.

Other material examined.

Bothriocera basalis . Costa Rica, La Tarde Ecolodge, Osa Peninsula , 16.VI.2021 (3 males, 2 females).

Bothiocera datuna . Coral Springs , FL, U.S.A., Tall Cypress Natural Area , 3.IV.2022 (1 male).

Bothriocera drakei . Milford , DE, U.S.A., Woods Haven , 6.VII.2021 (1 male).

Bothriocera maculata . Coral Gables , FL, U.S.A., Montgomery Botanical Center , 16.V.2022 (2 males).

Bothriocera transversa . Coral Gables , FL, U.S.A., Montgomery Botanical Center , 16.V.2022 (5 males, 2 females).

Figures

FIGURE 2. Adult male habitus of Bothriocera harthi sp. n.: A) dorsal view and B) lateral view; scale bar = 1 mm.

FIGURE 3. Adult male Bothriocera harthi sp. n.: A) head, pronotum and mesonotum in dorsal view, B) head in frontal view and C) head, pronotum and mesonotum in lateral view; scale bar = 1 mm.

FIGURE 4. Figure 4. Adult male Bothriocera harthi sp. n. forewing; black text = vein, italic text = crossvein, green text = cell.

FIGURE 5. Adult male Bothriocera harthi sp. n. terminalia; (A) lateral view, (B) ventral view and (C) dorsal view.

FIGURE 6. Bothriocera harthi sp. n. aedeagus; (A) right lateral view, (B) left lateral view, (C) dorsal view and (D) ventral view.

FIGURE 7. Bothriocera harthi sp. n. aedeagus line art; (A) right lateral view, (B) left lateral view, (C) dorsal view and (D) ventral view.

FIGURE 8. Adult male Bothriocera drakei; A) dorsal habitus, B) terminalia left lateral view, C) terminalia ventral view, D) aedeagus dorsal view, E) aedeagus ventral view, F) aedeagus right lateral view and G) aedeagus left lateral view, scale = 1 mm.

FIGURE 9. Adult male Bothriocera maculata; A) dorsal habitus, B) terminalia left lateral view, C) terminalia ventral view, D) aedeagus dorsal view, E) aedeagus ventral view, F) aedeagus right lateral view and G) aedeagus left lateral view, scale = 1 mm.

FIGURE 10. Adult male Bothriocera transversa; A) dorsal habitus, B) terminalia left lateral view, C) terminalia ventral view, D) aedeagus dorsal view, E) aedeagus ventral view, F) aedeagus right lateral view and G) aedeagus left lateral view, scale = 1 mm.

FIGURE 11. Adult male Bothriocera datuna; A) dorsal habitus, B) terminalia left lateral view, C) terminalia ventral view, D) aedeagus ventral view, E) aedeagus left lateral view, F) aedeagus dorsal view and G) aedeagus right lateral view, scale = 1 mm.

FIGURE 12. Adult male Bothriocera basalis; A) dorsal habitus, B) terminalia left lateral view, C) terminalia ventral view, D) aedeagus dorsal view, E) aedeagus ventral view, F) aedeagus left lateral view and G) aedeagus right lateral view, scale = 1 mm.

FIGURE 13. Maximum likelihood phylogenetic tree based on 1,000 replicates; (A) COI gene, (B) 18S rRNA gene, (C) H3 gene, and (D) 28S gene; scale bar = percent nucleotide difference.

FIGURE 14. Maximum likelihood phylogenetic consensus tree based on 1,000 replicates for concatenated COI, 18S, 28S, and H3 sequence data.

Tables

TABLE 3. Biometric data for Bothriocera harthi sp. n.

Character	Male (n =2)		Female (n =2)
	Range	Average ± SE	Range	Average ± SE
Body length, with wings	5.21–5.21	5.21±0.0	5.25–5.25	5.25±0.0
Body length, no wings	3.28–3.28	3.28±0.0	3.31–3.32	3.31±0.1
Forewing length	4.30–4.30	4.30±0.0	4.61–4.61	4.61±0.0
Vertex length	0.23–0.23	0.23±0.0	0.25–0.25	0.25±0.0
Vertex width, basal margin	0.66–0.66	0.66±0.0	0.67–0.67	0.67±0.0
Vertex width, distal margin	0.52–0.52	0.52±0.0	0.52–0.52	0.52±0.0
Pronotum length, midline	0.05–0.05	0.05±0.0	0.05–0.05	0.05±0.0
Mesonotum length, midline	1.09–1.09	1.09±0.0	1.09–1.09	1.09±0.0
Mesonotum width	1.06–1.06	1.06±0.0	1.07–1.07	1.07±0.0
Frons width, dorsal margin	0.56–0.56	0.56±0.0	0.56–0.56	0.56±0.0
Frons width, clypeal suture	0.49–0.49	0.49±0.0	0.50–0.50	0.50±0.0
Frons width, widest	0.72–0.72	0.72±0.0	0.72–0.72	0.72±0.0
Frons width, narrowest	0.49–0.49	0.49±0.0	0.50–0.50	0.50±0.0
Frons length, midline	0.61–0.61	0.61±0.0	0.61–0.61	0.61±0.0
Clypeus length	0.43–0.43	0.43±0.0	0.43–0.43	0.43±0.0

TABLE 4. Pairwise comparison showing estimates of evolutionary divergence between sequences based on the COI gene for Bothriocera harthi sp. n. demonstrating intrageneric (orange) and intergeneric (blue) variability; the number of base differences per site from between sequences are shown. Standard error estimate(s) are shown above the diagonal and were obtained by a bootstrap procedure (1000 replicates). 1 2 3 4 5 6 7 8 9 10 11 1 Bothriocera harthi sp. n. 0.020 0.022 0.010 0.020 0.020 0.025 0.027 0.024 0.025 0.028 2 Bothriocera datuna 0.207 0.021 0.017 0.019 0.021 0.026 0.025 0.024 0.023 0.028 3 Bothriocera basalis 0.212 0.189 0.019 0.019 0.020 0.025 0.023 0.025 0.020 0.027 4 Bothriocera drakei 0.061 0.156 0.176 0.016 0.017 0.023 0.024 0.023 0.023 0.027 5 Bothriocera maculata 0.186 0.166 0.179 0.136 0.016 0.021 0.025 0.024 0.023 0.027 6 Bothriocera transversa 0.191 0.198 0.190 0.150 0.140 0.024 0.025 0.024 0.022 0.025 7 Haplaxius crudus 0.244 0.256 0.254 0.223 0.195 0.232 0.023 0.022 0.022 0.019 8 Myxia belinda 0.298 0.274 0.245 0.265 0.243 0.257 0.251 0.026 0.023 0.023 9 Nymphocixia unipunctata 0.252 0.247 0.266 0.246 0.235 0.250 0.202 0.279 0.023 0.023 10 Oecleus mackaspringi 0.234 0.250 0.188 0.209 0.216 0.211 0.212 0.233 0.226 0.024 11 Melanoliarus maidis 0.295 0.293 0.278 0.283 0.266 0.243 0.158 0.232 0.224 0.243

		1	2	3	4	5	6	7	8	9	10	11
1	Bothriocera harthi sp. n.		0.020	0.022	0.010	0.020	0.020	0.025	0.027	0.024	0.025	0.028
2	Bothriocera datuna	0.207		0.021	0.017	0.019	0.021	0.026	0.025	0.024	0.023	0.028
3	Bothriocera basalis	0.212	0.189		0.019	0.019	0.020	0.025	0.023	0.025	0.020	0.027
4	Bothriocera drakei	0.061	0.156	0.176		0.016	0.017	0.023	0.024	0.023	0.023	0.027
5	Bothriocera maculata	0.186	0.166	0.179	0.136		0.016	0.021	0.025	0.024	0.023	0.027
6	Bothriocera transversa	0.191	0.198	0.190	0.150	0.140		0.024	0.025	0.024	0.022	0.025
7	Haplaxius crudus	0.244	0.256	0.254	0.223	0.195	0.232		0.023	0.022	0.022	0.019
8	Myxia belinda	0.298	0.274	0.245	0.265	0.243	0.257	0.251		0.026	0.023	0.023
9	Nymphocixia unipunctata	0.252	0.247	0.266	0.246	0.235	0.250	0.202	0.279		0.023	0.023
10	Oecleus mackaspringi	0.234	0.250	0.188	0.209	0.216	0.211	0.212	0.233	0.226		0.024
11	Melanoliarus maidis	0.295	0.293	0.278	0.283	0.266	0.243	0.158	0.232	0.224	0.243

TABLE 7. Pairwise comparison showing estimates of evolutionary divergence between sequences based on the 28S gene for Bothriocera harthi sp. n. demonstrating intrageneric (orange) and intergeneric (blue) variability; the number of base differences per site from between sequences are shown. Standard error estimate(s) are shown above the diagonal and were obtained by a bootstrap procedure (1000 replicates).

		1	2	3	4	5	6	7	8	9	10	11
1	Bothriocera harthi sp. n.		0.003	0.002	0.002	0.013	0.002	0.026	0.025	0.031	0.029	0.029
2	Bothriocera basalis	0.005		0.003	0.002	0.013	0.003	0.025	0.026	0.030	0.029	0.028
3	Bothriocera drakei	0.001	0.004		0.001	0.013	0.002	0.026	0.025	0.030	0.029	0.029
4	Bothriocera transversa	0.002	0.003	0.001		0.013	0.002	0.026	0.025	0.030	0.029	0.028
5	Bothriocera maculata	0.028	0.030	0.029	0.030		0.013	0.037	0.037	0.042	0.040	0.040
6	Bothriocera datuna	0.001	0.005	0.001	0.002	0.029		0.026	0.026	0.030	0.029	0.029
7	Oecleus mackaspringi	0.059	0.058	0.059	0.058	0.079	0.059		0.017	0.016	0.014	0.018
8	Myxia belinda	0.058	0.058	0.057	0.057	0.079	0.058	0.040		0.023	0.019	0.020
9	Nymphocixia unipunctata	0.068	0.068	0.068	0.068	0.089	0.067	0.036	0.050		0.015	0.024
10	Haplaxius crudus	0.065	0.065	0.065	0.064	0.084	0.064	0.032	0.043	0.034		0.021
11	Melanoliarus maidis	0.065	0.063	0.064	0.063	0.085	0.065	0.041	0.046	0.056	0.048