Haplaxius dougwalshi, Bahder & Barrantes Barrantes & Zumbado Echavarria & Mou & Helmick & Bartlett, 2020

Bahder, Brian W., Barrantes Barrantes, Edwin A., Zumbado Echavarria, Co. A., Mou, De-Fen, Helmick, Ericka E. & Bartlett, Charles R., 2020, A new species of planthopper in the genus Haplaxius (Hemiptera: Auchenorrhyncha: Fulgoroidea: Cixiidae) on palms in Costa Rica and a new country record for Haplaxius skarphion, Zootaxa 4767 (4), pp. 543-552 : 546-551

Haplaxius dougwalshi sp. n.

( Figures 2–7 )

Type locality. La Selva Biological Station, Heredia, Costa Rica

Diagnosis. This species is distinguished by a combination of bright orange coloration, facial color pattern of an orange frons with white arch traversing width of frons resulting in a trapezoidal shape dorsad of frontoclypeal suture, large, incurved processes on lateral margins of the pygofer and aedeagus with curved spine near midlength of shaft.

Description. Color. General body color is uniformly orange with abdominal sternites, middle and hind legs paler orange ( Fig. 2 ). Frons with median white patch merging with lateral white patches, reaching frontoclypeal suture, orange patch dorsad of frontoclypeal suture trapezoidal in shape ( Fig. 3A ). Forewing veins pale orange ( Fig. 2 ). Structure. Body length males (n= 14): 3.16–3.19 mm with wings; 2.53–2.56 mm without wings; females (n =9): 3.19–3.20 mm with wings; 2.56–2.58 mm without wings.

Head. In frontal view, frons narrowed between eyes, expanding at ventral margin, transverse carina present separating frons and vertex ( Fig. 3 ). Fastigium rounded in lateral view ( Fig. 3 ). In dorsal view, frons concave, vertex widest posteriorly, narrowing to almost half width at base ( Fig. 3 ). Vertex length males: 0.33–0.34 mm; females 0.35 mm. Vertex width at hind margin males: 0.21–0.22 mm; females: 0.22–0.23 mm. Vertex width at distal margin males: 0.11–0.12 mm; females: 0.12–0.13 mm. Frons length males: 0.50–0.51 mm; females: 0.51–0.52 mm. Frons dorsal width males: 0.10–0.11 mm; females: 0.11–0.12 mm. Frons width at widest portion males: 0.36–0.37 mm; females: 0.37–0.38 mm. Frons frontoclypeal margin width males: 0.29–0.30 mm; females: 0.31–0.32 mm. Clypeus length males: 0.39–0.40 mm; females: 0.40–0.41 mm.

Thorax. Carinae extending to lateral margin in frontal view ( Fig. 3 ). Mesonotum tricarinate, lateral margins subparallel ( Fig. 3 ). Pronotum length at midline males: 0.05–0.06 mm; females: 0.06–0.07 mm. Mesonotum length at midline males: 0.47–0.48 mm; females: 0.48–0.49 mm. Mesonotum width males: 0.65–0.66 mm; females: 0.66– 0.67 mm. Fore wings transparent, clouded posterior to pseudostigma ( Fig. 4 ), with conspicuous tubercles along main veins ( Fig. 4 ). Apex of clavus just beyond midlength of wing, PCu+A1 reaching wing margin before claval apex. Fork of CuA just basad of RA(+ScP)+RP fork, both just basad of wing midlength; MP forked from ScP+R in basal forth of wing; forewing branching pattern: RA 1-branched, RP 3, MP 5, CuA 4 (or 3 with icu crossvein); icu joining apex of clavus. PCu joining with A1 at midpoint of clavus. Forewing length males: 2.58–2.60 mm; females: 2.60–2.62 mm.

Terminalia. Pygofer in lateral view widest ventrad, narrowing dorsally, posterior margin with large process on median posterior margin ( Fig. 5A ). In ventral view, pygofer opening bearing an elongate rounded lobe, approximately 2x long as wide ( Fig. 5B ); in caudal view, pygofer with lateral processes, medially angled ( Fig. 6). Gonostyli in lateral view angled upwards with acute apex, strongly sinuate on dorsal margin with basal curve strong, resulting in constriction of gonostyli, appearing spoon-like ( Fig. 5A ). In ventral view, appearing clubbed with inner and outer margins of clubbed apex irregularly sinuate ( Fig. 5B ), inner margin strongly sinuate, outer margin weakly sinuate ( Fig. 5B ). Aedeagus asymmetrical and complex ( Fig. 7 A–C); aedeagal shaft with small spine at midlength on right lateral margin, strongly curved (almost 90° angle) ( Fig. 7B ) and larger spine at apex, angled dorsad and slightly distad on left lateral margin, nearly extending to base of aedeagus, approximately 2x length and thickness of small spine. Flagellum (endosoma) with one dorsal spine angled distad toward right lateral margin, terminating approximately at base of small aedeagal spine ( Fig. 7C ) and one larger, ventral spine, arching ventrad, approximately twice length of dorsal flagellar spine, terminating approximately at same point as smaller aedeagal spine ( Fig. 7B ). Anal segment in lateral view with parallel dorsal and ventral margins, approximately 5X long at apex as wide ( Fig. 5B ), apex angled slightly downward, median process angled ventrad approximately 2/3 length distad of pygofer, median process of anal segment about equal size to lateral process of pygofer ( Fig. 5 ).

Plant associations. Pata de gallo palm ( Asterogyne martiana ), Arecaceae .

Distribution. Costa Rica (Heredia, Puerto Viejo de Sarapiqui)

Etymology. The specific name given is an honorarium in reference to the lead authors professor and chair during graduate school (Ph.D.), Dr. Doug Walsh at Washington State University, who’s signature mustache resembles the color pattern observed on the frons of the novel taxon.

Material examined. Holotype male “ Costa Rica, Heredia / La Selva Biological Station / 26.V.2019 / Sweeping Astrogyne martiana / Coll.: B.W.Bahder // Holotype / Haplaxius dougwalshi ” ( FLREC) ; Paratypes, La Selva Biological Research Station [26. V.2019] (13 males, 9 females, FLREC and FSCA).

Other Haplaxius species collected.

Haplaxius skarphion – “ Costa Rica, Limón / Gandoca, nr. Panama / B.W.Bahder; 14. V.2018 / aspirated from coconut palm” (2 males) ( Fig. 8 )

Sequence data. For the COI locus, 683 bp of the 5-prime region of the gene were obtained for Haplaxius dougwalshi sp. n. (GenBank Accession No. MT 080284 ) and was compared to the corresponding region for Haplaxius crudus , Nymphomyndus carribea and Oecleus mackaspringi . There was strong bootstrap support (88) for H. dougwalshi sp. n. resolving next to H. crudus relative to the other two genera of oecleines included ( Fig. 9A ) and based on the pairwise comparison, was 12% different than H. crudus , 16.4% different than N. caribbea , and 17% different from O. mackaspringi ( Table 1). For the 18S locus, 1,358 bp were obtained for H. dougwalshi sp. n. (GenBank Accession No. MT 002395 ) and was compared to the corresponding region for H. crudus , N. caribbea , Myxia belinda , Myndus taffini , and O. mackaspringi . There was strong bootstrap support for H. dougwalshi resolving in the genus Haplaxius (98) relative to other genera of oecleines ( Fig. 9B ). Additionally, H. dougwalshi sp. n. was 0.6% different from H. crudus for the 18S gene whereas it was on average, 3.0% different from the other oecleine genera included, ranging from 1.8% to 5.1% different ( Table 2).

Remarks. The general form of the aedeagus of H. dougwalshi sp. n. is similar to many species illustrated by Kramer (1979). While the presence of a spine on the aedeagus before the apex is not unique to H. dougwalshi sp. n. (shared with H. fulvus (Osborn) , H. pusillus (Van Duzee) , H. viridis (Ball) , H. jamaicae (Kramer) , H. simplicatus Caldwell , and H. vilbastei (Kramer)) , the spine on the shaft in H. dougwalshi sp n. is distinct in being positioned further basad than the other species and strongly curved, whereas the other taxa possess a spine that is either straight or only slightly curved. The lateral processes of the pygofer of H. dougwalshi sp. n. is unique and easily diagnoses it, being much longer and narrower than any other species of Haplaxius . Other taxa that also a possess a similar process include H. glyphis (Kramer) , H. gnophos (Kramer) , H. neopusillus (Kramer) , and H. pusillus (Van Duzee) ; however, in these taxa the base of the process is broad whereas in H. dougwalshi sp. n., the base is strongly narrowed, producing a very well-defined process.

While the features of the genitalia are the most reliable to distinguish H. dougwalshi sp. n., the white patterns observed on the frons also appear distinct, along with the uniform orange coloration. In addition, there is molecular support for H. dougwalshi sp. n. belonging to Haplaxius based on both the COI and 18S loci.

Figures

FIGURE 2. Adult habitus Haplaxius dougwalshi sp. n.; (A) body dorsal view male (B) body lateral male, scale = 1 mm.

FIGURE 3. Adult Haplaxius dougwalshi sp. n.; (A) male head frontal view, (B) male head, pronotum, and mesonotum dorsal view (C) male head, pronotum, and mesonotum lateral view, scale = 1 mm.

FIGURE 4. Forewing venation of Haplaxius dougwalshi sp. n.

FIGURE 5. Male terminalia of Haplaxius dougwalshi sp. n.; (A) lateral view, (B) ventral view, and (C) dorsal view.

FIGURE 8. Adult male of Haplaxius skarphion; (A) body lateral view, (B) frontal view of head and prothorax, (C) ventral view of terminalia, (D) lateral view of terminalia, (E) left lateral view of aedeagus, and (F) right lateral view of aedeagus, scale bar = 1 mm.

FIGURE 9. Maximum Likelihood phylogenetic tree based on COI sequence data (A) and 18S sequence data (B). Branch support was assessed using 1,000 bootstrap replicates.

FIGURE7. Aedeagusof adult male Haplaxiusdougwalshi sp. n.; (A) leftlateral view,(B) rightlateral view, and(C) dorsalview.