Description of Nesophrosyne Melemele Sp. N., an Endemic Hawaiian Leafhopper (Hemiptera: Cicadellidae: Delto- Cephalinae: Opsiini

NProceediesophrosyngsn eof melemele the Hawaiia sp. nn., E anntomological endemic Hawaiian Society leaf (2017)hopper 49:47–50 47

Description of Nesophrosyne melemele sp. n., an Endemic Hawaiian Leafhopper (Hemiptera: Cicadellidae: Delto- cephalinae: Opsiini) Associated with Myoporum sandwicense (Scrophulariaceae)

Michael D. Denis1 and Gordon M. Bennett1,2 1Department of Plant and Environmental Protection Sciences, University of Hawaii at Manoa; 2Corresponding author, [email protected]

Abstract. A new species of the endemic Hawaiian leafhopper genus, Nesophrosyne (Hemiptera: Cicadellidae: Deltocephalinae: Opsiini), is described: Nesophrosyne melemele sp. n. Morphological and molecular characters were used to delineate this species. Nesophrosyne melemele is endemic to Kauai and strictly associated with the endemic plant species, Myoporum sandwicense (Scrophulariaceae).

The endemic Hawaiian leafhopper ge- Kirkaldy 1907, 1910). Recent molecular nus, Nesophrosyne (Cicadellidae: Delto- systematic work provided a clearer delin- cephalinae: Opsiini), is descended from a eation of morphologically and genetically single colonization event from the western distinct Nesophrosyne species (Bennett rim region of the Pacific Ocean (Bennett and O’Grady 2012). Here we describe one and O’Grady 2012). Nesophrosyne likely of these new species from Kauai that relies shares a common ancestor with the Oro- exclusively on the endemic Hawaiian host- sius genus (also placed in Opsiini), which plant, Myoporum sandwicense (Scrophu- is widely distributed throughout Asia, lariaceae). This species was previously Australia, and African continents (Ghauri assigned a morphotaxon designation with 1966, Fletcher et al. 2017). The Hawai- the provisional identifier, “N. sp. 126”. ian diversity is derived from an adaptive Phylogenetic analysis further revealed that radiation of species specializing broadly this new Kauai species is sister to the other on the endemic Hawaiian flora (Zimmer- Nesophrosyne that specialize on the same man 1948, Bennett and O’Grady 2013). host-plant across the archipelago (Bennett Recently, Bennett and O’Grady (2011, and O’Grady 2012). 2012) conducted a taxonomic review and molecular survey of the Nesophrosyne, de- Materials and Methods scribing eight new species representing the Insect specimens were field collected first newly described species in the genus under native invertebrate collection and in over sixty years. This work revealed that locality access permits furnished by the the genus comprises well over 200 species; State of Hawaii, Department of Land however, only about a third of this diver- and Natural Resources. Specimens were sity is currently described. Furthermore, obtained by sweep netting host-plants many existing species names are question- and preserved directly in 95% EtOH. able and difficult to identify, as historical Specimens were measured and described workers based some descriptions on im- using a Zeiss Discovery-8 dissection mi- matures, females, or only a few individuals croscope and photographed with Olympus (reviewed by Zimmerman 1948, Bennett SZX10 and Nikon D7100. Stacked images and O’Grady 2011; also see Osborn 1935, were merged with Helicon software (Heli- 48 Denis and Bennett consoft). For description of the genitalia, while N. giffardi interrupta is darker in specimens had whole abdomens dissected. color with three articulated pale claval Genitalia capsules were cleared by incuba- markings (see Zimmerman 1948). Oth- tion in 10% potassium hydroxide at room erwise, the remainder of the forewing is temperature overnight. After description distinctly dark, verging on black, in both and measurements, cleared genitalia were subspecies. As has been found for all Ne- stored in glycerol within plastic vials sophrosyne species investigated to date, pinned and deposited with specimens. both are single-island endemics restricted Abdomens were used for DNA extrac- to Hawaii island (Bennett and O’Grady tion (published in Bennett and O’Grady 2012). Compared with material examined 2012: Accession Numbers: JX417555 here for the Kauai taxon, this new spe- [Cytochrome Oxidase II], JX433111 cies is consistently and almost entirely [Cytochrome Oxidase I], JX433284 [16S yellow in coloration. Genetic data further rRNA], JX433489 [Nuclear Histone 3], demonstrates that it is indeed a distinct and JX433698 [Nuclear Wingless]). Spe- morphotaxon restricted to Kauai; it was cies description followed that established previously given the provisional name by Bennett and O’Grady (2011), which “N. sp. 126” (see supplemental material used the morphological nomenclature out- in Bennett and O’Grady 2012). lined in Dietrich (2005) and Oman (1949). All examined material was subsequently Taxonomy deposited in the University of Hawaii Nesophrosyne melemele Denis & Ben- Insect Museum, Honolulu, Hawaii, USA. nett Diagnosis. Length: male = 2.88 mm, Results and Discussion female = 3.58 mm. Yellow colored species. Delimitation of a new species associ- Head and vertex rounded. Frontoclypeus ated with Myoporum sandwicense. The with a dark narrow (stylized) heart-shaped designation of a new species status was marking. Yellow saddle mark present. Ae- established based on previous taxonomic deagus with pronounced distal processes, and genetic work that reviewed the de- hooking laterally to almost 1/3 length of scribed material for the Nesophrosyne aedeagal arms. genus (see Bennett and O’Grady 2011, Description. Dorsum (Figure 1A): Zimmerman 1948). The species described Yellow colored species. Head rounded here does not match existing descriptions with vertex weakly produced. Pronotum and was previously determined to be new predominately yellow with posterior edge with a combination of morphological and light brown, extending 1/2 pronotal length molecular characters. Only a single sub- and tapering towards lateral edges. Scu- species, N. giffardi interrupta Osborn, tellum yellow; centrally split by diffuse has been described as associated with dark vertical triangle that is variable in the host-plant, M. sandwicense (reviewed width and length; further subdivided by by Zimmerman 1948). Although no host thin bowed dark horizontal line curving information was originally given for N. posteriorly. Forewing cells hyaline to giffardi giffardi Kirkaldy, it is also re- infuscate, darkest forms with cells dark stricted to M. sandwicense (Bennett and brown colored; veins darkly colored O’Grady 2012). Nesophrosyne giffardi throughout; outer anteapical cell present. giffardi is morphologically and geneti- Clavus with diffuse yellow saddle mark cally distinct from N. giffardi interrupta. with anterolateral margins fading to dark The former has a completely pale clavus, brown; yellow color broadening towards Nesophrosyne melemele sp. n., an endemic Hawaiian leafhopper 49

Figure 1. Nesophrosyne melemele sp. n. endemic to Kauai: (A) dorsal and ventral habitus, (B) male pygofer lateral view, (C) connective dorsal view, (D) style dorsal view, (E) aedeagus posterior view, and (F) aedeagus lateral view.

anterior claval edges. posterior appendages with microsetae. Venter (Figure 1A): Face with clypellus Connective close in length to styles with yellow. Gena and lorum yellow. Antennal posterior edge notched; anterior arms sockets yellow. Frontoclypeus broadly bowed and widely splayed with ends hook- dark or brownish; vertex and edges pale, ing mesad. tapering towards posterior base creating Distribution: Hawaiian Islands, Kauai, a stylized heart-shaped mark on face. Polihale Beach. Elevation: 3 meters Thoracic plates dark. Legs pale. Ventral Measurements: Body: male (n = 7) = abdominal segments pale with infused 2.89 mm (2.7–3.0 mm); female (n = 5) = dark bands along anterior edges. 3.58 mm (3.2–4.0 mm). Genitalia: (n = 4): Genitalia (Figure 1B–F): Pygofer pygofer = 0.45 mm (0.44–0.45 mm); style bluntly rounded with posterior apex rising = 0.34 mm (0.33–0.35 mm); connective to near dorsal edge; dorsal edge flat; ven- = 0.27 mm (0.26–0.28 mm); aedeagus tral lobe reduced with ventral edge angled lateral length = 0.28mm (0.27–0.30 mm); towards apex; 7 macro setae irregularly aedeagus posterior height = 0.26 mm distributed. Subgenital plate with dense (0.24–0.28 mm). setae distributed along its lateral edges. Type material: Holotype: 1 male, Aedeagus with paired aedeagal arms Hawaiian Islands, Kauai, Polihale State widely splayed, rising conspicuously Park. GPS: 22.29222222, 159.95972222. higher than central apodeme; gonopore Elevation: 3 m. Collected on 5-Jan-2010. preapical; apical processes elongate, ex- Host plant: Myoporum sandwicense. Coll. tending just above gonopore and hooking G.M. Bennett. Deposited at the University to approximately 1/3 length of aedeagal of Hawaii Insect Museum, Honolulu, Ha- arms. Style with apophysis curved, widen- waii, USA. ing at base; preapical lobe with flat lateral Paratype material: 7 males and 5 fe- and posterior edges, appearing angular; males with the same locality information articulating arm short and triangular; and collection date as the holotype mate- 50 Denis and Bennett rial (see above). Additional material was logeny, origin, and species diversity of also deposited in the University of Hawaii native Hawaiian leafhoppers (Cicadellidae: Insect Museum, Honolulu, Hawaii, USA. Nesophrosyne). Mol. Phylogenet. Evol. 65: Etymology: Melemele is the Hawaiian 705–717. Bennett, G.M., and P.M. O’Grady. 2013. word for yellow. It was chosen to describe Historical biogeography and ecological the predominately yellow body coloration opportunity in the adaptive radiation of observed in male and females of this spe- native Hawaiian leafhoppers (Cicadellidae: cies. Nesophrosyne). J. Biogeog. 40: 1512–1523. Notes: Host plant: Myoporum sand- Dietrich, C.H. 2005. Keys to the families of wicense. Previous work referred to this Cicadomorpha and subfamilies and tribes species under the provisional designation of Cicadellidae (Hemiptera: Auchenorrhyn- as “N. sp. 126” (Bennett and O’Grady cha). Fla. Entomol. 88: 502–514. Fletcher, M., H. Löcker, A. Mitchell, and D. 2012). Gopurenko. 2017. A revision of the genus Orosius Distant (Hemiptera: Cicadellidae) Acknowledgments based on male genitalia and DNA barcod- We thank two anonymous reviewers ing. Austral. Entomol. 56: 198–217. for their comments that greatly improved Ghauri, M.S.K. 1966. Revision of the genus this work. We thank Meng Mao, Kristen Orosius Distant (Homoptera: Cicadelloi- Poff, and Jared Bernard for their help dea). Brit. Mus. Occas. Pap. 18: 231–252. with edits on this manuscript. We also Kirkaldy, G.W. 1907. Biological notes on the thank Michelle Au and Xiushuai Yang for Hemiptera of the Hawaiian Isles No. 1. Proc. Hawaiian. Entomol. Soc. 1: 135–161. their technical support. Finally, we thank Kirkaldy, G.W. 1910. Supplement to Hemip- Patrick O’Grady and Dan Polhemus for tera. pp. 531–599. In D. Sharp (ed.), Fauna early advising and help on developing this Hawaiiensis. Cambridge: Cambridge Uni- project. versity Press. Oman, P.W. 1949. The nearctic leafhoppers Literature Cited (Homoptera: Cicadellidae): a generic clas- Bennett, G.M., and P.M. O’Grady. 2011. sification and checklist. Mem. Entomol. Soc. Review of the native Hawaiian leafhopper Washington. 1: 1–253. genus Nesophrosyne (Hemiptera: Cicadel- Osborn, H. 1935. Cicadellidae of Hawaii. lidae: Deltocephalinae) with descriptions Bernice. Bish. Mus. B. 134: 1–62 of eight new species associated with Brous- Zimmerman, E.C. 1948. Insects of Hawaii. saisia arguta. Zootaxa. 2805: 1–25. Vol. 4. Honolulu: University of Hawaii Bennett, G.M., and P.M. O’Grady. 2012. Press. Host-plants shape insect diversity: phy-