Research Article |
Corresponding author: Shusuke Shimamoto ( 1202shusuke@gmail.com ) Academic editor: Jader Oliveira
© 2023 Shusuke Shimamoto, Seidai Nagashima, Hiroshi Nagano, Tadashi Ishikawa.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Shimamoto S, Nagashima S, Nagano H, Ishikawa T (2023) A remarkable new species of the flat bug genus Nesoproxius (Hemiptera, Aradidae), the first Oceanian representative with brachyptery. ZooKeys 1146: 147-163. https://doi.org/10.3897/zookeys.1146.96029
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A new flat bug species, Nesoproxius kishimotoi sp. nov., from the Oceanian region (Ogasawara Islands, Japan) is described. It is the first brachypterous representative in the genus Nesoproxius. The sexual dimorphism, nymph, and habitat are also described for the first time in this genus. A key to the species of Nesoproxius is also provided.
Carventinae, nymph, oceanic island, taxonomy, the Ogasawara (Bonin) Islands
The Ogasawara (Bonin) Islands, in the northernmost part of Micronesia, are among the Pacific islands that make up the Oceanian biogeographic region. These subtropical oceanic islands belonging to Japan are registered as a UNESCO World Natural Heritage site because of abundant endemic species with unique evolutionary patterns (
Many undetermined species have recently been discovered, with some of them having been described as new endemic species (
Nesoproxius Usinger & Matsuda, 1959, a genus within the flat bug subfamily Carventinae, was originally established as a subgenus of Proxius Stål, 1873 and then upgraded to its current rank by
Most of the data were obtained from field surveys conducted by the first author in three of the Ogasawara Islands (Chichijima, Anijima, and Ototojima islands) during 2021 and 2022. These surveys which were part of a biodiversity monitoring program in a series of green anole control projects executed by the Ministry of the Environment in Japan. The remaining analyzed specimens were provided by our collaborators. Dried specimens were used for morphological observations, which were performed using a stereoscopic microscope (Olympus SZX7 and Leica M165C). All measurements were performed using a micrometer eyepiece and provided in millimeters. Illustrations were made using a stereoscopic microscope (Leica M165C) with the aid of a drawing tube (Figs
Photographs of the specimens were taken using a digital camera (Canon EOS 5D Mark IV) with a Canon MP-E 65 mm f/2.8 1–5× macro lens (Figs
Nesoproxius
Usinger & Matsuda, 1959: 113 (as subgenus of Proxius); upgraded to the generic rank by
Nesoproxius was previously diagnosed as a macropterous genus (
Carventus
sp.–
Holotype
: ♂, “Japan, Ogasawara Islands, Ototojima Island, southwest of Ainosawa, 27.1587°N, 142.1894°E, alt. ca 160 m, 11.VII.2021, Shusuke Shimamoto” (
Paratypes
(5 ♂ 12 ♀): Japan, Ogasawara Islands: Chichijima Island: 1 ♀, Renju-dani, 7.III.1999, Toshio Kishimoto (
Nymphs (2 spec.): Japan, Ogasawara Islands: Ototojima Island: 1 spec. (fourth instar), same data as holotype (
This new species is the only brachypterous species in this genus, and it can be distinguished from all other Nesoproxius species by a combination of the following characters: body length approximately 3.0–3.5 mm; incrustation of body surface ocher; head vertex only slightly longitudinally raised; pronotum with only a slightly convex median ridge; scutellum trapezoidal without a median ridge; and abdomen with a relatively smooth margin.
Male (holotype) (Figs
Pronotum 1.9 times as wide as its length on midline, 1.3 times as long as head (excluding neck) on midline; anterior lobe strongly incrusted, with median ridge weakly inflated and slightly projected anteriad, and with four pairs of ovate smooth depressions; anterior margin slightly arched forward beyond collar at lateral one-third; anterolateral angles rounded, not projected beyond collar; lateral margins of anterior lobe convex and sinuate; posterior lobe weakly incrusted; lateral margins of posterior lobe convex anteriorly, then posteriorly concave; posterior margin weakly projected posteriorly. Scutellum trapezoidal, 0.4 times as long as its basal width, widely incrusted and elevated along lateral margins, with lateral margins straight and apex slightly rounded; median ridge thinly incrusted, slightly elevated basally; lateral incrusted fields isosceles triangular. Metanotum slightly visible behind apex of scutellum in dorsal view. Hemelytron reaching basal part of mediotergite I+II; corium reaching basal half of scutellum, projected laterally beyond lateral margin of metanotum, with posterolateral angle reflexed; hemelytral membrane rugose.
Abdomen 1.4 times as long as its maximum width, with subparallel lateral margins. Mediotergite I+II mostly covered with incrustation, provided with a pair of smooth depressions laterally; mediotergites III–VI fused, weekly elevated longitudinally on midline, mostly covered with four inner pairs and three outer pairs of incrustations; inner paired incrustations each with a round smooth depression, and outer paired incrustations reaching lateral margins of respective mediotergites; mediotergite VII covered with incrustations anteriorly and laterally. Dorsal laterotergites mostly covered with incrustations, each with two round callous spots and callous outer anterolateral angle; dorsal laterotergite II+III slightly protruding at middle (posterolateral angle of original dorsal laterotergite II) and at posterolateral angle; posterolateral angles of dorsal laterotergites IV–VI not protruding; outer margin of dorsal laterotergite VI slightly angulated posteriorly; dorsal laterotergite VII posteriorly protruding and subangular, reaching level of tip of paratergite VIII in dorsal view, not reaching level of tip of pygophore. Sternite I+II covered with incrustation; sternites III–VI reticulately incrusted with small to large callosities; sternite VI with a pair of circular humps medially; sternite VII less incrusted, elevated posteromedially, with a pair of subtriangular humps medially. Paratergite VIII rhomboid, angulated posteriorly, reaching level of basal two-thirds of pygophore. Spiracles II–V ventral, spiracles VI and VII lateral, visible in dorsal view, spiracle VIII dorsolateral, visible in dorsal view.
Pygophore
(Figs
Female (Figs
Variation
(Fig.
Nesoproxius kishimotoi sp. nov., paratypes A, B habitus, dorsal view A male B female C, D habitus, ventral view C male D female E left antenna, male F, G hemelytra, dorsal view F male G female H, I apical part of abdomen, dorsal view H male I female J, K apical part of abdomen, ventral view J male K female.
Measurements [in mm, ♂ (holotype and paratypes; n = 5), holotype in parentheses / ♀ (paratypes; n = 5)]. Body length 2.85–3.06 (2.88) / 3.06–3.47; head length 0.48 (0.48) / 0.48–0.57, width across eyes 0.55–0.57 (0.57) / 0.50–0.61; length of antennae 0.69–0.72 (0.72) / 0.70–0.80; pronotum length 0.61–0.64 (0.64) / 0.61–0.70, width 1.07–1.16 (1.11) / 1.11–1.20; scutellum length 0.32–0.36 (0.32) / 0.30–0.55, width 0.61–0.80 (0.80) / 0.68–0.93; abdomen length 1.55–1.64 (1.64) / 1.55–1.84, width 1.18–1.30 (1.27) / 1.30–1.41; pygophore length 0.23–0.25 (0.23), width 0.32–0.34 (0.32).
Nymph
(Figs
Fourth instar. Generally similar to fifth instar but body generally dark gray, both sides of head beige; body length smaller, 2.6 mm; setae arising from margin of body relatively longer than fifth instar.
This new species is the first one to exhibit a brachypterous condition in Nesoproxius; all specimens examined showed brachypterous features, and none exhibited an apterous or macropterous condition. Even excluding the characteristics of brachypterous wings, this new species can be easily distinguished from other Nesoproxius species by the relatively low development of the median ridges on the pronotum and scutellum, as well as the relatively smooth abdominal margin. The unique characteristics of this new species may have been acquired through the long-term isolation in the Ogasawara Islands, which are far from New Guinea, the center of the geographic distribution of the genus.
In this study, we also clarified for the first time that sexual dimorphism in this Nesoproxius species is manifested in the pattern of incrustations, particularly those on mediotergite VII. Previous studies have described and illustrated this characteristic; however, all known species have been described based on one or two individuals, most of which were females (
Moreover, this is the first time that nymphal stages have been described for Nesoproxius species. The body of the nymph is covered with sparse pubescence on the dorsal surface; however, it does not show the incrustations found in adults. In addition, as setae on the body margin are longer in 4th instar than in 5th instar nymphs, they possibly are relatively longer in younger instars.
The specific name is after Toshio Kishimoto, the first collector of this species.
(Fig.
This new species, endemic to the Ogasawara Islands, represents the northernmost occurrence reported for Nesoproxius, which is far from the distribution of its congeners, and it is the first representative in this genus from the Oceanian region.
(Figs
Habitats and living individuals of Nesoproxius kishimotoi sp. nov. A, B Habitat in Ototojima Island C, D decayed fallen branches of Schima wallichii mertensiana, of which the type specimens were collected E adult male, dorsal view F ditto, dorsolateral view G adult female, feigning death H fourth instar nymph, dorsolateral view.
Nesoproxius kishimotoi sp. nov. was collected from the undersurface of decayed fallen branches of Schima wallichii mertensiana on the forest floor. Both adults and nymphs moved very slowly and frequently feigned death with folded legs and antennae. As the adults and nymphs were found together on the same branches, they all seem to inhabit the same cluster; however, their habitat range seems to be limited and scattered. The reason for this is not clear; however, it is possible that the severe damages to the soil ecosystem caused by predation by alien nemertines in the Ogasawara islands (
1 | Small species, less than 3.5 mm | 2 |
– | Larger species, over 4.0 mm | 5 |
2 | Median ridge of scutellum clearly elevated as a T-shape | 3 |
– | Median ridge of scutellum slightly elevated basally or clearly elevated longitudinally | 4 |
3 | Median ridge of scutellum contiguous with lateral incrusted fields posteriorly | N. constrictus Kormilev, 1978 |
– | Median ridge of scutellum not contiguous with lateral incrusted fields posteriorly | N. gracilis Kormilev, 1968 |
4 | Anterior margin of pronotum straight; anterior angles of pronotum not projected; scutellum triangular, with median ridge clearly elevated along midline wholly | N. minutus Usinger & Matsuda, 1959 |
– | Anterior margin of pronotum sinuate; anterior angles of pronotum projected beyond collar; scutellum trapezoidal, with median ridge slightly elevated mediobasally | N. kishimotoi sp. nov. |
5 | Median ridge of pronotum strongly inflated, overlapping with base of head | 6 |
– | Median ridge of pronotum slightly inflated, not overlapping with base of head | 8 |
6 | Spiracle VIII lateral | N. malayensis Kormilev, 1983 |
– | Spiracle VIII dorsal | 7 |
7 | Median ridge of vertex subtriangular; median ridge of pronotum truncate posteriorly | N. vietnamensis Kormilev, 1968 |
– | Median ridge of vertex ovate; median ridge of pronotum angulate posteriorly | N. yoshimotoi Kormilev, 1970 |
8 | Pronotum hexagonal; posterior angle of abdominal segment VII of female not reaching tip of paratergite | N. hexagonalis Kormilev, 1968 |
– | Pronotum subrectangular or trapezoidal; posterior angle of abdominal segment VII of female reaching or exceeding tip of paratergite | 9 |
9 | Pronotum subrectangular, without a projection on lateral margin; posterior angle of abdominal segment VII of female not produced into a long spine | N. punctulatus Kormilev, 1968 |
– | Pronotum trapezoidal, with a projection on lateral margin slightly before middle; posterior angle of abdominal segment VII of female produced into a long spine | N. angulatus Kormilev, 1968 |
We are very thankful to Jader Oliveira (University of São Paulo, São Paulo, Brazil), Ernst Heiss (Tiroler Landesmuseum, Innsbruck, Austria), Hélcio Gil-Santana (Instituto Oswaldo Cruz, Rio de Janeiro, Brazil), and an anonymous reviewer for the critical reading of the manuscript and for giving valuable comments. We are grateful to the Kanto Regional Environment Office, Ministry of the Environment, Japan (Saitama, Japan) for allowing us to use the specimens collected from the Ogasawara Islands by the project of the Ministry of the Environment in Japan. We sincerely thank Toshinobu Matsumoto (Japan Wildlife Research Center, Tokyo, Japan), Toshio Kishimoto (Museum of Natural and Environmental History, Shizuoka, Japan), and Hiroaki Kojima (