Tijdschrift voor Entomologie 159 (2016) 209–216
A new mirine plant bug genus (Heteroptera: Miridae: Mirinae: Mirini), with two confirmed species from Nepal and Taiwan Tomohide Yasunaga
A new mirine plant bug genus Gotoshinomiris is proposed to accommodate two Asian representatives, G. formosacolus, herein described from Taiwan as new to science, and G. ptilophallus (Yasunaga & Duwal) known from Nepal and transferred from Mahania Poppius as a new combination. This intriguing new genus, at first reminiscent of Mahania, appears most closely related to Orientocapsus Yasunaga & Schwartz based on the genitalic structures. A hypothesis is presented that places Gotoshinomiris in an intermediate phylogenetic position between the Orientocapsus lineage (including Philostephanus Distant) and the Mahania lineage (with Castanopsides Yasunaga and Liocapsus Poppius). A key is provided to aid in identification of the six related genera. Keywords: Heteroptera; Miridae; new genus; new species; new combination; Nepal; Taiwan Tomohide Yasunaga, Research Associate, Division of Invertebrate Zoology, American Museum of Natural History, New York, NY 10024, USA. [email protected]
Introduction Material and methods The present paper documents the finding of a plant Type specimens are deposited in the American Mu- bug from Taiwan, which represents a new genus seum of Natural History, New York, USA (AMNH); and species, Gotoshinomiris formosacolus, in the tribe the National Institute of Agro-Environmental Sci- Mirini of the subfamily Mirinae. This intriguing ences, Tsukuba, Japan (NIAES); the National Muse- genus is presumed to occupy an intermediate phylo- um of Natural Science, Taichung, Taiwan (NMNS); genetic position between Orientocapsus Yasunaga & and the T. Yasunaga Collection, Nagasaki, Japan Schwartz and Mahania Poppius, based on the char- (TYCN). acters of the male and female genitalia. A matrix code label was attached to the holotype The new species G. formosacolus was found within specimen of the new species; these labels uniquely the central montane zone dominated by decidu- identify each specimen, and are referred to as ‘unique ous broadleaf forests in Taiwan. A second species specimen identifiers’ (USIs). The USI codes [e.g., described from Nepal, Mahania ptilophalla Ya- AMNH_PBI 000123] comprise an institution and sunaga & Duwal, was confirmed as a member of project code (AMNH_PBI) and a unique number Gotoshinomiris . A key is also provided to distinguish (000123). These data were digitized on the Arthro- Gotoshinomiris and five additional related genera, pod Easy Capture (formerly the Planetary Biodiver- all of which were not treated in the latest compre- sity Inventory) database maintained by the AMNH hensive work on the Mirini by Chérot & Malipatil (http://research.amnh.org/pbi/) and are also search- (2016). able on the ‘Heteroptera Species Pages’ (http:// research.amnh.org/pbi/heteropteraspeciespage/).
Tijdschrift voor Entomologie 159: 209–216, Table 1, Figs 1–21. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 30 December 2016. DOI 10.1163/22119434-15903002 Downloaded from Brill.com09/25/2021 11:37:38PM via free access
All measurements are in millimeters (mm). Termi- narrowly keeled along apical margin (Fig. 10). nology of the genitalia principally follows Yasunaga Endosoma (Figs 11–14, 17) heavily sclerotized, & Duwal (2008) and Yasunaga & Schwartz (2007). with apical sclerite bearing bundle of hair-like spines Abbreviations used to indicate the male and female (APS, sensu AS by Yasunaga & Schwartz 2007), genitalic structures (Figs 8–21 below) correspond to spatula-shaped lobal sclerite extending from ven- those mentioned in the generic description. tral secondary gonopore (SLS), and membranous, apically spinulate secondary lobe (MSL); secondar y gonopore (SGP) small, thick-rimmed, situated at Taxonomy middle. Female genitalia (Figs 18–21, Table 1): Gotoshinomiris Yasunaga, gen. n. Basically symmetrical and similar to Orientocap- Diagnosis. Distinguished from the related mirine sus in general shape. Bursa copulatrix (Figs 18, 19) genera, Castanopsides Yasunaga, Mahania Poppius, with combined structure of vermiform gland and Orientomiris Yasunaga & Schwartz, and Philo- roof of genital chamber partly sclerotized, dorsally stephanus Distant (and some superficially similar with perimeter surrounded by sclerotization (SPGC: Macrolygus Yasunaga and Tricholygus Poppius, see sclerotized perimeter of combined vermiform gland Chérot & Malipatil 2016), by the following com- and dorsal wall of genital chamber, sensu Yasunaga bination of characters: moderate to relatively large & Schwartz 2007) and ventrally with paired inner size (6.40–7.70 mm total length); castaneous to lateral projections (MPGC: medial process project- chocolate brown general coloration; brown simple ing into genital chamber from sclerotized portion of setae uniformly distributed on pronotum, scutel- posterolateral margin of combined vermiform gland lum and hemelytron; smooth head without mesal and roof of genital chamber, loc. cit.); genital cham- sulcus or basal carina; almost linear antennal seg- ber dorsally with a pair of medial keels (MEK, loc. ment II much longer than basal width of prono- cit.) dividing each lateral oviduct. Sclerotized ring tum; short, slender labium, not exceeding apex of (SCR) enlarged, elliptical, narrowed inward; dorsal mesocoxa; and shape of the male and female geni- labiate plate (DLP) rather widened. Posterior wall of talia. The key below will help distinguish Gotoshi- bursa copulatrix (Fig. 20) with bilobate, spinulate, nomiris from these similar-appearing taxa (see also symmetrical dorsal structure (DSS); interramal lobe Table 1). (IRL) developed, slender, densely spinulate; lateral Description. Body generally brown-castaneous, lobe (LLB) well-sclerotized, long, horn-like, tapered elongate, subparallel-sided, moderate to large in size apically. (6.40−7.70 mm); dorsal surface rather shining, with uniformly distributed, brown or pale brown, semi- Type species. Gotoshinomiris formosacolus erect or reclining, simple setae. Head shiny brown, Yasunaga, n. sp. almost smooth, with sparsely distributed, silky, short Etymology. Named after the late Mr. Shin Goto setae, without a median sulcus or basal carina; head (or Gotoh), a naturalist, who collected most of the across eyes subequal in length to antennal segment I, specimens of the type species of this new genus, about half as wide as base of pronotum; vertex rather combined with the generic name Miris Fabricius; wide. Antenna relatively long; segment I about as masculine. thick as II; segment II almost linear, much longer Distribution. Nepal and Taiwan. than basal width of pronotum; segments III and IV Discussion. The most consistent features of this filiform. Labium short, slender, not exceeding apex new genus are exhibited predominantly in the male of mesocoxa. Pronotum shining, finely and shallow- and female genitalia as described above and listed in ly punctate, with silky, reclining setae; calli weakly Table 1. Most of the external characters are shared by swollen; collar narrow, about as thick as base of an- the related genera, Castanopsides, Liocapsus, Maha- tennal segment II; scutellum flat. Hemelytron largely nia, Orientocapsus and Philostephanus (see Yasunaga chocolate brown, shining but weakly shagreened or 1998a; Yasunaga & Schwartz 2007; and Yasunaga & roughened, with uniformly distributed, brown, Duwal 2008 for detailed information on these taxa). simple, semierect or reclining setae; cuneus concol- Gotoshinomiris is also reminiscent of certain mem- orous without pale base. Legs long; tibial spines pale bers of Creontiades-Megacoelum-Orientomiris group; brown; short; meta-tarsomere I subequal in length however, the similarity is only superficial (cf. Yasu- to II, shorter than III. Male genitalia (Figs 7–17, naga 1998b; Chérot & Malipatil 2016). Table 1): Parameres as in Figs 8, 9, 15, 16; right The present careful evaluation of characters sug- paramere slender and elongate, with moderately gests that Gotoshinomiris is posited to be sister to (not strongly) flattened inner surface (Figs 9, 16); Mahania or Orientocapsus and constitutes a com- left paramere with a prominent, ‘dorsobasal projec- pact Asian genus including two Oriental elements, tion (DBP)’ on sensory lobe and apically hooked G. formosacolus n. sp. from Taiwan and G. ptilophallus hypophysis (Figs 8, 15). Phallotheca almost smooth, (Yasunaga & Duwal) from Nepal. The distribution Downloaded from Brill.com09/25/2021 11:37:38PM via free access
Table 1. Principal characters in the genitalia for six related mirine genera.
Characters Castanopsides Mahania Gotoshinomiris Orientocapsus Philostephanus Liocapsus
1 Dorso-basal moderate moderate prominent absent absent weak projection of left paramere 2 Process at apex of present weak weak present present present phallotheca 3 Spicule on endosoma present present absent absent absent present 4 Apical sclerite absent simple apex with bundle simple but simple but absent of spines modified modified 5 Inner elongate absent absent absent present absent absent sclerite 6 Gonoporal sclerite moderate moderate prominent (SLS) moderate variable absent 7 Female genital symmetrical symmetrical weakly weakly strongly weakly chamber asymmetrical asymmetrical asynmetrical asymmetrical 8 MPGC absent absent present present present absent 9 SPGC absent absent present present present absent 10 MEK* absent absent present present present* present* 11 Origin of vermiform conventional conventional anterior to lateral posterior anterior anterior gland oviduct 12 Interramal lobe conventional conventional elongate conventional elongate** conventional 13 Horned lateral lobe absent absent present absent present*** absent 14 Dorsal structure single lobe single lobe bilobate bilobate bilobate single lobe
Abbreviations as mentioned in generic description. * MEK replaced by medial sclerite of combined vermiform gland and roof of genital chamber (MSGC in Philostephanus) or toughened projection of dorsal sac (Liocapsus), sensu Yasunaga & Schwartz (2007). ** Strong intraspecific variation present inPhilostephanus. *** Possessed by three species in Philostephanus. of each species seems to be disjunct; however, a are shared by Orientocapsus and/or Philostephanus. similarly fragmented distribution pattern is also Currently without a wider survey of numerous mir- known in some mirid taxa and other insect taxa ine taxa, it is difficult to determine which characters (cf. Yasunaga 2000; 2001). It must be recognized are undoubtedly synapomorphic or homoplasious; that the vast modern environment in continental therefore, I herein refrain from a definitive cladistic China lying between the Himalayan region (includ- analysis. ing N Indochina) and easternmost Asia (in particular There is a strong likelihood that the species of Japanese Ryukyus and Taiwan) has undergone rapid Gotoshinomiris are associated with deciduous broad- urbanization and industrialization, which have seri- leaf trees, particularly Quercus spp. (Fagaceae); ously disturbed and decreased the forest zones — G. ptilophallus was collected from Quercus inflores- habitats of wildlife there, which could most probably cences in Kathmandu, Nepal. The similarity of their result in disjunction of the originally continuous dis- habitat and probable host preferences may imply a tribution patterns. relationship of Gotoshinomiris with Castanopsides and The systematic position of Mahania, seemingly Liocapsus. most similar to Gotoshinomiris, has not been eluci- dated well. The present discovery of both male and Key to Gotoshinomiris and related genera female specimens of G. formosacolus revealed that 1. Dorsal surface highly polished, oily shiny, this new species most probably occupies an interme- impunctate and very smooth; vestiture on diate position between the Mahania–Castanopsides dorsum absent or only sparsely distributed; la- lineage (Yasunaga & Duwal 2008) and the Ori- bium very short, not surpassing apex of procoxa entocapsus–Philostephanus lineage (Yasunaga & nor reaching base of mesocoxa ������������������������ Schwartz 2007). Liocapsus, superficially similar to ���������������������������������������������Liocapsus Poppius Philostephanus, is now presumed to be rather closer – Dorsal surface more or less shining, but with to the former lineage than to the latter, in having punctures at least on pronotum and scutellum; the shortened labium and an independent spicu- one or both of pronotum and hemelytron usu- lum on the endosoma. Among the characters in ally with uniformly distributed simple setae Table 1, three characters in the male genitalia (1, (if dorsum glabrous, labium always exceeding 4, 6) are assumed to be autapomorphic for Gotoshi- apex of mesocoxa, and/or hemelytron speck- nomiris, whereas the majority of apomorphic states led); labium reaching or exceeding apex of of characters in the female genitalia (7–10, 12, 14) mesocoxa ������������������������������������������������������ 2 Downloaded from Brill.com09/25/2021 11:37:38PM via free access
2. Head smooth, without mesal sulcus or basal Recreation Area, 23.06 N, 120.75 E, 12–13 May carina; labium shorter, not exceeding apex of 1989, S. Gotoh (AMNH_PBI 00380469) (NMNS). mesocoxa ������������������������������������������������������ 3 Paratypes: Taiwan, 1♂ 1♀, same data as for the – Head with one or both of longitudinal mesal sul- holotype (TYCN); 1♀, Taichung [current Nantou cus and basal transverse carina on vertex; labium Country], Puli Township, 24.00 N 121.00 E, 15–18 longer, exceeding apex of mesocoxa ���������������� 4 May 1989, S. Gotoh (AMNH); 1♂, Taitung, near 3. Body smaller (total length less than 6.70 mm in Takaolao (detailed locality unknown), 1 May 1985, ♂/ 7.80 mm in ♀); length of antennal segment A. Saito (NIAES). II greater than basal with of pronotum; prono- Diagnosis. Recognized by the characters men- tum pubescent ��������������Gotoshinomiris n. gen. tioned in the generic diagnosis. Distinguished from – Body larger (total length more than 7.70 mm G. ptilophallus by its larger size, yellowish-brown bas- in ♂/ 8.30 mm in ♀); antennal segment II al 2/3 of antennal segment III, and almost entirely shorter than basal with of pronotum; pr onotum pale pronotum, in addition to the isolated, insular glabrous �������������������������������Mahania Poppius distribution. 4. Pronotal collar thicker than base of antennal seg- Description. Body generally brown, with uniformly ment II; cuneus brown or paler, usually with dark darkened hemelytron; dorsal surface weakly shining, apex (if cuneus almost entirely darkened, pro- with uniformly distributed, brown, simple setae. notal collar yellow or yellowish brown); endo- Head shiny pale brown, almost smooth. Antenna soma largely membranous, with an independent dark brown; segment II much longer than basal spiculum and a median sclerite extending from width of pronotum, slightly longer than metafe- distal end of seminal duct; female genital cham- mur; basal 3/4 of segment II in ♀ reddish brown; ber largely membranous, without any noticeable basal 2/3 of segment III yellowish brown; base of sclerotization ����������������� Castanopsides Yasunaga segment IV in ♀ pale brown. Labium shiny pale – Pronotal collar as thick as or narrower than base brown, reaching apex of mesocoxa; apical half of seg- of antennal segment II; cuneus largely brown, ment IV reddish brown. Pronotum shining, almost dark brown to fuscous, with narrowly pale an- uniformly pale brown, in ♀ sometimes tinged with terior margin (if cuneus uniformly fuscous, red; pleura widely darkened, partly shagreened; scent hemelytron and pronotum including collar efferent system grayish brown; scutellum chocolate also concolorously fuscous); endosoma heavily brown, shallowly and transversely rugose. Hemely- sclerotized, lacking independent spiculum but tron chocolate brown; embolium and outer mar- various lobal-sclerites; female genital chamber gin of cuneus pale reddish brown in ♀; membrane with distinct sclerotization ���������������������������� 5 including veins and areolar cells smoky brown. All 5. Labium shorter than metafemur [except for coxae and legs yellowish brown, partly tinged with O. picinus (Lu & Zheng) known only from red; apical half of each tarsomere III darkened. Ab- Yunnan, China]; endosoma with inner elon- domen wholly chocolate brown, in ♀ mottled with gate sclerite and apical sclerite (possibly ho- pale brown portions. Other structures including mologous to APS as in Fig. 14); female genital male and female genitalia as mentioned in generic chamber nearly symmetrical, dorsally with a description. sclerotized, median keel dividing each lateral Measurements. ♂/♀: Total body length 6.50– oviduct; vermiform (spermathecal) gland ex- 6.60/7.22–7.70; width of head across eyes 1.07– tending from posterior end of genital chamber 1.11/1.15–1.16; width of vertex 0.39–0.42/0.46– ����������������� Orientocapsus Yasunaga & Schwartz 0.49; lengths of antennal segments I–IV: 0.98–1.01, – Labium as long as or longer than metafemur; 2.59–2.80, 1.29–1.38, ?/1.05–1.18, 2.72–2.85, endosoma principally composed of four scler- 1.47, 0.86; length of labium 1.71–1.96/2.08– ites: apical sclerite (homologous to APS), 2.16; mesal pronotal length including collar lateral sclerite, gonoporal sclerite (= SLS in 1.29–1.35/1.47–1.50; basal pronotal width 2.15– Gotoshinomiris) and ventral sclerite (terminol- 2.21/2.37–2.41; maximum width across hemelytron ogy sensu Yasunaga & Schwartz 2007); fe- 2.37–2.45/2.74–2.94; and length of metafemur, male genital chamber heavily sclerotized and tibia and tarsus 2.42–2.45, 3.65–3.68, 0.68–0.74/ more or less asymmetrical; vermiform gland 2.62–2.77, 3.92–3.95, 0.75–0.76. extending from anterior to roof of genital Etymology. Named for its occurrence in Taiwan, chamber ������������������������Philostephanus Distant formerly called Formosa. Distribution. Taiwan (central montane zones). Gotoshinomiris formosacolus Yasunaga, n. sp. Figs 1–4, 7–14, 18–21 Gotoshinomiris ptilophallus (Yasunaga & Type material. Holotype ♂. Taiwan, Kaohsiung Duwal), n. comb. City, Taoyuan District, Tengihih National Forest Figs 5, 6, 15–17 Downloaded from Brill.com09/25/2021 11:37:38PM via free access
Figs 1–6. Gotoshinomiris species, habitus images. — 1, G. formosacolus, holotype male; 2, same, paratype male; 3, 4, same, paratype female; 5, 6, G. ptilophallus, holotype male (alive).
Mahania ptilophalla Yasunaga & Duwal, 2008: with red; dorsal surface widely chocolate brown; la- 411 (n. sp.); Schuh 2012–2014: web catalog bium pale brown, reaching base of mesocoxa; api- (http://research.amnh.org/pbi/catalog/references. cal half of segment IV dark brown; pronotal collar php?id=103671). – Holotype ♂. Nepal, Kath- creamy yellow; pleura uniformly pale brown; hem- mandu, Nagarjun (1,600–1,800 m alt.), 27.74 N elytron partly with silky, reclining setae; extreme base 85.26 E, on Quercus sp., 16 May 2005, T. Yasunaga (knee) of meso- and metatibia with a sanguineous (AMNH) [examined]. spot; dorsobasal protuberance (DBP) of left param- ere flattened (Fig. 15); hypophysis of right param- Diagnosis. Recognized primarily by the following ere blunt-tipped (Fig. 16); endosomal apical sclerite characters: Body generally brownish, partly tinged (APS) with feather-like, hairy appendages (Fig. 17). Downloaded from Brill.com09/25/2021 11:37:38PM via free access
Figs 7–17. Male genitalia of Gotoshinomiris formosacolus (7–14) and G. ptilophallus (15–17). — 7, pygophore; 8, 15, left paramere; 9, 16, right paramere; 10, phallotheca; 11–14, 17, endosoma. Scale bars 0.2 mm. Abbreviations as mentioned in generic description.
A detailed description was provided by Yasunaga & Discussion. This Nepalese species, described from a Duwal (2008). single male specimen, was placed in Mahania Pop- Measurements: Holotype (♂). Body length 6.40; pius, based on the overall similarity and the form head width across eyes 1.15; vertex width 0.36; of the male genitalia (e.g., principal coloration and lengths of antennal segments I–IV 0.96, 2.71, 1.08, structure of dorsal surface; short labium; somewhat 0.53; labial length 1.58; mesal pronotal length in- flattened right paramere; developed sensory lobe of cluding collar 1.20; basal pronotal width 2.11; width the left paramere; and a narrow keel of phallothe- across hemelytron 2.40; and lengths of metafemur, ca). Yasunaga & Duwal (2008) also considered the tibia and tarsus 2.16, 3.48, 0.72. highly modified form of the endosoma as simply an Distribution. Nepal (Kathmandu Valley). autapomorphy. However, some characters in the male Downloaded from Brill.com09/25/2021 11:37:38PM via free access
Figs 18–21. Female genitalia of Gotoshinomiris formosacolus. — 18, 19, bursa copulatrix; 20, posterior wall; 21, left sclerotized ring and its adjacent structures. Scale bars 0.2 mm. Abbreviations as mentioned in generic description.
genitalia (e.g., left paramere with DBP, and endoso- Museum of Natural History, Smithsonian Institu- ma with hairy or spine-like APS and distinct SLS) are tion, Washington, DC, USA) for improving the man- now unequivocally recognized as synapomorphies uscript with important comments and suggestions. shared with G. formosacolus. Therefore, ptilophallus is transferred to Gotoshinomiris from Mahania. References Acknowledgements Chérot, F. & M.B. Malipatil, 2016. A review of Adelphoco- Special thanks are due to Mrs Michiko Gotoh, wife ris-Creontiades-Megacoelum complex (Hemiptera: Het- eroptera: Miridae: Mirini), with descriptions of two of the late Mr Shin Gotoh (Tanabe City, Wakayama new genera and four new species. — Zootaxa 4126: Pref., Japan) for providing opportunity to access and 151–206. examine valuable specimens. I also thank Dr Ram Schuh, R.T., 2002–2014. On-line Systematic Catalog of Keshari Duwal (Kyushu University, Fukuoka, Japan) Plant Bugs (Insecta: Heteroptera: Miridae). — Avail- for sharing her ideas and Dr Y. Nakatani (NIAES) for able from: http://research.amnh.org/pbi/catalog/. the loan of material. Thanks are extended to Dr M.D. Yasunaga, T., 1998a. Revision of the mirine genus Schwartz (Agriculture & Agri-Food Canada, Ottawa, Castanopsides Yasunaga from the eastern Asia (Heter- Ontario, Canada) and Dr T.J. Henry (Systematic optera: Miridae). — Entomologica Scandinavica 29: Entomology Laboratory, ARS-USDA, c/o National 99–119. Downloaded from Brill.com09/25/2021 11:37:38PM via free access
Yasunaga, T., 1998b. Revision of the mirine genus assumed sister genus Mahania Poppius from Nepal, Creontiades Distant and allies from Japan (Heteroptera: with a new synonymy of the genus Liocapsus Poppius Miridae). Part III. Neomegacoelum gen. n. and exotic (Heteroptera, Miridae, Mirinae). — In: S. Grozeva & new taxa, new synonymy and new combinations. — N. Simov (Eds), Advances in Heteroptera Research, Entomological Science 1: 63–70. Festschrift in Honor of 80th Anniversary of Mi- Yasunaga, T., 2000. The mirid subfamily Cylapinae (Het- chail Josifov, pp. 403–417. Pensoft Publishers Sofia– eroptera: Miridae), or fungal inhabiting plant bugs in Moscow, Bulgaria. Japan. — Tijdschrift voor Entomologie 143: 183–209. Yasunaga, T. & M.D. Schwartz, 2007. Revision of the mir- Yasunaga, T., 2001. Family Miridae Hahn, plant bugs. ine plant bug genus Philostephanus Distant and allies — In: T. Yasunaga, M. Takai & T. Kawasawa (Eds), A Heteroptera (Miridae: Mirinae: Mirini). — Tijdschrift Field Guide to Japanese Bugs, Vol. II, pp. 1–96, 111– voor Entomologie 150: 100−180. 351. Zenkoku Noson Kyoiku Kyokai, Publishing Co. Ltd., Tokyo, Japan. [In Japanese] Received: May 22, 2016 Yasunaga, T. & R.K. Duwal, 2008. New species of the Accepted: October 3, 2016 mirine plant bug genus Castanopsides Yasunaga and its
Downloaded from Brill.com09/25/2021 11:37:38PM via free access