THE MIRID SUBFAMILY CYLAPINAE (HETEROPTERA: MIRIDAE), OR FUNGAL INHABITING PLANT BUGS IN JAPAN

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1 TOMOHIDE YASUNAGA Hokkaido University of Education, Sapporo THE MIRID SUBFAMILY CYLAPINAE (HETEROPTERA: MIRIDAE), OR FUNGAL INHABITING PLANT BUGS IN JAPAN Yasunaga, T., The mirid subfamily Cylapinae (Heteroptera: Miridae), or fungal inhabiting plant bugs in Japan. Tijdschrift voor Entomologie 143: , figs [ISSN ]. Published 1 December The plant bug subfamily Cylapinae of Japan is revised. This subfamily comprises species with a specialized biology, most of which are known to be associated with fungi. Sixteen species are recognized in six genera of three tribes. A new genus Yamatofulvius (type species Y. miyamotoi sp. n.) is proposed to accommodate three unique new species; its phylogeny is discussed. Eleven new species are described: Bothriomiris capillosus, Bothriomiris gotohi, Cylapomorpha michikoae, Fulvius niveonotatus, Peritropis hasegawai, Peritropis insularis, Peritropis iriomotensis, Peritropis takahashii, Yamatofulvius laevigatus, Yamatofulvius miyamotoi, Yamatofulvius sinuicornis. Fulvius dimidiatus Poppius, F. tagalicus Poppius and Peritropis advena Kerzhner are reported from Japan for the first time. A key is provided to distinguish Japanese tribes, genera and species of the Cylapinae. The zoogeography of the Japanese Cylapinae is also discussed. Correspondence: Tomohide Yasunaga, Biological Laboratory, Hokkaido University of Education, Ainosato 5-3-1, Sapporo , Japan. yasunaga@atson.sap.hokkyodai.ac.jp Key words. Heteroptera; Miridae; Cylapinae; revision; new genus; new species; Japan. The Cylapinae form a unique plant bug subfamily within the Miridae. The majority of the species in this group are known to be associated with fungi. The faunae of the subfamily have been poorly documented, due to the specialized habit and habitat, and in part to the remarkable speed and agility of both immature and mature forms, which makes their capture by hand challenging (Wheeler 1994). Only 14 species have been known to occur in the Palearctic Region (Kerzhner & Josifov 1999), and the Japanese fauna has been represented by Fulvius anthocoroides (Reuter) and Punctifulvius kerzhneri Schmitz of the tribe Fulviini (Carvalho 1956, Yasunaga et al. 1999). Continuing efforts by the author and colleagues to clarify the Japanese cylapine fauna have resulted in the recognition of 16 species in six genera. Of these, eleven species were found to be undescribed, and three other species were not previously recorded from Japan. In addition, three unique undescribed species could not be accommodated by any known genus. This paper documents the Japanese cylapine fauna comprehensively, with descriptions of eleven new species. A new genus related to Punctifulvius Schmitz, Yamatofulvius, is diagnosed and described with a discussion on its phylogeny. Fulvius dimidiatus Poppius, F. tagalicus Poppius and Peritropis advena Kerzhner are recorded from Japan for the first time. All species are diagnosed, and the biology is briefly documented for most species. A key is provided to distinguish the Japanese tribes, genera and species. The zoogeography of the Japanese Cylapinae is discussed. The female genitalia of the Cylapinae had not been documented sufficiently in previous works, probably due to the fragile, widely membranous and hardly visible structure. Therefore, in this paper, illustrations and descriptions of the female genitalia are provided for eight species that were represented by sufficient material for dissection. MATERIAL AND METHODS About 300 dried specimens of 28 species (including exotic ones) were examined. Depositories of the material are abbreviated as follows: BMNH Department of Entomology, the Natural History Museum, London, UK ELKU Kyushu University Entomological Collection, Fukuoka, Japan HUES Hokkaido University of Education, Sapporo, Japan 183

2 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, Figs Cylapomorpha michikoae. 1, Female adult; 2, the same, frontal view; 3, male adult, left lateral view; 4, final instar nymph. NIAS NSMT USNM ZMAS ZMUF National Institute of Agro-Environmental Sciences, Tsukuba, Ibaraki, Japan Department of Zoology, National Science Museum, Tokyo, Japan United States National Museum of Natural History, Washington, D. C. Zoological Institute, Russian Academy of Sciences, St. Petersburg Zoological Museum, University of Helsinki, Finland Nymphs of five species were also examined; they are preserved in 80% ethyl alcohol in small vials (HUES). Photographs presented in this paper were made with an Olympus OM-System (OM-4Ti 35mm camera with T10 Ringflash, Auto Extension Tube, and either Zuiko 50mm/f3.5 or 38mm/f2.8 Macro Lens). Because cylapines are extremely agile and quickly sprint away, most photos of live material were taken from specimens which had been weakly anaesthetized with chloroform vapour, or which had been in the refrigerator for a while. Many photos were made and offered by my colleagues M. Takai (figs. 1, 2, 18-20, 31-35, 55-56, 77) and Y. Nakatani (fig. 21). All measurements in the text are given in millimetres. In the synonymic lists, only selected references are cited; for detailed lists see the catalogues of Schuh (1995) and Kerzhner & Josifov (1999). New distributional records for known species are each indicated by an asterisk (*) after the name of a region. Terminology of the genitalia The following abbreviations are used in figures of the genitalia for appropriate indication: AS, anterior sac; DLP, dorsal labiate plate; DS, dorsal sac; IRS, interramal sclerite; LA, lateral arm; MP, mesal sclerotized 184

3 YASUNAGA: Japanese Cylapinae plate (of posterior wall of bursa copulatrix); OC, oviductus communis; OL, oviductus lateralis; PT, phallotheca; SR, sclerotized ring; VLP, ventral labiate plate; VLV, valvifer. The terms for the female genitalia mainly follow Kullenberg (1947). This eminent work was published in German, and many of his terms are translated into English. Several new terms (e.g. LA, MP) are used to tentatively indicate structures of the female genitalia because Kullenberg did not treat any species of the Cylapinae. As my examination is limited to the Japanese species, however, some terms widely used in other subfamilies (e.g. IRS, SR) are not perfectly applicable to cylapines. Therefore, these terms are also tentative, and much broader observations on many other cylapine taxa are required to determine a definitive homology between cylapine female genitalia and those of other subfamilies. Checklist of Cylapinae in Japan Tribe Bothriomirini Kirkaldy, 1906 Genus Bothriomiris Kirkaldy, 1902 Bothriomiris capillosus sp. n. Bothriomiris gotohi sp. n. Tribe Cylapini Kirkaldy, 1903 Genus Cylapomorpha Poppius, 1914 Cylapomorpha michikoae sp. n. Tribe Fulviini Uhler, 1886 Genus Fulvius Stål, 1862 Fulvius anthocoroides (Reuter, 1875) Fulvius dimidiatus Poppius, 1914 Fulvius niveonotatus sp. n. Fulvius tagalicus Poppius, 1914 Genus Peritropis Uhler, 1891 Peritropis advena Kerzhner, 1972 Peritropis hasegawai sp. n. Peritropis insularis sp. n. Peritropis iriomotensis sp. n. Peritropis takahashii sp. n. Genus Punctifulvius Schmitz, 1978 Punctifulvius kerzhneri Schmitz, 1978 Genus Yamatofulvius gen. n. Yamatofulvius laevigatus sp. n. Yamatofulvius miyamotoi sp. n. Yamatofulvius sinuicornis sp. n. long, sometimes much longer than body, not significantly thickened; hemelytra rather tough; tarsi linear, 2- or 3-segmented; claws long, usually with a subapical tooth; parempodia simple, setiform; pulvilli absent; male vesica simple, widely membranous. Carvalho (1952) recognized three tribes, Bothriomirini, Cylapini, and Fulviini in the Cylapinae. However, Schuh (1995) did not use these tribes because monophyly of the latter two tribes cannot be explained by current membership as subsequently mentioned by Gorczyca & Eyles (1997). On the other hand, since Japanese cylapines are undoubtedly grouped into any of the three tribes, I here use them for the classification of the Japanese taxa. But additional tribes or further subtribe level divisions are evidently required to correctly classify many tropical members that are still in great need of investigation. Most Japanese members of the Cylapinae are associated with fungi of the Polyporaceae, such as Coriolus spp., on branches, trunks, decaying woods, or rotten logs under dark and humid forests. Some authors reported that several species prey on larvae of beetles, although many species actually feed on fungi (Wheeler 1994). Most cylapines appear to be nocturnal. The movement of cylapines is very quick. Since they usually do not fly away but cleverly and rapidly sprint away when chased, it is usually difficult to collect sufficient specimens. Such specialized habit and habitat are liable to hinder investigations. Particularly, in tropical rain forests, there are many unknown cylapines. A univoltine life cycle is assumed for temperate and cold temperate species, but collection records suggest that species from the Ryukyus, or subtropical islands, appear to have two or more generations annually. Cylapini Kirkaldy, 1906 Members of this tribe are distinguished from those of other tribes by the elongate oval body with moderate to large size, flattened, impunctate, usually shagreened dorsum, vertical, strongly and anteriorly flattened head, and remarkably long, gracile antenna. In Japan and adjacent regions, no species of this tribe has been reported previously. During this study, an undescribed species of the genus Cylapomorpha was discovered from southwestern Japan. SYSTEMATIC PART Subfamily Cylapinae Kirkaldy, 1903 As mentioned by Gorczyca & Eyles (1997), Schuh (1974) and Schuh & Slater (1995), this subfamily is recognized by the following combination of characters: Dorsum usually heavily punctate; head often elongate anteroposteriorly or dorsoventrally; antenna Cylapomorpha Poppius Cylapomorpha Poppius, 1914: 124 (n. gen.), type species: C. gracilicornis Poppius 1914: 125, original designation; Schuh 1995: 23 (cat.). This is a small genus hitherto known from three species of the Seychelles, Philippines, New Guinea and Pacific islands, and characterized by the elongate oval body, anteriorly flattened, vertical head, dorsally 185

4 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, 2000 Figs Male (5-6) and female (7-8) genitalia of Cylapomorpha michikoae. 5, Genital segment with parameres, dorsal view; 6, apical part of vesica; 7, bursa copulatrix; 10, posterior wall of bursa copulatrix. Scale bars: 0.1 mm. projected eyes, tumid, very short antennal segment I, extremely elongate segment III that is much longer than II, and short, narrowed pronotum. A detailed description of the external structure was provided by Poppius (1914). The present discovery of a species from the temperate zone of Japan proper represents the northernmost distributional record for the genus. Cylapomorpha michikoae sp. n. (figs. 1-9) Type material. Holotype, Sanpo Shrine, Mengawa, Ohtoh Vil., Wakayama Pref., 14.vii.1998, M. & S. Gotoh (HUES). Paratypes: Honshu: 1 1, same data as for holotype (HUES). Kyushu: 1 1, Mt. Osuzu, Miyazaki Pref., 9.viii.1970, S. Tawara (NIAS); 1 1, same locality, 22.viii.1999, M. Takai (HUES). Ryukyus: 3 3, Okunirindo, Ohgimi Vil., Okinawa Is., vi.1990, T. Yasunaga; 1, Ohsittai, Nago C., Okinawa Is., 26.vi.1999, T. Yasunaga & M. Takai (HUES); 1 1, Nosoko, Ishigaki Is., 22.v.& 5.vi.1999, K. Takahashi (HUES). Diagnosis. Readily recognized by the characters mentioned in the generic diagnosis, and brown general coloration partly with reddish tinge, yellow extreme apex of antennal segment II, a dark, square mark on the corium (fig. 1), and mesally upturned posterior margin of abdominal tergum VIII and a thumb-like, mesial process of the genital segment in male (fig. 3). Allied to C. gracilicornis Poppius from the Philippines, from which this new species can be distinguished by the pale head, yellow apex of the antennal segment II, shorter antennal segment III that is less than twice as long as II, and a fuscous, square, mesial mark on the corium. The final instar nymph of C. michikoae is recognized by the external structure and coloration similar to those exhibited in the adult (fig. 4). Description. Body generally brown, partly tinged with red, elongate oval, subparallel-sided; dorsal surface shagreened, with uniformly distributed, dark brown or brown setae. Head pale brown, shagreened, triangular in frontal view (fig. 2), with several, dark, short setae; eyes produced dorsally; vertex with a pair of oblique sutures divergent anteriad, and a mesial, rounded, convex area accompanied with a short, longitudinal, mesal sulcation; frons concave mesad. Antenna dark brown; segment I and sometimes basal part of II pale brown or brown; extreme apex of segment II yellow; lengths of segments I-IV ( / ): / , / , / , / Rostrum reddish brown, reaching abdominal sternum VIII; segment IV infuscate, short, less than half length of segment II or III. Pronotum reddish brown, with a V- shaped pale, mesal mark anteriad, bearing dark setae, shallowly and transversely rugose posteriad; calli distinct; collar present, about twice as thick as antennal segment II; mesoscutum reddish brown, with a pair of 186

5 YASUNAGA: Japanese Cylapinae lateral, oblique carinae; scutellum brown, with a pale, longitudinal, rounded keel mesally; pleura brown, widely pruinosed; propleuron and epimeron sometimes darkened; ostiolar peritreme pale brown. Hemelytra brown, partly tinged with red, shagreened and roughened, with uniformly distributed, brown, reclining setae; corium with a fuscous, square mark mesially; cuneus reddish brown, with pale, semitransparent base. Coxae yellow; legs brown; apex of each tibia pale brown; lengths of metafemur, tibia and tarsus ( / ): / , / Abdomen shiny dark reddish brown, usually with widely pale ventromesial region; posterior mesal margin of tergum VIII distinctly upturned. Male genitalia (figs. 5, 6): Dorsal surface of genital segment with a mesal, thumb-like projection. Parameres with some sensory setae; left paramere semi-circularly curved; right paramere curved subbasally. Vesica with bilobate, wide, thin, densely sutured sclerite, and inner, spinulate, narrow membranous areas. Female genitalia (figs. 7-8): Sclerotized ring absent but a narrow, dorsal labiate plate present; oviductus lateralis slender. Posterior wall of bursa copulatrix with a wide, continuous, laterally warped interramal sclerite. Dimensions. / : Body length / ; head width including eyes / ; head length /0.39; head height / ; vertex width / ; rostral length / ; mesal pronotal length / ; basal pronotal width / ; width across hemelytra / Etymology. Named after Mrs. Michiko Gotoh who kindly helped with my investigation on cylapines in the Kii Peninsula; a noun in the genitive case. Distribution. Japan (Honshu: southern part of Kii Peninsula, Kyushu, Ryukyus: Okinawa, Ishigaki and Iriomote Islands). Biology. This unique cylapine inhabits rotten logs grown with polyporaceous fungi under dark, humid, evergreen broad-leaved forest, and the general coloration of both adults and nymphs is cryptic, very harmonious with the habitat (fig. 4). A univoltine life cycle is assumed for C. michikoae, and the newly emerged adults are found from May in the subtropi- Figs Female metatarsi of the Japanese cylapines. 9, Cylapomorpha michikoae; 10, Fulvius anthocoroides; 11, F. niveonotatus; 12, Peritropis advena; 13, Punctifulvius kerzhneri; 14, Yamatofulvius laevigatus; 15, Y. miyamotoi; 16, Y. sinuicornis; 17, Bothriomiris gotohi. Scale bar: 0.1 mm. 187

6 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, Figs Fulvius spp. 18, F. anthocoroides, female; 19, the same, left lateral view; 20, F. tagalicus, male; 21, F. niveonotatus, holotype female; 22, F. dimidiatus, female. cal Ryukyus to July in the temperate Kii Peninsula. Discussion. The male of this new species has the upturned posteromesal margin of the abdominal tergum VIII and a mesal, thumb-like process on the genital segment (figs. 3, 5). Although Carvalho (1956) and Carvalho & Lorenzato (1978) described and illustrated the male genitalia of Cylapomorpha pacifica Carvalho, 1956, they do not refer to the abdominal segments VIII and IX. So I currently cannot recognize whether these unique forms exhibited in C. michikoae are generic characters or not. I collected a few individuals representing an undescribed species of Cylapomorpha from Malaysia. Unfortunately, they are females. Fulviini Uhler, 1886 The Fulviini are primarily distinguished from other tribes by the rather elongate body, and long, conical head. Judging from the various forms exhibited in the dorsal vestiture and punctation, tarsal segmentation, and genitalia, however, this tribe appears not to be a monophyletic group. A global revision is re- 188

7 YASUNAGA: Japanese Cylapinae quired to correctly redefine the Fulviini. In addition to the above mentioned characters, a single or paired, narrow-rimmed sclerotized ring and distinct interramal sclerite on the posterior wall of bursa copulatrix are recognized in the female genitalia in most Japanese fulviine members. Fulvius Stål Fulvius Stål, 1862: 322 (n. gen.), type species: F. anthocorides Stål, 1862: 322, monotypic; Schuh 1995: 25 (cat.); Kerzhner & Josifov 1999: 8 (cat.). This genus is recognized by the small and rather slender body with principally dark coloration, shagreened, almost impunctate and relatively robust dorsum, long head, short antenna, narrow pronotum, 2- segmented tarsi, flattened and apically hooked sensory lobe of the left paramere, and peculiar bundle of slender sclerites on the vesica. Further diagnostic characters for external structure were provided by Poppius (1909), Reuter (1895), and Wagner (1974). The tarsi of Fulvius were referred to as 3-segmented by some authors (e.g., Carvalho 1956), but at least the Japanese species have rather short, 2-segmented tarsi that enable us to distinguish Fulvius from other fulviine genera in Japan (figs. 10, 11). Fulvius anthocoroides (Reuter) (figs. 10, 18, 19, 23-26) Teratodella anthocoroides Reuter, 1875a: 77, 1875b: 8 (n. sp.). Lectotype, FRANCE: gc, Rouen (ZMUF) [examined]. Fulvius anthocoroides Kerzhner, 1997: 116 (list); Kerzhner & Josifov, 1999: 8 (cat.). Fulvius brevicornis Reuter, 1895: 138, unnecessary replacement name for F. anthocoroides Reuter, 1875 [not preoccupied by F. anthocorides Stål, 1862 (see Kerzhner & Josifov 1999; Schuh (1995: 26, cat.) incorrectly cited F. anthocoroides Stål that should be read as anthocorides )]; Carvalho 1956: 6 (redesc.); Miyamoto & Yasunaga 1989: 158 (list); Schuh 1995: 27 (cat.). Fulvius samoanus Knight, 1935: 203 (n. sp.). Holotype, SAMOA: Malouleilei, iv.1924, Buxton & Hopkins (BMNH) [examined]. Diagnosis. Easily recognized by the fuscous body with four remarkable white areas on the hemelytra. A redescription of the external structure as well as a description of the male parameres were provided by Carvalho (1956, under a junior synonym, brevicornis). Male genitalia (figs ): Left paramere curved at right angle, with hypophysis flattened, twisted at base, terminated in apical hook; hypophysis of right paramere slender. Vesica with bundle of apical small sclerotized processes and mesial flat sclerite. Female genitalia (figs. 26, 27): Bursa copulatrix widely membranous; anterior sac with a few, small, pointed process. Interramal sclerite symmetrically subtriangular, with double-margined anterior edge, constricted mesally. Dimensions. / : Body length / ; head width including eyes / ; head length / ; vertex width / ; lengths of antennal segment I / , II / , III / , IV / ; rostral length / ; mesal pronotal length / ; basal pronotal width / ; width across hemelytra / ; lengths of metafemur / , tibia / , tarsus / Distribution. Japan (Ogasawara (Bonin): Chichijima & Hahajima* Isls., Ryukyus*: Okinawa & Ishigaki Isls.), Taiwan, France (introduced from Senegal with shipment of logs), Pan-tropical. Biology. Reuter (1875b) described this mirid from France, based on a specimen collected on a log imported from Senegal. In Japan, Mr. Takahashi (pers. comm.) found this species under withered grasses on Ishigaki Island, and I collected several specimens by sweeping withered branches of broad-leaved trees on Hahajima Island. The collection records suggest that F. anthocoroides has two or more generations per year. Material examined. JAPAN: Ogasawara (Bonin), Hahajima Is.: 2 2, Kuwanokiyama, 16.vii.1991, T. Yasunaga (HUES); 1, Mt. Chibusa, 15.vii.1991, T. Yasunaga (HUES); 1, same locality, 1.ii.1997, T. Matsumoto (HUES); 2, Miyukihama, 17.vii.1991, T. Ueno (HUES). Ryukyus: 1, Yona, Kunigami Vil., Okinawa Is., 19.x.1987, M. Tomokuni (NSMT); 1, Mt. Yarabudake, Ishigaki Is., 22.i.1998, K. Takahashi (HUES); 1, Takeda, Ishigaki Is., 2.vii.1998, K. Takahashi (HUES). SRI LANKA: 1, Peradeniya, xii.1910, A. Luther (det. as F. brevicornis by Poppius, ZMUF). SAMOA: 1, Malouleilei,iv.1924, Buxton & Hopkins (BMNH). FRANCE: 1, gc, Rouen (ZMUF). Fulvius dimidiatus Poppius (fig. 22) Fulvius dimidiatus Poppius, 1909: 33 (n. sp.); 1915: 50 (n. rec. from Taiwan); Schuh 1995: 27 (cat.); Kerzhner & Josifov 1999: 8 (cat.). Diagnosis. Recognized by the rather elongate body, pale apical 1/6 of antennal segment II, whitish anterior half of the corium, and a creamy yellow spot occupying the lateral apex of the corium and apex of the embolium. Closely related to F. ussuriensis Kerzhner, from which it can be distinguished by the darker antenna with the pale apex of the segment II, and wider, pale, anterior regions of the corium and clavus (fig. 22). Poppius (1909) offered further description of the external structure. The male and female genitalia were not examined as merely a single specimen 189

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, 2000 Figs Male (23-25, 28-30) and female (26-27) genitalia of Fulvius anthocoroides (23-27) and F. tagalicus (28-30). 23 & 28, right paramere; 24 & 29, left paramere; 25 & 30, vesica; 26, bursa copulatrix; 27, posterior wall of bursa copulatrix. Scale bars: 0.1 mm. of each sex was available during this study. Dimensions. : Body length 3.3; head width including eyes 0.55; head length 0.65; vertex width 0.27; lengths of antennal segment I 0.38, II 0.90, III 0.44, IV 0.51; rostral length 2.47; mesal pronotal length 0.50; basal pronotal width 1.01; width across hemelytra 1.24; lengths of metafemur 1.31, tibia 1.62, tarsus I did not measure the male at ZMUF. Distribution. Japan* (Tsushima Is.), Taiwan, Malaysia. Biology. Unknown. Discussion. At first I considered this cylapine to be conspecific with F. ussuriensis Kerzhner, 1972 described from the Russian Primorskij Territory, because many continental animals that are not distributed in Japan proper were reported from Tsushima Island. But subsequent observations on a male identified as F. dimidiatus by Poppius himself (ZMUF) and the holotype male and a paratype female of F. ussuriensis (ZMAS) revealed that the specimen from Tsushima fits the former species having hitherto been known from Taiwan and W. Malaysia. Material examined. JAPAN: 1, Kofunakoshi, Tsushima Is., Nagasaki Pref., 3.ix.1968, I. Fujiyama (NIAS). TAIWAN: 1, Formosa, Kosempo, 7.vii.1911, H. Sauter (ZMUF). Fulvius tagalicus Poppius (figs. 20, 28-30) Fulvius tagalicus Poppius, 1914: 128 (n. sp.); Schuh 1995: 29 (cat.); Kerzhner & Josifov 1999: 8 (cat.). Syntypes, PHILIPPINES: Los Baños, P. I. Baker (ZMUF) [2 examined]. Diagnosis. Recognized by the brownish body, yellowish white apex of the antennal segment II, almost unicolorous dark corium, clavus and femora, and a creamy yellow band on apex of the embolium. Poppius (1914) offered further description of the external structure. 190

9 YASUNAGA: Japanese Cylapinae Male genitalia (figs ): Parameres resembling those of F. anthocoroides, but apical part of right sensory lobe has a small process. Vesica with a distinct, semi-circularly curved, slender sclerite, and bundled sclerites. Female genitalia: Not examined; no female was available in this study. Dimensions. : Body length ; head width including eyes ; head length ; vertex width ; lengths of antennal segment I , II , III , IV ; rostral length ; mesal pronotal length ; basal pronotal width ; width across hemelytra ; lengths of metafemur , tibia , tarsus Distribution. Japan* (Ryukyus: Ishigaki Is.), Taiwan, Philippines, Indonesia. Biology. According to Dr. Takahashi, the specimen shown in fig. 20 was collected by beating withered branches. Material examined. JAPAN, Ryukyus: 1, Nagura, Ishigaki Is., 22.x.1998, K. Takahashi (HUES). TAIWAN: 1, Antsun, 14.viii.1941, H. Hasegawa (NIAS). PHILIPPINES: 2, Los Baños, P. I. Baker (ZMUF). Fulvius niveonotatus sp. n. (figs. 11, 21) Type material. Holotype, Maesato-dam, Ishigaki Is., Ryukyus, 9.v.1998, light trap, Y. Nakatani (HUES). Paratype: 1, Takeda, Ishigaki Is., 24.v.1999, K. Takahashi (HUES). Diagnosis. Easily distinguished from other congeners by the fuscous, suboval, relatively wide body, white apical 1/3 of antennal segment II with the dark extreme apex, widely white mesal part of the pronotum, and a circular, white mark occupying the apices of the corium and embolium and basal half of the cuneus (fig. 21). Description. Female: Body generally dark brown, suboval; dorsal surface shagreened, with dark, short, reclining setae except on white regions bearing pale setae. Head dark brown, almost glabrous; vertex, tylus and frons whitish brown; vertex with a weak, longitudinal, mesal sulcus. Antenna dark brown; apical 1/3 of segment II yellow, with dark apex, incrassate; lengths of segments I-IV: , , , Rostrum dark brown, reaching abdominal sternum VIII. Pronotum yellowish white mesally and fuscous laterally, with a pair of tumid, wide calli, divided by a narrow, longitudinal, mesal furrow; collar present, very narrow, about as thick as antennal segment III; scutellum unicolorous fuscous, shagreened, almost glabrous; pleura dark brown, shagreened, with yellow ostiolar peritreme. Hemelytra dark brown, shagreened, with a white, circular mark occupying posterior apices of corium and clavus and basal half of cuneus; embolium between anterior 1/3 and 1/2 pale brown. Coxae creamy yellow, except for dark brown procoxa; legs pale brown; apical part of each femur dark reddish brown; lengths of metafemur, tibia and tarsus: , , Abdomen dark brown, with ventral surface mesally pale brown. Male: Unknown. Female genitalia: Not examined. Dimensions. : Body length ; head width including eyes ; head length ; vertex width ; rostral length ; mesal pronotal length ; basal pronotal width ; width across hemelytra Etymology. From Latin niveus (snowy white) combined with notatus (mark), referring to the noticeable white marks on the dorsum; an adjective. Distribution. Japan (Ryukyus: Ishigaki Is.). Biology. Only two females of this new species were collected by light traps, and no information is available on the biology. Peritropis Uhler Peritropis Uhler, 1891: 121 (n. sp.), type species: P. saldaeformis Uhler, 1891, monotypic; Schuh 1995: 33 (cat.); Gorczyca & Eyles 1997: 226 (diag.); Kerzhner & Josifov 1999: 9 (cat.). Easily recognized by the oval and rather flat body, brown to fuscous, impunctate and roughened or shagreened dorsum speckled with numerous, small, pale spots, narrow and conical head, presence of the mesial pale ring or rounded spot on the antennal segment II (exceptionally obliterated in P. insularis), characteristic shape of the pronotum that is strongly carinate laterally, short legs, and 2-segmented tarsi with the tarsomeres II each divided by a line in consequence that the tarsi at first sight seem 3-segmented (fig. 12; Gorczyca & Eyles (1997) referred to this form as pseudotrisegmented ), and elongate right paramere usually longer than the left paramere. Gorczyca & Eyles (1997) provided further diagnosis for external characters. Twenty-nine species of Peritropis have been known from all zoogeographical regions of the world (Gorczyca 1998a, b, 1999, Gorczyca & Eyles 1997). Most of them were reported from warm temperate zone, subtropics, and tropics. In Japan, no definitive record was associated with this unique genus; a few previous records were only cited as Peritropis sp. The present study adds four new species to science and P. advena Kerzhner to the Japanese fauna. 191

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, Figs Adults (31-34) and final instar nymph (35) of Peritropis spp. 31, P. advena, female; 32, P. iriomotensis, holotype female; 33, P. takahashii, holotype male; 34 & 35, the same, female (same individual). Peritropis advena Kerzhner (figs. 12, 31, 36, 37, 45, 46) Peritropis advena Kerzhner, 1972: 279 (n. sp.); Kerzhner 1988: 790 (key); Schuh 1995: 33 (cat.); Gorczyca & Eyles 1997: 229 (list); Kerzhner & Josifov 1999: 9 (cat.). Holotype, RUSSIA: Primor je, S. Ussuri, viii.1911, A. Tsherskij (ZMAS) [examined]. Peritropis sp. Endo et al. 1998: 17 (list). Diagnosis. Recognized by the rather fuscous body, a pale, rounded, mesial spot of the antennal segment II (fig. 31), pale coxae, almost wholly dark femora, and shape of the parameres. Kerzhner (1972) provided further descriptions of the external structure and male genitalia. Male genitalia (figs 36, 37): Right paramere semicircularly curved, with a small, mesial process. Vesical spiculum weak. Female genitalia (figs. 45, 46): Bursa copulatrix widely membranous, with a pair of narrow-rimmed sclerotized rings; oviductus lateralis broad. Posterior wall of bursa copulatrix narrowed, with a continuous, slender interramal sclerite and trapezoidal mesal sclerotized plate. Dimensions. / : Body length / ; head width including eyes / ; head length / ; vertex width

11 YASUNAGA: Japanese Cylapinae Figs Male (36-44) and female (45-46) genitalia of Peritropis advena (36, 37, 45 & 46), P. hasegawai (38 & 39), P. insularis (40 & 41) and P. takahashii (42-44). 36, 38, 40 & 42, Right paramere; 37, 39, 41 & 43 left paramere; 44, vesical sclerotized appendages; 45, bursa copulatrix; 46, posterior wall of bursa copulatrix. Scale bars: 0.1 mm. 0.32/0.36; lengths of antennal segment I / , II / , III / , IV / ; rostral length / ; mesal pronotal length / ; basal pronotal width / ; width across hemelytra / ; lengths of metafemur / , tibia / , tarsus / Distribution. Japan* (Hokkaido, Honshu, Shikoku), Russian Primorskij Territory. This species has the northernmost distribution among the Japanese congeners. Biology. This species is associated with polyporaceous fungi on decaying woods or rotten logs under deciduous forests, and apparently has a univoltine life cycle. Both mature and immature forms are nocturnal, observed to have hidden between folds of a fungus, Coriolus sp., in the daytime. Material examined. 38 specimens (HUES, NSMT, ZMAS) collected between Jul. 18 and Sep. 20 from the following localities: JAPAN: Hokkaido: Kyushu Univ. Exp. Forest, Ashoro T., Tokachi; Aoyama, Tobetsu T., Ishikari; Maruyama, Sapporo C., Ishikari. Honshu: Daibosatsu Pass, Yamanashi Pref.; Ikenokuchi, Tsuruga C., Fukui Pref. Shikoku: Mt. Takashiro, Tokushima Pref.; Mt. Fukumi, Ehime Pref. RUSSIA, Primor je: Kedrovaja Pad Nature Reserve, Khasanskij Dist.; S. Ussuri. 193

12 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, 2000 Peritropis hasegawai sp. n. (figs. 38, 39) Type material. Holotype, Tsuta Onsen (hot spa), 8.vii.1953, H. Hasegawa (NIAS). Paratypes: 1 2, same data as for holotype (NIAS); 1, Yagen spa, Simokita, Aomori Pref., 20.ix.1971, M. Tomokuni (NSMT); 2, Higashi-Tanzawa, Kanagawa Pref., 25.viii.1996, T. Nakamura (HUES). Diagnosis. Closely allied to P. advena, from which this new species can be distinguished by the generally paler and larger body, a pale, continuous mesial ring of the antennal segment II, dark procoxa, and a distinct, basal, thumb-like process of the left paramere sensory lobe (fig. 39). Description. Body generally brown, oval; dorsal surface shagreened, speckled with numerous pale spots and pale portions, with sparsely distributed, short, silky setae, very similar to that of P. advena. Head speckled, glabrous; vertex with a rather broad, longitudinal, mesal sulcus. Antenna dark brown; segment II with a pale, continuous, mesial ring that is not interrupted ventrally; lengths of segments I-IV ( / ): / , / , 0.30/0.30, /0.30. Rostrum dark brown, exceeding apex of metacoxa; apices of segments II and III sometimes pale. Pronotum with somewhat notched posterior margin; apex of scutellum yellow; pleura dark brown, partly speckled with yellow. Hemelytra and membrane with numerous, pale, circular spots. Procoxa dark brown; meso- and metacoxae yellow, with dark bases; femora and tibiae dark brown; every tibia with two pale spots at base; apical half of each tibia pale with a mesial obscure annulation; tarsi pale brown; lengths of metafemur, tibia and tarsus ( / ): /1.29, / , / Abdomen dark brown. Male genitalia (figs. 38, 39): Right paramere weakly curved, with a small, mesial process; hypophysis of left paramere broad, with a distinct, thumb-like projection basally. Vesical spiculum short, weak. Female genitalia: Not examined. Dimensions. / : Body length / ; head width including eyes / ; head length / ; vertex width / ; rostral length /2.25; mesal pronotal length / ; basal pronotal width / ; width across hemelytra / Etymology. Named after Mr. Hitoshi Hasegawa who first collected this species; a noun in the genitive case. Distribution. Japan (northern Honshu). Biology. Unknown. Discussion. In the mostly tropical or subtropical genus Peritropis, this new P. hasegawai, P. advena Kerzhner and P. husseyi Knight occur in cold temperate zone. The latter is a Nearctic species, and has the northernmost distribution for the genus in Alaska (Gorczyca & Eyles 1997). Peritropis insularis sp. n. (figs. 40, 41) Type material. Holotype, Mt. Yuwandake, Amami-Oshima Is., Ryukyus, 6.v.1977, M. Sakai (NSMT). Paratypes: Ryukyus: Amami-Oshima Is.: 1, same data as for holotype (NSMT); 1, same locality & date, A. Oda (NSMT); 1, same data as for holotype, except date 5.v.1977 (NSMT); 1, Daikuma, Naze City, 1.v.1977, M. Sakai (NSMT). Okinawa Is.: 1, Hiji~Hiji Fall, Kunigami Vil., 22.x.1987, M. Sakai (NSMT). Diagnosis. Distinguished from other congeners by the remarkably shortened, small body, and unicolorous dark, incrassate antennal segment II. Description. Body generally brownish, short, ovoid; dorsal surface shagreened, speckled with numerous, small, pale, circular spots, very sparsely furnished with silky, short setae. Head pale brown, speckled; vertex with a pair of dark spots contiguous to inner margin of each eye. Antenna dark reddish brown, tumid (segments III and IV mutilated in all available specimens); segment II sometimes with a tiny, pale, circular spot ventrally; lengths of segments I and II ( / ): / , / Rostrum dark reddish brown, exceeding apex of metacoxa. Pronotum darkened along lateral margins of calli; apex of scutellum pale; pleura widely dark reddish brown, partly speckled with yellow portions. Hemelytra almost uniformly provided with pale spots; membrane dark greyish brown, with rather sparsely distributed, pale spots. Procoxa and all femora dark reddish brown; meso- and metacoxa pale brown, partly tinged with red; tibiae widely dark reddish brown basally, each with 2 basal, pale spots, pale mesial ring and pale apical part; tarsi pale brown; lengths of metafemur, tibia and tarsus ( / ): 0.90/ , 1.20/ , 0.30/ Abdomen dark reddish brown, speckled laterally. Male genitalia (figs. 40, 41): Right paramere elongate, with long, straight hypophysis. Female genitalia: Not examined. Dimensions. / : Body length ; head width including eyes 0.66/ ; head length 0.54/ ; vertex width / ; rostral length?/ ; mesal pronotal length / ; basal pronotal width 1.20/ ; width across hemelytra 1.35/ Etymology. From Latin insularis (of an island), 194

13 YASUNAGA: Japanese Cylapinae Figs Male genitalia of Peritropis iriomotensis. 47, Right paramere; 48, left paramere; 49, vesica. Scale bars: 0.1 mm. referring to its occurrence on subtropical islands of the Ryukyus; an adjective. Distribution. Japan (Ryukyus: Amami-Oshima & Okinawa Isls.). Biology. Unknown. Peritropis iriomotensis sp. n. (figs. 32, 47-49) Type material. Holotype, Mt. Sonaidake, Iriomote Is., Ryukyus, 19.iv.2000, T. Befu (HUES). Paratypes: 1, same data as for holotype (HUES); 1, Nadara Riv., Iriomote Is., 11.v.1999, K. Takahashi (HUES). Diagnosis. Allied to Taiwanese P. pusillus Poppius and other Japanese members, from which this new species is distinguished by the whitish head, long, widely pale antennal segment I, and pale base of the segment II (fig. 32). Description. Body oval, flat; dorsal surface ochreous, shagreened, speckled with dark brown regions and numerous pale spots, with sparsely distributed, silky, short, upright setae. Head widely whitish ochreous, with paired, lateral, dark spots on vertex and frons, longer than mesal pronotum; vertex somewhat darkened in, with a narrow, longitudinal, mesal sulcus; jugum, lorum and tylus partly infuscate. Antenna ochreous; segment I with a subapical, reddish brown ring continuing to a lateral stripe in, much longer than width of vertex; segment II obscured apically in, annulated with 4 brown rings in ; segments III and IV dark brown; lengths of segments I- IV ( / ): 0.45/ , 1.13/ , 0.42/ ,?/ Rostrum chocolate brown, reaching abdominal segment VIII. Pronotum dark brown, with many pale, circular spots, short, trapezoidal; calli speckled, swollen; apex of scutellum creamy white; pleura less speckled, widely reddish brown, or sanguineous, with creamy yellow margins. Inner basal corner of cuneus dark brown; membrane sombre greyish brown, with pale, circular spots. Coxae and legs whitish yellow; apex of each femur yellow, with a small, sanguineous mark; profemur widely dark brown; apical half of mesofemur dark brown; metafemur with a mesial, larger dark band and a subapical, narrower band; pro- and mesotibiae each with two obscure annulations; lengths of metafemur, tibia and tarsus ( / ): 1.17/ , 1.62/ , 0.42/ Abdomen widely dark brown, partly margined by yellow areas. Male genitalia (figs ): Parameres each with a distinct, rounded protuberance at base; right paramere not strongly curved, with a mesial teeth. Vesica widely membranous, minutely spinulate, with a slender sclerite dentate at apex. Female genitalia: Not examined. Dimensions. / : Body length 3.5/ ; head width including eyes 0.65/ ; head length 0.60/ ; vertex width 0.29/ ; rostral length 2.22/ ; mesal pronotal length 0.50/ ; basal pronotal width 1.08/ ; width across hemelytra 1.52/ Etymology. Named after the type locality, Iriomote Island; an adjective. Distribution. Japan (Ryukyus: Iriomote Is.). Biology. Unknown. Peritropis takahashii sp. n. (figs , 42-44) Type material. Holotype, Maesato-dam, Ishigaki Is., Ryukyus, 24.ii.1999, K. Takahashi (HUES). Paratypes: Ryukyus, Ishigaki Is.: 1, Mt. Banna, 16.ii.1998, M. Sugimoto (HUES); 1, same data as for holotype except for date, 8.iv.1999 (HUES). 195

14 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, 2000 Figs , Male (50-52) and female (53 & 54) genitalia of Punctifulvius kerzhneri. 50, Right paramere; 51, left paramere; 52, vesical spiculum; 53, bursa copulatrix; 54, posterior wall of bursa copulatrix. Scale bars: 0.1 mm. Diagnosis. Allied to the preceding new species, from which P. takahashii can be distinguished by the smaller size, darker head and antenna, shorter antennal segment I, narrow, pale, apical spot of scutellum with the darkened extreme apex, smaller, irregularshaped pale spots on the hemelytra, and pale basal half of the profemur. The final instar nymph is recognized by the reddish brown dorsum with many pale, small spots, dark antenna with yellowish white rings of apices of the segments I-III and mesial segment II, reddish femora with yellow bases, and clearly annulated tibiae (fig. 35). Description. Body generally dark brown, oval; dorsal surface shagreened, speckled with pale regions and numerous spots, sparsely furnished with pale, short setae. Head dark brown, speckled; longitudinal mesal sulcus on vertex wide and shallow. Antenna dark brown; segment I with pale base and apex, about as long as width of vertex; segment II with pale, mesial ring and extreme apex, in with pale spots basally; lengths of segments I-IV ( / ): 0.30/ , 0.93/0.93, 0.27/0.27, 0.31/ Rostrum dark brown, reaching abdominal sternum VII or VIII. Pronotum dark brown, usually paler mesally; scutellum widely darkened, with pale apex but extreme apex dark; pleura widely reddish brown or sanguineous. Pale spots on hemelytra small, irregularshaped, often connected. Coxae and legs creamy yellow; femoral apex narrowly pale, with sanguineous spots; apical halves of pro- and metafemora and apical 1/3 of mesofemur dark reddish brown; every tibia with 3 dark annulations; lengths of metafemur, tibia, and tarsus ( / ): 0.99/0.99, 1.38/1.38, 0.32/0.33. Abdomen dark brown. Male genitalia (figs ): Right paramere shortened in consequence that the parameres seem symmetrical. Vesica with a distinct spiculum and a V- shaped sclerite. Female genitalia: Not examined. Dimensions. / : Body length 3.3/ ; head width including eyes 0.68/0.69; head length 0.59/0.60; vertex width 0.29/0.33; rostral length 1.80/1.95; mesal pronotal length 0.48/0.51; basal pronotal width 1.12/1.16; width across hemelytra 1.32/1.35. Etymology. Named after Dr. Keiich Takahashi who enthusiastically collected and offered many cylapine specimens for this study; a noun in the genitive case. 196

15 YASUNAGA: Japanese Cylapinae Figs Female adults (55 & 57) and final instar nymphs (56 & 58) of Punctifulvius kerzhneri (55-56) and Yamatofulvius miyamotoi (57-58). Distribution. Japan (Ishigaki Is.). Biology. Unknown. Dr. Takahashi collected this species by beating branches. Punctifulvius Schmitz Punctifulvius Schmitz, 1978: 190 (n. gen.), type species: P. kerzhneri Schmitz, 1978, original designation; Kerzhner 1988: 779 (key); Schuh 1995: 35 (cat.); Kerzhner & Josifov 1999: 9 (cat.). This genus is currently represented by a single species restricted to temperate zone and cold temperate zone of the eastern Eurasia, and is recognized by the fuscous, rather shagreened dorsum provided with the very short setae, distinct punctures on the dorsum, head and thoracic pleurites, and unique shape of the parameres. The basal processes of the paramere sensory lobes and a pair of circular, minutely spinulate areas on the dorsal sac are considered as autapomorphies for Punctifulvius. The external appearance is similar to that of Bironiella Poppius, 1909 of the Indo-Pacific, but the relationships between these two genera are only superficial, as the male genital structure significantly differs from each other. 197

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, 2000 Fig. 59. Cladogram showing the phylogenetic relationships of Yamatofulvius and Punctifulvius. Numbers of apomorphic states of characters corresponding to those mentioned in the text. Punctifulvius kerzhneri Schmitz (figs ) Punctifulvius kerzhneri Schmitz, 1978: 192 (n. sp.); Kerzhner 1988: 791 (key); Schuh 1995: 35 (cat.); Yasunaga et al. 1999: 2 (list); Kerzhner & Josifov 1999: 9 (cat.). Holotype, RUSSIA: Primor je, zap. Kedrovaja Pad, 22.viii.1963, I. M. Kerzhner (ZMAS) [examined]. Diagnosis. Recognized by the generally fuscous, oval body, distinctly punctate dorsum, head and thoracic pleurites, creamy yellow coxae, and pale brown femora each with two obscure, apical annulations. A detailed description including the male genitalia was provided by Schmitz (1978). The final instar nymph is easily recognized by the principally white general coloration, darkened apical parts of the antennal segments II and III, and tibiae, pale reddish brown, somewhat speckled pronotum, mesonotum and wing pads, and the sanguineous bands on the abdominal terga (fig. 56). Male genitalia (figs ): Parameres tumid; left paramere strongly curved, with widened hypophysis and a hooked, basal process on sensory lobe; right paramere membranous inward, with a small, pointed process at base. Vesica widely membranous, with a single spiculum apically. Female genitalia (figs. 53, 54): A wide, crescent, thin-rimmed sclerotized ring present; anterior sac spherical when expanded, with a pair of circular, minutely spinulate areas. Posterior wall of bursa copulatrix with widened, trapezoidal mesal sclerotized plate and narrow interramal sclerite. Dimensions. / : Body length / ; head width including eyes / ; head length / ; vertex width / ; lengths of antennal segment I / , II / , III / , IV / ; rostral length / ; mesal pronotal length / ; basal pronotal width / ; width across hemelytra / ; lengths of metafemur / , tibia / , tarsus / Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu), Korea, Russia (S. Primor je). Biology. This common cylapine is found on polyporaceous fungi on rotten logs or bark in deciduous forests. Some individuals were also collected under evergreen broad-leaved forests. A univoltine life cycle is assumed for this species, and the late instar nymphs are found from late May to early June in southern Japan. The 3rd to final nymphs were observed to have been aggregated on blackish polyporaceous fungi at a shaded, deciduous forest in Chitose City, Hokkaido in early July. Material examined. 132 specimens (HUES, ZMAS) collected between Jun. 15 and Sep. 23 from the following localities: JAPAN: Hokkaido: Aoyama, Tobetsu T., Ishikari; Chitose C., Ishikari; Hokkaido Univ. Exp. Forest., Takaoka, Tomakomai C., Iburi; Inada T., Obihiro C., Tokachi; Kyushu Univ. Exp. Forest, Ashoro T., Tokachi. Honshu: Towadako T. & Iwasaki Vil., Aomori Pref.; Oze, Tokura, Gunma Pref.; Mt. Gomadan, Wakayama Pref.; Ryujin Vil. and Ohtoh Vil., Wakayama Pref.; Tsubakio, Wakayama Pref.; Matsubara T., Takahashi C., Okayama Pref.; Ibara C., Okayama Pref.; Kamiishi T., Hiroshima Pref. Shikoku: Befu, Monobe Vil., Kochi Pref.; Ohishi & Yoshinobu, Motoyama T., Kochi Pref. Kyushu: Mt. Kurodake, Mts. Kuju, Oita Pref.; Momiki, Izumi Vil., Kumamoto Pref. RUSSIA: Kedrovaja Pad Nature Reserve, Khasanskij Dist., S. Primor je (holotype & subsequently identified 1 1, ZMAS). Yamatofulvius gen. n. Diagnosis. Allied to Punctifulvius Schmitz, from which this new genus is distinguished by the shiny brownish dorsum, shiny, impunctate head, and long antennal segment I much longer than width of vertex. The conspicuous sexual dimorphism is exhibited in this unique genus, and the male adults have the following features quite differing from the females: Body significantly smaller; antennal segment II incrassate, more or less flattened, partly curved or winding (figs. 60, 63, 64); clypeus with short but distinct spines. Description. Body oval to elongate; dorsal surface generally brown, shining, with silky, short pubescence. Head conically projected anteriorly, rather shagreened, with sparse, short, silky setae, lacking noticeable punctures; vertex with a longitudinal, mesal sulcation, not carinate basally; clypeus with paired, short spines. Antenna long, slender; segment I long, much longer than width of vertex; segment II incrassate, partly flattened and winding; segment III 198

17 YASUNAGA: Japanese Cylapinae more or less curved. Rostrum reaching genital segment. Pronotum trapezoidal, polished, usually punctate; calli impunctate, with a distinct, longitudinal, mesal sulcation; collar present; scutellum almost or completely impunctate, somewhat shagreened. Hemelytra shining, punctate, with silky, short setae; cuneus impunctate, smooth. Meso- and metafemoral trichobothria very long, some of them even longer than diameter of each femur; tarsi long, 3-segmented. Male genitalia: Left paramere roundly and strongly curved, with broadened, flattened, apically hooked hypophysis, lacking dorsal process; right paramere flat, straight. Vesica widely membranous, with a sclerotized region laterally. Female genitalia: A single, narrow-rimmed, widened sclerotized ring present; interramal sclerite divided mesally, oblique; mesal sclerotized plate reduced, small. Type species. Yamatofulvius miyamotoi Yasunaga, new species. Etymology. Yamato (an antique name of Japan) combined with the cylapine generic name Fulvius Stål; gender masculine. Discussion. This new genus is considered to be a sister genus of Punctifulvius, from which it can be distinguished by the characters diagnosed above. The remarkable sexual dimorphism is not exhibited in Punctifulvius, and the unique shape of the antennal segment II and distinct clypeal spines in male are considered to represent autapomorphy for the present new genus. In the meantime, judging from the similarity in the general appearance of the final instar nymph and general shape of the parameres and bursa copulatrix, members of the two genera appear to be derived from the common ancestor, and to have been speciated in eastern Eurasia. Yamatofulvius is currently represented by three Japanese members inhabiting polyporaceous fungi on decaying woods or rotten logs under dark, humid evergreen broad-leaved forests in southwestern Japan. The inferred phylogeny is shown in fig. 59, on the basis of evaluation of the following ten characters and hand-calculated cladistic analysis. Numbers in the parentheses indicate character status (0 = plesiomorphic state; 1-2, apomorphic state). 1. Body size: Moderate or small [0]; elongate [1]. 2. Male antennal segment II: Slender, cylindrical [1]; incrassate, partly flattened and curved [2]. 3. Male clypeus: Smooth [0]; bearing spines [1]. 4. Pronotal calli: Punctate [0]; impunctate [1]. 5. Scutellum: Uniformly punctate [0]; sparsely punctate [1]; smooth [2]. 6. Left paramere: moderately curved [0]; strongly, roundly curved [1]. 7. Hypophysis of left paramere: Not strongly widened [0]; distinctly widened and flattened [1]. 8. Vesica: Without spiculum [0]; with one spiculum [1]; with two spiculi [2]. 9. Vesical lateral sclerotized region: Absent [0]; present [1]. 10. Mesal, wide sclerotized ring on bursa copulatrix: Absent [0]; present [1]. Yamatofulvius laevigatus sp. n. (figs. 14, 60-62, 66-69) Type material. Holotype, Hiji~Hiji Fall, Kunigami Vil., Okinawa Is., Ryukyus, 22.x.1987, M. Sakai (NSMT). Paratypes: 3 3, same data as for holotype (NSMT); 1, Okunirindo, Ohgimi Vil., Okinawa Is., 25.vi.1999, T. Yasunaga (HUES). Diagnosis. Easily recognized by the polished, dark reddish brown dorsum with the very sparsely distributed, silky pubescence and shallow punctures (figs. 60, 62), and impunctate head and thoracic pleura. The male is much smaller than female. Description. Body dark reddish brown and oval, tiny in male; dorsal surface highly polished, with very sparsely distributed, silky pubescence and shallow punctures. Head weakly shagreened, almost glabrous; vertex pale reddish brown along inner margin of eye; male clypeus with a pair of spines basally. Antenna dark brown; segment I pale reddish brown basally; male segment II thickened, compressed and curved at basal 1/3; lengths of segments I-IV ( / ): / , / , 0.75/ , / Rostrum pale brown, reaching genital segment, or slightly exceeding it in male. Pronotum weakly shagreened, with narrowly yellow posterior margin connected with a pale, posterior, mesal line, impunctate and almost glabrous; mesoscutum and scutellum fuscous, smooth; pleura dark brown, impunctate, shagreened or pruinosed except for shiny propleuron. Hemelytra shiny chocolate brown, shallowly, sparsely punctate except on cuneus, with sparsely distributed, silky, suberect pubescence; inner margin of clavus and basal margin of cuneus pale reddish brown; membrane sombre greyish brown, with fuscous veins. Coxae dark brown; procoxa and apical parts of male meso- and metacoxae creamy yellow; legs pale reddish brown; femora somewhat darker; lengths of metafemur, tibia and tarsus ( / ): / , / , / Abdomen dark reddish brown. Male genitalia (figs ): Parameres similar to those of miyamotoi; left paramere strongly curved, with broadened hypophysis. Vesica with developed lateral sclerotized area. Female genitalia (fig. 69): Sclerotized ring rather small; anterior sac with a small, sclerotized process. Interramal sclerite subtriangular. 199

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, Figs Male (60, 61, 63 & 64) and female (62 & 65) adults of Yamatofulvius laevigatus (60-62), Y. miyamotoi (63) and Y. sinuicornis (64-65). Dimensions. / : Body length / ; head width including eyes / ; head length / ; vertex width / ; rostral length / ; mesal pronotal length / ; basal pronotal width / ; width across hemelytra / Etymology. From Latin laevigatus (smooth, polished), referring to the polished, shiny dorsum; an adjective. Distribution. Japan (Ryukyus: Okinawa Is.). Biology. I collected a female of Y. laevigatus on a rotten log grown with fungi under dark, humid, evergreen broad-leaved forest. In such humid forests on Okinawa Island, a dangerous venomous snake, Trimeresurus flavoviridis (Hallowell), occurs frequently, and because of this, it is not easy to investigate the biology of this cylapine any further. 200

19 YASUNAGA: Japanese Cylapinae Yamatofulvius miyamotoi sp. n. (figs. 15, 57, 58, 63, 70-73) Type material. Holotype, Mt. Kenashi, Shinjo Vil., Okayama Pref., Honshu, 8.viii.1994, Wesco Co. Res. (HUES). Paratypes: Honshu: 1, Miyake Is., Tokyo, 8.viii.1956, H. Yamazaki (NIAS); 1 1, Mengawa, Ohtoh Vil., Wakayama Pref., 15.vi.1998, T. Yasunaga & S. Gotoh (HUES); 1, same locality, 2.vii.1998, S. Gotoh (HUES); 1 1, same locality, 14.vii.1998, S. Gotoh (HUES); 1 1, same data as for holotype (HUES); 2 1, same data, except for date, 10.viii.1994 (HUES). Shikoku: 2, Teragawa, Hongawa Vil., Kochi Pref., 11.x.1995, M. Takai (HUES); 3, Yoshinobu, Motoyama T., Kochi Pref., 14.viii.1998, M. Takai (HUES); 1, Motoyama T., 22.viii.1998, M. Takai (HUES); 1 2, Befu, Monobe Vil., Kochi Pref., 15.viii.1998, M. Takai (HUES); 1, Higashi-iya Vil., Tokushima Pref., 8.viii.1998, M. Takai (HUES); 1, Mt. Fukumi, Ehime Pref., 21.ix.1970, M. Tomokuni (NSMT). Kyushu: 1, Mt. Wakasugi, Fukuoka Pref., 22.viii.1940, H. Hasegawa (NIAS); 2 1, Mt. Kurodake, Mts. Kuju, Oita Pref., 15.ix.1985, S. Nomura (HUES); 1, Mt. Ichibusa, Kumamoto Pref., 2.viii.1974, M. Sakai (NSMT); 2 1, Mt. Osuzu, Miyazaki Pref., 21.viii.1999, M. Takai (HUES). Diagnosis. Recognized by the shiny dark reddish brown dorsum mottled with pale regions, and punctate posterior part of the pronotum, mesoscutum and thoracic pleura. The final instar nymph is similar in general appearance to that of Punctifulvius kerzhneri, from which it is easily distinguished by the more tumid body, dark antenna with a mesial, pale ring on the segment II, white pronotum, and widely darkened bases of the femora (fig. 58). Description. Body generally dark brown, elongate oval ( ) or oval ( ). Dorsal surface shiny, somewhat reddish, mottled with ochreous or brown regions, uniformly furnished with silky, short, suberect setae. Head ochreous, partly and symmetrically infuscate, with sparsely distributed, silky, short setae and fine punctures; tylus, jugum and lorum widely darkened; clypeus with two distinct spines. Antenna dark brown, with a pale ring at mesial part of segment II; antennal segment II incrassate, compressed and curved at middle; segment III slightly arched; lengths of segments I-IV ( / ): / , / , / , / Rostrum shiny dark reddish brown; segment I widely pale brown. Pronotum dark brown, with symmetrical pale portions, uniformly punctate; calli rather tumid, shagreened, impunctate; mesoscutum shagreened, punctate; scutellum fuscous, shagreened, with very sparsely distributed, weak punctures; pleura dark reddish brown, somewhat roughened, uniformly punctate; ostiolar peritreme whitish yellow. Hemelytra dark brown, shining, symmetrically mottled with pale areas; basal margin of cuneus yellowish white; membrane dark greyish brown. Coxae and legs pale brown; procoxa dark brown; femora dark reddish brown, each with a subapical, pale band; lengths of metafemur, tibia and tarsus ( / ): / , / , / Abdomen dark reddish brown. Male genitalia (figs ): Left paramere broad, circularly curved, with widened hypophysis. Vesica mostly membranous, with a narrow, lateral sclerotized area. Female genitalia (fig. 73): Sclerotized ring wide, weakly present. Posterior wall of bursa copulatrix with rectangular, mesally divided interramal sclerite. Dimensions. / : Body length / ; head width including eyes / ; head length / ; vertex width / ; rostral length / ; mesal pronotal length / ; basal pronotal width / ; width across hemelytra / Etymology. Named after Dr. Syoiti Miyamoto in honour of his great contribution to the clarification of the Japanese heteropteran fauna; a noun in the genitive case. Distribution. Japan (Honshu, Shikoku, Kyushu). Biology. This new species is collected from polyporaceous fungi on decaying woods or rotten logs under dark, humid forests. This cylapine appears to have bivoltine life cycles as the newly emerged adults are found in June and October. The adults of the second generation are assumed to hibernate. Yamatofulvius sinuicornis sp. n. (figs. 16, 64, 65, 74-77) Type material. Holotype, Shiromizu Valley, 500 m, Mt. Koshiragadake, Kumamoto Pref., Kyushu, 20.vii.1996, T. Saigusa (HUES). Paratypes: 12 6, same data as for holotype (HUES). Diagnosis. Easily recognized by the elongate body especially in (fig. 65), conspicuously winding antennal segment II (fig. 64), carinate posterolateral margin of the pronotum, and pale mesoscutum. Description. Body generally brown, elongate; dorsal surface shining, with uniformly distributed, silky, short, suberect setae and punctures on pronotum and hemelytra. Head dark brown, shagreened, elongate, slightly longer than wide; vertex narrowly pale brown along inner margin of eye; clypeus with two pairs of small spines. Antenna pale brown, partly tinged with red, long; apical part of segment II, and 201

20 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, 2000 Figs Male (66-68, 70-72, 74-76) and female (69, 73 & 77) genitalia of Yamatofulvius laevigatus (66-69), Y. miyamotoi (70-73) and Y. sinuicornis (74-77). 66, 70 & 74, Right paramere; 67, 71 & 75, left paramere; 68, 72 & 76, vesica; 69 & 77, bursa copulatrix; 73, posterior wall of bursa copulatrix. Scale bars: 0.1 mm. 202

21 YASUNAGA: Japanese Cylapinae whole III and IV dark brown; segment II conspicuously winding; segment III arched; lengths of segments I-IV ( / ): / , / , / , / Rostrum pale brown, reaching or slightly exceeding posterior margin of abdominal sternum VII. Pronotum shiny pale brown, with a pair of wide stripes mesially and darkened lateral carina; calli impunctate, somewhat shagreened; mesoscutum fuscous, with a pair of pale spots; scutellum fuscous, somewhat shagreened, impunctate; pleura dark brown, punctate, widely shagreened except shiny propleuron; ostiolar peritreme yellow. Hemelytra shiny dark brown; clavus, anterior part of corium and basal 1/3 of cuneus pale brown; membrane sombre greyish brown. Coxae and legs yellowish brown; metafemur in with two obscure annulations, in somewhat darkened apically; tarsi long; lengths of metafemur, tibia and tarsus ( / ): / , / , / Abdomen unicolorous fuscous. Male genitalia (figs ): Parameres densely furnished with long sensory setae. Vesica with a widened sclerotized area and two distinct spiculi. Female genitalia (fig. 77): Bursa copulatrix wide, with weakly sclerotized ventral labiate plate; sclerotized ring rather large. Dimensions. / : Body length / ; head width including eyes / ; head length / ; vertex width / ; rostral length / ; mesal pronotal length / ; basal pronotal width / ; width across hemelytra / Etymology. From Latin sinuo (bent, curve, wind) combined with cornis (horn, or antenna), referring to the conspicuously bent or winding male antennal segment II; an adjective. Distribution. Japan (Kyushu). Biology. This unique cylapine was collected on a huge rotten log covered with polyporaceous fungi under shaded, evergreen broad-leaved forest. Bothriomirini Kirkaldy, 1906 Members of this tribe are easily recognized by the tumid, oval body, shiny, heavily punctate dorsum usually with dense pubescence, and shortened antenna and rostrum. The Bothriomirini is considered to form a monophyletic group, judging from the similar external characters exhibited by its current members (e.g., Bakeriola Bergroth, Dashymenia Poppius, Dashymeniella Poppius). However, no further study on these genera has been made since the original descriptions, and much broader survey on the poorly documented characters in the genitalia is required to correctly redefine these closely related genera. Bothriomiris Kirkaldy Bothriomiris Kirkaldy, 1902: 270 (n. gen.), type species: Bothriomiris marmoratus Kirkaldy, 1902 (= Capsus dissimulans Walker, 1873), original designation; Schuh 1995: 20 (cat.); Kerzhner & Josifov 1999: 10 (cat.). This genus is easily recognized by the fuscous, oval, tumid body, distinct pubescence and punctures on the shiny dorsum, shortened rostrum not exceeding apex of the mesocoxa, more or less arched scutellum, and mesial knob-like projection of the ostiolar peritreme. Bothriomiris has hitherto been known by four species from Taiwan, New Guinea and the Oriental Region. The present study adds two new species, representing the northernmost distributional record for the genus. Bothriomiris capillosus sp. n. (figs. 80, 81) Type material. Holotype, Shirahama, Iriomote Is., Ryukyus, Japan, 7.vi.1998, K. Takahashi (HUES). Paratype: 1, Fenchihu, 1,400 m, Nantow Hsien, Taiwan, 7-8.vii.1965, T. Nakane (NSMT). Diagnosis. Allied to Taiwanese B. lugubris (Poppius, 1915), from which it can be distinguished by the shorter antennal segment I, entirely fuscous scutellum, polished, impunctate epimeron and episternum, a distinct mesial knob of the ostiolar peritreme, a clear, white, rounded mark at the posterior apex of the each corium (fig. 80), almost entirely dark greyish brown membrane without mesial pale regions, and unicolorous reddish brown procoxa. Description. Female: Body generally fuscous, suboval; dorsal surface shiny blackish brown, with a pair of white spots on apical part of hemelytra, densely punctate, furnished with silky, long, upright pubescence. Head brown, subvertical, with silky, upright setae; vertex distinctly carinate basally; tylus infuscate. Antenna dark brown; segment I paler, much shorter than width of vertex; segment II subequal to width of head including eyes; segments III and IV mutilated; lengths of segments I and II: , Rostrum shiny chocolate brown, shortened, reaching apex of procoxa; segments I and IV very short. Pronotum tumid, roundly declivous laterally; calli swollen, heart-shaped, with punctures smaller and sparser than those on disk; scutellum swollen; propleuron punctate; epimeron and episternum highly polished, only with sparsely distributed, fine punctures; ostiolar peritreme dark greyish brown, with a distinct, shiny, mesial knob. Hemelytra rounded laterally; mesial apex of corium with a creamy white spot; membrane dark 203

22 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 143, Figs Male (78) and female (80-82) adults, and 4th instar nymph (79) of Bothriomiris spp. 78 & 79, B. gotohi; 80, B. capillosus, holotype; 81, the same, paratype, left lateral view; 82, a specimen identical with B. lugubris from Taiwan. greyish brown, semitransparent, narrowly pale along inner margin of cuneus, with densely distributed, short setae. Coxae and legs dark reddish brown; lengths of metafemur, tibia, and tarsus: , , Abdomen dark reddish brown, ventrally with densely distributed, brown, straight, suberect setae. Female genitalia: Not examined. Dimensions. : Body length ; head width including eyes ; head length ; head height ; vertex width ; rostral length ; mesal pronotal length ; basal pronotal width ; width across hemelytra Etymology. From Latin capillosus (hairy), referring to the densely pubescent body; an adjective. 204