Heteroptera, Miridae, Cylapinae), with Descriptions of Five New Species1

Second report on the Japanese cylapine plant bugs (Heteroptera, Miridae, Cylapinae), with descriptions of five new species1

T. YASUNAGA & S. MIYAMOTO

Abstract: Additional records and descriptions of Japanese cylapines, or fungal-inhabiting plant bugs, are provided. The following five new species are described from southwestern subtropical islands of Japan: Bothriomiris ernsti nov.sp., B. yakushima nov.sp., Euchilofulvius lepidopterus nov.sp., Fulvius nakatai nov.sp., and Yamatofulvius minutus nov.sp. Clearly detected functions of two autapomorphies previously assumed for Yamatofulvius are documented and discussed. Keys are provided to distinguish every Bothriomiris, Ful- vius and Yamatofulvius species occurring in Japan. A checklist of Japanese cylapines is given, with updated distributional records. Male and/or female genitalic structures are described for the first time for Bothriomiris capillosus and Fulvius tagalicus.

Key words: checklist, Cylapinae, Heteroptera, Japan, mating behavior, Miridae, new record, new species.

Introduction phies assumed for Yamatofulvius by YASUNA- GA (2000) was demonstrated through field Cylapinae, one of basal plant bug sub- observations in subtropical broadleaved for- families, consists predominantly of fungal- est on Okinawa Island. The functions ap- inhabiting members, most of which live on parently are correlated with the specialized fungous-covered rotten logs or decaying mating behavior within Yamatofulvius woods in dark, humid forests, particularly in species, as mentioned in the generic Discus- the warm temperate zone, subtropics and sion. tropics. Due to the specialized habitat and The present paper documents five new remarkable agility, hand collecting is the species. Brief comments and/or revised keys best method for capture. Cylapines usually to Japanese species are also provided for the do not take flight but quickly run away four genera, Bothriomiris, Euchilofulvius, Ful- when disturbed. Only some members of Ful- vius and Yamatofulvius, containing these vius are known to be occasionally attracted new species. The function of the autapo- to light at night. morphies for Yamatofulvius is discussed. The In a revisional work by the first author male of B. capillosus is reported for the first (YASUNAGA 2000), 16 species in six genera time. Descriptions of male and/or female were documented from Japan, with a key to genitalic structures are also for the first time all known Japanese genera and species, and provided for Bothriomiris capillosus and Ful- a cladistic analysis for the newly established vius tagalicus. genus, Yamatofulvius. However, subsequent All measurements in the text are in mil- efforts by TY and several enthusiastic col- limeters (mm). Synonymic lists for known leagues have revealed the presence of five taxa are not always cited here as comprehen- hitherto unknown species. In addition, evi- sive catalogues are now available (SCHUH dence for the function of two autapomor- 1995; KERZHNER & JOSIFOV 1999). Terminol-

Denisia 19, zugleich Kataloge 1This paper was written to celebrate the 70th birthday of Dr. Ernst Heiss, a father-like mentor of the first author as der OÖ. Landesmuseen well as a long time friend of the second author. Neue Serie 50 (2006), 721–735

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ogy of the female genitalia mainly follows Peritropis insularis YASUNAGA 2000 – Japan KULLENBERG (1947) and YASUNAGA (2000), (Amami-Oshima & Okinawa Isls.). but some terms are tentatively used in this Peritropis iriomotensis YASUNAGA 2000 – Japan paper since cylapine female genital structures (*Ishigaki & Iriomote Isls.). have not been sufficiently studied. Habitus Peritropis takahasii YASUNAGA 2000 – Japan (Ishi- and genitalic structure of each species are gaki Is.). shown by digital photographic images. Genus Punctifulvius SCHMITZ 1978

Punctifulvius kerzhneri SCHMITZ 1978 – Japan Checklist of Japanese Cylapinae (Hokkaido, Honshu, Shikoku, Kyushu), Korea, Russia (S. Primorskij Territory). [New distributional record indicated by an aster- isk (*)] Genus Yamatofulvius YASUNAGA 2000 Yamatofulvius laevigatus YASUNAGA 2000 – Japan Tribe Bothriomirini KIRKALDY 1906 (Okinawa Is.). Genus Bothriomiris KIRKALDY 1902 Yamatofulvius minutus nov.sp. – Japan (Amami- Oshima Is.). Bothriomiris capillosus YASUNAGA 2000 – Distrib- Yamatofulvius miyamotoi YASUNAGA 2000 – Japan ution: Japan (Ishigaki* & Iriomote Isls.). (Honshu, Shikoku, Kyushu, *Yakushima Is.). Bothriomiris ernsti nov.sp. – Japan (Ishigaki Is.). Yamatofulvius sinuicornis YASUNAGA 2000 – Japan Bothriomiris gotohi YASUNAGA 2000 – Japan (Kyushu, *Yakushima & *Amami-Oshima Isls.). (Honshu (S. Kii Peninsula), Shikoku, Kyushu, Miyake & Okinawa Isls.). Genus Bothriomiris KIRKALDY Bothriomiris yakushima nov.sp. – Japan (Yakushi- Bothriomiris KIRKALDY 1902: 270 (n.gen.), type ma Is.). species: Bothriomiris marmoratus KIRKALDY 1902 Tribe Cylapini KIRKALDY 1906 (= Capsus dissimulans WALKER 1873), original designation; SCHUH 1995: 20 (cat.); KERZHNER & Genus Cylapomorpha POPPIUS 1914 JOSIFOV 1999: 10 (cat.); YASUNAGA 2000: 203 Cylapomorpha michikoae YASUNAGA 2000 – Japan (diag.); 2001: 124 (diag.). (Honshu (S. Kii Peninsula), Kyushu, Okinawa & Diagnosis. Recognized by the generally Ishigaki Isls.), Taiwan. fuscous, oval, tumid body, distinct pubescence and punctures on the shiny dorsum, Tribe Fulviini UHLER 1886 wide vertex with a narrow, basal transverse Genus Euchilofulvius Poppius 1909 carina, shortened rostrum not exceeding Euchilofulvius lepidopterus nov.sp. – Japan (Oki- apex of the mesocoxa, more or less arched nawa Is.). scutellum, mesial knob-like projection of the ostiolar peritreme, and hook of the left Genus Fulvius STÅL 1862 paramere base. Fulvius anthocoroides (REUTER 1875) – Japan (Ogasawara (Bonin) Isls., Ishigaki Is., Iriomote Discussion. On the basis of the antennal Is.); widely distributed in Old World subtropics structure and male genitalia, four Japanese and tropics, and Neotropical Region; a record Bothriomiris species can be divided into two (original description) from France was the result subgroups. One group contains B. capillosus of accidental introduction with the shipment of and B. yakushima nov.sp., characterized by logs from Senegal. the shortened, broadened antennal segment Fulvius dimidiatus POPPIUS 1914 – Japan (W. Hon- shu*, Tsushima Is.), Taiwan, Peninsular Malaysia. I and II, arched scutellum, highly polished Fulvius nakatai nov.sp. – Japan (Okinawa Is.). epimeron and episternum, developed Fulvius niveonotatus YASUNAGA 2000 – Japan thumb-like process on the male genital seg- (Ishigaki Is.). ment, and highly sclerotized vesical ap- Fulvius tagalicus POPPIUS 1914 – Japan (Ishigaki pendages and female bursa copulatrix, Is.), Taiwan. whereas in the other two species, the anten-

Genus Peritropis UHLER 1891 nal segments I and II are longer and slen- derer, the scutellum is rather flat, the Peritropis advena KERZHNER 1972 – Japan (Hokkaido, Honshu, Shikoku), Russian Far East epimeron and episternum are distinctly (Primorskij Territory). punctate, the male genital segment lacks Peritropis hasegawai YASUNAGA 2000 – Japan (N. such process, the vesica is not heavily scle- Honshu) rotized, only with a slender, tubular exten-

722 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at sion, and the female bursa copulatrix is the corium, developed thumb-like processes widely membranous. Further subdivision on the male genital segment, and highly may be required for the classification of the sclerotized appendages of the vesica. YA- members currently placed in Bothriomiris. SUNAGA (2000) provided further diagnosis However, there are several related genera and description of the adult female. (e.g., Bakeriola BERGROTH, Dasymenia POP- Description. Male: Similar in overall ap- PIUS, Dasymeniella POPPIUS), the genitalic pearance to female, but body narrower and structures of which have been poorly exam- an apical, white semicircular spot at apex of ined or reported. A comprehensive revision, corium smaller. Lengths of antennal seg- including critical examination of the geni- ments I-IV: 0.51, 1.85, 0.68, 0.80. Lengths talia, is desirable to correctly redefine all re- of metafemur, tibia and tarsus: 2.23, 2.67, lated genera. 0.82. Male genitalia (Fig. 3A-C): Genital segment (A) with a pair of developed Key to Japanese species thumb-like processes, the right lateral one [Revised couplet 3 of a key provided by smaller. Left paramere (B) semicircularly curved, with weak basal hook. Vesica (C) YASUNAGA (2000: 206)] with highly sclerotized appendages, and 1 Antennal segment I long, more than half ventrally with a flattened, claw-like sclerite. length of segment II; epimeron and Female genitalia (Fig. 4A, B): Bursa episternum distinctly punctate as in dor- copulatrix (A, B) with asymmetrical sclero- sum ...... 2 tized structure; posterior wall of bursae al- – Antennal segment I short, equal to or less most membranous, without distinct sclero- than 1/3 length of segment II; epimeron tized area. and episternum highly polished, only = with weak minute punctures ...... 3 Measurements. : Body length 6.6; head width including eyes 1.66; vertex width 2 Body broader, with maximum width more 0.81; basal pronotal width 2.81; and width than 3.5 mm (=)/ 4 mm (Y); rostrum across hemelytron 3.28. shorter, not reaching apex of mesocoxa; apical pale spot on corium orange yellow, Biology. See the following species. clear; Ishigaki Island . . . B. ernsti nov.sp. Discussion. This species is characterized – Body narrower than 2.5 mm (=)/ 3 mm by having a pair of developed thumb-like (Y); rostrum longer, reaching apex of processes on the male genital segment and mesocoxa; apical pale spot on corium ob- highly sclerotized appendages on the vesica. scure whitish brown; north of Okinawa Based on similarity of the male genitalia, Island ...... B. gotohi the closest relative of B. capillosus appears to 3 Rostrum reaching but not exceeding apex be B. yakushima nov.sp. of procoxa; subtropical inhabitant of Ishi- gaki and Iriomote Island . . . B. capillosus Bothriomiris ernsti nov.sp. – Rostrum exceeding apex of procoxa, (Figs 1C, 2B, G, 3D-F; 4C, D) reaching middle of mesocoxa; warm tem- Diagnosis. Recognized by the broad, perate zone inhabitant of Yakushima Is- rounded body, rough, distinct punctures on land ...... B. yakushima nov.sp. the dorsum, clear orange yellow spot on the apex of corium, and broad exocorial area, in Bothriomiris capillosus YASUNAGA addition to being currently known only (Figs 1A, B, 2A, F, 3A-C, 4A, B) from Ishigaki Island. Bothriomiris capillosus YASUNAGA 2000: 203 (n.sp.); 2001: 124 (diag.). Description. Body generally shiny black, Diagnosis. Recognized by the heavily ovoid, broad; dorsal surface punctate, with pubescent dorsum, short antennal segment uniformly distributed, silvery, semierect se- I, rather thickened antennal segment II, en- tae. Head rather brownish, shining, almost tirely fuscous scutellum, polished, impunc- impunctate; vertex wide; frons shallowly tate epimeron and episternum, a clear, and transversely wrinkled. Antenna dark white, rounded mark at the posterior apex of brown, generally short and slender; segment

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Fig. 1: Japanese Bothriomiris species. I and basal part of II slightly paler; lengths of thoracic pleura shiny fuscous, punctate as in (A) B. capillosus, male; (B) ditto, female; segments I-IV (=/Y): 1.03/ 1.03, 1.78/ 1.74, pronotum; ostiolar peritreme dark grayish (C) B. ernsti nov.sp., female; (D) B. gotohi, male; (E) fungal rotten log, on which B. 1.01/ 0.94, 1.19/ 1.23. Rostrum dark reddish brown, pruinosed, with medial knob pru- ernsti nov.sp. (F) and B. capillosus brown, short, exceeding apex of procoxa but inosed, relatively flattened. Hemelytron (G) co-occurred, found in subtropical forest not reaching apex of mesocoxa. Pronotum wide; a subtriangular spot at apex of corium on Ishigaki Island of the Ryukyus, Japan (late September, 2002). heavily punctured, roundly carinate or clear orange yellow; exocorial area broad- keeled laterally; calli swollen, finely punc- ened and flattened; membrane somber gray- tate, somewhat convex medially; scutellum ish brown, semitransparent, with whitish flat, punctate, with an orange yellow apex; veins. Coxa and leg shiny chocolate brown;

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Fig. 2: Japanese Bothriomiris species. (A, F) B. capillosus; (B, G) B. ernsti nov.sp.; (C, H) B. gotohi; (D, H, I) B. yakushima nov.sp.; (A-E) dorsal habitus; (F-I) left lateral view; (A-D, F-I) male; (E) female.

tarsus paler basally; lengths of metafemur, 5== 15YY, same locality as for holotype, tibia and tarsus (=/Y): 2.34/ 2.67, 2.81/ 22.vii.2002, T. Nakata; 3==, same data as for 3.28, 0.96/ 1.07. Abdomen shiny dark holotype. brown. Male genitalia (Fig. 3D-F): Genital Etymology. Named in honor of Dr. Ernst segment (D) smooth, without thumb-like Heiss; a noun in genitive case. process. Left paramere (E) rather strongly Biology. This new species co-occurred curved, with a blunt hooked process basally with B. capillosus on Ishigaki Island. TY ob- and a distinct, pointed process at apex; right served both adults and immatures of these paramere tiny (D). Vesical tubular process two species on heavily fungous-covered rot- short, almost straight (F). Female genitalia ten logs in dark, humid subtropical rain for- (Fig. 4C, D): Bursa copulatrix (D) and pos- est in September, 2002 (Fig. 1E-G). Habitat terior wall of bursae (C) mostly membra- preferences of the two species are similar to nous, lacking sclerotized structures. those of B. gotohi. Although collection Measurements. =/Y: Body length 7.0/ records suggest that B. capillosus has at least 8.0; head width including eyes 1.98/ 2.13; a bivoltine life cycle, additional data are vertex width 1.01/ 1.07; basal pronotal needed to determine the actual number of width 2.83/ 3.28; and width across heme- generations. lytron 3.49/ 4.18. Discussion. The new species is morpho- Holotype =, JAPAN: Ryukyus, Ishigaki Is., Mt. logically most similar to B. gotohi, as both Nosokodake, 29.ix.2002, T. Yasunaga. Paratypes: species have a similar, tubular sclerotized ex-

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Fig. 3: Male genitalia of Bothriomiris species. (A-C) B. capillosus; (D-F) B. ernsti nov.sp.; (G) B. gotohi; (H-J) B. yakushima nov.sp.; (A, D, H) genital segment with parameres in dorsal view; (B, E, G, I) left paramere; (C, F, J) vesica.

tension on the vesica. Bothriomiris ernsti legs, relatively long rostrum exceeding apex nov.sp. can be distinguished from B. gotohi of the procoxa, polished epimeron and epis- by the characters mentioned in the above ternum, a single thumb-like process on left key and Diagnosis. These two species are side of the male genital segment, and weak- now shown to be allopatric. ly curved left paramere with a subapical process, in addition to being currently Bothriomiris yakushima nov.sp. known only from Yakushima Island, south of (Figs 2D, E, I, 3H-J) Kyushu. Diagnosis. Recognized by the generally Description. Body blackish brown, oval; pale brown head, antennal segment I and dorsal surface shining, distinctly punctate,

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Fig. 4: Female genitalia of Bothriomiris species. (A-B) B. capillosus; (C-D) B. ernsti nov.sp.; (A) Bursa copulatrix in right lateral view; (B) ditto, in left lateral view; (C) posterior wall of bursae in frontal view; (D) vaginal vestibule in frontal view.

with uniformly distributed, silky, suberect mere (I) not strongly curved, with a subapi- pubescence. Head pale brown, shining, al- cal, small, pointed process. Vesica (J) with most impunctate, with sparsely distributed, distinct sclerites; ventral triangular sclerite silky, erect setae; vertex narrowly margined short. basally; buccula chocolate brown. Antenna Measurements. =/Y: Body length 7.2/ fuscous, except for segment I pale brown; 7.7; head width including eyes 1.80/ 1.76; segment II cylindrical; segments III and IV vertex width 0.82/ 0.82; basal pronotal filiform; lengths of segments I-IV (=/Y): width 3.04/ 3.12; and width across heme- 0.60/ 0.62, 2.07/ 1.91, 1.03/ 0.94, 1.01/ ?. lytron 3.28/ 3.63. Rostrum brown, reaching middle of mesocoxa; apical part of segment IV darker. Holotype =, JAPAN: Kagoshima Pref., Yakushi- Pronotum continuously margined laterally; ma Is., Ohkawa Timber Road, 8.viii.2002, T. scutellum relatively arched; pleura polished, Nakata. Paratype: 1Y, same data as for holotype. Etymology. Named after Yakushima Is- only with sparsely distributed, minute punc- land, type locality of this new species, a tures, except for propleuron punctate as in well-known World Natural Heritage site; a pronotum; ostiolar peritreme grayish brown, noun in apposition. shagreened, with shiny, chocolate brown medial knob. Hemelytron with a white spot Discussion. This new species apparently at apex of corium; = embolium pale brown; is a close relative of B. capillosus, from which membrane smoky brown. Coxa and leg pale it can be distinguished by the characters di- brown; tarsus darkened; lengths of metafe- agnosed above. Both species are allopatri- mur, tibia and tarsus (=/Y): 2.67/ 2.67, cally distributed on these islands. Between 2.97/ 3.28, 0.92/ 0.98. Abdomen dark Yakushima and Ishigaki islands are several chocolate brown. Male genitalia (Fig. 3H- other islands that have a rich fauna (e.g., J): Genital segment (H) with a single Amami-Oshima, Tokunoshima, Tokara), thumb-like process on its left side; left para- and additional congeners may occur there.

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Fig. 5: Japanese Euchilofulvius and Fulvius species. (A-E) E. lepidopterus nov.sp.; (F-H) F. nakatai nov.sp.; (A, C-E) male; (B, F-H) female; (A-B, D, F, H) dorsal habitus; (C, G) left lateral view; (E) scutellum and hemelytron.

Further investigations on these islands are shagreened dorsum, shortened antennal seg- required to clarify distribution patterns of ments III and IV, subapically toothed claws, Japanese Bothriomiris. and the short hypophysis and rather tumid sensory lobe of the left paramere. Further di- Genus Euchilofulvius POPPIUS agnostic characters and redescription were provided by GORCZYCA (1999). Euchilofulvius POPPIUS 1909: 28, type species: Eu- chilofulvius tibialis POPPIUS 1909: 28, monotypic; Discussion. This Indo-Pacific genus pre- GORCZYCA 1999: 3 [as Eucholofulvius]. viously included five tropical species and is Diagnosis. Recognized by the scale-like divided into two subgenera, Euchilofulvius setae covering the rather flattened body, s.str. and Lepidofulvius. Our discovery of a

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species belonging to the nominotypical sub- Description. Body generally somber pale Fig. 6: Male (A-G, I-N) and female (H) genus from Okinawa Island of Japan repre- brown, elongate oval; dorsal surface sha- genitalia of Euchilofulvius (D-H), Fulvius (A- C) and Yamatofulvius (I-N) species. (A-C) F. sents the northernmost distributional record greened or pruinosed, with uniformly or in dimidiatus; (D-H) E. lepidopterus nov.sp.; (I- for the genus. part densely distributed, silvery, scale-like L) Y. minutus nov.sp.; (M-N) Y. miyamotoi; setae. Head shagreened, longer than wide, (A, E, K, N) right paramere; (B, F, J, M) left Euchilofulvius lepidopterus nov.sp. with densely distributed, silvery, scale-like paramere; (C, G, L) vesica; (D, I) male genital segment in dorsal view. (Figs 5A-E, 6D-H) setae; eye small; vertex somewhat swollen, Diagnosis. Recognized by the somber with a shallow, mesal depression. Antenna pale brown general coloration, shagreened somber brown; segments I and II tinged with dorsum with uniformly distributed, silvery, red; segments III and IV short, filiform; scale-like setae, and short rostrum not ex- lengths of segments I-IV (=/Y): 0.50/ 0.58, ceeding apex of the metacoxa. 1.15/ 1.05, 0.38/ 0.35, 0.50/ 0.45. Rostrum

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pale brown, reaching but not exceeding antennal segment II that is less than the apex of metacoxa. Pronotum trapezoidal, basal pronotal width, and the hemelytron carinate laterally, with densely distributed, that lacks any red portion. silvery, scale-like setae; calli swollen, with a Biology. Unknown. longitudinal, mesal sulcation; collar about as broad as antennal segment II; mesoscu- Genus Fulvius REUTER tum dark brown, tumid; scutellum dark Fulvius STÅL 1862: 322 (n.gen.), type species: brown, flat, shallowly and transversely ru- Fulvius anthocoroides STÅL 1862: 322, monotypic; gose, with yellowish apex; pleura generally SCHUH 1995: 25 (cat.); KERZHNER & JOSIFOV chocolate brown, shagreened, with sparsely 1999: 8 (cat.); YASUNAGA 2000: 189 (diag.); distributed, silvery, scale-like setae. Heme- 2001: 125 (diag.). lytron somber brown, shagreened, with uni- Diagnosis. Recognized by the small, formly distributed, silvery, scale-like setae; rather slender, anthocorid-like body with median part of corium and basal margin of the principally dark coloration, usually sha- cuneus pale brown; membrane somber gray- greened or pruinosed, almost impunctate ish brown, with a pale, semitransparent spot dorsum, long head, short antenna, long ros- contiguous to apex of cuneus. Coxa and leg trum sometimes reaching the genital seg- brown, partly tinged with red; trochanter, ment, narrow, slender pronotum, 2-seg- apical part of tibia and tarsus yellowish mented tarsi, and flattened, apically hooked brown; lengths of metafemur, tibia and tar- sensory lobe of the left paramere. sus (=/Y): 1.35/ 1.38, 1.83/ 1.94, 0.35/ 0.36. Discussion. The included species and in- Abdomen reddish brown, relatively shining. terspecific variation suggest that Fulvius re- Male genitalia (Fig. 6, D-G): Genital seg- quires subdivisions. Although some workers ment (D) trapezoidal. Left paramere (F) C- have proposed several species groups for Ful- shaped, with tumid sensory lobe and short, vius, we refrain from using any of them until flattened hypophysis. Right paramere (E) with an inner apical tooth at base of short the genus is revised on a worldwide basis. hypophysis. Vesica (G) weakly sclerotized Definitive treatments are beyond the scope but lacking noticeable bundles of sclerites. of this study. Female genitalia (Fig. 6H): Posterior wall of The habitus of most members of Fulvius bursae with weakly sclerotized interramal is suggestive of certain anthocorid species. sclerites. Such anthocorid-like body shape may imply Measurements. =/Y: Body length 3.65/ a convergence in food habits of Fulvius 3.96; head width including eyes 0.69/ 0.71; species: predaceous rather than myco- vertex width 0.30/ 0.35; mesal pronotal phagous. Recently, our colleague Dr. K. length including collar 0.69/ 0.88; basal Takahashi reported immatures of Fulvius an- pronotal width 1.14/ 1.35; and width across thocoroides from a henhouse in Ogasawara hemelytron 1.31/ 1.64. (Bonin) Islands; they developed into adults when provided with larvae of Tribolium con- Holotype =, JAPAN: Ryukyus, Okinawa Is., Na- fusum JAQUELIN DU VAL (Coleoptera) as go City, Katsuyama, 17.vii.2002, T. Nakata. prey (TAKAHASHI et al. 2001). Further, most Paratypes: 46 specimens same data as for holo- cylapines are presumably nocturnal but do type, and 9 specimens, Okinawa Is., Ohgimi Vil., not migrate far from their typical habitats. Shanashiro, 18.vii.2002, T. Nakata. In Japan, only Fulvius species were collected Etymology. From Greek, lepidopterus (= by light traps, and no other cylapines have scaled wing), referring to the scale-like setae been attracted to light. It seems to us that of this new species; an adjective. the biology of Fulvius differs from that of Discussion. No other congener has been other cylapines, as suggested by WHEELER found in neighboring regions of Japan. In (2001). the key provided by GORCZYCA (1999), the present species may key to E. antennatus GORCZYCA from South Sulawesi, Indonesia, but E. lepidopterus nov.sp. can be distinguished from it by the larger body, shorter

Key to Japanese species dles of sclerites on the vesica are also found in two other Japanese species, F. antho- [Revised couplets 6-8 of a key provided coroides REUTER and F. tagalicus POPPIUS (see by YASUNAGA (2000: 207)] YASUNAGA 2000). 1 Head white; pronotum fuscous, with a Fulvius nakatai nov.sp. (Fig. 5F-H) widely white mesial region ...... F. niveonotatus Diagnosis. Recognized by the shiny fuscous head and pronotum, light apical 3/5 of – Head and pronotum entirely dark brown the antennal segments II, long, filiform an- ...... 2 tennal segments III and IV, and four pale 2 Clavus unicolorously somber brown . . . . spots on the hemelytron...... F. tagalicus Description. Female: Body elongate, – Clavus brown or fuscous, with paler or parallel-sided; dorsal surface shining, dark whitish basal region ...... 3 brown, with uniformly distributed, simple, 3 Head and pronotum polished, shining; silky, semierect pubescence. Head shiny fus- apical 3/5 of antennal segment II pale . . cous, a little longer than width including ...... F. nakatai nov.sp. eyes. Antenna dark brown; apical 3/5 of seg- – Head and pronotum more or less sha- ment II brownish white; segments III and IV greened or pruinosed, not distinctly shin- long, filiform; lengths of segments I-IV: ing; less than apical 1/3 of antennal seg- 0.56, 1.13, 0.78, 1.13. Rostrum shiny dark ment II pale ...... 4 brown, elongate, reaching genital segment; 4 Head as long as, or shorter than width in- segments II and III paler. Pronotum shiny cluding eyes; apical 1/3 of antennal seg- fuscous, with a longitudinal, mesal sulcus ment II pale ...... F. anthocoroides between roundly swollen calli; collar about – Head longer than width including eyes; as thick as base of antennal segment III; apical 1/6 of antennal segment II pale . . scutellum shiny fuscous, slightly arched; ...... F. dimidiatus pleura chocolate brown; ostiolar peritreme dark grayish brown. Hemelytra dark brown; Fulvius dimidiatus POPPIUS (Fig 6 A-C) apices of corium and embolium, and base Fulvius dimidiatus POPPIUS 1909: 33 (n.sp.); and apex of clavus creamy brown, partly SCHUH 1995: 27 (cat.); KERZHNER & JOSIFOV tinged with red. Coxa, and trochanter pale 1999: 8 (cat.); YASUNAGA 2000: 189 (diag.); brown; femur chocolate brown; tibia and 2001: 126 (diag.). tarsus yellowish brown; lengths of metafe- Diagnosis. Recognized by the rather mur, tibia and tarsus: 2.00, 2.25, 0.81. Ab- elongate body, pale apical 1/6 of antennal domen uniformly dark brown. Male: Un- segment II, whitish anterior half of the cori- known. um, and a creamy yellow spot occupying the lateral apex of the corium and apex of the Measurements. Y: Body length 4.3; embolium. A detailed description of the ex- head width including eyes 0.75; vertex ternal structure was given by POPPIUS width 0.34; mesal pronotal length including (1909). collar 0.65; basal pronotal width 1.25; and width across hemelytron 1.69. Description. Male genitalia (Fig. 6A-C): Parameres distinctly pubescent; right para- Holotype Y, Japan, Ryukyus, Okinawa Is., Nago City, Katsuyama, 17.vii.2002, T. Nakata. mere sensory lobe with a small process at base of slender, apically tapered hypophysis Etymology. Named after Mr. Tadafumi (A); left paramere curved at right angle, Nakata, who collected and offered us many with a protuberance near base of sensory cylapine specimens including the holotype lobe (B). Vesica with bundled slender scle- of this new species; a noun in genitive case. rites subapically (C). Discussion. We consider this new species Discussion. An examination of male most closely allied to Fulvius sauteri, de- genital structure suggests this species is a scribed from Taiwan by POPPIUS (1915). nominotypical member of Fulvius. The sim- Based on comparison with the holotype of F. ilar general shape of the parameres and bun- sauteri, deposited in the Zoological Museum,

Fig. 7: Yamatofulvius species. (A) Y. minutus nov.sp., female; (B) Y. sinuicornis, female; (C-K) Y. laevigatus; (C-D) male adult; (E) female adult; (F) male final instar nymph; (G-I) male adult riding female final instar nymph; (J) male adult continuing to ride female just after emerging; (K) male copulating with still teneral female.

Fig. 8: Dorsal habitus (A-B) and left lateral view (C-D) of Yamatofulvius minutus nov.sp. (A-C) and Y. miyamotoi (D); (A, C-D) male; (B) female. Abbreviation: CS, clypeal spine.

University of Helsinki, Finland, F. nakatai 8C, D) and the curved second antennal seg- nov.sp. can be distinguished by the highly ments (Fig. 7D, 8A), until the females polished head and pronotum, the uniformly emerge. Soon after a female would emerge, dark antennal segment I, the darker anten- the male began to copulate with the still nal segment II that is darkened at nearly whitish, teneral female (Fig. 7J, K). There- basal half, and the triangular, much narrow- fore, the clypeal spines, and curved or bent er apical pale spot on the exocorium. antennal segments II, found only in adult Biology. Unknown. males of Yamatofulvius, can be regarded as specialized modifications for the peculiar Genus Yamatofulvius YASUNAGA mating behavior. Yamatofulvius YASUNAGA 2000: 198 (n.gen.); No male adults were observed to ride fe- 2001: 128 (diag.). male final instars when the population den- Diagnosis. Characterized by synapomor- sity was low. The adult males presumably phies such as the brown general coloration, need to reserve immature females only when more or less flattened and curved male an- populations are larger. Unfortunately, we tennal segment II, male clypeal spines, impunctate pronotal calli, and lateral sclero- have not seen aggregations of other Yamato- tized region of the vesica. Adult males are fulvius species because they are not so com- significantly smaller than females. monly found and their habitats are usually restricted. Further observations are required Discussion. YASUNAGA (2000) assumed to determine whether all Yamatofulvius the four above-mentioned synapomorphies members exhibit similar mating behavior. for Yamatofulvius. During subsequent field- Only one other species of Orthotylinae, Fal- work carried out by Dr. K. Takahashi and conia intermedia (DISTANT), in the diverse TY in Okinawa Island of the Ryukyus, a pe- family Miridae appears to exhibit similar culiar mating behavior was observed for Y. mating behavior (WHEELER 2001). laevigatus (Fig. 7C-K), which implies actual functions of two of the synapomorphies. Both mature and immature forms of this species sometimes aggregate on rotten logs or decaying wood covered with polypora- ceous fungi in dark, humid subtropical forests. In such aggregations adult males were frequently observed to ride final instar females (Fig. 7G-I). These males apparently occupy their mates so as not to be robbed by other males. The riding males hold young females tightly, using the clypeal spines (Fig.

Key to Japanese species dark chestnut brown, with paler spots at basal 1/3 of corium and inner corner of [Revised couplets 14-15 of a key pro- clavus (near apex of scutellum); cuneus red- vided by YASUNAGA (2000: 207)] dish brown, basally with a creamy yellow, se- 1 Pronotum and thoracic pleura polished, mi-circular mark; membrane smoky brown, almost impunctate ...... Y. laevigatus with a semitransparent spot contiguous to apex of cuneus. Coxa and leg dark brown, – Posterior part of pronotum and thoracic partly tinged with red; apex of coxa and pleura punctate ...... 2 trochanter yellowish; apex of each femur, 2 Body oval or suboval; head shorter than subapical ring of metafemur and each tarsus width including eyes; pronotum not cari- pale brown; lengths of metafemur, tibia and nate laterally; mesoscutum with punc- tarsus (=/Y): 1.50/ 1.66, 1.76/ 2.06, 0.63/ tures ...... 3 0.73. Abdomen dark castaneous brown. – Body elongate; head about as long as, or Male genitalia (Fig. 5I-L): Genital segment slightly longer than wide; pronotum with (I) somewhat bent right. Left paramere (J) distinctly carinate posterolateral margin; C-shaped; right paramere (K) not strongly mesoscutum impunctate . . . Y. sinuicornis excavated apically. Vesica (L) with a horn- 3 Antennal segment II dark, with a pale like sclerite and a flat, apically toothed scle- ring medially ...... Y. miyamotoi rite. – Antennal segment II almost uniformly Measurements. =/Y: Body length 3.6/ brownish, without distinct pale parts . . . 4.3; head width including eyes 0.85/ 0.94; ...... Y. minutus nov.sp. vertex width 0.40/ 0.44; mesal pronotal Yamatofulvius minutus nov.sp. length including collar 0.73/ 0.88; basal (Figs 5I-L, 7A, 8A-C) pronotal width 1.28/ 1.50; and width across hemelytron 1.56/ 2.03. Diagnosis. Recognized by the small size, dark castaneous brown general coloration, Holotype =, JAPAN: Ryukyus, Amami-Oshima unicolorously dark brown, short, and slight- Is., Kinsakubaru, 11.ix.2002, T. Nakata. == ly bent antennal segment II that lacks a no- Paratypes: 3 1Y, same data as for holotype. ticeable mesial pale ring, and unique vesica Etymology. From Latin, minutus (= with the apically toothed sclerotized ap- small, tiny), referring to comparatively pendage. small size of this new species; an adjective. Description. Body generally dark casta- Discussion. This new species appears to neous brown, oval but a little elongate; dor- be sister to Y. miyamotoi YASUNAGA, from sal surface shining, punctate, with uniform- which it is easily distinguished by the small- ly distributed, simple, short setae. Head er size, uniformly dark antennal segment II, slightly shagreened, with sparsely distrib- and possession of the characteristic vesical uted, simple setae; vertex with a pale spot sclerites. These species are currently consid- mesally; frons sparsely punctate; = clypeus ered allopatric. with a small spine at middle. Antenna dark Biology. Unknown. brown; segment I reddish brown; segment II weakly curved or bent in =, slightly longer than basal width of pronotum; segments III Acknowledgements and IV filiform; lengths of segments I-IV We are grateful to the following individ- (=/Y): 0.65/ 0.56, 1.13/ 1.06, 0.73/ 0.75, uals for offering invaluable specimens 0.88/ 1.00. Rostrum shiny chocolate brown, and/or supporting fieldwork: Mr. M. Takai reaching base of genital segment. Pronotum (Nankoku City, Kochi); Dr. K. Takahashi slightly pale at basal angles, uniformly punc- (Tsukuba City, Ibaraki); Mr. T. Nakata (JIR- tate, not carinate laterally, with a longitudi- CAS, Subtropical Station, Ishigaki City, nal, mesal sulcus between calli; collar a lit- Okinawa). Mr. Takai was kind enough to tle broader than antennal segment III; provide superb photographs of living indi- mesoscutum and scutellum punctate; pleura viduals of Yamatofulvius minutus and Y. dark reddish brown, densely punctate; ostio- sinuicornis. We are also much indebted to lar peritreme creamy yellow. Hemelytron Dr. I.M. Kerzhner (Zoological Institute,

Russian Academy of Sciences, St. Peters- SCHUH R.T. (1995): Plant bugs of the world (Insecta: burg) and Dr. L. Hulden (Zoological Muse- Heteroptera: Miridae). Systematic catalog, distributions, host list and bibliography. — um, University of Helsinki, Finland) for The New York Entomological Society, New their kind permission to access related ma- York: xii + 1-1329. [Continuing updated cata- terial including type series. Thanks are ex- logue available on webpage: http://re- tended to Dr. W. Rabitsch for kindly invit- search.amnh.org/pbi/catalog/] ing us to this commemorative volume, and STÅL C. (1862): Hemiptera mexicana enumeravit also to Dr. A.G. Wheeler Jr. (Clemson speciesque novas descripsit (continuatio). — Univ., SC, USA) and Dr. J. Gorczyca (Sile- Stettin. Entomol. Zeit. 23: 289-325. sian University, Katowice, Poland) for re- TAKAHASHI K., OHBAYASHI T. & N. SOTA (2001): Investi- viewing the manuscript and providing in- gation of stored-product insect pests and their natural enemies in Chichijima Island, valuable comments and suggestions. Ogasawara (Bonin), Japan. — Jpn. J. Ent. (N. S.) 3: 97-103. [In Japanese, with English ab- Zusammenfassung stract] WHEELER A.G. Jr. (2001): Biology of the Plant Bugs Neue Funde und Beschreibungen japa- (Hemiptera: Miridae). Pests, Predators, Op- nischer Cylapinae (an Pilzen lebende portunists. — Cornell University Press, Ithaca Weichwanzen) werden mitgeteilt. Die fol- & London: xv + 1-507. genden fünf neuen Arten werden von den YASUNAGA T. (2000): The mirid subfamily Cylapinae südwestlichen subtropischen Inseln Japans (Heteroptera: Miridae), or fungal inhabiting beschrieben: Bothriomiris ernsti nov.sp., B. plant bugs in Japan. — Tijdschr. v. Entomol. 143: 183-209. yakushima nov.sp., Euchilofulvius lepidopterus nov.sp., Fulvius nakatai nov.sp. und Yamato- YASUNAGA T. (2001): Family Miridae HAHN, plant bugs. — In: YASUNAGA T., TAKAI M. & T. KAWA- fulvius minutus nov.sp. Die Funktion zweier SAWA (Eds), A Field Guide to Japanese Bugs II. – bereits früher vermuteter – autapomorpher Terrestrial Heteropterans. — Zenkoku Noson Merkmale wird für Yamatofulvius dokumen- Kyoiku Kyokai Publ. Co. Ltd., Tokyo: 112-276. tiert und diskutiert. Bestimmungsschlüssel [In Japanese] werden für alle japanischen Arten der Gat- tungen Bothriomiris, Fulvius und Yamatoful- vius zur Verfügung gestellt. Eine Checkliste der japanischen Cylapinae mit aktualisier- ten Verbreitungsangaben wird präsentiert. Männliche und/oder weibliche Genital- merkmale werden erstmals für Bothriomiris capillosus und Fulvius tagalicus beschrieben.

References

KERZHNER I.M. & M. JOSIFOV (1999): Miridae HAHN, 1833. — In: AUKEMA B. & C. RIEGER (Eds), Cata- logue of the Heteroptera of the Palaearctic Region. Vol. 3, Cimicomorpha II. Netherlands Entomol. Soc., Amsterdam: 1-576. Address of the Authors: KIRKALDY G.W. (1902): Memoir upon the Rhyn- Dr. Tomohide YASUNAGA chotal family Capsidae Auctt. — Entomolo- gist 35: 243-272, pls. 5-6. Natural History Museum Institute of Science & Technology KULLENBERG B. (1947): Über Morphologie und Funk- tion des Kopulationsapparats der Capsiden Tribhuvan University (TU) und Nabiden. — Zool. Bidr. Från Upps. 24: Swayambhu, Kathmandu, Nepal 217-418, pls. 1-22. c/o JICA (Japan International Cooperation POPPIUS B. (1909): Zur Kenntnis der Miriden-Unter- Agency) Nepal Office familie Cylapina REUT. — Acta Soc. Sci. fenn. Helsingf. 37 (4): 1-46. P. O. Box 450, Kathmandu, Nepal E-Mail: [email protected] POPPIUS B. (1915): H. Sauter’s Formosa=Ausbeute: Nabidae, Anthocoridae, Termatophylidae, Dr. Syôiti MIYAMOTO Miridae, Isometopidae und Ceratocombidae (Hemiptera). — Archiv für Naturg. 80A (8): 1- Sawaraku, Iikura 3-34-4 80 (1914). Fukuoka 814-0161, Japan

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