Anthocoris confusus Reuter, 1884

Yamada, Kazutaka & Yasunaga, Tomohide, 2021, The tribe Anthocorini in Japan (Hemiptera: Anthocoridae): descriptions of new species, review of distribution and bionomics, Acta Entomologica Musei Nationalis Pragae 61 (2), pp. 375-426 : 394-396

Anthocoris confusus Reuter, 1884

( Figs 7C–D , 9B , 10C–D , 12B , 13D–F , 15B , 16D , 20 )

Anthocoris confusus Reuter, 1884:194 . Lectotype (Pඣඋංർൺඋඍ 1970:738): J, France, Vosges (MNHN).

Anthocoris confusus var. funestus Horváth, 1896:329 . Holotype:unsexed, Germany, Bavaria, Neu-Ulm (HNHM).

Anthocoris confusus ab. aterrimus Gravestein, 1945: 122 . Holotype: ♀, The Netherlands, Prov. Limburg, Geulem (= Geulhem) (ZMAN → RMNH). Junior primary homonym of Anthocoris aterrimus J. Sahlberg, 1878 , unavailable name.

Anthocoris confusus f. gravesteini Stichel, 1958: 22. New name for aterrimus Gravestein, 1945 , unavailable name.

Anthocoris confusus f. pallipes Péricart,1972: 135. Holotype:J, Georgia, Antsal-Or River , Signakhi Distr. ( ZMAS).

Anthocoris confusus (selected references): Eൽඐൺඋൽඌ (1890): 236 (in key, redescription); LൾඍHංൾඋඋඒ & Sൾඏൾඋංඇ (1896): 243 (listed, taxonomic status); LංඇൽൻൾඋG (1927): 21 (listed); Hൺඅൻൾඋඍ (1935): 257 (records, life cycle, habitat); Pඋඈർඍൾඋ (1946): 77 (record); Sൺඇൽඌ (1957): 300 (description of immature stages); SඍංർHൾඅ (1958): 21–22 (in key, diagnosis); Hංඎඋൺ (1959): 6, pls. 3–4 (record, distribution, figures); SඍංർHൾඅ (1960): 357 (listed, distribution); Aඇൽൾඋඌඈඇ (1962): 75 (bionomics); Aඇൽൾඋඌඈඇ & Kൾඅඍඈඇ (1963): 439 (record); Jඈඌංൿඈඏ (1964): 85 (record); KൾඋඓHඇൾඋ (1964): 698 (in key); Hංඅඅ (1965): 245 (bionomics); Sൾඋඏൺൽൾං (1967):215 (distribution); Jඈඌංൿඈඏ (1970): 839 (distribution); Pඣඋංർൺඋඍ (1972): 114, 134–135 (in key, redescription, figures); Cඈඎඅංൺඇඈඌ & Oඌඌංൺඇඇංඅඌඌඈඇ (1976):150 (listed, distribution); Eඅඈඏ (1976): 370–371, 376 (in key, figure, diagnosis); Eඅඈඏ & KൾඋඓHඇൾඋ (1977): 209 (record, bionomics); Hൾංඌඌ (1977): 42–43 (record, distribution, habitat); Hඈൻൾඋඅൺඇൽඍ (1977):65 (listed); Kൾඅඍඈඇ (1978): 35, 43, 71, 82, 92 (in key, redescription,figures, photo); Fඈඋൽ (1979): 55 (listed, distribution); Dඋඈඌඈඉඈඎඅඈඌ (1980): 171 (listed); Gංൽൺඒൺඍඈඏ et al. (1980): 90 (habitat); Öඇൽൾඋ (1982): 17, 79, 92–93 (in key, redescription, prey, habitat, distribution); Rඈ෡ർൺ & Pඈඉඈඏ (1982): 129 (listed); Sർඎൽൽൾඋ (1986): 64 (record); Hൾඇඋඒ (1988): 14 (catalogue, distribution); KൾඋඓHඇൾඋ (1988): 773 (in key, figure); Bඈඌආൺඇඌ & Pඣඋංർൺඋඍ (1989): 41 (distribution, records); Mංඒൺආඈඍඈ & YൺඌඎඇൺGൺ (1989): 165 (listed, distribution); Pඣඋංർൺඋඍ & Hൺඅඉൾඋංඇ (1989): 92 (listed); ZHൾඇG & Bඎ (1990): 24 (listed); RൾංർHඅංඇG & Gൾඋൾඇൽ (1994): 282 (listed); VංඇඈKඎඋඈඏ & KൺඇඒඎKඈඏൺ (1995): 12 (listed); Pඣඋංർൺඋඍ (1996): 111 (catalogue, distribution); PඎඍඌHKඈඏ & PඎඍඌHKඈඏ (1996): 16 (distribution); Cൺඋൺඉൾඓඓൺ (1997): 201 (records, distribution); LඎKൺඌHඎK (1997): 9 (distribution); Cඈඎඅං-

ൺඇඈඌ (1998): 25 (listed, distribution); Pඋඈඍංම (1998): 42 (records, distribution, habitat); Kඈඇൽඈඋඈඌඒ (1999): 138 (listed); Nඈඇඇൺංඓൺൻ (1999): 55 (distribution); Hඎൺ (2000): 198 (listed, distribution); Lൺඍඍංඇ (2000): 613 (bionomics); Bඎ & ZHൾඇG (2001): 120, 155 (in key, redescription, figures); YൺඌඎඇൺGൺ (2001b): pl. 85, 281 (photo, diagnosis, habitat, phenology); Hඈൿൿආൺඇඇ & Mൾඅൻൾඋ (2003): 243 (distribution); GඈGൺඅൺ (2004): 248 (records); Cඈඎඅංൺඇඈඌ (2005): 18 (record, distribution); Lൾඐංඌ et al. (2005): 61 (taxonomic history, record); RൺൻංඍඌർH (2005): 35 (listed); Aൻൽ- Rൺൻඈඎ & GHൺHൺඋං (2006): 43 (listed as predator of whitefly, associated plant); Öඇൽൾඋ et al. (2006): 28 (listed, distribution); WൺർHආൺඇඇ et al. (2006): 186 (distribution, habitat, photo); Bඋඒඃൺ & Kආൾඇඍ (2007): 32 (record); Kൾ & Bඎ (2007):91–92 (figure, description of female genitalia); Hඈඋඍඈඇ & Lൾඐංඌ (2009): 610 (summary of N. American records); GHൺHൺඋං et al. (2009): 46 (catalogue, distribution); Sදൽൾඋආൺඇ & Rංඇඍൺඅൺ (2009): 18 (record); Nൾංආඈඋඈඏൾඍඌ (2010): 21 (records); VංඇඈKඎඋඈඏ et al. (2010): 56 (catalogue, distribution); GHൺHൺඋං et al. (2011): 4 (record); FൺඋඓൺඇൾH et al. (2011): 338 (record, photo); HൾർKආൺඇඇ & BඅදർHඅංඇGൾඋ (2011):108 (records); Hඈඋඍඈඇ & Lൾඐංඌ (2011):1262, 1265, 1268–1270, 1273 (description of male and female genitalia, photo, figures); Kආൾඇඍ & Kൾඃඏൺඅ (2011): 38 (listed); MൺඅൾඇඈඏඌKප et al. (2011): 144 (record, prey, habitat, associated plant); Rංඇඍൺඅൺ & Rංඇඇൾ (2011): 170 (diagnosis, habitat, distribution, photo); Cඎඉඉൾඇ (2012):159 (listed); Kආൾඇඍ & BൺŃൺෞ (2012): 509 (record, distribution);AඎKൾආൺ et al. (2013a):84 (catalogue, distribution); EඌൾඇൻൾKඈඏൺ (2013): 56–57 (habitat, distribution); Hൺඈ & Mൺ (2013): 36 (listed); Jൾඋංඇංම- Pඋඈൽൺඇඈඏංම & Pඋඈඍංම (2013): 171 (record, distribution, prey); JඎඇG et al. (2013): 422 (record, diagnosis, distribution, habitat); AඅൻඋൾർHඍ et al. (2015): 24 (listed); Bൺඅඅൺඅ & Yൺආൺൽൺ (2016): 191 (record); Pඋඈඍංම (2016): 366 (listed); Yൺආൺൽൺ et al. (2016): 422 (catalogue, distribution); JඎඇG & Lൾൾ (2017): 36, 70 (diagnosis, redescription, associated plant, distribution, photo, figure); Oඌඍඈඏൺඇ et al. (2017): 453 (distribution, habitat, prey); AඎKൾආൺ et al. (2018): 22 (record); Bൺඅඅൺඅ et al. (2018):209 (catalogue, distribution);MඈඋKൾඅ et al. (2018): 229 (record); Mඈඎඅൾඍ et al. (2018): 135, 139 (listed, distribution, in key); Rඒൺඇ (2018): 26 (listed); VංඇඈKඎඋඈඏ (2019): 21 (record); Gඈඎඅൺ et al. (2020): 12 (listed); VංඇඈKඎඋඈඏ (2020): 39 (catalogue, distribution).

Material examined. JAPAN: HඈKKൺංൽඈ: 1♀, Horokanai-cho, Uen-nai

Dam, 20.viii.2000,T.Yasunaga ( TYCN); 1J ( Figs 12B , 13D–F ), Sappo-

ro-shi, Hitsujigaoka, 15.v.1992, collector unknown ( TKPM). HඈඇඌHඎ:

Tochigi Pref.: 1 J ( Figs 7C–D ), Nikko-shi, Yumoto, 16.xi.2006, S.

Maehara ( TKPM); 1 ♀, same locality, 18.vi.2010, S. Maehara ( TKPM);

1 J, same locality, 4.ix.2017, S. Maehara ( TKPM). Yamanashi Pref.: 1

♀ ( Fig. 15B ), Hokuto-shi, Kanayamadaira, 27.ix.2007, T. Ban ( TKPM).

– FRANCE: 1 ♀, Forêt de Fontainebleau, 20.viii.1966, J. Péricart

( MNHN). THE NETHERLANDS: 2 JJ, Leiden near ‘Naturalis’,

19.vii.2006, K. Yamada ( TKPM). INDIA: HංආൺർHൺඅ PඋൺൽൾඌH: 2 JJ

(one in Fig. 9B , one in Figs 10C–D ) 2 ♀♀, unknown date and collector

( TKPM).

Differential diagnosis. Recognized by the following combination of characters: head and pronotum ( Figs 7C–D ) blackish brown; antennal segments I, III, and IV blackish brown, segment II pale brown or yellowish brown and darkened at base and apex ( Figs 7C–D ); hemelytra ( Figs 7C , 16D ) more or less brown or reddish brown at base, a strip along the inner embolioum dull or pruinose, the outer portion of the embolium shiny; membrane ( Fig. 7C ) smoky dark brown, with three whitish markings on basal and innermost portions and area behind apex of cuneus; tibiae ( Figs 7C–D ) mostly pale brown; ostiolar peritreme ( Fig. 9B ) straight, almost same width throughout, curved slightly anteriad at apex; abdominal sternum II ( Fig. 12B ) with a pair of small triangle-shaped membranous areas along posterior margin. Most similar in appearance to Palaearctic A. minki Dohrn, 1860 and European A. simulans Reuter, 1884 , but distinguished from both by antennal segment II mostly pale brown and darkened at base and apex (in A. minki , pale yellow with fuscous tinge at apex; in A. simulans , uniformly darkened) and each tibia mostly pale brown (in A. minki , reddish brown to blackish brown; in A. simulans , blackish brown). The shape of the paramere is also most similar to both above mentioned species by the possession of a subapical projection. However, these species might not be clearly separated by relying solely on the form of the paramere.

Redescription. Male genitalia ( Figs 10C–D , 13D–F ): Pygophore ( Fig. 13D ) turbinate, longer than wide, covered with 8–12 long, stout setae intermixed with short, suberect setae along outer margin and on posteroventral surface, of which the longest are approximately half length of pygophore; mid-dorsal surface very hirsute with suberect setae; paramere ( Figs 10D , 13E–F ) lamellate, slender, acute at apex, subapically with a sharply defined projection, with a few very short, erect setae on median portion, strongly curved at base in posterolateral view; longitudinal groove lacking.

Female genitalia ( Fig. 15B ): Copulatory tube fused on middle of intersegmental membrane between sterna VII and VIII, approximately 1.5 mm in length, very thin and almost same width from base to apex (junction to trunk of conductive tissue), with few distinct twists at apex and base; trunk of conductive tissue pronounced.

Measurements [mm; JJ (n = 2) / ♀♀ (n = 3)]. Body length 4.50–4.60 / 4.33–4.63; head length (excl. neck) 0.53–0.56 / 0.50–0.56; head width across eyes 0.57–0.58 / 0.55–0.57; vertex width 0.30–0.31 / 0.30–0.33; length of antennal segments I – 0.18–0.19 / 0.16–0.20, II – 0.56–0.57 / 0.53–0.55, III – 0.32–0.35 / 0.31–0.33, and IV – 0.40 / 0.36–0.39; length of labial segments II – 0.15–0.17 / 0.15–0.16, III – 0.56–0.57 / 0.52–0.59, and IV – 0.30–0.31 / 0.30–0.34; anterior pronotal width 0.48–0.50 / 0.46–0.48; mesal pronotal length 0.60–0.63 / 0.60–0.62; basal pronotal width 1.32–1.35 / 1.30–1.35; length of embolial margin 1.40–1.44 / 1.35–1.48; length of cuneal margin 0.82–0.84 / 0.73–0.80; maximum width across hemelytra 1.45–1.49 / 1.45–1.55.

Bionomics. Anthocoris confusus is found on various broad-leaved trees, including Acer , Betula , Alnus , Salix , Populus , Quercus , Fagus , Carpinus, Ulmu s, Fraxinus , and Tilia (Pඣඋංർൺඋඍ 1972). This species was collected from Fagus grandifolia Ehrhart and Tilia sp. in North America (Lൺඍඍංඇ 2000, Hඈඋඍඈඇ & Lൾඐංඌ 2009) and in Japan, from Cerasus sargentii (Rehder) H. Ohba var. sargentii , Betula platyphylla Sukaczev , and Alnus japonica (Thunb.) Steud. (YൺඌඎඇൺGൺ 2001b). A specimen from Tochigi Prefecture, Japan, was collected from under the bark of Betula ermanii Cham. in November (Satoshi Maehara, pers. comm.). In South Korea (Seoul), hibernating specimens were discovered under the bark of Zelkova serrata (JඎඇG et al. 2013).

Distribution. Japan: Hokkaido (Hංඎඋൺ 1959); Honshu*: Tochigi, Yamanashi. We present the first record from Honshu ( Fig. 20 ); however, it is rare. The species is widely distributed throughout the Palaearctic Region (Pඣඋංർൺඋඍ 1996) and is also introduced into North America (Aඇൽൾ- උඌඈඇ & Kൾඅඍඈඇ 1963, Hඈඋඍඈඇ & Lൾඐංඌ 2009).

Summary of known records: ASIA: Armenia (Pඣඋං- ർൺඋඍ 1972). Azerbaijan (Gංൽൺඒൺඍඈඏ et al. 1980, Pඣඋංർൺඋඍ 1996). China: Nei Mongol, Hebei, Sichuan (ZHൾඇG & Bඎ 1990, Bඎ & ZHൾඇG 2001). Georgia (Pඣඋංർൺඋඍ 1972). India: Himachal Pradesh (Bൺඅඅൺඅ & Yൺආൺൽൺ 2016, Bൺඅඅൺඅ et al. 2018). Iran (Lංඇඇൺඏඎඈඋං & Hඈඌඌൾංඇං 2000, Oඌඍඈඏൺඇ et al. 2017). Israel (Pඣඋංർൺඋඍ & Hൺඅඉൾඋංඇ 1989). Kazakhstan: Asian part (Pඣඋංർൺඋඍ 1972, EඌൾඇൻൾKඈඏൺ 2013). Korea: Chungcheongnam-do, Seoul (JඎඇG et al. 2013, JඎඇG & Lൾൾ 2017). Mongolia: (Pඣඋංർൺඋඍ 1972, Eඅඈඏ & KൾඋඓHඇൾඋ 1977). Russia: West and East Siberia, Far East (LංඇൽൻൾඋG 1927, VංඇඈKඎඋඈඏ et al. 2010). Turkey: Asian part (Pඣඋංർൺඋඍ 1996, Öඇൽൾඋ et al. 2006). Turkmenistan (Pඣඋංർൺඋඍ 1996). EUROPE: Albania (Jඈඌංൿඈඏ 1970). Andorra (Pඣඋංർൺඋඍ 1996). Austria (Hൾංඌඌ 1977, RൺൻංඍඌർH 2005). Belgium (SඍංർHൾඅ 1958, Bඈඌආൺඇඌ & Pඣඋංർൺඋඍ 1989). Bosnia and Herzegovina (Pඣඋංർൺඋඍ 1972). Bulgaria (SඍංർHൾඅ 1958, Jඈඌංൿඈඏ 1964). Belarus (Pඣඋංർൺඋඍ 1972, LඎKൺඌHඎK 1997). Croatia (HඈඋඏගඍH 1896, Pඣඋං- ർൺඋඍ 1972). Czech Republic (Hඈൻൾඋඅൺඇൽඍ 1977, Kආൾඇඍ & BൺŃൺෞ 2012). Denmark (Pඣඋංർൺඋඍ 1972). Estonia (Cඈඎඅංൺඇඈඌ 2005). Finland (Rൾඎඍൾඋ 1884, AඅൻඋൾർHඍ et al. 2015). France (Rൾඎඍൾඋ 1884, Pඣඋංർൺඋඍ 1972). Germany (HඈඋඏගඍH 1896, Hඈൿൿආൺඇඇ & Mൾඅൻൾඋ 2003). Greece (SඍංർHൾඅ 1958, Dඋඈඌඈඉඈඎඅඈඌ 1980). Hungary (Pඣඋංർൺඋඍ 1972, Kඈඇൽඈඋඈඌඒ 1999). Ireland (Hൺඅൻൾඋඍ 1935). Italy (Rൾඎඍൾඋ 1884, Sൾඋඏൺൽൾං 1967). Latvia (Pඣ- උංർൺඋඍ 1996). Lithuania (Sදൽൾඋආൺඇ & Rංඇඍൺඅൺ 2009). Luxembourg (Pඣඋංർൺඋඍ 1996). Moldavia (Pඣඋංർൺඋඍ 1996). Montenegro (SඍංർHൾඅ 1958, Pඋඈඍංම 2016). The Netherlands (Rൾඎඍൾඋ 1884, Gඋൺඏൾඌඍൾංඇ 1945, AඎKൾආൺ et al. 2018). North Macedonia (WൺGඇൾඋ 1962). Norway (SඍංർHൾඅ 1958, Cඈඎඅංൺඇඈඌ 1998). Poland (SඍංർHൾඅ 1958). Romania (Pඣඋංർൺඋඍ 1972, Rඈ෡ർൺ & Pඈඉඈඏ 1982). Russia: South, Central and North European Territories (SඍංർHൾඅ 1958, Nൾංආඈඋඈඏൾඍඌ 2010). Serbia (Pඋඈඍංම 1998). Slovakia (Hඈൻൾඋඅൺඇൽඍ 1977, Pඣඋංർൺඋඍ1996, Bඋඒඃൺ & Kආൾඇඍ 2007). Slovenia (GඈGൺඅൺ 2004). Spain (SඍංർHൾඅ 1958, Gඈඎඅൺ et al. 2020). Sweden (SඍංർHൾඅ 1958, Pඣඋංർൺඋඍ 1972). Switzerland (SඍංർHൾඅ 1958, HൾർKආൺඇඇ & BඅදർHඅංඇGൾඋ 2011). Ukraine (PඎඍඌHKඈඏ & PඎඍඌHKඈඏ 1996). United Kingdom (Eൽඐൺඋൽඌ 1890, Sൺඇൽඌ 1957). NORTH AFRICA: Tunisia (Pඣඋංർൺඋඍ 1972). NORTH AMERICA: Canada: Nova Scotia, Newfoundland, British Columbia (Pඋඈർඍൾඋ 1946, Hඈඋඍඈඇ & Lൾඐංඌ 2009). USA: Maine, New York, North Carolina, Michigan, Tennessee, Washington (Lൾඐංඌ et al. 2005, Hඈඋඍඈඇ & Lൾඐංඌ 2009).

Figures

Fig. 7. Habitus of Anthocoris spp., dorsal (A, C, E, G, I, K) and lateral (B, D, F, H, J, L) views. A–B – A. chibi Hiura, 1959, male; C–D – A. confusus Reuter, 1884, male; E–F – A. japonicus Poppius, 1909, male; G–H – A. kalopanacis Kerzhner, 1977, female; I–J – A. nemoralis (Fabricius, 1794), male; K–L – A. takahashii Hiura, 1959, male. Scale bars: 1.0 mm.

Fig. 9. Scanning electron micrographs of ostiolar peritreme and evaporatorium of Anthocoris spp, left lateroventral view. A – A. chibi Hiura, 1959, female; B – A. confusus Reuter, 1884, male (from India); C – A. japonicus Poppius, 1909, female; D – A. kalopanacis Kerzhner, 1977, female; E – A. takahashii Hiura, 1959, female; F – A. miyamotoi Hiura, 1959, female; G – A. venustus sp. nov., paratype, male. Abbreviations: eva ‒ evaporatorium; mf ‒ median furrow; op ‒ ostiolar peritreme; sca ‒ supracoxal area.

Fig. 10. Scanning electron micrographs of male genitalia of Anthocoris spp. A–B – A. chibi Hiura, 1959; C–D – A. confusus Reuter, 1884 (from India); E–F – A. japonicus Poppius, 1909.A, C, E – pygophore with paramere, dorsal (A, C) and laterodorsal (E) views; B, D, F – paramere, dorsal (B, D) and laterodorsal (F) views. Abbreviations: end ‒ endosoma; prm ‒ paramere; pyg ‒ pygophore.

Fig. 12. Abdominal sterna II–III of Anthocoris spp., male, ventral view (setae omitted). A – A. chibi Hiura, 1959; B – A. confusus Reuter, 1884; C – A. japonicus Poppius, 1909; D – A. nemoralis (Fabricius, 1794); E – A. takahashii Hiura, 1959; F – A. miyamotoi Hiura, 1959; G – A. venustus sp. nov., paratype. Abbreviations: ma ‒ membranous area; st2–3 ‒ abdominal sternum II to III. Scale bars: 0.2 mm.

Fig. 13. Male genitalia of Anthocoris spp. A–C – A. chibi Hiura, 1959; D–F – A. confusus Reuter, 1884; G–I – A. japonicus Poppius, 1909; J–L – A. nemoralis (Fabricius, 1794); M–O – A. takahashii Hiura, 1959. A, D, G, J, M – pygophore with paramere (ejaculatory bulb omitted), dorsal view; B–C, E–F, H–I, K–L, N–O – paramere, two different views. Abbreviations: prm ‒ paramere; pyg ‒ pygophore. Scale bars: 0.1 mm.

Fig. 15. Female genitalia of Anthocoris spp., dorsal view.A – A. chibi Hiura, 1959; B – A. confusus Reuter, 1884; C – A. japonicus Poppius, 1909; D – A. nemoralis (Fabricius,1794); E – A.takahashii Hiura, 1959; F – A. miyamotoi Hiura, 1959; G – A. venustus sp. nov., paratype.Abbreviations: ct ‒ copulatory tube; ism ‒ intersegmental membrane; sp ‒ sperm pouch; t ‒ trunk of conductive tissue (C–D, trunk of conductive tissue dissolved). Scale bars: 0.1 mm.

Fig. 16. Habitus images of living individuals of Japanese Anthocorini spp. A – Acompocoris brevirostris Kerzhner, 1979, male; B – same, female; C – Tetraphleps aterrima (J. Sahlberg, 1878); D – Anthocoris confusus Reuter, 1884; E – A. takahashii Hiura, 1959; F – A. venustus sp. nov. (courtesy of M. Takai).

Fig. 20. Distribution of Anthocoris chibi Hiura, 1959 (circles) and A. confusus Reuter, 1884 (squares) in Japan. Filled symbols represent records based on specimens examined by us, open symbols represent literature records.