Elasmucha dorsalis ( Jakovlev, 1876 )

Tsai, Jing-Fu & Rédei, Dávid, 2015, The identity of Acanthosoma vicinum, with proposal of a new genus and species level synonymy (Hemiptera: Heteroptera: Acanthosomatidae), Zootaxa 3936 (3), pp. 375-386 : 377-381

Elasmucha dorsalis ( Jakovlev, 1876)

Acanthosoma haematogaster (non Burmeister, 1835): Uhler (1860: 224). Misidentification ( Uhler 1861: 286).

Acanthosoma vicinum Uhler, 1861: 286 . Lectotype ( Kumar 1974: 47) (♂): China: Hong Kong; USNM! New subjective synonym.

Elasmostethus dorsalis Jakovlev, 1876: 106 . Lectotype (Kerzhner 1972: 219) (♂): Russia: Ussuri [River]; ZMAS.

Clinocoris stali Sahlberg, 1878: 16 [as Ståli ]. Holotype (♀): Russia: Mt. Takmak near Krasnoyarsk; NHRS. Synonymized by Jakovlev (1883: 110).

Elasmostethus davidi Fallou, 1891: 7 . Syntype (s): China: “Pékin” [= Beijing]; MNHN. Synonymized by Bergroth (1892: 262).

Elasmucha signoreti (non Scott, 1874): Horváth (1909: 631) and subsequent authors. Misidentification (Kerzhner 1972: 218, Kwon et al. 2001: 377, Vinokurov et al. 2010: 228).

Elasmostethus dorsalis: Puton 1886: 15 (catalogue, distribution), Jakovlev 1893: 289 (records), Lethierry & Severin 1893: 256 (catalogue, distribution), Puton 1899: 18 (catalogue, distribution).

Clinocoris dorsalis: Jakovlev 1883: 110 (synonymy), Reuter 1885: 37 (generic placement, in key, figures), Bergroth 1892: 262 (synonymy), Oshanin 1906: 172 (catalogue, distribution), Tang 1935: 360 (catalogue, distribution).

Elasmucha dorsalis: Reuter 1891: 172 , 177 (record, bionomics), Kirkaldy 1909: 175 (catalogue, distribution), Kiritshenko 1910: 177 (records), Oshanin 1910: 69 (catalogue, distribution), Hoffmann 1932: 10 (listed), Hoffmann 1935: 103 (catalogue, distribution), Hoffmann 1948: 8 (catalogue, distribution), Lindberg 1951: 4 (type material), Miyamoto 1957: 72 (ovariole number), Miyamoto 1959: 98 (description and figure of larva, phenology, host plant), Miyamoto 1962: 81 (redescription, photo, habitat, host plant, distribution), Petrova 1966: 200 (listed), Kobayashi 1971: 8 (photo, host plant, rearing), Lee 1971: 223 (in key, redescription, photo, distribution, host plant), 503 (catalogue, record, distribution), Hori 1972: 502 (salivary enzymes), Kerzhner 1972: 215 (in key), 218 (type material, diagnostic characters, figures, distribution, bionomy), Kiritshenko & Kerzhner 1972: 398 (records, distribution, habitat), Kerzhner 1973: 88 (listed), Kawasawa & Kawamura 1975: 238 (listed, host plants), Kiritshenko & Vinokurov 1975: 86 (record, distribution, host plant), Petrova 1975: 103 (in key), 108 (redescription, figure, host plant, records, distribution), Hoberlandt 1977: 512 (record, distribution), Hsiao & Liu 1977: 164 (in key, redescription, figure, photo, distribution), Josifov & Kerzhner 1978: 165 (records, host plant), Kerzhner 1978: 53 (records, distribution, host plant, bionomics), Liu 1979: 56 (records), 58 (diagnostic characters), Vinokurov 1979: 155 (in key, figures), Zhang et al. 1980: 33 (records, phenology), Kerzhner 1981: 9 (records), Watanabe 1982: 416 (record), Chao 1982: 51 (listed, distribution), Wu 1984: 38 (listed), Chen 1985: 124 (listed), Li 1985: 116 (redescription, records, as dorsails [inadvertent error]), Zhou 1985: 16 (redescription, photo, distribution), Kim & Chang 1987: 97 (listed), Kanyukova 1988: 915 (in key, figures), Nonnaizab 1988: 223 (in key), 225 (redescription, figures, host plant, distribution), Zheng et al. 1988: 3 (records), Kudo et al. 1989: 75 (behaviour), Miyamoto & Yasunaga 1989: 188 (listed, distribution), Chen 1990: 108 (diagnosis, record, host plant, phenology), Kudo 1990a: 431 (behaviour), Kudo 1990b: 156 (behaviour), Kudo 1990c: 188 (behaviour, ovarium, photos), Zheng & Gao 1990: 18 (record, host plant), Lee & Kwon 1991: 58 (listed, distribution), Park & Josifov 1991: 100 (record), Tachikawa 1991: 64 (reproductive behaviour), 125 (listed, distribution), Tomokuni 1992: 160 (records), Zhang & Sie 1992: 250 (redescription, figure, records, host plant, distribution), Lin & Zhang 1993: 134 (redescription, figure, distribution, host plants), Liu et al. 1993: 7 (record), Sun 1993: 126 (listed), Tomokuni 1993: 238 (diagnostic characters, photos, host plants, bionomy, distribution), Xiong & Liu 1993: 201 (record, distribution), Xu 1993: 49 (listed), Lee & Kwon 1994: 78 (listed), Chen et al. 1995: 207 (listed, host plant), Lin 1995: 64 (redescription, larva, host plants, phenology, distribution), Schuh & Slater 1995: 217 (behaviour), Vinokurov & Kanyukova 1995: 195 (in key, figures, host plants), Kwon et al. 1996: 116 (listed), Bu & Zheng 1997: 209 (redescription, figure, record, distribution), Kerzhner & Marusik 1997: 27 (listed), Cai et al. 1998: 232 (records), Kwon & Huh 1998: 73 (redescription, photos, distribution), Lei & Zhou 1998: 43 (listed), Lin et al. 1999: 99 (redescription, figure, host plants, distribution), Nonnaizab 1999: 78 (listed, distribution, host plants), Hua 2000: 167 (listed, distribution, host plant), Hori 2000: 16 (salivary enzymes), Kudo 2000: 137 (behaviour), Chen & Liu 2001: 279 (listed, distribution), Chen et al. 2001: 45 (listed), Fang & Wu 2001: 61 (listed, host plant, distribution), Kudo 2001: 209 (ecology), Kwon et al. 2001: 377 (catalogue, distribution), Li & Bu 2001: 125 (record), Nakamura et al. 2001: 50 (records), Derzhansky et al. 2002: 362 (type material), Kikuchi & Fukatsu 2003: 6085 (gut symbiotic bacteria), Meng 2003: 37 (listed), Hayashi & Ozaki 2004: 247 (listed, records), Kerzhner et al. 2004: 236, 240 (records), Kobayashi & Tachikawa 2004: 274 (listed), 301 (description and figures of egg and larval instars, distribution, host plants, phenology, bionomics), Vinokurov & Gromova 2004: 23 (listed, distribution, host plant), Liu & Wang 2005: 273 (records, distribution), Xie & Bu 2005: 185 (redescription, distribution), Averensky et al. 2006: 130 (listed, habitat, host plant), Bao & He 2006: 362 (listed, host plants), Costa 2006: 284 (host plant, behaviour, reproductive strategy), Göllner- Scheiding 2006: 174 (catalogue, distribution), Kanyukova & Marusik 2006: 173 (listed), Kinota 2006: 390 (photo, habitat, host plants), Ren et al. 2006: 274 (listed, record), Shen et al. 2006: 24 (listed, host plants, distribution), Xue & Bu 2006: 226 (redescription, distribution), Roth et al. 2006: 1163 (behaviour), Enju 2007: 96 (listed, distribution), Guo et al. 2007: 344 (listed, host plants), Li & He 2007: 350 (listed), Peng et al. 2007: 243 (listed), Wang et al. 2007: 59 (records), Zhang et al. 2008: 801 (listed), Zhao et al. 2008: 456 (listed), Kikuchi et al. 2009: 6, 22 (16S rRNA of gut symbiotic bacteria), 11 (mtCOI), 16 (reproductive behaviour), Liu et al. 2009: 71 (record, listed), Yamamoto et al. 2009: 35 (host plants), Ye 2009: 54 (record, host plants), Kanyukova & Kerzhner 2010: 559 (records, distribution), Vinokurov et al. 2010: 228 (bibliography and distribution in Asian part of Russia), Babichev & Vinokurov 2011: 36 (record, host plant), Jia et al. 2011: 392 (listed, distribution), Jiang et al. 2011: 51 (listed), Xu et al. 2011: 22 (listed), Yang et al. 2011: 126 (listed, photo, distribution, host plants), Prado & Zucchi 2012: 3 (in figure, gut symbiotic bacteria), Trumbo 2012: 83 (behaviour), Yamamoto & Hayashi 2012: 514 (diagnosis, bionomics, distribution), Aukema et al. 2013: 431 (catalogue, distribution), Esenbekova 2013: 230, 306 (habitat, host plant, phenology, overwintering, distribution), Hanelová & Vilímová 2013: 434, 445 (behaviour), Huang et al. 2013: 139 (records, distribution), Liang et al. 2014: 377 (listed), Yasuda 2014: 80 (listed, distribution).

Elasmucha signoreti (misidentification): Horváth 1909: 631 (synonymy), Oshanin 1912: 18 (catalogue, distribution), Lindberg 1921: 47 (records), Esaki 1932: 1592 (redescription, habitus, distribution), Doi 1933: 92 (record), Doi 1938: 91 (listed), Tanaka 1942: 661 (listed, distribution), Tsherepanov & Kiritshenko 1962: 18 (records, habitat), Kulik 1965: 157 (distribution, habitat, phenology), Petrova 1966: 200 (listed, habitat).

Acanthosoma vicinum: Stål 1876: 115 (as of doubtful identity, listed), Kirkaldy 1909: 181 (as of doubtful identity, catalogue, distribution), Hoffmann 1932: 10 (as of doubtful identity, listed), Hoffmann 1935: 105 (as of doubtful identity, catalogue, distribution), Hsiao & Liu 1977: 175 (listed), Hua 2000: 166 (listed, distribution).

Acanthosoma vicina: Lethierry & Severin 1893: 260 (as of doubtful identity, catalogue, distribution).

Cyphostethus vicinum : Wu 1933: 217 (listed, distribution).

Grossaria vicinum : Kumar 1974: 47, 50 (generic placement, type material, figures).

Grossaria vicina : Göllner-Scheiding 2006: 178 (catalogue, distribution).

Type material examined. Acanthosoma vicinum . Lectotype (♂): “ Acanthosoma \ vicinum \ Uhler. \ HongKong \ Roger’s. Exp.” [handwritten]”, “PRUhler \ Collection” [printed], “TypeNo. [printed] \ 69812 [handwritten] \ U.S. N.M. [printed]” [red square], “LECTO- \ TYPE ” [printed circle outlined in dark blue]; with right antennal segments III–IV, apical third of scutellum, right mid leg, and some tarsal segments lacking; genitalia preserved in glass microvial with glycerol pinned with the specimen; abdomen of another conspecific male mounted on card and pinned with the specimen (it is not considered as part of the type series); deposited in USNM ( Figs. 1–5 ).

Additional specimens examined. About 300 specimens were examined in connection with the present study, they are not listed in detail but their localities are marked under Distribution with exclamatory points.

Diagnosis. Recognized within the genus based on the following combination of characters: humeri with a pair of distinct but relatively short, posterolaterally directed, apically blunt processes which are reddish to black with coarse black punctures ( Figs. 1–2 ); metapleuron with narrowly separated, occasionally confluent punctures; laterotergites of abdominal segments VI and VII yellow, unpunctured, posterolateral angle of laterotergite VII narrowly, posterior margin of laterotergite VII broadly infuscate; abdominal venter with coarse, black punctures sublaterally ( Fig. 2 ). The shape of the genital capsule of the male is diagnostic; it was figured, among others, by Kerzhner (1972), Hsiao & Liu (1977), Vinokurov (1979), Kanyukova (1988), Nonnaizab (1988) and Vinokurov & Kanyukova (1995). Female terminalia are illustrated in Fig. 6 and compared with those of two other morphologically similar species, E. signoreti Scott, 1874 ( Fig. 7 ), E. putoni Scott, 1874 ( Fig. 8 ) and E. laeviventris Liu, 1979 ( Fig. 9 ). The species can readily be identified using the redescriptions and illustrations of any of the above listed works.

Distribution. A Manchurian-Temperate Northwest Pacific element, distributed all over East Asia from the Russian Far East to the temperate parts of China and Japan, entering the Oriental Region only marginally.— RUSSIA (bibliography provided by Vinokurov et al. 2010). West Siberia ( Vinokurov et al. 2010); East Siberia: Zabaikalsky Region: “Gorbitza”! [= Gorbitsa] (HNHM); Far East Territory: Amur Region: Raddefka! [= Radde] (HNHM), Ussuri Region! (HNHM).— KAZAKHSTAN. Almatinskiy Zapovednik: Issyk valley! (HNHM); East, Southeast ( Esenbekova 2013).— MONGOLIA ( Kiritshenko & Kerzhner 1972, Hoberlandt 1977).— JAPAN. Hokkaido: Samani District: Apoi Area!, Hidaka: Mt. Apoi!, Shikotsuko!, Tokatidake!, Shari District: Shiretoko!, Mt. Taisetsu!, Sapporo: Jozankei!, Hoheikyo!, Noboribetsu!, Nemuro: Kenebetsu!, Kitami City: Takinoue!, Tokachi: Ashoro-cho!, Shintoku!, Mashike: Mt. Syokanbetsu-dake! (SEHU); Rebun Is.! (SEHU); Honshu: Toyama Pref.: Toyama!, Aomori Pref.: Towada!, Iwate Pref.: Mt. Hayatine!, Yamanashi Pref.: Mt. Daibosatsu!, Totigi Pref.: Nasu, Mt. Minamigassan! (SEHU); Shikoku; Kyushu; Rishiri Is. ( Miyamoto & Yasunaga 1989, Yamamoto & Hayashi 2012).— KOREA (bibliography provided by Kwon et al. 2001). North: Kangwon Prov.: Mt. Kumgang! (HNHM), South Pyongan: Za-mo san! (HNHM); Central ( Kwon et al. 2001).— CHINA. Ningxia: Mt. Liupan!, Jingyuan! (NKUM); Gansu: Wen County: Xiangjiaba!, Kang County: Heimaguan!, Yuzhong County: Mt. Xinglong! (NKUM); Inner Mongolia: Alxa Zuoqi! (NKUM); Heilongjiang: Mudanjiang!, Wuying!, Zhangzhi: Mt. Mao’er! (NKUM); Liaoning: Yingkou! (NKUM); Hebei: Xinglong: Mt. Wuling!, Yu County! (NKUM); Tianjin: Fu County! (NKUM); Shaanxi: “Weitzeping”! [= Xi’an], Zhouzhi!, Feng County! (NKUM); Shanxi: Hongdong County!, Nanshui! (NKUM); Henan ( Cai et al. 1998); Hubei: Lichuan!, Shennongjia!, Wufeng County! (NKUM); Sichuan: Chengdu!, Ma’erkang [= Ba’erkang]!, Xiaojin!, Baoxing! (NKUM); Chongqing: Nanchuan! (NKUM), Hunan ( Hua 2000); Anhui: Huangshan! (NKUM); Zhejiang ( Hsiao & Liu 1977, Chen 1985, Chen et al. 1995, Ye 2009); Jiangxi ( Peng et al. 2007); Fujian ( Hsiao & Liu 1977, Lin et al. 1999); Guizhou: Daozhen: Dashahe Nature Reserve! (NKUM); Guangxi: Xing’an: Mt. Mao’er! (NKUM); Guangdong ( Chen et al. 2001); Hong Kong! (USNM).

Discussion. The identity of Elasmucha dorsalis was clarified by Kerzhner (1972) based on the reexamination of syntypes; the author designated a lectotype and provided illustrations of diagnostic characters, particularly the male genital capsule. There has been an unequivocal agreement about the identity of this species among subsequent authors. Reexamination of the lectotype of Acanthosoma vicinum (including its male genitalia removed by a previous worker but preserved in good condition) in connection with the present study revealed that it is conspecific with E. dorsalis , and as a consequence the following new species level subjective synonymy is proposed: Elasmucha vicina ( Uhler, 1861) , comb. nov. = Elasmucha dorsalis ( Jakovlev, 1876) , syn. nov.

There is a considerable number of modern works referring to this biological species under the junior name E. dorsalis , and more than 100 of them were published in the immediately preceding 50 years. Besides of numerous faunistic records the bibliography of the species includes papers providing redescriptions, photos or illustrations of diagnostic characters of adults ( Petrova 1975, Hsiao & Liu 1977, Vinokurov 1979, Zhou 1985, Kanyukova 1988, Nonnaizab 1988, Zhang & Sie 1992, Lin & Zhang 1993, Tomokuni 1993, Lin 1995, Vinokurov & Kanyukova 1995, Bu & Zheng 1997, Lin et al. 1999, Yamamoto & Hayashi 2012) and immatures ( Lin 1995, Kobayashi & Tachikawa 2004), host plant records and other information on various aspects of its bionomics ( Josifov & Kerzhner 1978, Kerzhner 1978, Zheng & Gao 1990, Lin & Zhang 1993, Tomokuni 1993, Chen et al. 1995, Lin 1995, Lin et al. 1999, Kobayashi & Tachikawa 2004, Yamamoto et al. 2009), agricultural importance ( Lin 1995) and even physiology ( Hori 1972). Particularly the remarkable reproductive behaviour of the species was a subject of a detailed research ( Kudo et al. 1989, Kudo 1990a – c, 2000, 2001); this information has been cited in comprehensive handbooks on general insect ecology and behaviour ( Costa 2006, Trumbo 2012). The species was selected to represent Acanthosomatidae in a cladistic analysis of pentatomoidean families ( Xu 1993). It was included in a molecular study on the phylogenetic relationships of various acanthosomatids and coevolution with their gut symbiotic bacteria ( Kikuchi et al. 2009); its mitochondrial gene of cytochrome oxidase subunit I (mtDNA-COI) is available from GenBank (AB368845.1). The species name has been cited in comprehensive reference books on Heteroptera ( Schuh & Slater 1995) , handbooks on agricultural and forestry entomology ( Wu 1984, Lin 1995, Hori 2000) and popular insect books and field guides targeting a wide public ( Kawasawa & Kawamura 1975, Tomokuni 1993, Kwon & Huh 1998, Kinota 2006, Yamamoto & Hayashi 2012).

In contrast with E. dorsalis , no authors were able to identify E. vicinum correctly since its original description ( Uhler 1861). As a consequence, much less papers used the senior specific epithet vicinum in combination with different generic names, and all but one of them were checklists and catalogues providing only secondary data based on the original description, most of them explicitly listing the species as of unknown identity ( Stål 1876, Lethierry & Severin 1893, Kirkaldy 1909, Hoffmann 1932, Wu 1933, Hoffmann 1935, Hsiao & Liu 1977, Hua 2000). The single exception is the contribution by Kumar (1974), who reexamined a single male syntype (designated as lectotype), provided highly inaccurate drawings of the pronotum and the metapleuron, and established a new monotypic genus, Grossaria, for it (see also discussion about the generic level synonymy above). As a result of the poor illustrations and the incorrect and misleading description based on the defective lectotype no subsequent authors could recognize this species after Kumar (1974) and the genus and the species have never been cited except the taxonomic catalogue by Göllner-Scheiding (2006).

We believe that changing the prevailing usage of the name of this species simply because of adherence to the Principle of Priority is undesirable and would threaten stability. The number and authorship of the works using the junior name meet the conditions of Article 23.9.1.2., but the senior name was used as a valid name after 1899 therefore the ‘automatic’ conservation of E. dorsalis under Article 23.9.2 is not possible. Therefore, no nomenclatural change is proposed in this paper, but an application has simultaneously been submitted to the International Commission on Zoological Nomenclature to give the specific name dorsalis precedence over the specific name vicinum under the provision of Article 23.9.3.

Figures

FIGURES 1 – 5. Lectotype of Acanthosoma vicinum Uhler, 1861 and its labels. Fig. 1, dorsal view; Fig. 2, ventral view; Fig. 3, labels; Fig. 4, abdomen of a conspecific male pinned with the specimen; Fig. 5, microvial with male genitalia of lectotype. Scales in mm.

FIGURES 6 – 9. Terminalia of females of Elasmucha species. Fig. 6, E. dorsalis (Jakovlev, 1876); Fig. 7, E. signoreti Scott, 1874; Fig. 8, E. putoni Scott, 1874; Fig. 9, E. laeviventris Liu, 1979. Scales in mm.