Ione cornuta Bate, 1864

Ione cornuta Bate, 1864: 668 (type locality: east side of Vancouver Island, British Columbia, Canada, infesting Callianassa longimana Stimpson, 1856 = Neotrypaea gigas (Dana, 1852)) .—Bate & Westwood, 1862: 253 (mention).— Giard & Bonnier, 1887a: 41, 63 (list), 77 (list).— Perrier, 1893: 1021 (mention).— Richardson, 1899a: 869 (list).— Richardson, 1899b: 338 (list).— Bonnier, 1900: 48, 61, 62, 222, 245–247 (discussion of characters).— Richard, 1900: 71 (list).— Richardson, 1900: 308 (key).— Gerstaecker & Ortmann, 1901: 185 (list), 266 (list).— Giard, 1904: 592 (mention).— Richardson, 1904: 75 (list), 77–78 (comparison with other Ione species).— Richardson, 1905: 504 – 505 (repeat of the description by Bate, 1864), 510 (mention).— Giard, 1913: 420 (mention).—Nierstrasz & Brender à Brandis, 1923: 80 (list).—Nierstrasz & Brender à Brandis, 1931: 180–182, figs. 57–59 (Vancouver, British Columbia, Canada, infesting unknown host).— Hiraiwa, 1933: 55 (mention).— Caspers, 1939: 243 (list).— Shiino, 1939a: 13 –16, fig. 2, 3 (Seto, Japan, infesting Nihonotrypaea japonica (Ortmann, 1891)) .— Hatch, 1947: 164 (list), 225 (characters, host list, distribution), pl. 9, figs. 113–115 (figures after Richardson, 1905).— Codreanu & Codreanu, 1956a: 577, 580, 583 (mention).— Codreanu & Codreanu, 1956b: 120 (mention).— Gooding, 1957: 698 (Hammond Bay, Nanaimo, British Columbia, Canada, first mention of this species infesting Callianassa californiensis Dana, 1854 = Neotrypaea californiensis (Dana, 1854)) .— Danforth, 1963a: 8, 33, 43 (list), pl. 8, figs. 1–3.— Shiino, 1964a: 30, 32 (mention).— Schultz, 1969: 316 (mention; erroneous statement that the species was never illustrated).— Danforth, 1970b: 10, 17, 47 (list), 79–80 (translation of description from Bonnier, 1900), 147 (key), fig. 22a–c.— Shiino, 1972: 8 (list).— Miller, 1975: 285, 286 (key), 305 (list), pl. 64, fig. 19.— Rudy & Rudy, 1979: 152 (mention).— Bird, 1982: 52, 54, 55 (mention).— Hart, 1982: 32 (mention).— Kozloff, 1983: 301 (mention).— Austin, 1985: 587 (list).— Rafi, 1985: 12 (list).— Bourdon, 1987: 342 (key, list).— Kim & Kwon, 1988: 205 –207, 220, fig. 4 (South Korea, infesting N. japonica).— Markham, 1992a: 3, table 1 (list).— Kozloff, 1993: 301 (mention).— Markham, 1995: 86 (mention).— Brusca et al., 2001: 7 (key), 20 (list), pl. 6, fig. 32.— Saito et al., 2000: 42 (list).— Markham, 2001: 197, 199, 200 (list).— Sadro, 2001: 176 (list).— Saito, 2002: 223 (list).— Itani, 2004b: 37, table 3 (Japan, infesting Nihonotrypaea harmondi (Bouvier, 1901), N. japonica, N. petalura (Stimpson, 1860)) .— Saito & Kinoshita, 2004: 1 –6, fig. 3 (prevalence in Tokyo Bay, Japan infesting N. japonica).— Espinosa-Pérez & Hendrickx, 2006: 237 (list).— Brusca et al., 2007: 512 (key), pl. 236B.— Pernet et al., 2008: 1127 –1129, 1133–1134, 137–1140 (Oregon, infesting N. californiensis).— Yu & An, 2008: 692 (list).—An et al., 2009: 235–237, fig. 6 (Shandong, Liaoning, and Zhejiang Provinces, China, infesting N. japonica, N. petalura, N. harmondi and Upogebia major (De Haan, 1841)) .— Campos et al., 2009: 1255 (List), 1256 (mention).— Espinosa-Pérez et al., 2009: 229 (list).—Williams & An, 2009: 121– 122 (mention).— Passarelli, 2010: 7, 8, 14, 20, 32, 35–38, 41, 45, 50, 52–54, 5–60 (California, infesting N. californiensis).— Dumbauld et al., 2011: 346 (prevalence in Washington, U.S.A. infesting N. californiensis).— Chapman et al., 2012: 1223, 1229, 1230 (mention).— Cohen, 2012: 2, 32, 46, 48 (mention).— Rudy and Rudy, 2013: 288 (mention).—Boyko et al., 2013: 499 (placement on phylogenetic tree).— Chapman & Carter, 2014: 351 –352 (mention).— Miura et al., 2014: 31 (table), 34–35, fig. 1 (Japan, infesting N. japonica).— Saito, 2016: 146 (mention).

Ione cornutus — Bate, 1866: 283 –284 (repeat of information from Bate, 1864).— Stebbing, 1893: 414 (list).

Jone [sic] cornuta — Kossmann, 1881b: 171, 181 (mention).

Ione brevicauda Bonnier, 1900: 48, 61, 222, 248–250, pl. 4 (type locality: San Francisco, California, U.S.A., infesting “ Callianassa ” sp.).— Giard, 1904: 592 (mention).— Richardson, 1905: 504 –507, fig. 553 (translation of the description of Bonnier, 1900).— Giard, 1913: 419 (mention).— Fee, 1927: 36 –37 (British Columbia, infesting N. gigas; placement into synonymy with I. cornuta).—Nierstrasz & Brender à Brandis, 1931: 182 (mention).— Caspers, 1939: 243 (list).— Menzies & Miller, 1954: 141, fig. 65 (illustration).— Danforth, 1963a: 43 (list).— Shiino, 1964a: 32 (mention).— Schultz, 1969: 316 (list; figure from Bonnier, 1900).— Román-Contreras, 2008a: 382 (mention).

Ione sp. Menzies & Miller, 1954: 141 (key), 153 (list).— Carlton & Kuris, 1975: 408 (mention).— Haig & Abbott, 1980: 580 (mention).— Lee & Miller, 1980: 543, fig. 21.10 (“ British Columbia to California,” infesting N. californiensis).— Kuris et al., 2007: 635 (mention).— Campos et al., 2009: 1255 (list).

“parasitic isopods occur on Callianassa ” Kozloff, 1973: 232 (occurrence in Puget Sound region on C. californiensis = N. californiensis).

“bopyrid isopod parasites” Poore, 2000: 151 (Japan, infesting Grynaminna tamakii Poore, 2000).

“another species of bopyrid isopod” Anonymous, 2003: 4, 3.

? “parasit bopyrid” Wardiatno, 2004: 81–84 (Japan, infesting N. japonica; see Remarks).

not Ione sp. Lamb & Hanby, 2005: 280, fig. AR19.A (= Orthione griffenis Markham, 2004, infesting Upogebia pugettensis (Dana, 1852)) .

not Ione sp. Lamb & Hanby, 2005: 280, fig. AR19.B (= gen. et sp. indet., infesting Acantholithodes hispidus (Stimpson, 1860)) .

Material examined. Mexico: Mature female (7.2 mm), mature male (3.3 mm), ex left branchial chamber of female Neotrypaea gigas (5.7 mm CL), ovigerous female (11.0 mm), mature male (6.0 mm), ex right branchial chamber of intersex N. gigas (8.0 mm CL), ovigerous female (9.0 mm), mature male (4.0 mm), ex left branchial chamber of female N. gigas (9.2 mm CL), ovigerous female (8.0 mm), mature male (4.5 mm), ex left branchial chamber of female N. gigas (6.0 mm CL), Bahia de Los Angeles, Baja California Norte, coll. F. Mantelatto, J. Cuesta & R. Robles, 6 Dec 2001 (ULLZ 10196 [3 [pairs]; USNM 1437637 ex ULLZ 10196 [1 pair; female incomplete, tissue removed for molecular analysis]). Japan: Ovigerous female (10.2 mm), mature male (5.2 mm), ex left branchial chamber of female Grynaminna tamakii (12.0 mm CL), Minami-Arima-cho sandflat, near south end of Shimabara Peninsula, Nagasaki Prefecture, Kyushu (32°37'N, 130°13'E), coll. A. Tamaki, 8 Jul 1998 (NSMT-Cr 12529).

Distribution. Japan, South Korea, China; British Columbia, Canada to California, U.S.A., and Gulf of California, Baja California Norte, Mexico.

Hosts. Grynaminna tamakii Poore, 2000, Nihonotrypaea harmondi (Bovier, 1901), N. japonica (Ortmann, 1891), N. petalura (Stimpson, 1860), Neotrypaea californiensis (Dana, 1854), N. gigas (Dana, 1852) (type host), Upogebia major (De Haan, 1841) .

Remarks. The records of this species from an upogebiid host (An et al. 2009) may seem questionable as I. cornuta is otherwise only known from callianassid hosts but the congener I. thoracica (Montagu, 1808) has been reported to infest both callianassid and upogebiid hosts in the Mediterranean (Markham 2001). However, there are no records of I. cornuta from the west coast of the United States on any upogebiid host. Many early records had the host cited as Callianassa longimana, a synonym of N. gigas . Although Campos et al. (2009) listed I. cornuta as a parasite of N. gigas (based on data from Richardson 1905) and “ Ione sp.” as a parasite of N. californiensis (based on data from Haig & Abbott 1980), they did not find infested hosts of either species from the Gulf of California. The Mexican specimens reported here therefore represent the first record of I. cornuta from the Gulf of California. The specimens from Grynaminna tamakii are the first bopyrids recorded from this host species.

The inclusion of “Nobuhiro & Kyoko 2004” in the synonymy list for this species by An et al. (2009) is an error as these are the first names of the authors Saito and Kinoshita (2004). The specimens studied by Wardiatno (2004) were never named or figured; they are likely either this species or Pseudione longicauda Shiino, 1937, as these are the only species known from Nihonotrypaea japonica .