Argeia pugettensis Dana, 1852 Figure 1

Argeia pugettensis Dana, 1852: 804–805 [Puget Sound, Washington, infesting Metacrangon munita (Dana, 1852)]; Dana, 1855: pl. 53, fig. 7a–d; Stimpson, 1857: 511 [Puget Sound, Washington, infesting M. munita]; Stebbing, 1893: 336, 337; Calman, 1898: 281; Richardson, 1899a: 868; Richardson, 1899b: 336–337; Bonnier, 1900: 48, 61, 171, 221, 327–328, fig. 55; Richardson, 1900: 308; Gerstaecker, 1901: 185; Richardson, 1904a: 45, 60–64, figs. 35–40 [Bering Sea to San Diego Bay, California, infesting Argis alaskensis (Kingsley, 1883), Argis crassa (Rathbun, 1899), Argis dentata (Rathbun, 1902), Argis lar (Owen, 1839), Argis ovifer (Rathbun, 1902), Crangon alaskensis Lockington, 1877, Crangon alba Holmes, 1900, Crangon dalli Rathbun, 1902, Crangon franciscorum angustimanus Rathbun, 1902, Crangon nigromaculata Lockington, 1877, Crangon propinquus Stimpson, 1860, M. munita, Neocrangon communis Rathbun, 1899]; Richardson, 1904b: 858; Richardson, 1905a: 220 [British Columbia and Alaska, infesting Argis dentata (Rathbun, 1902), C. alaskensis, N. communis ]; Richardson, 1905b: 544–550, figs 586–597 [same material as Richardson, 1904a]; Richardson, 1909: 122 [Japan and Korea, infesting Argis sp.]; Chopra, 1923: 476–478, 480; Nierstrasz and Brender à Brandis, 1923: 87; Fee, 1927: 16, 40 [British Columbia, infesting Lissocrangon stylirostris (Holmes, 1900), M. munita]; Nierstrasz and Brender à Brandis, 1929: 16 [California, infesting Crangon sp.]; Fraser, 1932: 64; Hiraiwa, 1933: 53; Shiino, 1933: 249, 277–279, fig. 11 [Japan, infesting unknown host]; Gurjanova, 1936a: 219– 221, fig. 138 [Kamchatka Peninsula and Bering Sea, infesting Argis spp., Crangon spp.]; Gurjanova, 1936b: 258; Shiino, 1937a: 299 [Japan, infesting Crangon affinis De Haan, 1849]; Ricketts and Calvin, 1939: 185, 281; Baer, 1946: 65; Hatch, 1947: 164, 224, pl. 9, figs. 110–112; Ricketts and Calvin, 1948: 185, 320; Baer, 1951: 69–70; Ricketts and Calvin, 1952: 239, 438; Shiino, 1952: 34; Menzies and Miller, 1954: 141, 153, fig. 65c, d; Shiino, 1958: 56, pl. 3, fig. 8 [Japan, infesting A. lar, C. affinis, Neocrangon sagamiensis (Balss, 1913)]; Oguro, 1961: 43–47, figs. 1–3, 5–7 [Japan, infesting C. affinis]; Danforth, 1963: 35, 52, 54, 60, 61, 63, 64, 68, pl. 2, fig. 1, pl. 4, figs. 4–6; Sindermann and Rosenfield, 1967: 351; Bourdon, 1968: 358; George and Strömberg, 1968: 253 [Washington, infesting A. dentata, C. alaskensis, Crangon franciscorum Stimpson, 1856, Eualus suckleyi (Stimpson, 1854), Mesocrangon munitella (Walker, 1898), M. munita, N. communis]; Ricketts and Calvin, 1968: 286, 488; Schultz, 1969: 323–324, figs. 516–517; Danforth, 1970: 41; 42, 56–57, 143, 150, fig. 5a–c; Sindermann, 1970: 171; Strömberg, 1971: 28; Kozloff, 1974: 148; Miller, 1975: 285, 287, 305, pl. 64, fig. 18; Markham, 1977: 112–114 [Washington and California, infesting C. alaskensis, C. franciscorum, Neocrangon resima (Rathbun, 1902)]; Rudy and Rudy, 1979: 122, 124, 126; Beck, 1980: 135; Bourdon et al., 1981: 502; Jay, 1985: i–vii, 1–63 [California, infesting C. franciscorum]; Ricketts et al., 1985: 328, 533; Markham, 1986: 156, 161; Bourdon, 1987: 342–343, fig. 17.23; Strathmann, 1987: 436; Kim and Kwon, 1988: 199, 201, 207–210, fig. 6 [Korea, infesting A. lar, C. affinis, possibly Crangon hakodatei Rathbun, 1902]; Jay, 1989: 68–76 [California, infesting C. franciscorum]; Sinder mann, 1990: 196; Brusca and Wilson, 1991: 148, fig. 2a; Feeney, 1995: 6; Trilles, 1999: 285, 292, 334, fig. 8.13C; Brusca et al., 2001: 20; Shields, 2001: 313; An, 2006: 61– 63, fig. 26 [China, infesting C. affinis, Crangon cassiope de Man, 1906, Crangon crangon (Linnaeus, 1758)]; Espinosa-Pérez and Hendrickx, 2006: 237; Chaplin-Ebanks and Curran, 2007: 73; Marin Jarrin, 2007: x, 54, 90–92 96, 100, 118 [Oregon, infesting Lissocrangon stylirostris (Holmes, 1900)]; An et al., 2008: 225–226 [China, infesting C. affinis, C. cassiope, C. crangon]; Liu: 2008: Marin Jarrin and Shanks, 2008: 691; 613–620 [Oregon, infesting L. stylirostris]; Romero-Rodríguez and Román-Contreras, 2008: 1207; Wicksten, 2008: 143; Espinosa- Pérez et al., 2009: 229; Williams and An, 2009: 122; Dumbauld et al., 2011: 337; Han and Li, 2010: 228; An, 2011: 130–133, figs. 5-1–5-3 [China, infesting C. affinis, C. cassiope, C. crangon]; Marin Jarrin and Shanks, 2011: 237; Wicksten, 2012: 106; Williams and Boyko, 2012: 7; Penha-Lopes et al., 2013: 35; Rudy and Rudy, 2013: 208, 210, 272; Sherman and Curran, 2013: 1336; Romero-Rodríguez and Román-Contreras, 2014: 470; Seo et al., 2014: 413–417, fig. 2 [Korea, infesting A. lar]; Cericola and Williams, 2015: 239.

Argeia pauperata Stimpson, 1857: 511 [San Francisco, California, infesting C. franciscorum]; Stebbing, 1893: 415; Calman, 1898: 281; Bonnier, 1900: 61, 171, 221, 328, 381; Gerstaecker, 1901: 185, 266; Richardson, 1905b: 551; Chopra, 1923: 476–478; Nierstrasz and Brender à Brandis, 1923: 87; Gifford, 1934: 1–20, pls. 1–5; [California, infesting C. franciscorum]; Danforth, 1963: 20, 28, 34; 68, 75, pl. 1, fig. 4, pl. 2, fig. 2, pl. 4, figs. 1–3; Schultz, 1969: 323; Danforth, 1970: 41, 56, 150, fig. 4d–f; Strömberg, 1971: 28; Miller, 1975: 287, 305; Nelson et al., 1986: 121–124.

Argeja [sic] pugettensis Cornalia and Panceri, 1861: 86, 16.

? Bopyrus sp. Bate, 1888: 485 [Japan, infesting C. affinis].

Argeia sp. Calman, 1898: 261, 281 [Washington, USA, infesting C. affinis].

Argeia depauperata [sic] Richardson, 1899a: 868; Richardson, 1899b: 336, 337; Richardson, 1900: 308.

Argeia (?) [spp.] Bonnier, 1900: 221.

Argeia Calmani Bonnier, 1900: 171, 221, 329, 381 [Washington, infesting C. affinis] (nomen nudum).

Argeia Pingi Yü, 1935: 52–53 [Hopei and Shantung Provinces, China, infesting C. cassiope].

Argaeia [sic] pugettensis — MacGinitie and MacGinitie, 1949: 266 [west coast of United States, infesting C. alaskensis]; MacGinitie and MacGinitie, 1968: 265–266.

MATERIAL EXAMINED: Infesting Crangon affinis De Haan, 1849. 2♀♀, ♂ (CIECR 007601), South Yellow Sea, Stn. 76, 123 ° 30′E, 27 ° 20′N, 45 m, 18 April 1975; 2♀♀, 2♂♂ (CIECR020801), South Yellow Sea, Stn. 208, 127 ° 00′E, 30 ° 30′N, 46 m, 12 July 1968; ♀, ♂ (CIECR104941), Yellow Sea, Stn. 10494, 122 ° 00′E, 35 ° 00′N, 52 m, 13 June 2004; 2♀♀, 2♂♂ (CIECR007602), South Yellow Sea, Stn. 76, 123 ° 30′E, 27 ° 20′N, 45 m, 10 June 1975; ♀, ♂ (CIECR030601), Yellow Sea, Stn. 306, 126 ° 00′E, 29 ° 30′N, 60 m, 13 November 1957; ♀ (CIECR350801), Yellow Sea, Stn. 3058, 120 ° 00′E, 35 ° 00′N, 81 m, 12 September 1992; ♀, ♂ (CIECR626301); South China Sea, Stn. 6263, 107 ° 30′E, 19 ° 00′N, 77 m, 8 March 1962; ♀, ♂ (CIECR008101), South Yellow Sea, Stn. 81, 122 ° 00′E, 27 ° 10′N, 31 m, 8 June 1975; ♀, ♂ (CIECR005101), South Yellow Sea, Stn. 51, 123 ° 20′E, 28 ° 30′N, 65 m, 18 April, 1975; ♀, ♂ (CIECR000101), South Yellow Sea, Stn. 1, 121 ° 00′E, 26 ° 30′N, 75 m, 29 April 1976; ♀, ♂ (CIECR590501), Shazikou, Qingdao, Shandong Province, 16 May 1959, coll. Fenxuan Zhang; 2♀♀, 2♂♂ (CIECR580901), South Yellow Sea, Stn. 81, 122 ° 00′E, 27 ° 10′N, 31 m, 20 September 1958; 2♀♀, 2♂♂ (CIECR303801), Yellow Sea, Stn. 3038, 123 ° 45′E, 36 ° 00′N, 71.5 m, 10 November 1958.

Infesting Crangon crangon (Linnaeus, 1758). 2♀♀, 2♂♂ (CIECR530401), Yantai Fish Market, Shandong Province, 20 April 1953; 2♀♀, 2♂♂ (CIECR570101), Shazikou, Qingdao, Shandong Province, 20 January 1957, coll. Shaowu Wang, Xiuxi Zhang and Zhengang Fan; ♀, ♂ (CIECR550401), Yantai, Shandong Province, 5 April 1955, coll. Engze Yang; ♀, ♂ (CIECR500301), Beidai He, Hebei Province, 5 March 1950.

Infesting Crangon cassiope de Man, 1906. 2♀♀, 2♂♂ (CIECR510401), 2♀♀, 2♂♂ (CIECR510402), Yantai, Shandong Province, 6 April 1951.

DESCRIPTION: (CIECR007601): Length 8.12 mm, pereon length 4.58 mm, maximal width 6.2 mm, head length 0.94 mm, head width 1.87 mm, distorted 36 ° (fig. 1A, B). All segments distinct. Head with narrow frontal lamina, anterior edge of head deeply bilobate (fig. 1A), eyes lacking. Antennule of three articles, antenna of five articles, terminally setose. Maxilliped (fig. 1C, D) with setose palp, plectron sharp, anterior article two times larger than posterior article. Barbula with two pairs of smooth falcate lateral projections on each side (fig. 1E).

Pereon broadest across third pereomere (fig. 1A). Coxal plates very reduced, only visible on first three pereomeres of long side. Dorsolateral bosses on first three pereomeres of longer side, and first four pereomeres on shorter side. Tergal projections on longer side extended, tapeing distally to sharp tip. Brood pouch open (fig. 1B). Oostegite 1 (fig. 1F, G) with nearly smooth internal ridge, posterolateral point lacking. Posterior pereopods larger than anterior pairs, ischium with enlarged tubercular lobe, dactyli blunt (fig. 1H, I). Pleon of six pleomeres, first five pleomeres with lateral plates and uniramous pleopods. Sixth pleomere without lateral plates, with uniramous uropods (fig. 1B).

DESCRIPTION: Male (CIECR007601): Len‐ gth 2.26 mm, maximal width across pereomere 4, 0.71 mm, head width 0.43 mm, head length 0.20 mm, pleonal length 0.55 mm. (fig. 1K, L). Head elliptical, separated from first pereomere (fig. 1K). Black eyes near posterior edge of head (fig. 1K). Antennule of three articles, antenna of four articles, terminally setose (fig. 1M). Pereomeres distinct, subequal in width, midventral projections lacking (fig. 1L). Dactyli of first pereopod larger others (fig. 1N). Pleon fused into single ovate piece, pleopods and uropods lacking; anal cone usually lacking (occasionally present) (fig. 1O).

VARIATIONS: The head of some females is so deeply biobed it looks like a pair of spheres united. One female specimen (CIECR530401) with an extremely bulging head, very extended, terminally sharp tergal projections (fig. 1J) and the surface of the lateral plates are crenulated. One male (CIECR510401) with anal cone on the terminal end of the pleon (fig. 1O).

HOSTS AND LOCALITIES: Infesting Argis alaskensis (Kingsley, 1883), Argis crassa (Rathbun, 1899), Argis dentata (Rathbun, 1902), Argis lar (Owen, 1839), Argis ovifer (Rathbun, 1902), Crangon affinis De Haan, 1849, Crangon alaskensis Lockington, 1877, Crangon alba Holmes, 1900, Crangon cassiope de Man, 1906, Crangon crangon (Linnaeus, 1758), Crangon dalli Rathbun, 1902, Crangon franciscorum Stimpson, 1856, Crangon franciscorum angustimanus Rathbun, 1902, Crangon hakodatei Rathbun, 1902, Crangon nigromaculata Lockington, 1877, Crangon propinquus Stimpson, 1860, Eualus suckleyi (Stimpson, 1854), Lissocrangon stylirostris (Holmes, 1900), Mesocrangon munitella (Walker, 1898), Metacrangon munita (Dana, 1852), Neocrangon communis Rathbun, 1899, Neocrangon resima (Rathbun, 1902), Neocrangon sagamiensis (Balss, 1913) (Crangonidae), Bering Sea to San Diego, Japan, Korea, China.

REMARKS: This species has an extremely wide range, occurring in both the northeast and northwest Pacific on at least 23 crangonid hosts. Specimens from Japan, Korea, and China have been well described, but northwestern Pacific specimens are much more poorly known. Richardson (e.g., 1905b) described the species in some detail, but a fuller redescription, especially of characters now known to be informative at the species level, is needed. An example of a possible difference between northwestern and northeastern Pacific specimens, based on literature review, is that the northwestern specimens appear to have moderate to pronounced bilobation of the head, while northeastern specimens have a nearly flat head in dorsal view (see, e.g., Gifford, 1934). Additionally, given the wide geographic range of the species, a molecular analysis of specimens from both sides of the Pacific should be undertaken to test the possibility of cryptic species.

There are three hosts of this species in China and one of these, Crangon cassiope, is the type host for Argeia pingi Yü, 1935, which was only very briefly described, not illustrated, and never reported subsequent to its original description. Markham (1977) thought A. pingi might be a synonym of A. pugettensis. Because this host has now been recollected and is infested with Argeia specimens that do not differ from A. pugettensis as currently defined, A. pingi is here considered a synonym of A. pugettensis. However, if future moleculer and/or morphological analyses show any differences warranting recognition of the northeastern and northwestern populations as distinct species, A. pingi would likely be the valid name for northwestern Pacific specimens, assuming that all northwestern Pacific specimens are conspecific. However, the type specimens of A. pingi appear to be lost and a neotype would be required.