identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
FF0187ECFF91CE5FEAE7F895FAF5F7A7.text	FF0187ECFF91CE5FEAE7F895FAF5F7A7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pugettia quadridens (De Haan 1837) Ohtsuchi & Kawamura 2019	<div><p>Pugettia quadridens (De Haan, 1837) sensu stricto</p><p>[Japanese name: Yotsuha-mo-gani]</p><p>(Figs. 2–13)</p><p>Pisa (Halimus) quadridens De Haan, 1837: pl. 24, fig. 2 [type locality: Japan].</p><p>Pisa (Menoethius) [sic.] quadridens .—De Haan 1839: 97–98 (in part).— Yamaguchi &amp; Baba 1993: 353, fig. 113 (in part); 2003: 38 (in part).</p><p>Menaethius quadridens .— Adams &amp; White 1849: 20.</p><p>Pugettia quadridens .— Sakai 1936: 88 (in part), pl. 20, fig. 2, text-fig. 37; 1965: 73 (in part?), pl. 32, figs. 4, 5.— Yamaguchi et al. 1976: 35; 1987: 13, pl. 4, fig. 3a, b.— Ikeda 1981: 15.— Takeda 1982: 120, 1 unnumbered figure.— Griffin &amp; Tranter 1986: 97 (in part).— Yamaguchi &amp; Baba 1993: 353, fig. 113 (in part); 2003, 38 (in part).— Muraoka 1998: 24 (in part).— Minemizu 2000: 209, 1 unnumbered figure; 2002: 209, 1 unnumbered figure.— Marumura &amp; Kosaka 2003: 32 (in part).— Ikeda &amp; Kuramochi 2004: 12, 2 unnumbered figures.— Takeda et al. 2006: 196 (list); 2011: 49, fig. 16-69.— Ng et al. 2008: 101 (list).— Yamaguchi &amp; Henmi 2008: 80, figs. 1a, 2a–c, 3a–c.— Nunomura 2010: 52 (in part).— Wicksten &amp; Stachowicz 2013: 359 (list).— Watanabe 2014: 41, 1 unnumbered figure.— Yoshizaki 2018: 30 (list), 48 (with 11 unnumbered figures), 173 (1 unnumbered figure of frontal region), 181 (1 unnumbered figure of left chela).</p><p>Pugetti [sic.] quadridens .— Sakai 1938: 255–257 (in part), text-fig. 28a, pl. 26, fig. 1.</p><p>Pugettia quadridens quadridens .— Sakai 1976: 196–197 (in part), text-fig. 103a, pl. 68, fig. 1.— Miyake 1983: 206 (list); 1998: 206 (list).— Wada 1995: 387, pl. 103, fig. 5.</p><p>Pugettia nipponensis .— Miyake 1983: 36, pl. 12, fig. 5; 1998: 36, pl. 12, fig. 5. [not Pugettia nipponensis Rathbun, 1932]</p><p>Pugettia quadridens pellucens .— Marumura &amp; Kosaka 2003: 32 (in part). [not Pugettia quadridens pellucens Rathbun, 1932]</p><p>? Menaethius guadridens [sic.].—Matsuura 1895: 23.</p><p>? Pugettia quadridens .— Stimpson 1857: 219; 1907: 24–25.— Miers 1879: 23 (in part).— Ortmann 1893: 43 (in part?).— Rathbun 1894: 71–72 (in part?); 1902: 28 (in part).— Doflein 1902: 655–656.— Terazaki 1903: 15–16, unnumbered figure.— Parisi 1915: 285–286.— Balss 1924: 24 (in part).— Urita 1926: 32.—Kikuchi 1931: 19.— Sakai 1934: 294.— Kamita 1935: 63.— Kim et al. 1979: 110.— Kim &amp; Kim 1982: 146.— Kim &amp; Chang 1985: 45.— Komai 1999: 86.</p><p>? Pugettia quadridens quadridens .— Kim 1985: 79–80.— Kim &amp; Kim 1986: 325.— Ito &amp; Honma 2001: 27</p><p>Material examined. Lectotype: male (CL 24.2 mm) (RMNH D 42298), Japan, coll. H. Bürger, 1825–1834.</p><p>Paralectotypes: 4 females (CL 16.9–23.6 mm) (RMNH D 42298), same data as lectotype .</p><p>Non-types: Japan. One female (18.9 × 15.1 mm) (CBM-ZC 14872), near low tidal mark, Ahnfeltia paradoxa turf, Misaki, Iwaki, Fukushima, hand collection, coll. N. Ohtsuchi, 2 Feb. 2017; 1 male (18.3 × 14.1 mm), 1 ovigerous female (17.0 × 13.3 mm) (CBM-ZC 14873), Gelidium elegans turfs, near low tidal mark, Oarai, Kashima Sea, hand collection, coll. N. Ohtsuchi &amp; S. Houki, 8 May 2012 ; 2 males (18.9 × 15.0, 19.6 × 15.0 mm), 2 ovigerous females (17.4 × 13.5, 18.0 × 15.3 mm), 1 female (with a rhizocephalan parasite, 17.6 × 13.6 mm) (CBM-ZC 14874), 1 male (19.4 × 15.1 mm), 1 ovigerous female (21.0 × 16.6 mm) (NSMT-Cr 26063), A. paradoxa turfs, intertidal, Inubosaki, Chiba, hand collection, coll. N. Ohtsuchi &amp; S. Houki, 9 May 2012 ; 1 male (18.5 × 14.2 mm) (NSMT-Cr 26064), intertidal, Futomi, Kamogawa, Chiba, Boso Peninsula, hand collection, coll. N. Ohtsuchi, 9 Apr. 2005; 1 female (14.3 × 10.4 mm) (TOYA Cr-20553), Shirahama, Shirahama-machi, Chiba, coll. N. Nunomura, 18 Apr. 1995; 1 male (19.5 × 15.3 mm) (CBM-ZC 7581), intertidal, Okinosima, Tateyama, Chiba, Boso Peninsula, coll. Y. Matsuzawa, Aug. 2003 (exmined in Ohtsuchi et al. 2014); 3 males (13.9 × 10.4–19.1 × 14.7 mm), 2 ovigerous females (15.4 × 11.1, 18.0 × 13.5 mm) (NSMT-Cr 7561), Kohyatsu, Tateyama, Chiba, Boso Peninsula, coll. student of Ochanomizu Univ. (examined in Ohtsuchi et al. 2014); 4 males (19.7 × 15.4–22.9 × 18.0 mm), 2 females (20.5 × 15.9, 22.2 × 17.5 mm), 2 ovigerous females (20.7 × 15.9, 21.9 × 17.4 mm) (OMNH-Ar 10699), 5 males (14.5 × 11.5–20.3 × 15.9 mm), 3 females (17.6 × 14.1–20.1 × 16.6 mm), 1 ovigerous female (20.6 × 16.2 mm) (CBM-ZC 14875), Grateloupia cornea turfs, near low tidal mark, Tsurugizaki, Miura Peninsula, hand collection, coll. N. Ohtsuchi, J. Hayakawa &amp; S. Houki, 26 Feb. 2012 ; 3 males (6.9 × 4.7–8.9 × 6.2 mm) (CBM-ZC 14876), 2–4 m, Ge. elegans turfs, Nagai, Yokosuka, Miura Peninsula, SCUBA +air-lifting sampler, coll. J. Hayakawa &amp; T. Onitsuka, 10 Oct. 2008 ; 1 female (10.5 × 7.3 mm) (CBM-ZC 14877), 2–4 m, Marginisporum crassissimum turfs, same locality as previous, SCUBA +air-lifting sampler, coll. N. Ohtsuchi &amp; J. Hayakawa, 5 Apr. 2010 ; 1 female (11.2 × 8.1 mm) (NSMT-Cr 26065), Sargassum fusiforme beds, near low tidal mark, Nagai, Yokosuka, Miura Peninsula, hand collection, coll. N. Ohtsuchi &amp; S. Houki, 24 Oct. 2010; 1 female (16.7 × 12.8 mm) (RUMF-ZC-4338), same locality and habitat as previous, hand collection, coll. N, Ohtsuchi, 19 Apr. 2016; 1 male (16.7 × 12.8 mm) (RUMF-ZC- 4974), same locality and habitat as previous, hand collection, coll. N. Ohtsuchi, 11 Mar. 2017; 1 male (14.9 × 10.2 mm) (NSMT-Cr 16481), 6 m, Hayama Beach, Miura Peninsula, Sagami Bay, coll. I. Soyama, 27 Jan. 2005 (examined in Ohtsuchi et al. 2014) ; 2 males (21.6 × 17.6, 23.8 × 18.6 mm) (NSMT-Cr 26061), intertidal, S. fusiforme beds, rocky beach behind Morito Shrine, Hayama, Miura Peninsula, hand collection, coll. N. Ohtsuchi, 11 Apr. 2009; 1 male (27.3 × 22.4 mm) (NSMT-Cr 26059), 1 ovigerous female (21.8 × 18.2 mm) (NSMT-Cr 26060), 3 males (16.7 × 13.1–27.4 × 23.3 mm), 1 ovigerous female (21.1 × 16.0 mm) (NSMT-Cr 26062), same habitat and locality as previous, hand collection, coll. N. Ohtsuchi, 29 Apr. 2010; 1 male (12.1 × 8.9 mm) (KPM-NH 104997), Hayamacho, Miura-gun; 1 male (21.0 × 17.3 mm) (WMNH-Na-Cr 0312), Oiso, Kanagawa, Sagami Bay, coll. S. Nagai, Sep. 1970 ; 1 males (13.6 × 10.2 mm), 1 female (with rhizocephalan parasite, 13.3 × 10.2 mm) (KPM-NH 104582), Manazuru-machi, Ashigara-shimogun, 1962; 6 males (10.1 × 7.2–27.4 × 21.7 mm), 1 female (16.9 × 12.3 mm), 3 ovigerous females (17.1 × 13.4–19.4 × 14.9 mm) (KPM-NH 104045), 1 male (19.3 × 15.7 mm) (KPM-NH 104188), Sagami Bay; 1 ovigerous female (17.7 × 12.6 mm) (NSMT-Cr 6062), Izu Ohshima I., coll. I. Soyama, 6 May 1979 ; 5 males (4.3 × 2.9–6.2 × 4.4 mm), 3 females (4.2 × 2.9–6.2 × 4.5 mm), 4 juveniles (NSMT-Cr 26066), near low tidal mark, Ge. elegans turfs, Hinodehama, Izu Ohshima I., hand collection, coll. N. Ohtsuchi, 10 Aug. 2006; 1 male (25.9 × 19.1 mm) (NSMT-Cr 11306), Himaga I., Mikawa Bay, Mie, coll. T. Nishikawa, 7 Apr. 1993 ; 2 males (22.1 × 17.0, 23.2 × 18.3 mm) (KPM-NH 104513), Wagu, Shima, Shima, Mie; 1 male (20.6 × 16.0 mm) (OMNH-Ar 9916), Wagu, Tousi-cho, Toba-shi, Mie, coll. T. Yamamoto, 11 Apr. 1982; Three males (20.3 × 17.0–26.8 × 21.7 mm) (KPM-NH 104476), Kii Nagashima, Kitamuro-gun, Mie; 1 male (10.7 × 7.4 mm) (NSMT-Cr 11645), Kushimoto, Kii Peninsula, Wakayama, coll. I. Soyama, 21 Jun. 1994; 1 ovigerous females (17.7 × 13.2 mm), 1 females (with rhizocephalan parasite, 18.7× 14.2 mm) (NSMT-Cr 4894), Shirahama, Kii Peninsula, Wakayama, coll. Y. Koyama, 1 Apr. 1972; 2 males (3.2 × 4.6, 6.0 × 4.3 mm), 1 female (5.2 × 3.8 mm), 3 juveniles (NSMT-Cr 26067), Ge. elegans turfs, 1 m, Engetsu I., Banshozaki, Seto, Shirahama, Wakayama, snorkeling+hand collection, coll. N. Ohtsuchi, 28 Aug. 2015 ; 2 males (20.7 × 16.0, 21.8 × 17.1 mm) (SMBL-Art 1079), Seto, Shirahama-cho, Nishimuro-gun, Wakayama, 15 Apr. 1936; 2 males (15.3 × 11.0, 17.3 × 13.2 mm), 2 females (18.3 × 14.4, 15.2 × 11.9 mm) (SMBL-Art 1094), same locality as previous, 18 Feb.1939; 5 males (19.0 × 15.2–16.3 × 12.6 mm), 2 ovigerous females (19.3 × 14.2, 14.2 × 10.8 mm), 2 females (with a rhizocephalan parasite, 16.9 × 13.2, 16.5 × 12.9 mm) (SMBL-Art 1091), Ezura, Shirahama-cho, Nishimuro-gun, Wakayama, Jul. 1939; 1 male (24.9 × 19.8 mm) (KPM-NH 104081), 1 male (25.0 × 19.7 mm) (KPM-NH 104364), 4 males (22.8 × 18.2–28.4 × 22.4 mm) (KPM-NH 104378), 2 males (23.6 × 19.1, 26.2 × 20.7 mm) (KPM-NH 104448), 1 female (with rhizocephalan parasite, 10.9 × 8.2 mm) (KPM-NH 104614), 1 female (25.0 × 14.6 mm) (KPM-NH 104957), Minabe, Hidaka-gun, Wakayama; 2 females (with rhizocephalan parasite, 14.0 × 10.0, 14.1 × 10.6 mm) (KPM-NH 104624), Sakai, Minabe, Hidaka-gun, Wakayama; 1 male (13.1 × 9.7 mm) (OMNH-Ar 6644), Kuroshima, Yura-cho, Hidakagun, Wakayama, 3 May 1984; 2 females (10.4 × 7.7, 9.6 × 7.7 mm) (OMNH-Ar 6576), Nishihiro, Hirokawa-cho, Arida-gun, Wakayama, coll. T. Yamashita, 19 Aug. 2000; 1 ovigerous female (15.7 × 12.2 mm) (OMNH-Ar 2392), Tagurazaki, Kada, Wakayama-shi, 18 May 1980.— 6 males (13.9 × 10.2–22.5 × 18.4 mm) (OMNH-Ar 9913), same locality as previous, coll. Fukui, 24 Mar. 1974; 2 ovigerous females (15.6 × 12.2, 12.7 × 10.0 mm) (OMNH-Ar 3932), Minami-Tarumi, Tomogashima, Kada, Wakayama-shi, 26 Apr. 1997; 1 male (27.1 × 22.3 mm) (OMNH-Ar 524), 1 male (23.1 × 17.9 mm) (OMNH-Ar 525), Jogasaki, Kada, Wakayama, coll. Y. Fukui &amp; N. Nunomura, 19 May 1974; 1 ovigerous female (19.6× 15.8 mm) (OMNH-Ar 2384), Okawa-touge-shita, Okawa, Wakayama-shi, 13 Jul. 1980; 1 ovigerous female (18.6 × 14.4 mm) (OMNH-Ar 6212), Myojinzaki, Misaki-cho, Osaka, coll. H. Ariyama, 29 Apr. 1994; 1 male (18.8 × 14.0 mm), 1 female (12.2 × 9.4 mm) (OMNH-Ar 2448), bedrock, Toyokunizaki, Misaki-cho, Sennan-gun, 17 Feb. 1980; 2 males (18.0 × 13.7, 20.8 × 16.6 mm) (OMNH-Ar 9911), Nagasaki, Misaki-cho, Sennangun, Osaka, coll. R. Yamanishi, 11 May 1978; 4 males (23.5 × 18.2–15.3 × 11.6 mm), 2 ovigerous females (15.5 × 11.9, 18.6 × 14.5 mm) (OMNH-Ar 6024), Jogasaki (?), Misaki-cho, Osaka, coll. H. Ariyama, 16 Apr. 1995; 1 male (16.5 × 12.7 mm) (OMNH-Ar 2779), Yamatojima, Iwaya, Awaji-cho, Tsuna-gun, Hyogo, 8 May 1982; 1 ovigerous female (15.7 × 12.7 mm) (OMNH-Ar 5953), off Shioya, Tsuna-cho, Tsuna-gun, Hyogo, 7.8 m, coll. R. Yamanishi, 9 May 1985; 1 male (20.6 × 16.0 mm), 1 ovigerous female (16.4 × 12.9 mm) (OMNH-Ar 9917), Takino-chaya, Tarumi, Kobe, coll. Y. Shibata, 13 Mar. 1960; 1 male (19.7 × 14.4 mm) (CBM-ZC 8516), ca. 5 m, Kurahashi I., Hiroshima, Seto Inland Sea, commercial trawler, coll. K. Kuramoto, 13 Apr. 2005; 1 male (18.5 × 14.7 mm) (OMNH-Ar 6500), Nakamichi, Aio-cho, Yoshiki-gun, Yamaguchi, coll. T. Watanabe, 4 May 2003; 1 female (with rhizocephalan parasite, 15.2 × 11.8 mm) (OMNH-Ar 6512), same locality and date as previous, coll. K. Hatooka; 1 female (16.7 × 12.4 mm) (KPM- NH 104006), Matsuyama, Ehime, 1967; 1 male (32.1 × 26.0 mm) (KPM-NH 104903), Tosa Bay; 1 male (13.3 × 9.4 mm) (TOYA Cr-3230), 1 female (8.4 × 6.2 mm) (TOYA Cr-3231), Yokata, Toyama-shi, Toyama, coll. N. Nunomura, 8 Sep. 1982; 1 male (11.7 × 9.0 mm) (OMNH-Ar 9919), Shibagaki Kaigan, Hakui-shi, Ishikawa, 5 Aug. 1974; 1 male (24.1 × 19.2 mm) (TRPM-AAr-0000498), off Tottori, 4 Aug. 1998 (photographed in Takeda et al. 2011); 3 males (11.9 × 8.5–18.9 × 14.3 mm), 1 male (with rhizocephalan parasite, 12.4 × 9.9 mm) (NSMT-Cr 6771), 1 male (10.0 × 7.2 mm), 1 female (11.8 × 8.5 mm) (NSMT-Cr 6772), Tsuji I., Amakusa Archipelago, Kumamoto, coll. Y. Fukuda, 18 Mar. 1980; 1 male (19.5 × 14.2 mm) (KMNH IvR 100009), Shiroiwazaki, Tomioka, Amakusa, Kumamoto, Amakusa Sea, coll. Baba, 6 Apr. 1933; 1 ovigerous female (14.4 × 10.7 mm) (KMNH IvR 100010), Nomozaki, Nagasaki, Nagasaki Peninsula, Amakusa Sea, setnet around the rocky reaf, coll. K. Matsubayashi; 1 male (15.5 × 11.2 mm) (ZMUC-CRU- 20233), Nagasaki, coll. James Jordan (examined in Griffin &amp; Tranter 1986); 1 male (13.0 × 9.1 mm) (KMNH IvR 100012), 1 ovigerous female (14.2 × 11.0 mm) (KMNH IvR 100013), 1 ovigerous female (17.5 × 13.3 mm) (KMNH IvR 100014), Tsuyazaki, Fukutsu, Fukuoka, Genkai Sea, coll. Sakai, 23 Jun. 1960; 1 male (14.2 × 10.0 mm) (WMNH- Na-Cr 0312-2), lower tidal line, Kume-jima I., Okinawa Prefecture, Ryukyu Islands, southern Japan, coll. S. Nagai, 18 Nov. 1992 .</p><p>China. One male (16.2 × 12.6 mm) (MBM 160503), off east Ping Tan I., Fuzhou, Fujian, coll. Fan Xu, 18 Mar. 1957 .</p><p>Redescription. Male. Full-grown males (15.3–32.1 mm PCL, including lectotype). Carapace (Figs. 2A, 3A) pyriform, 1.2–1.4 longer than width (PCL/CW = 1.3±0.0, N = 52), surface smooth to naked eyes but closely covered with microscopic, apically flattened setae; gastric, cardiac, branchial, intestinal regions unclearly separated from each other. Gastric region (Figs. 2C, 3C) moderately elevated, sometimes with oblique row of dense hooked setae on either side of midline (Figs. 2A, C, 3A, C, 12B, C); mesogastric, metagastric, protogastric region on both sides each with rudimentary protuberance (Figs. 2A, 3A). Hepatic, cardiac, branchial regions (Figs. 2C, 3C) moderately elevated; mesobranchial region weakly elevated, not higher than gastric region, with two rudimentary tubercles apically, mesial one larger than lateral one (Figs. 2A, 3A); metabranchial region slightly elevated, without tubercles. Intestinal region (Figs. 2A, C, 3A, C) slightly elevated, fused with cardiac region.</p><p>Pseudorostral spines (Figs. 2A, 3A, C) short, length 0.2–0.3 of post-pseudorostral carapace length (PRL/PCL = 0.2±0.0, N = 33), each with two rows of dense, hooked setae on proximal half dorsally, single row of simple, long setae on proximal half mesially; lateral margins subparallel or slightly divergent. Preorbital spine (Figs. 3A, 4A, B) elongated, acuminate at tip, directed anterolaterally (Fig. 4A). Supraorbital eave (Figs. 3A, 4A, B) extended laterally, distinctly concave on lateral margin, weakly truncated on posterior end. Orbital hiatus (Figs. 2A, 3A, 4A) small, triangular concavity. Postorbital lobe (Figs. 2A, C, 3A, C, 4A) small, triangular, much shorter than preorbital spine, weakly compressed dorsoventrally, directed anterolaterally, incurved distally. Hepatic lobe (Figs. 2A, C, 3 A–C, 4A) not demarcated from hepatic region, broad, triangular, almost three times longer than postorbital lobe (HpL/PoL = 2.8±0.3, N = 9), compressed dorsoventrally, directed anteriorly, obtuse at tip, directed anterolaterally. Anterolateral carapace margin (Figs. 2A, 3A) often with short rows of hooked setae (Figs. 3C, 12B, C); lateral surface inferior to anterolateral margin with 1–3 (2 in general) spines (Figs. 2A, C, 3A, C). Epibranchial spine (Figs. 2A, 3A) slightly longer than, or as long as postorbital lobe, distinctly shorter than hepatic lobe, directed anterolaterally, slightly incurved, obtuse at tip, positioned at posterior 0.4 of postorostral carapace length (ESL/PCL = 0.4±0.0, N = 10), confluent to posterolateral carapace margin basally. Posterolateral carapace margin (Figs. 2A, 3A) faintly convex. Posterior carapace margin (Figs. 2A, 3A) projected roundly.</p><p>Subhepatic region (Fig. 3B) expanded anterolaterally, fully exposed in ventral view, sometimes with group of sparse, hooked setae. Pterygostomian region (Fig. 3B) not particularly inflated, with 3–5 (4 in general) papiliform tubercles along pleural suture. Anterolateral angle of buccal frame moderately produced anteriorly (Fig. 3C), not overlapped by anterolateral angle of merus of third maxilliped when closed.</p><p>Basal antennal article (Figs. 2B, D, 4B) smooth on surface, bearing low, blunt longitudinal ridge mesial to midline; distolateral angle moderately produced into small spine directed anterolaterally; lateral margin thickened dorsoventrally in distolateral part (Fig. 3C), bearing to posterior orbital margin, with low tubercle basally. Antennal peduncle (Fig. 4D) consisting of two articles; penultimate article (Figs. 2D, 4D) weakly compressed on lateral half, distal end as broad as proximal end; ultimate article (Fig. 4D) two-thirds of penultimate article in length, weakly compressed, slightly broadened distally.</p><p>Third maxilliped (Figs. 3B, 4E) smooth on surface. Ischium with shallow, broad median depression, lateral margin shallowly concave. Merus with dilated, moderately upturned anterolateral angle. Exopod more than half of ischium in maximum width, gradually narrowed in distal half, mesial margin with subacute angle on distal one-third (Fig. 4E).</p><p>Chelipeds (Figs. 2A, B, 3A, B) similar in size and shape. Ischium (Fig. 3B) strongly swollen ventrally in distal half; mesial margin obtusely ridged, without dentation; distolateral lobe distinct, compressed, rounded apically. Merus (Fig. 5 D–G) prismatic, length 2.5 longer than width (2.5±0.1, N = 11); dorsal surface (Fig. 5 D–F) with broad, longitudinal keel with 3–4 (3 in general) low, lamellar teeth, distalmost lowest; outer surface (Fig. 4 E–G) rugose, with unarmed, blunt longitudinal ridge; ventral surface (Fig. 5F, G) with blunt ridge bearing 2–3 (2 in general) low, broad teeth; inner surface (Fig. 5D, E, G) unarmed, irregularly rugose, with narrow, longitudinal ridge proximally ended in indistinct lobe; distal margin (Fig. 5 E–G) with 2 prominent knobs at articulation with carpus (outer knob larger than inner), prominent, obliquely erect, subrectangular lobe with short, acute projection on upper side. Carpus (Fig. 5 A–C) moderately inflated, with indistinct ridge on dorsal surface (Fig. 5A); outer margin obtusely ridged, divided into 2 lobes by broad concavity (Fig. 5A, B); ventral surface (Fig. 5C) smooth; inner margin obtusely crested, divided into broad, distal lobe and prominent, proximal lobe (Fig. 5A). Chelae (Fig. 6A) almost twice longer than high (ChL/ChH = 2.0±0.1, N = 29); palm strongly expanded, upper margin obtusely ridged, lower margin poorly defined; immovable fingers with subpentagonal teeth; movable finger uniformly dentate on distal two-thirds, with 2 large, isolated teeth on proximal one-third; fingers widely gaped in proximal two-thirds when closed (Figs. 2B, 6A).</p><p>Ambulatory legs (Figs. 3A, 7 A–C) decreasing in length posteriorly, surface generally smooth to naked eyes but closely covered with microscopic, apically-flattened setae. Meri subcylindrical, each with rudimentary, upper distal tubercle (Fig. 7A), more than five times longer than height in P2 (5.2±0.5, N = 11), more than three times in P3 (3.5±0.3, N = 11). Carpi each with faint, medial depression on extensor surface, most distinct in P2 (Fig. 7B). Propodi weakly flattened in P2 and P3, subcylindrical in P4 and P5, each with setal tufts on proximal 0.8 on flexor margin in P2, 0.6 in P3–5 (Fig. 7A, C). Dactyli each with two rows of large, calcareous spines on flexor surface (Fig. 7A, C).</p><p>Thoracic sternites (Figs. 2B, 3B, 8A, B) smooth on surface, with shallow, broad depression on second to fourth sternites on both sides (Fig. 8A); second sternite with pair of small depression anteriorly (often continuous to shallow depression on second to fourth sternites); third, fourth sternites faintly ridged medially (Fig. 8A); sterno-pleonal cavity without long setae on anterolateral margins (Fig. 8A).</p><p>Pleon (Figs. 2B, 3B, 8B) with six pleomeres and telson; third to sixth pleomeres fixed, with distinct suture. Third pleomere broadest, lateral margins arcuate; fourth pleomere trapezoid, shorter than fifth in midline length; fifth pleomere trapezoid; sixth pleomere rectangular, 0.6 of third pleomere in proximal width (0.6±0.0, N = 22); telson triangular.</p><p>Shaft of G1 (Fig. 9 A–D) straight, trilobate in distal one-sixth; dorsal lobe elongate triangular, more than twice longer than ventral lobe, weakly curved inwards (Fig. 9D); ventral lobe triangular, with subacute tip (Fig. 9B, C); mesial lobe as long as ventral lobe, projecting nearly perpendicular to dorsal lobe, strongly curled downwards (Fig. 9B, D); hiatus between dorsal, mesial lobes wide (Fig. 9D); mesial, lateral margins from dorsal to ventral lobe clearly concave medially; lateral margin higher than mesial margin, dilated in median part (Fig. 9 B–D). Shaft of G2 (Fig. 9E, F) stout, slightly narrowed distally, truncated apically; apex with relatively large, triangular, subacute projection.</p><p>Adolescent males (9.9–20.3 mm PCL) (Fig. 10A, B). Chelae (Fig. 6B) proportionally longer (ChL/ChH = 2.4±0.0, N = 11) than in full-grown specimens (Table 1), not gaped, both fingers uniformly dentate on cutting margin (Figs. 6B, 10A, B). Pleon subtriangular (PW6/PW3 = 0.6±0.0, N = 8) (Fig. 10B). G1 apically trilobate as in full-grown males.</p><p>Immature males (&lt;6.2 mm PCL) (Fig. 11A, B). Carapace relatively slender (PCL/CW = 1.4±0.1, N = 7) (Table 1). Chela similar to adolescent specimens in dentation on both fingers, but more slender (ChL/ChH = 2.7±0.1, N = 5) than in adolescent specimens (Table 1, see also Fig. 11A, B). Pleon subtriangular (PW6/PW3 = 0.7±0.0, N = 5) (Fig. 11B). G1 incompletely folded (Fig. 9I, J), otherwise folded but provided with undeveloped mesial lobe projecting anterolaterally (Fig. 9G, H).</p><p>Female. Full-grown females (12.7–22.2 mm PCL, including paralectotypes). Carapace (Fig. 3D, F) similar to males in general proportion (PCL/CW = 1.3±0.0, N = 27; HpL/PoL = 2.7±0.1, N = 12; ESL/PCL = 0.4±0.0, N = 12) (Student t -test, p&gt; 0.05); hepatic, mesobranchial regions more elevated than in males (Fig. 3D, F); anterolateral margins, protogastric, subhepatic regions usually with oblique rows of hooked setae (Fig. 3 D–F). Cheliped merus slightly slenderer than in males (length/height = 2.6±0.1, N = 7) though there was no significant difference (Student t -test, p = 0.12); chelae (Fig. 6C) smaller, more slender than in full-grown males (ChL/ChH = 2.6±0.1, N = 17; Student t -test, p &lt;0.01), both fingers uniformely dentated on cutting edges, not gaped when closed. Tufts of few elongate setae sometimes on midline of gastric region, midpoint of metabranchial region, apex of epibranchial spines, summits of cardiac and intestinal regions (Fig. 13C). Pleon (Figs. 3E, 13D) with six pleomeres and telson, expanded (PW6/PW3 = 1.6±0.1, N = 10). Gonopores (Fig. 8D, E) comma-shaped, nearly circular in lateral twothirds, elongate in mesial one-third.</p><p>Adolescent females (9.6–16.9 mm PCL) (Fig. 10C, D). Chela more slender than in adolescent males (ChL/ChH = 2.6±0.0, N = 5; Student t -test, p = 0.02) (Fig. 10D), both fingers uniformly dentate as in full-grown individuals. Pleon ovate (PW6/PW3 = 1.2±0.1, N = 3) (Fig. 10D).</p><p>Immature females (&lt;6.2 mm PCL) (Fig. 11C, D). Carapace relatively slender (PCL/CW = 1.4±0.0, N = 3) (Table 1). Chela slender (ChL/ChH = 2.7±0.0, N = 3) (Table 1, see also Fig. 11D). Pleon generally rectangular, with subtriangular telson, lateral margins subparralel (PW6/PW3 = 1.2±0.1, N = 3) (Table 1, see also Fig. 11D).</p><p>Variations. Carapace gets broader (PCL/CW decreases from 1.4 to 1.2) in relation to size growth in both sexes (Table 1; see also Figs. 3, 10, 11, 12). Gastric, mesobranchial, and metabranchial regions are less elevated in smaller specimens, often missing tubercles or protuberances on each surface. Mesobranchial region is rarely elevated (Fig. 12C). Supraorbital eave rarely straight on lateral margin (Fig. 12B). Hepatic lobes are sometimes laminate (Fig. 12 B–D), relative length against postorbital lobe (HpL/PoL) increases from 2.0 to 3.6 in relation to size growth (Fig. 39). Groups of hooked setae on either side of gastric region and anterolateral margin of carapace, as well as elongate setae on the tops of mesobranchial, intestinal regions, are likely to be reduced in large specimens in both sexes (Figs. 3A, 13A vs. Figs. 9A, 10A, 12 A–C). Ambulatory legs often with a few, elongate setae on upper margins of meri, upper, lower surfaces of carpi in small specimens of both sexes (Figs. 11, 13F).</p><p>Size. Largest male: 32.1 × 26.0 mm; largest female: 20.9 × 16.1 mm; smallest ovigerous female: 11.1 × 8.9 mm (Ohtsuchi et al. 2018; this study).</p><p>OS, ontogenetic stage; N, sample size; PCL, postrostral carapace length; CW, maximum carapace width; PRL, length of pseudorostral spine; ChL, chela length; ChH, chela height; PW3, PW6, proximal width of pleomere 3 and 6, respectively. Means in the same row followed by different small letters are significantly different from each other (Tukey-Kramer HSD test or Steel-Dwass test, p &lt;0.05). Means in the same line followed by diffferent capital letters are significantly different from each other (Tukey-Kramer HSD test or Steel-Dwass test, p &lt;0.05). Differently colored boxes indicate significant difference between sexes (Student t -test or Welch two samples t -test, p &lt;0.05).</p><p>Coloration in life. Carapace (Fig. 13A, D, F) generally dark red, reddish brown, or dark green, each simlar to general coloration of collected habitats (Fig. 13C); small specimens with some blotches and/or dense speckles (Fig. 13F); female specimen often with irregular speckles, mottling even in full-grown specimens (Fig. 13D); patterning reduced in males especially in large specimens (Fig. 13A). General coloration of thorax, pleon, chelipeds, ambulatory legs similar to carapace in males (Fig. 13B), often slightly reduced in third maxilipeds, thoracic plastron, flexor surface of cheliped meri, ambulatory leg meri in females (Fig. 13E). Cheliped meri, palm often with scale-like pattering in full-grown males (Fig. 13B). Ambulatory legs with whitish band on the joint of each articulation and the distal parts of dactyli. See also Miyake (1983, 1998: pl. 12, fig. 5, as P. nipponensis), Wada (1995: pl. 103, fig. 5), Minemizu (2000, 2002: 209, unnumbered figure), Watanabe (2014: unnumbered figure), and Yoshizaki (2018: 48, 173, 181, unnumbered figures).</p><p>Distribution. Pacific coast of Japan, from Iwaki, Fukushima to Osaka Bay, including Izu Ohshima Island; Seto Inland Sea; coast of Sea of Japan from Toyama Bay to Nagasaki; East China Sea from Amakusa Archipelago to southeast coast of China mainland (Fuzhou), including Kume-jima Island, Ryukyu.</p><p>Habitat. Intertidal to 8 m depth. Commonly found in various kind of macroalgal communities developed on rocky reefs, such as brown algal beds ( Sargassaceae: Sargassum fusiformes, S. hemiphyllum, and S. hornei; and Dictyotacea: Padina alborescens) (Fig. 13C), red algal turfs ( Phyllophoraceae: Ahnfeltiopsis paradoxa; Gelidiaceae: Gelidium elegans; and Halymeniaceae: Grateloupia cornea), and sometimes articulated coralline algal turfs ( Corallinaceae: Corallina pilulifera, C. crassissima) (Sato &amp; Wada 2000; Ohtsuchi et al. 2018; this study).</p><p>Decorating materials. Pieces of various macroalgae, mainly of Sargassum spp. or branched red algae on the coast of Shirahama, Kii Peninsula, and Nagai, Sagami Bay (Sato &amp; Wada 2000; this study).</p><p>Ecological notes. This species was suggested as herbivorous (Sato &amp; Wada 2000), but we found they infrequently feed on the sea urchin Hemicentrotus pulcherrimus (A. Agassiz, 1864) under captive conditions (N. Ohtsuchi pers. obs.). Juveniles and smaller individuals usually inhabit subtidal turfs of various branched red algae e.g. gelidiaceans and Grateloupia cornea, and the majority move to Sargassum beds near low tidal marks on the coast of Miura and Kii Peninsulas (Ohtsuchi et al. 2018). On the other hand, the individuals of all the ontogenetic stages were collected sympatrically from dense turfs of red algae ( Ahnfeltia paradoxa, Gelidium elegans, and G. cornea) near low tidal mark on the rocky shore coast of Iwaki (Fukushima, Pacific coast of northeast Japan), Oarai (Ibaraki, Pacific coast of east Japan), and Tsurugizaki (Kanagawa, Miura Peninsula, Pacific coast of east Japan).</p><p>Remarks. De Haan (1837, 1839) did not designate any type when the species was described. The larger male in the syntype materials (RMNH D 42298) was subsequently designated as the lectotype by Yamaguchi &amp; Baba (1993: 353). The lectotype (Fig. 2 A–D) is a large male with enlarged chelipeds, viz. full-grown male. It is remarkable that the movable finger has two large, isolated teeth subproximally (Fig. 2B), which is also shown in the plate of De Haan (1837: pl. 24, fig. 2, as Pisa (Halimus) quadridens). On the other hand, the smaller male, which was designated as paralectotype by Yamaguchi &amp; Baba (1993: fig. 113a), is considerably different from the lectotype in many morphological characters, and it should have been attributed to P. intermedia instead (Fig. 2E, F).</p></div>	https://treatment.plazi.org/id/FF0187ECFF91CE5FEAE7F895FAF5F7A7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Ohtsuchi, Naoya;Kawamura, Tomohiko	Ohtsuchi, Naoya, Kawamura, Tomohiko (2019): Redescriptions of Pugettia quadridens (De Haan, 1837) and P. intermedia Sakai, 1938 (Crustacea: Brachyura: Epialtidae) with description of a new species. Zootaxa 4672 (1): 1-68, DOI: 10.11646/zootaxa.4672.1.1
FF0187ECFF80CE6AEAE7F9CEFDB0F1C3.text	FF0187ECFF80CE6AEAE7F9CEFDB0F1C3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pugettia intermedia Sakai 1938	<div><p>Pugettia intermedia Sakai, 1938</p><p>[Japanese name: Yotsuha-modoki]</p><p>(Figs. 2E, F, 14–24)</p><p>Pisa (Menoethius) [sic.] quadridens . —De Haan, 1839: 97–98 (in part, as paralectotype).— Yamaguchi &amp; Baba 1993: 353, fig. 113; 2003: 38 (in part, as paralectotype). [not Pisa (Halimus) quadridens]</p><p>Pugettia minor .— Shen 1937: 287–289, text-fig. 5a, d, f, g. [not Pugettia minor Ortmann, 1893]</p><p>Pugettia quadridens intermedia Sakai, 1938: 258, pl. 36, fig. 2; 1965, 72, pl. 32, fig. 3; 1976: 197, text-fig. 103b [type locality: Simoda (= Shimoda, Izu Peninsula)].— Miyake 1983: 206 (list); 1998: 206 (list).— Ariyama 1995: fig. 2, 3a–c, 4.</p><p>Pugettia similis .— Sakai 1976: 200, text-fig. 107b [not Pugettia similis Rathbun, 1932]</p><p>Pugettia intermedia .— Griffin &amp; Tranter 1986: 93–95, fig. 28a, b.— Yamaguchi et al. 1987: 13, pl. 4, fig. 2.— Muraoka 1998: 24 (in part), tbl. 1 (not holotype).— Marumura &amp; Kosaka 2003: 32 (in part).— Ng et al. 2008: 101 (list).— Yamaguchi &amp; Henmi 2008: 80, figs. 1b.— Wicksten &amp; Stachowicz 2013: 359 (list).</p><p>Pugettia quadridens .— Yamaguchi &amp; Baba 1993: 353, fig. 113; 2003: 38 (in part).— Muraoka 1998: 24 (in part).— Nabeshima 2011: 124, 1 unnumbered figure; 2013: 124, 1 unnumbered figure [not Pisa (Halimus) quadridens De Haan, 1837]</p><p>Pugettia quadridens pellucens . — Marumura &amp; Kosaka 2003: 32 (in part) [not Pugettia quadridens pellucens Rathbun, 1932] Pugettia vulgaris . — Yang et al. 2015: 203–206, figs. 1E, F, 3. [not Pugettia vulgaris Ohtsuchi, Kawamura &amp; Takeda, 2014]</p><p>? Pugettia quadridens intermedia .— Ikeda 1981: 15 (in part).— Kim &amp; Kim 1986: 325.</p><p>? Pugettia intermedia .— Watanabe 2014: 41, unnumbered figure.</p><p>Material examined. Paratype: 1 male (13.9 × 12.5 mm) (KPM-NH 124160), Shimoda, Izu Peninsula, coll. Misago. [not holotype though listed in Muraoka (1998: 24, table 1)]</p><p>Non-types: Japan. One male (RMNH D 42298), Japan, coll. H. Bürger, 1825–1834 (as one of paralectotypes of Pisa (Menaehius) quadridens De Haan, 1839); 1 male (15.9 × 12.0 mm) (CBM-ZC 2574), 50–60 m, off Takeoka, Uchibo, Boso Peninsula, gill-net, coll. T. Komai, 1 May 1996 ; 1 male (26.5 × 20.9 mm) (CBM-ZC 5128), 30–40 m, same locality as previous, gill-net, coll. T. Komai, 1 Feb. 1998; 1 ovigerous female (14.1 × 10.9 mm) (KPM-NH 110375), 1 ovigerous female (12.9× 10.1 mm) (KPM-NH 110376), Sagami Bay; 2 males (28.0 × 23.1, 28.4 × 23.8 mm) (WMNH-Na-Cr 0314-2), Himaga I., Aichi, coll. S. Nagai; 1 male (25.0 × 19.7 mm) (KPM-NH 104634), same locality as previous; 1 male (20.2 × 16.9 mm) (KPM-NH 104520), Shima, Mie; 1 male (9.5 × 6.4 mm), 3 females (6.8 × 4.8–9.5 × 6.8 mm) (WMNH-Na-Cr 313-1), 30 m, Shionomisaki, Kii Peninsula, Wakayama, coll. S. Nagai ; 3 males (3.9 × 2.7–12.1 × 8.9 mm) (OMNH-Ar 9928), Nishihiro, Hirogawa-cho, Arida-gun, Wakayama, coll. T. Yamahsita, 19 Aug. 2000 ; 1 male (18.4 × 13.8 mm) (KPM-NH 104049), Minabe, Hidaka-gun, Wakayama (examined in Sakai 1976) ; 1 male (13.4 × 9.8 mm) (KPM-NH 110363), Sakai, Minabe, Hidaka-gun, Wakayama ; 1 male (23.4 × 19.6 mm) (OMNH-Ar 10700), 1 male (13.6 × 9.7 mm) (OMNH-Ar 10701), 1 ovigerous female (20.0 × 15.6 mm) (OMNH-Ar 10702), 1 ovigerous female (15.7 × 12.3 mm) (OMNH-Ar 10703) Yura Fishery Port, Yura, Sumoto, Hyogo, 2 m, trap, coll. T. Watanabe, 3 Jun. 2017 ; 1 ovigerous female (19.5 × 14.9 mm) (OMNH-Ar 6188), 34°21´N– 135°06´E, near large fishing bank, 35–55 m, off Misaki-cho, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=135.1&amp;materialsCitation.latitude=34.35" title="Search Plazi for locations around (long 135.1/lat 34.35)">Sennan-gun</a>, Osaka, coll. H. Ariyama, 26 Mar. 1996 ; 4 males (18.4 × 14.0–26.7 × 21.1 mm), 3 ovigerous females (16.6 × 13.1–22.9 × 19.3 mm) (OMNH-Ar 6217), Nisikiminami-machi, Hannan roku-ku, Kaizuka-shi, Osaka, coll. H. Ariyama, 28 Mar. 1995 ; 1 male (22.9 × 18.9 mm) (OMNH-Ar 6117), off Sakai-shi, Osaka, stone dredge net, coll. H. Ariyama, 29 May 2000 ; 2 males (26.3 × 21.8, 28.1 × 24.2 mm) (OMNH-Ar 6199), from Kobe-shi, Hyogo, to Sakai-shi, Osaka, Osaka Bay, stone dredge net, coll. H. Ariyama, 15 Feb. 1995 (examined in Ariyama 1995) ; 1 female (28.5 × 23.6 mm) (OMNH-Ar 6208), 2 km off the coast of Kobe-shi to Ashiya-shi, Hyogo, Osaka Bay, coll. H. Ariyama, 8 Feb. 1994 (examined in Ariyama 1995) ; 1 ovigerous female (23.7 × 19.4 mm) (OMNH-Ar 6198), inner part of Osaka Bay, coll. H. Ariyama, 16 Mar. 1998 ; 1 male (19.1 × 15.3 mm), 1 ovigerous female (26.4 × 20.9 mm) (OMNH-Ar 6207), same locality as previous, coll. H. Ariyama, 22 Feb. 1996; 1 male, (21.6 × 16.1 mm) (OMNH-Ar 6037), off Nankou, Suminoe-ku, Osaka-shi, Osaka, coll. H. Ariyama, 4 Mar. 1993 (examined in Ariyama, 1995) ; 3 males (5.1 × 3.8–7.9 × 5.7 mm), 2 females (7.3 × 5.1, 8.8 × 6.3 mm) (OMNH-Ar 6025), near Osaka Aquarium Kaiyukan, Osaka, coll. H. Ariyama, Jun. 1998 ; 1 ovigerous female (11.2 × 8.7 mm) (OMNH-Ar 3805), subtidal zone, Maiko, Tarumi-ku, Kobe-shi, Hyogo, coll. R. Yamaguchi, 15–17 Jun. 1993 ; 1 female (15.3 × 11.6 mm) (OMNH-Ar 9922), Shizuki, Awaji-shi, Hyogo, coll. Y. Nakajima &amp; Y. Fukui, 30 Jun. 1974 ; 1 ovigerous female (14.7 × 11.5 mm) (OMNH-Ar 10704), off Karakoto, Kurashiki, Okayama, 0.3 m, muddy sand bottom, coll. T. Watanabe, Apr. 2017 ; 1 male (19.5 × 15.3 mm) (KPM-NH 104142), Tosa Bay; 1 female (21.3 × 17.0 mm) (TOYA Cr 10297), 20 m, off Mizuhashi, Toyama-shi, Toyama Bay, coll. N. Miyamoto, 1 Jul. 1990 ; 2 males (16.8 × 13.0, 21.1 × 16.6 mm), 3 ovigerous females (19.3 × 15.8–26.4 × 21.8 mm) (NSMT-Cr 26068), Maizuru Fisheries Research Station, Kyoto University, Maizuru, Tango Sea, coll. K. Sakemi, 13–17 Mar. 2013 ; 2 males (11.3 × 8.3, 18.1 × 14.3 mm) (TRPM-AAr-0000499), Makiya, Iwami, Tottori, coll. Y. Hirata, 20 Sep. 2009 ; 1 female (with rhizocephalan parasite, 16.7 × 12.8 mm) (KMNH IvR 100011), Nomozaki, Nagasaki, Nagasaki Peninsula, Amakusa Sea, setnet around the rocky reef, coll. K. Matsubayashi, Apr. 1961 ; 1 male (10.8 × 8.1 mm) (KMNH IvR 100008), Futae, Amakusa Archipelago, trawl net, coll. K. Baba, 26 Feb. 1935 ; 1 ovigerous female (15.4 × 11.4 mm) (OMNH-Ar 4271), Koujiro, Kunimi-cho, Nagasaki, 16 May 1999 .</p><p>East China. One female (21.3 × 18.7 mm) (MBM 160490), around the Qingdao No. 2 high school, coll. Baoling Wu, 23 Dec. 1963; 1 male, (22.9 × 18.2 mm) (MBM 160493), No. 1 Bathing Beach, Qindao, coll. Yang &amp; Zhang, 5 Dec. 1956 ; 1 male (26.2 × 20.7 mm), 1 ovigerous female (22.0 × 17.5 mm) (ZMUC-CRU-20234), Formosa Strait, 23°08'N 117°30'E, 43 m, coll. Kapt. Svenson, 23 Jan. 1912 (figured by Griffin &amp; Tranter 1986); 2 females (7.3 × 5.1, 6.1 × 4.2 mm) (ZMUC-CRU-20235), Formosa Strait, 23°57'N 118°33'E, 50 m, April 1897 (examined in Griffin &amp; Tranter 1986).</p><p>Redescription. Male. Full-grown males (18.4–28.6 mm PCL). Carapace (Fig. 15A) pyriform, 1.3 longer than width (PCL/CW = 1.3±0.1, N = 18), surface microscopically tomemtose, sparsely with elongate setae; gastric, cardiac, branchial, intestinal regions moderately separated from each other. Gastric region (Fig. 15A, C) moderately elevated, always anteriorly with oblique row of hooked setae on either side of midline, posterior end positioned posterior to hepatic lobe basis (Figs. 15C, 24C, D); mesogastric, metagastric, protogastric region on both sides each with distinct protuberance (Figs. 15A, C, 24A). Hepatic region not markedly elevated, sparsely with long setae. Cardiac, branchial regions (Figs. 15A, C, 24C, D) moderately elevated; mesobranchial region elevated, as high as gastric, cardiac regions, with 2 distinct, obtuse tubercles apically, mesial one larger than lateral one; metabranchial region faintly elevated, with low, obtuse tubercle apically. Intestinal region moderately elevated, separated from cardiac region, with obtuse protuberance apically (Figs. 15C, 24C, D). Tufts of a few elongate setae on 2 medial protuberances of gastric region, midpoint of epibranchial region, apex of epibranchial spines, summits of cardiac, intestinal regions (Figs. 15A, C, 24D).</p><p>Pseudorostral spines (Fig. 15A) relatively long, length 0.2–0.3 of post-pseudorostral carapace length (PRL/ PCL = 0.3±0.0, N = 17), each with two rows of sparse, hooked setae on proximal 0.8 dorsally, nearly connected to rows on protogastric region, single row of simple, long setae on proximal 0.8 mesially; lateral margins divergent anteriorly. Preorbital spine (Figs. 15A, C, 16A) triangular, acute at tip, with row of slender setae on mesial margin, compressed dorsoventrally, directed anterolateraly. Supraorbital eave (Figs. 15A, 16A, B) moderately extended laterally, slightly concave on lateral margin, distinctly truncated on posterior end (Fig. 16A). Orbital hiatus (Figs. 15A, B, 16A) large, deep, subrectangular sulcus in dorsal view. Postorbital lobe (Figs. 15A, B, 16A) elongate, triangular, subequal to or slightly shorter than supraorbital spine, compressed dorsoventrally, directed anterolateraly, slightly incurved distally. Hepatic lobe (Figs. 15A, 16A) faintly demarcated from gastric region, elongate, twice or more than twice longer than postorbital lobe (HpL/PoL = 2.1±0.2, N = 8), directed anterolaterally, acuminate at tip, directed anterolaterally, lateral margin bent outwards in distal half; concavity between postorbital, hepatic lobes extended as low, lamellar plate. Anterolateral carapace margin (Fig. 15A, C) with broad patch of sparse, hooked setae; lateral surface inferior to anterolateral margin with 2–5 (3 in general) subacute tubercles. Epibranchial spine (Fig. 15A) large, as long as postorbital lobe, distinctly shorter than hepatic lobe, directed laterally, slightly incurved distally, acuminate at tip, positioned at posterior 0.3 of post-pseudorostral carapace length (ESL/PCL = 0.3±0.0, N = 8), distinct from posterolateral carapace margin at base. Posterolateral carapace margin (Fig. 15A) faintly convex. Posterior carapace margin (Fig. 15A) projected roundly.</p><p>Subhepatic region (Fig. 15B) narrowly exposed in ventral view, with row of sparse, hooked setae. Pterygostomian region not markedly inflated, with 4–7 (4–5 in general) subacute tubercles along pleural suture. Anterolateral angle of buccal frame (Figs. 15C) produced anteriorly, not overlapped by anterolateral angle of merus of third maxiliped when closed, subrectangular in lateral view.</p><p>Basal antennal article (Figs. 15B, 16B) smooth on surface, bearing low, blunt longitudinal ridge mesial to midline; distolateral angle produced into small, subacute spine directed anterolaterally; lateral margin extended laterally, sinuous, truncate on proximal end, distinct from posterior orbital margin, with strong tubercle basally (Fig. 16B). Antennal peduncle (Fig. 16D) consisting of two articles; penultimate article (Fig. 16D) generally subcylindrical, with lateral, mesial margins carinate over entire length, distal end almost twice broader than proximal end; ultimate article (Fig. 16D) two-thirds of penultimate article in length, flattened dorsoventrally, slightly broadened distally; penultimate, ultimate articles with few, noticeably long setae on distomesial angle.</p><p>Third maxilliped (Figs. 15B, 16E) tomentose. Ischium with broad median groove, lateral margin nearly straight. Merus with dilated, upturned anterolateral angle. Exopod almost half of ischium in maximum width, immediately narrowed in distal one-third, compressed in lateral half, mesial margin with blunt angle on distal one-third (Fig. 16E).</p><p>Chelipeds (Figs. 15A, B, 17, 18) equal in size, similar in shape. Ischium (Fig. 15B) weakly swollen ventrally in distal half; mesial margin acutely ridged, cut into 4–6 teeth by faint sinus, distalmost one distinct; distolateral lobe distinct, compressed, rounded apically. Merus (Fig. 17 D–G) prismatic, length more than twice longer than height (2.3±0.2, N = 4); dorsal surface (Fig. 17 D–F) with narrow, longitudinal keel with 3–6 (3 in general) lamellar teeth, distalmost largest, directed anteriorly; outer surface (Fig. 17 E–G) faintly rugose, with blunt, logitudinal ridge, with 4–7 low tubercles; ventral surface (Fig. 17D, F, G) with blunt ridge bearing three low teeth; inner surface (Fig. 17D, E, G) irregularly rugose, with blunt, longitudinal ridge teminated in subtriangular, proximal lobe; distal margin (Fig. 17 E–G) with 2 prominent knobs at articulation with carpus (outer knob as large as inner), prominent, obliquely erect, triangular lobe terminated in acuminate tip on upper side. Carpus (Fig. 17 A–C) moderately inflated, with blunt ridge bearing 3–4 tubercles on dorsal surface (Fig. 16A); outer margin obtusely ridged, devided into 4 lobes, both terminal lobes broader than two median lobes (Fig. 17A, B); ventral surface with two tubercles (Fig. 17B, C); inner margin thinly crested, irregularly dentate, with round, proximal lobe (Fig. 17A). Chelae (Fig. 18A) more than twice longer than height (ChL/ChH = 2.2±0.2, N = 15); palm strongly expanded, upper margin obtusely ridged, lower margin poorly defined; both fingers bearing broad, subpentagonal teeth along distal two-thirds (proximal 2 or 3 broad), gaped widely in proximal half when closed (Fig. 18A).</p><p>Ambulatory legs (Figs. 15A, B, 19) decreasing in length posteriorly, surface generally tomentose. Meri subcylindrical, each with distinct, upper distal tubercule (Fig. 19A), almost six times longer than height in P2 (5.8±0.5, N = 5), more than four times in P3 (4.4±0.1, N = 3), with tufts of elongate setae on extensor surface, upper, lower flexor margins. Carpi each with deep, medial depression on extensor surface, with tufts of elongate setae on upper, lower flexor margins (Fig. 19B). Propodi weakly flattened, moderately narrowed in flexor half, each with pair of cluster of elongate setae on proximal half of extensor margin (Fig. 19A, C). Dactyli each with two rows of low, calcareous spines on flexor surface in P3–5, unarmed in P2 (Fig. 19A, C).</p><p>Thoracic sternites (Fig. 20A, B) tomentose on surface, with deep, narrow, rectangular depression on second to fourth sternites on both sides (Fig. 20A); first sternite with broad, median depression; second sternite with pair of small depression anteriorly; third to fourth sternites obutusely ridged medially (Fig. 20A); sterno-pleonal cavity sparsely with long setae on anterolateral margin (Fig. 20A).</p><p>Pleon (Fig. 20B) with six pleomeres and telson; third to sixth pleomeres fixed, with distinct suture. Third pleomere broadest, lateral margins oblique; fourth, fifth pleomeres trapezoid; sixth pleomere rectangular, dilated on distolateral angle, 0.7 of third pleomere in proximal width (0.7±0.0, N = 4); telson triangular.</p><p>Shaft of G1 (Fig. 21A, E) sinuous, trilobate in distal one-fifth; dorsal lobe elongate, triangular, more than twice longer than ventral lobe, weakly curved inwards (Fig. 21 B–D); ventral lobe triangular, with subacute tip, pointing upwards (Fig. 21B, C, F, G); mesial lobe elongate, as long as, or longer than dorsal lobe, directed anteriorly, crossing with dorsal lobe at tip (Fig. 21 A–C); hiatus between dorsal, mesial lobes narrow (Fig. 21D); mesial, lateral margins from dorsal to ventral lobes concave in median part; lateral margin as high as mesial margin, dilated (Fig. 21 B–D, F, G). Shaft of G2 (Fig. 21H, I) stout, narrowed distally, truncated apically; apex with elongate, finger-like projection.</p><p>Adolescent males (12.1, 16.8 mm PCL) (Figs. 14A, B, 22A, B). Chelae (Fig. 18C) proportionally shorter (ChL/ ChH = 2.6±0.1, N = 9) than in full-grown males (Table 1), not gaped, both fingers uniformly dentate on cutting margin (Figs. 18C, 23A, B). G1 apically trilobate as in full-grown males.</p><p>Immature males (&lt;7.5 mm PCL) (Fig. 23A, B). Carapace relatively slender (PCL/CW = 1.4±0.1, N = 3). Chela more slender (ChL/ChH = 2.9±0.1, N = 3) than in adolescent specimens (Table 1); dentation on both fingers similar to adolescent specimens (Fig. 23B). G1 incompletely folded, otherwise folded but provided with short, mesial lobe pointing upwards (Fig. 21J, K).</p><p>Female. Full-grown females (11.2–28.5 mm PCL). Carapace (Fig. 15D, F) similar to males in general proportion (PCL/CW = 1.3±0.1, N = 18; HpL/PoL = 2.2±0.2, N = 7; ESL/PCL = 0.3±0.0, N = 7) (Student t -test, p&gt; 0.05; hepatic, mesobranchial regions more elevated than in males (Fig. 15D, F). Pseudorostral spine shorter than in males (PRL/PCL = 0.2±0.0, N = 13) (Table 1, see also Fig. 15F) though there was no significant difference (Student t -test, p &lt;0.10). Cheliped merus more slender than in males (2.8±0.2, N = 5; Student t -test, p = 0.01); chelae (Fig. 18D) smaller, more slender than in full-grown males (ChL/ChH = 3.0±0.2, N = 15; Student t -test, p &lt;0.01), both fingers uniformely dentated on cutting edges, not gaped when closed. Pleon (Fig. 15E) with six pleomeres and telson, expanded (PW6/PW3 = 1.5±0.1, N = 5), fringed with short setae densely. Gonopores (Fig. 20D) generally elongate.</p><p>Adolescent females (15.3 mm PCL) (Fig. 22C, D). Chela slender (ChL/ChH = 2.7, N = 1), both fingers uniformly dentate as in full-grown individuals. Pleon rounded triangular, narrowed toward semicicular telson, lateral margins divergent (PW6/PW3 = 1.3, N = 1).</p><p>Immature females (&lt;8.8 mm PCL) (Fig. 23C, D). Carapace relatively slender (PCL/CW = 1.4, N = 2). Chela slender (ChL/ChH = 2.9, N = 2). Pleon subtriangular, narrowed toward triangular telson, lateral margins arcuate (PW6/PW3 = 1.1–1.3, N = 2).</p><p>Variations. Carapace tends to be broader (PCL/CW decreases from 1.4 to 1.1) in relation to size growth in both sexes (Table 1; see also Figs. 14, 15, 22–24); the surface is often densely covered with short setae, especially in females (Figs. 15D, F, 23C, 24D). Gastric, mesobranchial, and metabranchial regions are less elevated, and the tuberculations on each region are less distinct in small specimens even in the same ontogenetic stage (Figs. 14, 22, 23, 24A, C). HpL/PoL increases from 1.4 to 2.6 in relation to size growth (Fig. 39). Both fingers of chela rarely do not contact in distal half in small full-grown individuals (Fig. 18B). Spinulation on flexor surface of ambulatory leg dactyli is often reduced or abladed in large specimens. Mesial lobe of G1 is variable in length.</p><p>Size. Largest male: 28.4 × 23.8 mm; largest female 22.0 × 17.5 mm; smallest ovigerous female 12.9 × 10.1 mm.</p><p>Coloration in life. Based on a few specimens from Sumoto, Osaka Bay. The carapace generally deep red or dark brown, variably with striking patterning by numerous whitish or dark-colored blotches, speckles (Fig. 25A, C, E). The thorax, pleon, flexor surface of chelipeds and ambulatory legs white, or whitish light brown (Fig. 25B, D). The meri of the cheliped with irregular dark brown band in small specimens (Fig. 25E). Chelae generally olive green, both fingers dark brown at basis (Fig. 25A, D). Ambulatory legs often with whitish band on meri and propodi. Coloration probably variable, see also Sakai (1965: pl. 32 fig. 3), and Nabeshima (2011, 2013: 124, unnumbered figure).</p><p>Distribution. Japan, Pacific coast from Boso Peninsula to Tosa Bay, Osaka Bay, Seto Inland Sea, Sea of Japan coast from off Oga Peninsula to Nagasaki, Goto Islands, Amakusa Archipelago; Gangwon-do, northeast coast of South Korea (Yang et al. 2015, as P. vulgaris); both coast of Formosa Strait (Griffin &amp; Tranter 1986); off Qindao and Jiaozhou Bay, north China (Shen 1937, as P. minor; this study).</p><p>Habitat. From intertidal to 40 m depth, among algal turfs on the rock reefs, boulder zone, scalop farm (Sakai 1938, 1976; Yang et al. 2015, as P. vulgaris). Codium fragile on muddy sand bottom in Okayama, Seto Inland Sea (T. Watanabe pers. comm.). See also Species comparisons.</p><p>Decorating materials and epibionts. Branched colonies of hydrozoans, algal pieces, detritus compounds, sometimes encrusted with sponges (Ariyama 1995; this study).</p><p>Ecological notes. Ecology of this species has not been investigated as far as we know.</p><p>Remarks. Pugettia quadridens intermedia was described by Sakai (1938) based on three males from Shimoda (Izu Peninsula), one male from Ise Bay, one male and one female from the coast of Gobo, and one male and one female from Wakayama (both Kii Peninsula). He therein designated a full-grown male with 19.5 mm PCL and 15.3 mm CW as the holotype with a photograph of the specimen (Sakai 1938: 259, pl. 36 fig. 2), but did not indicate its locality. Subsequently, Sakai (1976) noted the type locality as “Shimoda (Sakai)”. This seems to imply that he had intended to designate a full-grown male specimen from Shimoda as the holotype. In the catalogue of major part of extant Sakai’s specimens, one male specimen from Shimoda (KPM-NH 124160: Fig. 14) was listed as the holotype (Muraoka 1998: 24, table 1). However, our re-examination revealed that this specimen is considerablly smaller (13.9 mm PCL and 12.5 mm CW) than Sakai’s measurements (Fig. 14 vs. Sakai 1938: pl. 36 fig. 2), and despite the lack of both chelipeds, it is likely not a full-grown specimen based on the results of morphometric analyses (Fig. 39). Although there were unfortunately no comments or discussions, Muraoka’s (1998) “ holotype ” is invalid and did not fulfill ICZN 1999, Article 74.6 since there were more than one specimen mentioned in the original description. KPM-NH 124160 is likely one of the three male specimens from “Simoda, the coast of M. B. S.” examined in Sakai (1938) because the specimen is preserved together with the label (Fig. 14C), and therefore, this specimen should be regarded as one of paratypes instead (ICZN 1999, Article 72.4.5). In this study, we could locate neither true holotype nor the other paratypes in the extant Sakai’s materials deposited in KPMNH, NSMT, and Shimoda Marine Research Center, University of Tsukuba (SMRC). However, we do not exclude the possibility that the other Sakai’s specimens we could not locate during this study can be found in another institute. Therefore, we here do not designate KPM-NH 124160 as lectotype.</p></div>	https://treatment.plazi.org/id/FF0187ECFF80CE6AEAE7F9CEFDB0F1C3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Ohtsuchi, Naoya;Kawamura, Tomohiko	Ohtsuchi, Naoya, Kawamura, Tomohiko (2019): Redescriptions of Pugettia quadridens (De Haan, 1837) and P. intermedia Sakai, 1938 (Crustacea: Brachyura: Epialtidae) with description of a new species. Zootaxa 4672 (1): 1-68, DOI: 10.11646/zootaxa.4672.1.1
FF0187ECFFB5CE65EAE7FC22FC58F542.text	FF0187ECFFB5CE65EAE7FC22FC58F542.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pugettia ferox Ohtsuchi & Kawamura 2019	<div><p>Pugettia ferox n. sp.</p><p>[New Japanese name: Oh-yotsuha-mo-gani]</p><p>(Figs. 26–38)</p><p>Pugettia quadridens .— Shen 1932: 49, pl. 2, fig. 2, text-figs. 26–30; 1937: 287–289, text fig. 5b, c, e, h.— Vinogradov 1950: 235, pl. 42, fig. 149.— Shen &amp; Dai 1964: 39, unnumbered figure.— Kobyakova 1966: 213, pl. 41, fig. 1.— Levin 1976: 54, fig. 107.— Griffin &amp; Tranter 1986: 97 (in part), fig. 28e, f.— Dai &amp; Yang 1991: 129–130, pl. 14(4), fig. 66(2).— Muraoka 1998: 24 (in part).— Marumura &amp; Kosaka 2003: 32 (in part).— Nunomura 2010: 52–53 (in part).— Lee et al. 2014: 47, figs. 3A, 4A.— Ko &amp; Lee 2015: 19, figs. 1B, 2A, B, pls. 11, 12 [not Pisa (Halimus) quadridens De Haan, 1837]</p><p>Pugetti [sic.] quadridens .— Sakai 1938: 255–257 (in part), text-fig. 28b. [not Pisa (Halimus) quadridens De Haan, 1837]</p><p>part).— Kim &amp; Kim 1998: 302–303, figs. 1, 2 [not Pugettia quadridens intermedia Sakai, 1938]</p><p>Pugettia quadridens quadridens .— Kim 1973: 529 (key), 530–532, pl. 53 fig. 194.— Kim &amp; Kim 1998: 303–304.— Miyake 1983: 35, pl. 12, fig. 2; 1998: 35, pl. 12, fig. 2.— Honma &amp; Muraoka 1992: 41, fig. 2E.— Ariyama 1995: 1, 3, fig. 3. [not Pisa (Halimus) quadridens De Haan, 1837]</p><p>Pugettia quadridens intermedia .— Ikeda 1981: 15 (in part).— Kim &amp; Kim 1998: 302–303, figs. 1, 2 [not Pugettia quadridens intermedia Sakai, 1938]</p><p>Pugettia intermedia .— Marumura &amp; Kosaka 2003: 32 (in part). [not Pugettia quadridens intermedia Sakai, 1938]</p><p>Pugettia pellucens .— Marumura &amp; Kosaka 2003: 32 (in part) [not Pugettia quadridens pellucens Rathbun, 1932]</p><p>Pugettia incisa .— Nunomura 2010: 52 (in part) [not Pisa (Halimus) incisa De Haan, 1837]</p><p>? Pugettia quadridens .— Miers 1879: 23 (in part).— Rathbun 1902: 28 (in part).— Doflein 1902: 655.— Balss 1924: 24 (in part).— Yokoya 1933: 148 (in part). [see Discussion]</p><p>Material examined. Holotype: male (40.0 × 32.1 mm) (NSMT-Cr 26069), 2–4 m, Akahama, Otsuchi Bay, Iwate, SCUBA +hand, coll. K. Nakamoto, 14 Feb. 2017.</p><p>Allotype: female (27.9 × 22.6 mm) (NSMT-Cr 26070), boulder zone, 5 m, off Horai-jima Islet, Otsuchi Bay, SCUBA +hand, coll. K. Fukuda, 27 Feb. 2013.</p><p>Paratypes: Three males (27.4 × 23.3–33.7 × 28.2 mm) , 1 female (21.8 × 17.7 mm) (RUMF-ZC-4975), same data as allotype; 1 male (30.7 × 26.3 mm) (NSMT-Cr 26071), boulder zones, 5 m, Akahama, Otsuchi Bay, Iwate, SCUBA +hand collection, coll. K. Nakamoto, 24 Jan. 2017 ; 3 males (5.5 × 3.8–25.1 × 20.1 mm), 1 female (17.2 × 13.6 mm) (NSMT-Cr 26072), Sargassum yezoense beds, 5 m, same locality and date as previous, SCUBA +hand, coll. N. Ohtsuchi ; 3 males (22.8 × 18.3–39.3 × 33.9 mm) (NSMT-Cr 26073), 1 male (40.3 × 33.5 mm) (CBM-ZC 14879), with Sargassum sp., 2 m, Akkeshi Bay, Hokkaido, baited trap, coll. S. Houki &amp; K. Fukuda, 1 Sep. 2012 ; 1 male (8.8 × 6.2 mm), 1 female (10.1 × 7.2 mm) (CBM-ZC 14878), red algal turfs, 4–5 m, Tomarihama, Oshika Peninsula, Miyagi, coll. N. Ohtsuchi, 24 Aug. 2011 ; 2 males (15.1 × 11.5, 19.8 × 14.7 mm), 3 females (15.4 × 11.6–22.8 × 18.3 mm), 1 ovigerous female (17.5 × 13.9 mm) (CBM-ZC 14880), near low tidal mark, algal turfs of A. paradoxa, Oarai, Kashima Sea, Ibaraki, hand collection, coll. N. Ohtsuchi &amp; S. Houki, 8 May 2012 .</p><p>Non-types: Japan. Eight males (12.7 × 8.9–32.0 × 22.3 mm), 6 females (15.1 × 11.0–28.6 × 22.8 mm), 1 ovigerous female (28.2 × 23.0 mm) (NSMT-Cr 11233), Soya Strait, Hokkaido, T. Miyauchi, 13 Jul. 1991; 2 males (20.0 × 15.8, 32.7 × 27.6 mm), 1 female (25.7 × 21.5 mm) (CBM-ZC 10791), Kabutoiwa, Shakotan Peninsula, Hokkaido, SCUBA diving, coll. T. Komai, 26 Aug. 2007 ; 1 male (39.4 × 33.4 mm) (NSMT-Cr 26076), Sakae-ura, Saroma Lake, Hokkaido, coll. S. Chiba, 9 May 2016 ; 2 females (28.7 × 23.5, 11.7 × 9.4 mm) (OMNH-Ar 10708), Notoro Lake, Hokkaido, coll. S. Chiba, 7 Oct. 2016 ; 2 males (6.2 × 4.4, 9.5 × 7.1 mm), 1 female (10.4 × 7.8 mm), 1 female (with a rhizocephalan parasite, 15.6 × 11.6 mm) (SMBL-Ar 1452), Abashiri, Hokkaido, coll. T. Yam, 12 Aug. 1960 ; 4 males (17.3 × 13.0–22.0 × 15.9 mm), 1 female (20.8 × 16.3 mm) (NSMT-Cr 3262), same locality as previous, coll. S. Sato, 30 Aug. 2008; 1 male (16.5 × 13.3 mm) (OMNH-Ar 10709), Cape Shiretoko, Hokkaido, coll. S. Chiba, 2 Nov. 2013 ; 1 male (24.5 × 18.3 mm) (CBM-ZC 10819), 2–5 m, Bunkichi Bay, Shiretoko Peninsula, Hokkaido, coll. T. Komai, 16 Sep. 2008 ; 2 males (33.3 × 27.7, 36.2 × 29.4 mm) (NSMT-Cr 10261), 2–3 m, Goyoumai, Kushiro, Hokkaido, coll. H. Hayashi, 7 Dec. 1977 ; 3 males (17.9 × 13.9–32.0 × 25.9 mm), 1 female (28.4 × 23.7 mm) (RUMF-ZC-4981), Off Aikappu, Akkeshi Bay, Hokkaido, epibenthic sled, coll. T. Yorisue, S. Hamano, H. Katsuragawa &amp; R. Yoshida, 9 Sep. 2015 ; 3 males (25.9 × 20.3–32.4 × 25.4 mm) (RUMF-ZC-4982), same locality and date as previous, trap, coll. T. Yorisue, S. Hamano, H. Katsuragawa &amp; R.Yoshida; 2 males (32.1 × 26.4, 36.2 × 29.3 mm) (RUMF-ZC-4983), pier of Akkeshi Marine Station, Akkeshi Bay, Hokkaido, trap, coll. R. Yoshida, 15 Sep. 2015 ; 3 females (9.7 × 6.6–11.0 × 8.3 mm) (SMBL-Ar 1453), Shirikishinai, Hokkaido, coll. T. Yam, 30 Aug. 1960 ; 1 male (24.2 × 22.9 mm) (WMNH-Na-Cr 0312-1), Hokkaido, coll. S. Nagai, Jul. 1981; 2 males (13.9 × 9.6, 21.8 × 17.8 mm), 2 females (11.2 × 8.3, 22.9 × 18.1 mm) (NSMT-Cr 17720), Miyako Bay, Iwate, coll. S. Koyama, 29 Aug. 1937 ; 1 female (28.0 × 21.7 mm), 3 ovigerous females (18.3 × 13.4–22.8 × 17.5 mm) (NSMT-Cr 17729), Funakoshi Bay, Shimohei, Iwate, coll. S. Koyama, 2–30 Aug. 1937 ; 1 male (13.2 × 8.9 mm) (NSMT-Cr 8389), 50 m, Otsuchi Bay, Iwate, coll. M. Takeda, 21 Jul. 1982 ; 4 males (8.3 × 5.8–19.1 × 13.8 mm), 1 female (13.7 × 9.5 mm), 1 juvenile (5.6 × 3.5 mm) (NSMT-Cr 8390), 38 m, Otsuchi Bay, Iwate, coll. M. Takeda, 21 Jul. 1982 ; 1 female (13.2 × 9.4 mm) (NSMT-8391), 17.5 m, Otsuchi Bay, coll. M. Takeda, 21 Jul. 1982 ; 1 male (9.2 × 7.0 mm) (OMNH-Ar 10706), Akahama, Otsuchi Bay, understory of Saccharina japonica var. diabolica beds, 2–4 m, SCUBA +air-lifting sampler, coll. M. Kodama, 26 Nov. 2015 ; 1 male (4.4 × 3.1 mm) (OMNH-Ar 10705), same locality as previous, understory of Phyllospadix iwatensis beds, 2–4 m, SCUBA +air-lifting sampler, coll. M. Kodama, 25 Feb. 2016 ; 1 male (5.4 × 3.7 mm) (OMNH-Ar 10707), same locality and habitat as previous, SCUBA +air-lifting sampler, coll. M. Kodama &amp; J. Hayakawa, 16 May 2017; 1 male (31.8 × 24.4 mm) (NSMT-Cr 17682), Ohya, Motoyoshi-cho, Motoyoshi-gun, Miyagi, coll. Ibayashi, 10 Mar. 1933 ; 1 female (24.9 × 18.9 mm) (NSMT-Cr 17674), Baba, Karakuwa-machi, Motoyoshi-gun, Miyagi, coll. S. Ohfuchi, 10 Mar. 1933 ; 1 female (9.9 × 7.3 mm) (RUMF-ZC-4978), red algal turfs, 4–5 m, Tomarihama, Oshika Peninsula, Miyagi, SCUBA +air-lifting sampler, coll. T. Kawamura &amp; H. Takami, 29 Jul. 2010 ; 1 female (12.0 × 9.2 mm) (RUMF-ZC-4979), same habitat and locality as previous, SCUBA +air-lifting sampler, coll. T. Kawamura &amp; H. Takami, 10 Nov. 2010; 1 male (6.7 × 4.8 mm) (RUMF-ZC- 4980), same habitat and locality as previous, SCUBA +air-lifting sampler, coll. T. Kawamura &amp; H. Takami, 24 Aug. 2011; 3 females (8.4 × 6.3–8.7 × 6.4 mm) (RUMF-ZC-4976), 1 female (10.1 × 7.2 mm), 4 ovigerous females (16.7 × 12.7–21.0 × 17.8 mm) (RUMF-ZC-4977), same habitat, locality, and sampling method as previous, coll. N. Ohtsuchi, 24 Aug. 2011; 3 males (22.1 × 17.8–25.5 × 21.5 mm), 1 female (19.9 × 16.0 mm), 1 ovigerous female (19.1 × 15.6 mm) (CBM-ZC 14881), 5 m, boulder zone, Tomarihama, Oshika Peninsula, SCUBA, coll. T. Kawamura &amp; N.-I. Won, 9 Jul. 2011 ; 9 males (15.7 × 12.4–25.2 × 19.6 mm), 1 female (12.9 × 9.7 mm), 1 ovigerous female (20.8 × 15.5 mm) (NSMT-Cr 17637), Nakanosaku, Ena, Iwaki, Fukushima, coll. S. Ohfuchi &amp; H. Kakuda, 3 Aug. 1932 ; 7 males (12.5×9.1–23.6× 19.3 mm) (NSMT-Cr 26075), near low tidal mark, Sargassum sp. beds on rocky reef, Nagasaki, Iwaki, Fukushima, hand collection, coll. N. Ohtsuchi, 2 Feb. 2017 ; 3 males (19.2 × 15.3–23.1 × 19.4 mm), 3 females (21.3 × 17.4–22.4 × 18.0 mm) (CBM-ZC 14882), near low tidal mark, Ahnfeltia paradoxa turf on rocky reef, Misaki, Iwaki, Fukushima, hand collection, coll. N. Ohtsuchi, 2 Feb. 2017 ; 2 females (31.3 × 24.4, 35.0 × 29.3 mm) (NSMT-Cr 19866), Onahama, Iwaki, Fukushima, coll. H. Kakuda, 20 Jun. 1931 ; 1 male (32.8 × 28.8 mm) (WMNH-Na-Cr 0312-1), Choshi, Boso Peninsula, 80 m, coll. T. Watanabe, 1988 ; 2 males (17.0 × 13.5, 20.7 × 17.2 mm) (CBM-ZC 14884), intertidal, Gelidium elegans turfs, Inubosaki Light, hand collection, coll. N. Ohtsuchi &amp; S. Houki, 9 May 2012 ; 1 male (27.3 × 22.3 mm) (CBM-ZC 2230), Tomiyama Fishery Port, Minami-Boso-shi, Chiba, rafts for aquaculture, hand collection, coll. T. Komai, 25 Dec. 1995 ; 4 males (26.5 × 21.6–18.1 × 13.1 mm), 5 ovigerous females (13.1 × 10.3–18.4 × 14.2 mm) (KPMNH- 110366 – 110374), Sagami Bay; 1 female (34.4 × 29.0 mm) (WMNH-Na-Cr 0314-1), Hayama, coll. S. Nagai, 6 May 1973 ; 1 male (17.2 × 13.6 mm) (NSMT-Cr 26074), near low tidal mark, S. fusiforme beds, rocky beach behind Morito Shrine, Hayama, Sagami Bay, hand collection, coll. N. Ohtsuchi, 29 Apr. 2009 ; 2 males (18.3 × 14.1, 11.2 × 8.4 mm) (OMNH-Ar 6294), Oura, Takeno-cho, Kinosakigun, Hyogo, coll. R. Yamanishi, 25 Jul. 1999 ; 1 male (30.4 × 24.9 mm) (OMNH-Ar 3048), Tanagawa, Misaki-cho, Sennan-gun, Osaka, 28 Jul. 1986 ; 1 male (17.8 × 13.1 mm) (OMNH-Ar 6023), Tanigawa, Misaki-cho, Sennan-gun, Osaka, coll. H. Ariyama, 24 May 1994 (figured in Ariyama 1995) ; 1 male (27.9 × 22.9 mm) (CBM-ZC 14885), ca. 50–80 m, off Kitaura Port, Oga Peninsula, Akita Prefecture, Northern Japan, gill-net, coll. M. Marumura, 23 Nov. 1994 ; 1 male (12.7 × 12.0 mm), 1 female (8.5 × 5.5 mm) (TOYA-Cr 10454), Nozaki, Notojima-cho, Ishikawa, coll. H. Nambu, 25 Jul. 1990 ; 2 males (16.0 × 12.2, 8.0 × 5.7 mm), 2 females (5.9 × 4.3, 8.2 × 5.9 mm) (TOYA-Cr 17320), Miyazaki Fishery Port, Miyazaki, Asahi-cho, Toyama, coll. H. Nambu, 11 Nov. 2008 ; 1 male (18.1 × 14.9 mm) (ZMUC-CRU-20232), Nagasaki, coll. James Jordan, 1 Jul. 1911 (figured in Griffin &amp; Tranter 1986).</p><p>North China. One ovigerous female (18.0 × 14.2 mm) (MBM 160494), Yantai, coll. Yang Jing, 1 Jul. 1957 ; 1 female (20.2 × 15.3 mm) (MBM 160497), Xishawang, Yantai, 22 Mar. 1975 ; 1 male (23.2 × 18.5 mm) (MBM 160514), exact locality unknown, 12 Feb. 1960; 1 male (20.3 × 15.5 mm), 1 female (20.4× 18.8 mm) (MBM 160515), Changxing Island, coll. Fangzeng Sun, 6 Oct. 1956 ; 4 males (10.1 × 7.4–15.8 × 12.1 mm) (MBM 160513), Kong tong Island, Yantai, Shanton, North China, 7 Apr. 1951 .</p><p>Description. Male. Full-grown males (18.1–40.5 mm PCL, including holotype and paratypes). Carapace (Fig. 26A) pyriform, 1.1–1.3 longer than width (PCL/CW = 1.2±0.0, N = 19), surface smooth to naked eyes but closely covered with microscopic setae; gastric, cardiac, branchial, intestinal regions unclearly separated from each other. Gastric region (Fig. 26C) moderately elevated, always anteriorly with oblique row of dense hooked setae on either side of midline (Figs. 26A, C, 37G); mesogastric, metagastric, protogastric region on both sides each with subacute protuberance (Figs. 26A, 37G). Hepatic, cardiac, branchial regions moderately elevated; cardiac region separated from branchial region on each side by irregularly rugose groove (Figs. 25C, 37G); mesobranchial regions elevated, as high as cardiac regions, with 2–3 (2 in general) obtuse tubercles apically, mesial one larger than lateral ones (Figs. 26A, 37G); metabranchial regions faintly elevated, with low, obtuse tubercle. Intestinal region (Fig. 26A, C) moderately elevated, with obtuse protuberance apically, separated from cardiac region.</p><p>Pseudorostral spines (Figs. 26A, 27A) short, length 0.1–0.2 of post-pseudorostral carapace length (PRL/PCL = 0.2±0.0, N = 20), each with two rows of dense, hooked setae on proximal two-third dorsally, single row of long setae on proximal 0.8 mesially; lateral margins subparallel. Preorbital spine (Figs. 26A, 28A, B) slender, generally compressed dorsoventrally, directed anterolaterally, acuminate at tip, with row of slender setae on mesial margin (Fig. 28A). Supraorbital eave (Figs. 26A, 28A, B) moderately extended laterally, lateral margin sinuous. Orbital hiatus (Figs. 26A, 28A) deep, rounded, triangular concavity. Postorbital lobe (Figs. 26A, 27A, 28A) small, triangular, shorter than preorbital spine, weakly compressed dorsoventrally, directed anteriorly or anterolaterally, weakly incurved distally. Hepatic lobe (Figs. 26, 27A, 28A) not demarcated from hepatic region, broad, triangular, immediately narrowed distally, more than twice longer than postorbital lobe (HpL/PoL = 2.4±0.3, N = 15), compressed dorsoventrally, directed anterolaterally, acuminate at tip, variably incurved distally (Fig. 37). Anterolateral carapace margin (Figs. 26A, 27A) always with rows of sparse, hooked setae; lateral surface inferior to anterolateral margin with 3–5 (3 in general) spines. Epibranchial spine (Fig. 26A) longer than postorbital lobe, shorter than hepatic lobe, directed anterolaterally, weakly incurved, acuminate at tip, positioned at posterior 0.4 of postorostral carapace length (ESL/PCL = 0.4±0.0, N = 18), confluent to posterolateral carapace margin basally. Posterolateral carapace margin (Fig. 26A) faintly convex. Posterior carapace margin (Fig. 26A) weakly projected roundly.</p><p>Subhepatic region (Fig. 26B) narrowly exposed in ventral view, with few, sparse, hooked setae. Pterygostomial region (Fig. 26B) not particularly inflated, with 4–5 (4 in general) papiliform tubercles along pleural suture. Anterolateral angle of buccal frame (Fig. 27A) produced anteriorly, not overlapped by anterolateral angle of merus of third maxilliped when closed, subrectangular in lateral view.</p><p>Basal antennal article (Fig. 28B) smooth on surface, bearing blunt, faintly granulate longitudinal ridge mesial to midline; mesial margin grooved; distolateral angle moderately produced into spine directed anterolaterally; lateral margin extended laterally, concave, sometimes faintly granulate (Fig. 28C), proximal end produced into triangular lobe, separated from posterior orbital margin, with subacute tubercle basally (Fig. 28B). Antennal peduncle (Fig. 28D) consisting of two articles; penultimate article (Fig. 28D) generally subcylindrical, broadened distally, with lateral margin bluntly carinate over entire length, distal end almost twice broader than proximal end; ultimate article (Fig. 28D) two-thirds of penultimate article in length, flattened dorsoventrally, slightly broadened distally; penultimate, ultimate articles with few, noticeably long setae on distomesial angle.</p><p>Third maxilliped (Figs. 26B, 28E) smooth on surface. Ischium with shallow, broad median depression, lateral margin nearly straight. Merus with dilated, faintly upturned anterolateral angle. Exopod almost half of ischium in maximum width, dilated laterally, immediately narrowed in distal two-fifth, mesial margin with blunt angle on distal one-third (Fig. 28E).</p><p>Chelipeds (Figs. 26A, B, 29, 30) equal in size, similar in shape. Ischium (Fig. 26B) weakly swollen ventrally in distal half; mesial margin obtusely crested, bearing 2–3 (3 in general) teeth, truncate in anterior end; distolateral lobe distinct, compressed, rounded apically. Merus (Fig. 29 D–G) prismatic, length 2.7 longer than height (2.7±0.1, N = 10); dorsal surface (Fig. 29 D–F) with broad, longitudinal keel bearing 3–6 distinct (3 in general) teeth, distalmost largest, directed anteriorly; outer surface (Fig. 29 E–G) faintly rugose, with blunt longitudinal ridge, with 2– 3 (3 in general) rudimentary teeth; ventral surface (Fig. 29F, G) with blunt ridge bearing 3–4 (3 in general) low, broad teeth; inner surface (Fig. 29D, E, G) depressed, irregularly rugose, with blunt, longitudinal ridge terminated in subtriangular, proximal lobe; distal margin (Fig. 29 E–G) with 2 prominent knobs at articulation with carpus (outer knob larger than inner), prominent, obliquely erect, subrectangular lobe with moderately long, acute projection on upper side. Carpus (Fig. 29 A–C) moderately inflated, blunt ridge bearing 2–3 (2 in general) low tubercles on dorsal surface (Fig. 29A); outer margin obtusely ridged, irregularly dentate (Fig. 29A, B); ventral surface (Fig. 29C) uneven; inner margin obtusely crested, irregularly dentate, with acute, proximal lobe (Fig. 29A). Chelae (Fig. 30A) more than twice longer than height (ChL/ChH = 2.3±0.1, N = 19); palm strongly expanded, upper margin obtusely ridged, lower margin poorly defined; immovable fingers with subpentagonal teeth along distal two-thirds (similar in size); movable finger with low, broad teeth (Figs. 26A, B, 30A); both fingers narrowly gaped in proximal half when closed (Fig. 30A).</p><p>Ambulatory legs (Figs. 26, 31) decreasing in length posteriorly, surface generally smooth to naked eyes but closely covered with microscopic setae. Meri weakly flattened, each with distinct, upper distal tubercle (Fig. 31A), more than six times longer than height in P2 (6.5±0.6, N = 10), more than four times in P3 (4.5±0.4, N = 10). Carpi each with shallow, medial depression on extensor surface (Fig. 31B). Propodi weakly flattened, moderately narrowed in flexor half, each with setal tufts on proximal 0.8 on flexor margin in P2, 0.6 in P3–P5 (Fig. 31A, C). Dactyli each with two rows of low, calcareous spines on flexor surface (Fig. 31A, C).</p><p>Thoracic sternites (Figs. 26B, 32A, B) smooth on surface, with shallow, broad depression on second to fourth sternite on both side; second sternite with pair of small depression anteriorly (often continuous to shallow depression on second to fourth sternites); third to fourth sternites obtusely ridged medially (Fig. 32A); sterno-abdominal cavity weakly ridged, without long setae on anterolateral margins (Fig. 32B).</p><p>Pleon (Figs. 26B, 32B) with six plomeres and telson; third to sixth pleomeres fixed, with distinct suture. Third pleomere broadest, lateral margin oblique; fourth pleomere trapezoid, as long as fifth in midline length; fifth pleomere trapezoid; sixth pleomere rectangular, dilated on distolateral angle, 0.6 of third pleomere in proximal width (0.6±0.0, N = 5); telson triangular.</p><p>Shaft of G1 (Fig. 33A, E) straight, trilobate in distal one-sixth; dorsal lobe elongate, triangular, with subacute tip, more than twice longer than ventral lobe, curved inwards, elevated basally to form incomplete lobate projection (Fig. 33 A–D); ventral lobe triangular, with subacute tip, pointing upwards (Fig. 33 A–D); mesial lobe as long as ventral lobe, projecting anteriorly obliquely, weakly twisted distally (Fig. 33A, B, D); hiatus between dorsal, mesial lobes moderately wide (Fig. 33D); mesial, lateral margins from dorsal lobe to ventral lobe concave; lateral margin lower than lateral margin of ventral lobe, with median dilation followed by lobule (Fig. 33 B–D). Shaft of G2 (Fig. 33H) stout, narrowed distally, truncated apically; apex with relatively large, acuminate process (Fig. 33I).</p><p>Adolescent males (11.2–32.4 mm PCL) (Fig. 35A, B). Chelae (Fig. 30B) proportionally longer (ChL/ChH = 2.8±0.2, N = 30) than in full-grown males (Table 1, see also Fig. 35B), not gaped, both fingers uniformly dentate on cutting margin (Figs. 30B, 35B). G1 trilobate apically as in full-grown males.</p><p>Immature males (&lt;9.2 mm PCL) (Fig. 36A, B). Carapace relatively slender (PCL/CW = 1.4±0.1, N = 5). Chela slenderer (ChL/ChH = 2.9±0.1, N = 3) than in adolescent males (Table 1, see also Fig. 36B), similar to adolescent specimens in dentation on both fingers. G1 incompletely folded, otherwise folded but but provided with rudimentary, mesial lobe projecting anterolaterally (Fig. 33J, K).</p><p>Female. Full-grown females (12.7–28.7 mm PCL, including allotype and paratypes). Carapace (Figs. 27B, 34A) similar to males in general proportion (PCL/CW = 1.3±0.0, N = 16; PRL/PCL = 0.2±0.0, N = 14; ESL/PCL = 0.4±0.0, N = 16) (Student t -test, p&gt; 0.05); mesobranchial region more elevated than in males (Figs. 27B, 34A, F versus Figs. 26A, 27A). Cheliped merus slenderer than in males (length/height = 2.9±0.2, N = 10; Student t -test, p &lt;0.01); chelae (Fig. 30C) much proportionally longer than in full-grown males (ChL/ChH = 2.8±0.2, N = 30; Student t -test, p &lt;0.01). Tufts of few elongate setae sometimes on midlines of protogastric region, midpoint of epibranchial region, apex of epibranchial spines, summits of cardiac and intestinal regions (Fig. 38C). Pleon (Fig. 34B) with six pleomeres and telson, expanded (PW6/PW3 = 1.4±0.1, N = 8). Gonopore (Fig. 32D, E) comma-shaped, suboval in lateral half, elongate in mesial half.</p><p>Adolescent females (9.9–16.9 mm PCL) (Fig. 35C, D). Chela slender (ChL/ChH = 2.8±0.0, N = 4), similar to full-grown individuals in dentation. Pleon ovate (PW6/PW3 = 1.3±0.1, N = 7).</p><p>Immature females (&lt;11.0 mm PCL) (Fig. 36C, D). Carapace relatively slender (PCL/CW = 1.4, N = 8) than in the other ontogenetic stages (Table 1, see also Fig. 36C). Chela slender (ChL/ChH = 2.8±0.1, N = 7). Pleon suboval (PW6/PW3 = 1.1±0.1, N = 9).</p><p>Variations. Carapace tends to be broader, from 1.4 to 1.1 in PCL/CW, in relation to size growth (Figs. 26, 34–38). Tuberculation on gastric, mesobranchial, and metabranchial regions more prominent in larger specimens in the same ontogenetic stage (Fig. 37A, D, E); tubercles on mesobranchial region count three, getting more prominent in larger specimens (Figs. 26, 37). HpL/PoL increases from 1.6 to 3.0 in relation to size growth (Fig. 39). Both fingers of chela sometimes do not contact in distal half (Fig. 37A). Ambulatory legs are often provided with sparse, long setae but they tend to be reduced in larger specimens (Figs. 26, 35–38). Spinulation on ambulatory leg dactyli often reduced in larger specimens probably due to ablasion (Fig. 31A, C). Hiatus between dorsal and mesial lobes of G1 variable in width; lobule on subdistal part of lateral margin of G1 variable in size and distinctness (arrowed in Fig. 33D, F).</p><p>Size. Largest male: 40.5 × 34.3 mm; largest female: 35.0 × 29.3 mm; smallest ovigerous female: 16.4 × 12.8 mm.</p><p>Coloration in life. The carapace is generally dark red, reddish brown or dark green (Fig. 38A, C, E, F); some specimens with some whitish or dark-colored blotches and/or dense speckles (Fig. 38A, E, F). The general coloration of the thorax, pleon, chelipeds, and ambulatory legs are white or yellowish brown (Fig. 38C, D). The coloration of chelipeds reduced in flexor surface; palms generally dark red to dark green, with scale-like patterning (Fig. 38A). Ambulatory legs with whitish band of variable width, on the joint of each articulation and the distal parts of dactyli (Fig. 38 A–E). Females similar to males in general coloration and patterning (Fig. 38C, D). See also Miyake (1983, 1998: pl. 12, fig. 2, as P. quadridens) and Ko &amp; Lee (2015: pl. 11, as P. quadridens).</p><p>Distribution. Sea of Okhotsk including Kuril Islands, southern Sakhalin coast. Japan, pacific coast from Hokkaido to Kii Peninsula and Osaka Bay; coast of Sea of Japan from Soya Strait to Nagasaki including Peter the Great Bay (Vinogradov 1950; Levin 1976; Griffin &amp; Tranter 1986; Honma &amp; Muraoka 1992; this study), Korean Peninsula (Lee et al. 2014), North China (Shen 1932, 1937; Dai &amp; Yang 1991).</p><p>Habitat. Various coastal environments, from the intertidal to 80 m depth: on turfs of small red algae e.g. Ahnfeltiopsis paradoxa, Gelidium elegans, Grateloupia cornea (Fig. 38F), and of brown algae, e.g. Sargassum confusum, S. yezoense, Stephanocystis hakodatensis (Fig. 38E), among kelp forest of Eisenia bicyclis, Saccharina japonica var. diabolica etc., among seagrass beds of Phyllospadix iwatensis on rocky turfs, boulders, large mussel reefs of Mytilus galloprovincialis Lamarck, 1819 on embankment, and rarely on mud flat.</p><p>Decorating materials and epibionts. Various combinations of pieces of red, brown algae, branched colonies of bryozoans or hydrozoans. Large specimens often encrusted by sponges, bryozoans (sometimes Lichenoporidae), hydrozoans, and barnacles.</p><p>Ecological notes. This species had been widely known to predate intensely on juveniles of Ezo Abalone, Haliotis discus hannai Ino, 1953, and sea urchins, Mesocentrotus nudus (A. Agassiz, 1864), and Strongylocentrortus intermedius (A. Agassiz, 1864) in captive condition (Shiraishi 1997; Agatsuma 2001; Hoshikawa 2003, each as P. quadridens; N. Ohtsuchi &amp; Y. Umezu pers. obs.) and field experiment (Kawai &amp; Agatsuma 1996, as P. quadridens). Laboratory observation confirmed they also predate on amphipods (H. Tamura pers. comm.) and hermit crab Pagurus middendorffii Brandt, 1851 (Matsuo et al. 2015, as P. quadridens).</p><p>Etymology. The species name ferox means “fierce” in Latin alluding to their positive carnivory (see above); used as an adjective.</p><p>Remarks. The present new species was discussed by Sakai (1938) as “a local variation” of Pugettia quadridens . Sakai (1938: 257, text-fig. 28) compared the specimens of P. quadridens from Iwate, Pacific coast of northeast Japan, with “typical” specimens from Shimoda, Pacific coast of southeast Japan, and regarded the former as a local variant of P. quadridens . Later, he again noted that specimens of P. quadridens from Akkeshi, Hokkaido, Pacific coast of northernmost Japan is “enormous in size, and postorbital and hepatic teeth are more or less fused together and the gastric, cardiac and intestinal regions are marked with a tubercle, while the branchial region are marked with two distinct tubercles” (Sakai, 1976: 196). Our specimens, which include many individuals from the coast of Iwate and Akkeshi, agreed with the morphological characters mentioned by Sakai (1938: 257): “the carapace markedly broader and the hepatic lobe more or less fused with the postocular tooth as seen in the case of P. incisa, and very often they form a plate-like expansion as in that species. The tubercles on each region are well marked and the movable finger of chelipeds is uniformly denticulated throughout its whole length, not being armed with the two isolated teeth of the typical form.”. In the present new species, however, the postorbital and hepatic lobes do not form plate-like expansions as seen in P. incisa (cf. Sakai 1938: text-fig. 27) throughout their ontogeny, though they are variably fused among individuals (Figs. 26, 34–38). This disagreement is probably due to Sakai’s confusion of P. quadridens sensu lato with P. incisa (see synonymy). Our direct comparison added more morphological differences between Sakai’s P. quadridens local variant and P. quadridens sensu stricto (See Species comparisons), and therefore, we have no hesitation to separate both as distinct species.</p></div>	https://treatment.plazi.org/id/FF0187ECFFB5CE65EAE7FC22FC58F542	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Ohtsuchi, Naoya;Kawamura, Tomohiko	Ohtsuchi, Naoya, Kawamura, Tomohiko (2019): Redescriptions of Pugettia quadridens (De Haan, 1837) and P. intermedia Sakai, 1938 (Crustacea: Brachyura: Epialtidae) with description of a new species. Zootaxa 4672 (1): 1-68, DOI: 10.11646/zootaxa.4672.1.1
