Resolution of the Portunusgladiator species complex: taxonomic status and identity of Monomiagladiator (Fabricius, 1798) and Monomiahaanii (Stimpson, 1858) (Brachyura, Decapoda, Portunidae) Author Windsor, Amanda M. Author Mendoza, Jose Christopher E. Author Deeds, Jonathan R. text ZooKeys 2019 858 11 43 http://dx.doi.org/10.3897/zookeys.858.33826 journal article http://dx.doi.org/10.3897/zookeys.858.33826 1313-2970-858-11 32E154398D0949E28DCC94C0C3AC6F7F 32E154398D0949E28DCC94C0C3AC6F7F Monomia haanii s. str. (Stimpson, 1858) Figs 1E, F, 2, 3 D-F , 4 E-H Portunus (Amphitrite) gladiator : De Haan 1833: 39; 1835: pl. 1 fig. 5, pl. A. Non Portunus gladiator Fabricius, 1798. Amphitrite haanii Stimpson, 1858: 38; 1907: 79; Ng et al. 2008 : 151 (synonymy), 156 (discussion). Neptunus (Amphitrite) gladiator : Ortmann 1893 : 73; Lanchester 1902 : 544; Parisi 1916 : 173; Balss 1922 : 107; Sakai 1934 : 303; 1936 : 129, pl. 36 fig. 3; 1939: 390, fig. 5a, pl. 47 fig. 3; Shen 1937: 101, fig. 2; Lin 1949 : 19. Non Portunus gladiator Fabricius, 1798. Portunus gladiator : Rathbun 1902 : 26, Miyake 1961 : 172; Sakai 1965 : 118, pl. 57 fig. 1; 1976: 341, fig 180a, pl. 120 fig. 1; Stephenson and Rees 1967 : 24; 1968 : 293; Takeda and Miyake 1968 : 551; Stephenson 1972a : 16, 39; 1972b : 135; Ko and Lee 2013 : 40, pl.-fig. 34. Non Portunus gladiator Fabricius, 1798. Portunus (Achelous) gladiator ; Rathbun 1910 : 36 (from Gulf of Siam). Non Portunus gladiator Fabricius, 1798. Portunus pseudoargentatus Stephenson, 1961: 109, Figs 2A, 3F, pl. 2 fig. 4, pl. 4 fig. F, pl. 5 fig. D; Stephenson and Rees 1967 b: 25; 1968: 294; Yang and Dai1994 : 137, fig. 13. Portunus haanii : Stephenson and Cook 1973 (in part): 429, figs 6 F-H , 7 F-H , 8 F-H , 9B; Dai and Yang 1991 : 223, fig. 120(2), pl. 27(4); Moosa 1996 : 521; Apel and Spiridonov 1998 (in part): 291; Ng et al. 2001 : 16 (list). Portunus (Monomia) gladiator : Sakai 1976 : 341, fig. 180, pl.120 fig. 1; Kim and Chang 1985 : 52. Not Portunus gladiator Fabricius, 1798. Portunus haanii : Miyake 1983 : 85, pl. 29 fig. 2; Takeda 1989 : 152 Portunus (Monomia) haanii Yamaguchi & Baba, 1993: 396, figs 137 A-C . Monomia pseudoargentata : Chertoprud et al. 2012 : 315, pl. 51 fig. H; Spiridonov et al. 2014 : 412, 427, fig. 3I, tab. 1. Monomia haanii : Ng et al., 2017 : 68 (list). Not Amphitrite media Stimpson, 1858: 39; 1907: 79, pl. 10 fig. 1. Not Monomia haanii , Chertoprud et al. 2012 : 314, pl. 51 fig. G (= Monomia gladiator (Fabricius, 1798)). Material examined. JAPAN: RMNH 379, lectotype, male, Japan, coll. P.F. von Siebold, 1823-1829 (photographs only); ZRC.1999.0084, 1 male, 1 female, off Hota, Uchibo, coast of Boso Peninsula, coll. T Komai, 22 Aug. 1997; USNM5255, 4 male, 1 female, coll. FC Dale and PL Jouy, Palos R/V; USNM26254, 1 male, 1 female, off Wakanoura, coll. DS Jordan and JO Snyder, 1900; USNM45882, 1 female, off Wakanoura, Kishu; USNM54519, 1 female, Yamagata Prefecture, coll. M Sasaki, Aug. 1917; USNM72540, 2 males, Enoshima, Bay of Jeddo, coll. ES Morse; USNM60250, 1 male, Toyama Bay, coll. M Sasaki, 1925; USNM112423, 1 male, Shimizu, Sugura, Albatross R/V, 14 Oct. 1906. AUSTRALIA: WAM-C7506, 1 male (holotype of Portunus pseudoargentatus Stephenson, 1961), Abrolhos Islands, Western Australia, coll. RW George, 11 May 1960; WAM-C34767, 1 female, Exmouth Gulf, Western Australia, coll. S Morrison and P Unsworth, 5 Nov. 2004; WAMC34900, 1 female, Exmouth Gulf, Western Australia, coll. S Morrison and P Unsworth, 12 Mar. 2004; WAM-C34938, 1 gravid female, Shark Bay, Western Australia, coll. S Morrison et al., 4 Oct. 2002; WAM-C5510, 1 male, Shark Bay, Western Australia, coll. S Morrison et al., 5 Mar. 2003; WAM-C44737, 1 female, Ningaloo Marine Park, Western Australia, coll. MP Salotti and SM Slack-Smith, 1 Feb. 2008. SOUTH CHINA SEA (FAO Area 61): USNM1421161, 1 male, USNM1421181, 1 male, USNM1421182, 1 male, USNM1421185, 1 male, USNM1421187, 1 male, USNM1421191, 1 male, USNM1421194, 1 male, USNM1421195, 1 male, USNM1421186, 1 male, USNM121202, 1 male, USNM1421204, 1 male, USNM1421206, 1 male. TAIWAN: UF29509, 1 female, Daxi Fishery Port; UF29511, 1 female, Daxi Fishery Port; UF29512, 1 female, Daxi Fishery Port; USNM1420827, 1 female, Daxi Fishery Port; USNM1420828, 1 male, Daxi Fishery Port; ZRC.1998.0186, 2 males, 1 female, Daxi Fishery Port, coll. PKL Ng, 3-4 Aug. 1996; ZRC.2016.0408, 4 males, 3 females, Daxi Fishery Port, coll. PKL Ng, 1 Jul. 2016. Diagnosis. Similar to Monomia gladiator except in the following morphological characters. Infraorbital margin granulate, terminating mesially in small triangular tooth, in line with rest of margin. Sixth pleomere (Fig. 2B, 3 D-F ) longer than wide, maximum width at base; anterior half of lateral margins convergent anteriorly, posterior half concave to straight, anterior and posterior halves separated by angular convexity. G1 (Fig. 3C, 4 E-H ) bent at 90° at midlength, tapering distally toward a minutely spatulate tip, slightly broader than immediate subdistal area. In fresh specimens, P5 propodus with dark purple spot on distal tip, distal one-third of P5 dactylus colored dark purple as well (Fig. 1E, F). Remarks. The primary morphological differences between Monomia haanii (Stimpson, 1858) and M. gladiator (Fabricius, 1798) have already been discussed in the Remarks for the latter species. Yamaguchi and Baba (1993) had previously reported on the type material of Amphitrite haanii Stimpson, 1858, listing several syntypes collected by P.F. von Siebold from Japan in the years 1823 to 1829, deposited in the then Rijkmuseum van Natuurlijke Historie (RMNH) in Leiden, and preserved either dry or in alcohol. From these they selected a lectotype, a young male (42 by 24 mm; RMNH 379) preserved in alcohol. No further or detailed descriptions of the material were provided. Although they did provide photographs of several specimens, the lectotype was photographed still inside the bottle (viz. Yamaguchi and Baba, 1993: fig. 137A) and no definitive or diagnostic features of its morphology could be discerned. The collections of the RMNH are now housed in the Naturalis Biodiversity Center, Leiden, and photographs of the lectotype (Fig. 2) were kindly provided to the authors by Dr. Charles Fransen. From these photographs, the diagnostic morphology of the 6th pleomere (Fig. 2B) and the 90-degree bend of the G1 (Fig. 2C) can already be observed even in such a young specimen. Furthermore, although the fresh coloration of this species was not recorded by either De Haan (1833) or Stimpson (1858) , later observations of topotypic material show that the purple spots on the P5 propodus and dactylus, as well as the more profuse but scattered spotting on the dorsal carapace, and the dark colored spines on the anterior margin of the P1 merus are consistently observed (Fig. 1E, F; also Sakai, 1976: pl. 120 fig. 1; Miyake, 1983: pl. 29 fig. 1). Stephenson and Cook (1973) had also previously synonymized Portunus pseudoargentatus Stephenson, 1961, under P. haanii on the basis of their similar morphology. We confirm that P. pseudoargentatus is a junior subjective synonym of Monomia haanii (Stimpson, 1858) sensu stricto on the basis of similarities in the dentition of the infraorbital margin, the shape of the male 6th pleomere (Fig. 3D), and the ~90° angle of the bend of the G1 at midlength (Fig. 4E; also Stephenson, 1961: fig. 2A). In a photograph of the presumably newly preserved holotype, the pigmented spots on the left P5 propodus and dactylus are still there (Stephenson, 1961: pl. 2 fig. 4), although these have since faded and can no longer be seen during the present examination of the holotype. Specimens morphologically identifiable as M. haanii comprise a highly-supported clade that includes specimens from Japan, the type locality of M. haanii , as well as the holotype of Portunus pseudoargentatus Stephenson, 1961, and the Vietnamese specimens referred to by Chertoprud et al. (2012) (JX398094) and by Koch et al. (2017) and Koch and Duris (2018) (KY524463, KY524464) as " Monomia pseudoargentata ". Both morphological and molecular phylogenetic analyses support the recognition of Monomia haanii as a full species. Molecular results also support a morphological basis for synonymy of P. pseudoargentatus Stephenson, 1961, under M. haanii . Figure 5. Maximum likelihood phylogram of three mitochondrial loci showing genetic distinction between M. gladiator and M. haanii along with three other members of the genus Monomia . Abbreviations: IOM = Institute of Oceanology and Museum, Nha Trang; MNHN = Museum National d'Histoire Naturelle, Paris; NHM = The Natural History Museum, London; UF = University of Florida Natural History Museum, Gainesville; UO = University of Ostrava, Ostrava; USNM = United States National Museum, National Museum of Natural History, Washington, D.C.; WAM = Western Australian Museum, Perth; ZMMU = Zoological Museum of the Moscow University, Moscow; ZRC = Zoological Reference Collection, Lee Kong Chian Natural History Museum, Singapore; CN = China; IN = India; JP = Japan; SG = Singapore; TH = Thailand; TW = Taiwan; VN = Vietnam. Figure 6. Neighbor joining phylogram of the barcode region of COI to visualize relationships between museum-vouchered reference specimens and 40 samples from four cans of pasteurized lump crabmeat (Product A-D ) labeled " Portunus haanii " and/or "red swimming crab"