Species Card of Copepod
Calanoida ( Order )
    Arietelloidea ( Superfamily )
        Heterorhabdidae ( Family )
            Heterostylites ( Genus )
Heterostylites longicornis  (Giesbrecht, 1889)   (F,M)
Syn.: Heterochäta longicornis Giesbrecht,1889; 1892 (p.373, 383, 773, Descr.F, figs.F);
Heterorhabdus longicornis : Giesbrecht & Schmeil, 1898 (p.116, Rem. F); Wolfenden, 1904 (p.124, fig.M); Esterly, 1905 (p.186, figs.M, Rem.F,M); 1906 a (p.77, fig.F); Pearson, 1906 (p.26); van Breemen, 1908 a (p.125); Farran, 1905 (p.45); 1908 b (p.67, Rem.); A. Scott, 1909 (p.131, Rem.); Wolfenden, 1911 (p.302); Farran, 1926 (p.283); Sewell, 1932 (p.301, Rem.); Wilson, 1932 a (p.134, figs.F,M); 1936 c (p.92); Oliveira, 1945 (p.191); Sewell, 1948 (p.503); Greze & al., 1985 (p.8); Lapernat, 2000 (tab.3); in CalCOFI regional list (MDO, Nov. 2013; M. Ohman, comm. pers.);
Heterochaeta zetesios Wolfenden, 1902 (p.367, Descr.M); Fowler, 1903 (p.124)
Ref.:
Sars, 1925 (p.238, figs.F,M); ? Farran, 1929 (p.209, 267); Sewell, 1932 (p.301, figs.F); Rose, 1933 a (p.207, figs.F,M); Jespersen, 1940 (p.55); Lysholm & al., 1945 (p.36); Sewell, 1947 (p.181); Farran, 1948 d (n°15, p.3, figsF,M); Davis, 1949 (p.60, figs.F,M); C.B. Wilson, 1950 (p.241); Brodsky, 1950 (1967) (p.358, Rem.F,M); Vervoort, 1957 (p.135, Rem.); Tanaka, 1964 a (p.23, figs.F); Vervoort, 1965 (p.122, Rem.); Owre & Foyo, 1967 (p.80, figs.F,M); Park, 1968 (p.561, Rem.); Bradford, 1970 a (p.359, figs.F,M); Gardner & Szabo, 1982 (p.378, figs.F,M); Roe, 1984 (p.358); Ali-Khan 1993 (p.120, figs.F,M); Razouls, 1994 (p.166, figs.F,M); Chihara & Murano, 1997 (p.817, Pl.117: F,M); Bradford-Grieve & al., 1999 (p.883, 945, figs.F,M); Bradford-Grieve,1999 b (p.81, figs.F,M, Rem., figs.176, 191); Park, 2000 (p.45, figs.F,M, Rem.); Vives & Shmeleva, 2007 (p.314, figs.F,M, Rem.)
Species Heterostylites longicornis - Plate 1 of morphological figuresissued from : T. Park in Bull. Scripps Inst. Oceanogr. Univ. California, San Diego, 2000, 31. [p.176, Fig.24].
Female: a, habitus (left side); b, forehead (dorsal); c, d, urosome (dorsal, left, respectively); e, f, genital somite (left, ventral, respectively).


Species Heterostylites longicornis - Plate 2 of morphological figuresissued from : T. Park in Bull. Scripps Inst. Oceanogr. Univ. California, San Diego, 2000, 31. [p.177, Fig.25].
Female: a, P5 (anterior).

Male: b, P5 (anterior); c, left P5 (with distal endopodal segments omitted), anterior; d, e, exopod of right P5 (anterior, postrior, respectively);f, exopod of left P5 (posterior).


Species Heterostylites longicornis - Plate 3 of morphological figuresissued from : J.M. Bradford-Grieve in The Marine Fauna of New Zealand: Pelagic Calanoid Copepoda. National Institute of Water and Atmospheric Research (NIWA). NIWA Biodiversity Memoir, 111, 1999. [p90, Fig.57].
Female: A, habitus (right lateral side); B, P5.

Male: C, P5.


Species Heterostylites longicornis - Plate 4 of morphological figuresissued from : O. Tanaka in Publs Seto Mar. Biol. Lab., 1964, XII (1). [p.24, Fig.185].
Female: a, habitus (dorsal); b, last thoracic segment and urosome (lateral left side); c, Md (mandibular palp); d, Md (biting edge); e, Mx2; f, P5.
Nota: The urosome segments and furca are in the proportional lengths as 38:16:13:11:22 (right) = 100; A1 over-reaches the furca by 7 terminal segments.

Immature male: g, P5.


Species Heterostylites longicornis - Plate 5 of morphological figuresissued from : J.M. Bradford in N.Z. Jl Mar. Freshw. Res., 1970, 4 (4). [p.360, Figs 60-62].
Female (off Kaikoura, New Zealand): 61, P5; 62, distal part of 2nd exopod of P5.
Male: 60, P5.
Scale bars represent 0.1 mm.


Species Heterostylites longicornis - Plate 6 of morphological figuresissued from : G.O. Sars in Résult. Camp. scient. Prince Albert Ier, Monaco, 1924. [Pl. LXVII, figs.1-16].
Female (from around Azores Islands & Bay of Biscay): 1, habitus (dorsal view); 2, idem (lateral view from left side); 3, left A1); 4, A2; 5, Md; 6, cutting edge of the manducatory plate; 7, Mx1; 8, Mx2; 9, Mxp.; 10, P1; 11, P3; 12, P5; 13, exopodite 2 of P5; 14, anal segment and furca.
Npta: A1 24-segmented, exceeding about 1/4 body length.

Male: 15, habitus (dorsal view); 16, P5.

Nota:
Nota: compare the four last segments of A1 with H. major ones.


Species Heterostylites longicornis - Plate 7 of morphological figuresissued from : R.B.S. Sewell in The John Murray Expedition, 1933-34, Scientific Reports, VIII (1), 1947. [p.166, Fig.44, B]. Process on the posterior aspect of the 2nd basal segment of P1.
Remarks: The shape of the hook-like or spine-like process shows some variation in different genera, but it is undoubtedly homologous throughout the whole series. The function of this organ is unknown.


Species Heterostylites longicornis - Plate 8 of morphological figuresIssued from : R.B.S. Sewell in Mem. Indian Mus., 1932, X (continued). [p.301, Fig.98, a-b].
Female (from N Indian): a, exopod of P2; b, exopod of P3.


Species Heterostylites longicornis - Plate 9 of morphological figuresissued from : C.O. Esterly in Univ. Calif. Publs. Zool., 1905, 2 (4). [p.186, Fig.40]; As Heterorhabdus longicornis.
Male (from San Diego Region): a, P3; b, right P5; c, left P5.


Species Heterostylites longicornis - Plate 10 of morphological figuresissued from : C.O. Esterly in Univ. Calif. Publs Zool., 1906, 3 (5). [Pl.13, Fig.74]. As Heterorhabdus longicornis).
Female (from San Diego, California): 74, P5.


Species Heterostylites longicornis - Plate 11 of morphological figuresissued from : S. Ali-Khan in Crustaceana, 1993, 65 (1). [p;121, Figs.1-6].
Female (from 24°09'N, 64°27'E): 1, habitus (dorsal); 2, urosome (lateral); 3, P5; 4, part of exopodite of P5 (enlarged); 5, P5 (from another specimen); 6, part of exopodite of P5 (enlarged; same specimen).

Nota: The female specimens agree in general with the previously descriptions. Exceptions noted include the left caudal ramus (4 times as long as wide), the 2nd basipodal segment of P5 (without any seta or spine), the 2nd exopodal segment of P5 (with a crown of 7 teeth), and the P5 of another specimen (atypical by having 3 extra teeth on the 1st exopodal segment).


Species Heterostylites longicornis - Plate 12 of morphological figuresissued from : S. Ali-Khan in Crustaceana, 1993, 65 (1). [p;122, Figs.7-13].
Male (from 24°09'N, 64°27'E): 7, 5th thoracic somite and urosome (dorsal); 8, forehead (dorsal); 9, Md (biting edge); 10, Mx1; 11, Mx2; 12, P1; 13, P5.


Species Heterostylites longicornis - Plate 13 of morphological figuresIssued from : W. Giesbrecht in Systematik und Faunistik der Pelagischen Copepoden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. – Fauna Flora Golf. Neapel, 1892. Atlas von 54 Tafeln. [Taf. 39 , Fig.44 ]. As Heterochäta longicornis.
Female: 44, habitus (dorsal).


Species Heterostylites longicornis - Plate 14 of morphological figuresissued from : H.B. Owre & M. Foyo in Fauna Caribaea, 1, Crustacea, 1: Copepoda. Copepods of the Florida Current. 1967. [p.17, Fig.51].
Female: 51, Mx2.


Species Heterostylites longicornis - Plate 15 of morphological figuresissued from : H.B. Owre & M. Foyo in Fauna Caribaea, 1, Crustacea, 1: Copepoda. Copepods of the Florida Current. 1967. [p.17, Fig.53].
Female: 53, Md (gnathobase).


Species Heterostylites longicornis - Plate 16 of morphological figuresissued from : H.B. Owre & M. Foyo in Fauna Caribaea, 1, Crustacea, 1: Copepoda. Copepods of the Florida Current. 1967. [p.19, Fig.63].
Female: 63, P5.


Species Heterostylites longicornis - Plate 17 of morphological figuresIssued from : W. Giesbrecht in Systematik und Faunistik der Pelagischen Copepoden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. - Fauna Flora Golf. Neapel, 1892. Atlas von 54 Tafeln. [Taf. 20 , Figs.14, 21 ]. As Heterochäta longicornis.
Female: 14, Mx2 (posterior view); 21, Basipodites 1 and 2 of Mxp (anterior view).


Species Heterostylites longicornis - Plate 18 of morphological figuresIssued from : W. Giesbrecht in Systematik und Faunistik der Pelagischen Copepoden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. - Fauna Flora Golf. Neapel, 1892. Atlas von 54 Tafeln. [Taf. 20 , Figs.25, 26 ]. As Heterochäta longicornis.
Female: 25, P3 (posterior view); 26, exopod of P5 (anterior view).


Species Heterostylites longicornis - Plate 19 of morphological figuresHeterostylites longicornis female: 1 - Dorsally, genital somite widest at middle.
2 - Ventrally, genital operculum much wider than 1/2 width of somite (Fig.24-f).


Species Heterostylites longicornis - Plate 20 of morphological figuresHeterostylites longicornis male: 1 - Basal lobe of right P5 long (Fig.25-b); basal lobe of left P5 relatively long and rounded.
2 - Basal lobe of left P5 relatively small (Fig.25-c).


Species Heterostylites longicornis - Plate 21 of morphological figuresIssued from : W. Giesbrecht in Systematik und Faunistik der Pelagischen Copepoden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. – Fauna Flora Golf. Neapel, 1892. Atlas von 54 Tafeln. [Taf.39, Fig.44]. As Heterochäta longicornis.
Female: 44, habitus (dorsal).


Species Heterostylites longicornis - Plate 22 of morphological figuresIssued from : C. Razouls in Ann. Inst. océanogr., Paris, 1994, 70 (1). [p.166]. Caractéristiques morphologiques de Heterostylites longicornis femelle et mâle adultes.
Terminologie et abbréviations: voir à Calanus propinquus.

Compl. Ref.:
Wilson, 1942 a (p.190); Sewell, 1948 (p.329, 504, 514: Rem., 525, 532, 533, 547, 549, 558, 567); C.B. Wilson, 1950 (p.241); Fagetti, 1962 (p.31); Grice, 1963 a (p.496); Unterüberbacher, 1964 (p.30); De Decker & Mombeck, 1964 (p.12); Furuhashi, 1966 a (p.295, vertical distribution in Oyashio/Kuroshio transitional area, Table 7); Grice & Hulsemann, 1967 (p.18); Fleminger, 1967 a (tabl.1); Morris, 1970 (p.2301); Vinogradov, 1968 (1970) (p.277); Itoh, 1970 a (tab.1); Park, 1970 (p.477); Deevey, 1971 (p.224); Roe, 1972 (p.277, tabl.1, tabl.2); Björnberg, 1973 (p.346, 387); Deevey & Brooks, 1977 (p.256, tab.2, Station "S"); Carter, 1977 (1978) (p.36); Dessier, 1979 (p.206); Kovalev & Shmeleva, 1982 (p.84); Guangshan & Honglin, 1984 (p.118, tab.); Stephen, 1984 (p.161, 169, Distribution vs thermocline & geographic); Greze & al., 1985 (p.9); Brenning, 1985 a (p.28, Table 2); Rudyakov, 1986 (tab.1); Brinton & al., 1986 (p.228, Table 1); Brenning, 1986 (p.13, Rem.); Wiebe & al., 1988 (tab.7); Heinrich, 1990 (p.18); Suarez & al., 1990 (tab.2); Suarez & Gasca, 1991 (tab.2); Gowing & Wishner, 1992 (tab.1); Suarez, 1992 (App.1); Shih & Young, 1995 (p.70); Padmavati & al., 1998 (p.349); Suarez-Morales & Gasca, 1998 a (p.110); Razouls & al., 2000 (p.343, tab. 5, Appendix); Madhupratap & al., 2001 (figs.4, 5); Holmes, 2001 (p.16); Lo & al., 2004 (p.89, tab.1); Kazmi, 2004 (p.228); Prusova & Smith, 2005 (p.75); Lavaniegos & Jiménez-Pérez, 2006 (tab.2, 4, Rem.); Cabal & al., 2008 (289, Table 1); Gaard & al., 2008 (p.59, Table 1, N Mid-Atlantic Ridge); Galbraith, 2009 (pers. comm.); Schnack-Schiel & al., 2010 (p.2064, Table 2: E Atlantic subtropical/tropical); Medellin-Mora & Navas S., 2010 (p.265, Tab. 2); in CalCOFI regional list (MDO, Nov. 2013; M. Ohman, comm. pers.); Williams R., 2013 (p.71, figs., Table 3.2, 3.4, 3.6, Appendix A, C; C & N composition, trophic level); Fierro Gonzalvez, 2014 (p.1, Tab. 3, 5, occurrence, abundance)
NZ: 22

Distribution map of Heterostylites longicornis by geographical zones
Loc:
Antarct. (Indian, Mc Murdo), sub-Antarct. (Indian), South Africa (E), Namibia, off St. Helena Is., off Ascension Is., Congo, G. of Guinea, off NW Cape Verde Is., Canary Is., Morocco, off Madeira, off Portugal, off W Cap Finisterre, Azores, off Amazon, off Orenoque, Barbada Is., Caribbean Sea, G. of Mexico, Caribbean, Florida, Sargasso Sea, off Bermuda: Station ‘’ S’’ (32°10’N, 64°30’W), Woods Hole, G. of Maine, S Iceland, Faroe Is., off Ireland (S & W), off W Scotland, North Sea, Bay of Biscay, Medit. (Alboran Sea, W Basin), G. of Oman, Arabian Sea, Pakistan, Natal, Indian, Bay of Bengal, Indonesia-Malaysia, China Seas (East China Sea, South China Sea), Taiwan (N: Mienhua Canyon), Japan (Izu), S Kuril Is., off British Columbia, Pacif. (central subtropical N-S), Guaymas Basin, San Diego, S California, Gulf of California, off W Mexico, Tehuantepc Bowl - Costa Rica Bowl, off Galapagos, off Peru, Pacif. (SE tropical), off Chile, Chile, Pacif. (W equatorial), E New Zealand, Tasman Sea
N: 85
Lg.:
(1) F: 3; (16) F: 2,7-2,3; M: 2,7; (17) F: 3,4; (22) F: 3,4-2,5; M: 3,5-3; (38) F: 3,5-2,94; M: 3,68-2,88; (45) F: 3-2,5; M: 3,25-3; (46) F: 3; (59) F: 4,3-3; M: 4,3-3; (72) F: 2,9-2,8; M: 2,9; (121) F: 3,05; (142) M: 3; (199) F: 2,89-2,58; M: 2,81; (313) F: 3,2-3,05; M: 3; (403) F: 3,2; M: 3,3; (432) F: 3,12-2,95; (824) F: 3,44-2,6; M: 3,4-2,56; (909) F: 2,9-3,4; M: 3,1; (1108) F: 2,9; {F: 2,30-4,30; M: 2,56-4,30}
Rem.: meso-bathypelagic. After Pearson (1906, p.26): vertical range down to 2.800 m.
Sampling depth (Antarct., sub-Antarct.) : 1000 m. Overall Depth Range in Sargasso Sea: 0-2000 m (Deevey & Brooks, 1977, Station "S");
Only the sizes and the relative length of the A1 permit to distinguish this species from Heterostylites major.
For Park (2000, p.46) certain identifications are not well ensured, leading to doubtful localisations. See remarks in Bradford-Grieve (1999 b, p.89).
For Itoh (1970 a, fig.2, from co-ordonates) the Itoh's index value of the mandibular gnathobase = 2050..
Last update : 29/01/2017
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