Species Card of Copepod
Calanoida ( Order )
    Arietelloidea ( Superfamily )
        Heterorhabdidae ( Family )
            Heterorhabdus ( Genus )
Heterorhabdus spinifrons  (Claus, 1863)   (F,M)
Syn.: Heterochaeta spinifrons Claus, 1863 (p.182, figs.F); Brady, 1883 (p.49, figs.F,M);
Heterochäta spinifrons (M) : Giesbrecht, 1892 (p.372, 382, 773, figs.M, non F);
Heterorhabdus (Heterorhabdus) spinifrons : Vervoort, 1965 (p.119, Rem.); Bradford-Grieve,1999 b (p.87, figs.F,M, Rem., figs.175, 191);
Ref.:
Giesbrecht & Schmeil, 1898 (p.114); Thompson & Scott, 1903 (p.235, 249); Esterly, 1905 (p.183, figs.F); 1906 a (p.77, fig.M); Farran, 1908 b (p.65); A. Scott, 1909 (p.130, Rem.); Wolfenden, 1911 (p.302); Pesta, 1920 (p.531); Lysholm & Nordgaard, 1921 (p.25); Sars, 1925 (p.227, figs.F, M); Farran, 1926 (p.282); 1929 (p.209, 264); Sewell, 1932 (p.300); Wilson, 1932 a (p.133, figs.F,M); Rose, 1933 a (p.202, figs.F,M); Farran, 1936 a (p.112); Jespersen, 1940 (p.55); Sewell, 1947 (p.179, fig.44, Rem.F); Farran, 1948 e (n°16, p.3, figs.F,M); Brodsky, 1950 (1967) (p.351, figs.F,M); Chiba & al., 1957 (p.308); 1957 a (p.11); Vervoort, 1957 (p.133, Rem.); Grice, 1962 (p.222, figs.F); Paiva, 1963 (p.59, figs.M); Tanaka, 1964 a (p.13, figs.F,M); Mazza, 1965 a (p.314, figs.F,M, juv.); Owre & Foyo, 1967 (p.79, figs.F,M); Mazza, 1967 (p.191, 202, figs.F,M, juv.); Shih & al., 1971 (p.40); Bowman, 1971 b (p.40, figs.F, Rem.); Razouls, 1972 (p.95, Annexe: p.76); Arcos, 1974 (p.223, figs.F,M); 1975 (p.22, figs.F,M); Björnberg & al., 1981 (p.648, figs.F,M); Gardner & Szabo, 1982 (p.372, figs.F,M); Roe, 1984 (p.358); Mazzocchi & al., 1995 (p.35, figs.F,M, Rem.); Nishida & Ohtsuka, 1996 (p.620, figs.); Chihara & Murano, 1997 (p.819, Pl.118,120: F,M); Ohtsuka & al., 1997 (p.579, figs.F); Bradford-Grieve & al., 1999 (p.883, 944, figs.F,M); Park, 2000 (p.92, figs.F,M, Rem.); G. Harding, 2004 (p.4, figs.F,M); Vives & Shmeleva, 2007 (p.308, figs.F,M, Rem.)
Species Heterorhabdus spinifrons - Plate 1 of morphological figuresissued from : T. Park in Bull. Scripps Inst. Oceanogr. Univ. California, San Diego, 2000, 31. [p.211, Fig.59].
Female (Atlantic specimen): a, habitus (left side); b, c, urosome (left, dorsal, respectively); d, e, forehead (left, dorsal, respectively); f, g, h, i, genital somite (dorsal, right, left, ventral, respectively); j, left A1 (ventral)


Species Heterorhabdus spinifrons - Plate 2 of morphological figuresissued from : T. Park in Bull. Scripps Inst. Oceanogr. Univ. California, San Diego, 2000, 31. [p.212, Fig.60].
Female: a, left A2 (posterior); b, first endopodal segment of left A2 (anterior); c, left Md (posterior); d, masticatory edge of left Md (posterior); e, masticatory edge of right Md (posterior); f, left Mx1 (posterior); g, left Mx2 (posterior); h, right Mxp (anterior); i, P1 (anterior); j, basipod of P1 (posterior); k, P2 (anterior); l, P3 (anterior); m, P4 (anterior).


Species Heterorhabdus spinifrons - Plate 3 of morphological figuresissued from : T. Park in Bull. Scripps Inst. Oceanogr. Univ. California, San Diego, 2000, 31. [p.213, Fig.61].
Female: a, P5 (anterior).

Male: b, urosome (dorsal); c, left A1 (dorsal); d, P5 (anterior); e, right P5 (endopod omitted), anterior; f, distal end of exopod of right P5 (lateral); g, exopod of right P5 (posterior); h, second exopodal segment of right P5 (anterior); i, distal exopodal segments of left P5 (anterior). dlb = distal lobe; plb = proximal lobe; tsp = terminal spiniform process.


Species Heterorhabdus spinifrons - Plate 4 of morphological figuresissued from : T. Park in Bull. Scripps Inst. Oceanogr. Univ. California, San Diego, 2000, 31. [p.214, Fig.62].
Female (specimen from off west coast of North America): a, urosome (left side); b, c, d, e, genital somite (left, right, dorsal, ventral, respectively).


Species Heterorhabdus spinifrons - Plate 5 of morphological figuresissued from : O. Tanaka in Publs Seto Mar. Biol. Lab., 1964, XII (1). [p.14, Fig.179].
Female: a, habitus (dorsal); b, forehead (left lateral side); c, last thoracic segment and genital somite (left lateral side); d, P5.
Nota: A1 exceeds the end of the furca by distal 2 segments.

Male: e, P5.


Species Heterorhabdus spinifrons - Plate 6 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. [p.87, Fig.55].
Female: A, genital somite (dorsal); B, idem (left lateral side); C, forehead (dorsal); D, anterior part of the head (left lateral side); E, Mx2 (lobe 4); F, P5. Nota: remark a short 3rd seta on lobe 4of Mx2.


Species Heterorhabdus spinifrons - Plate 7 of morphological figuresIssued from: M.G. Mazzocchi, G. Zagami, A. Ianora, L. Guglielmo & J. Hure in Atlas of Marine Zooplankton Straits of Magellan. Copepods. L. Guglielmo & A. Ianora (Eds.), 1995. [p.37, Fig.3.3.1].
Female: A, habitus (dorsal); B, genital somite (lateral left side); C, P5.
Nota: A1 exceeds caudal rami by about 2 segments. Proportional lengths of urosomites and furca 42:15:12:12:19 = 100. Caudal rami asymmetrical; left ramus longer, bearing very long strong inner marginal seta.

Male: D, habitus (dorsal); E, P5.
Nota: Proportional lengths of urosomites and furca 23:19:16:11:8:23 = 100


Species Heterorhabdus spinifrons - Plate 8 of morphological figuresIssued from: M.G. Mazzocchi, G. Zagami, A. Ianora, L. Guglielmo & J. Hure in Atlas of Marine Zooplankton Straits of Magellan. Copepods. L. Guglielmo & A. Ianora (Eds.), 1995. [p.39, Fig.3.3.3].
Female (SEM preparation): A, urosome (ventral); B, genital somite (lateral left side); C and D, detail of integumental cone-like pores on either side of genital aperture (arrow in B).
Bars: A-B 0.100 mm; C 0.010 mm; D 0.005 mm.


Species Heterorhabdus spinifrons - Plate 9 of morphological figuresIssued from: M.G. Mazzocchi, G. Zagami, A. Ianora, L. Guglielmo & J. Hure in Atlas of Marine Zooplankton Straits of Magellan. Copepods. L. Guglielmo & A. Ianora (Eds.), 1995. [p.40, Fig.3.3.4].
Male (SEM preparation): B, posterior margins of urosome (dorsal); C, asymmetrical caudal rami (dorsal).
Bars: B 0.050 mm; C 0.100 mm.


Species Heterorhabdus spinifrons - Plate 10 of morphological figuresIssued from: M.G. Mazzocchi, G. Zagami, A. Ianora, L. Guglielmo & J. Hure in Atlas of Marine Zooplankton Straits of Magellan. Copepods. L. Guglielmo & A. Ianora (Eds.), 1995. [p.41, Fig.3.3.5].
Male (SEM preparation): A, protuberance on forehead ending with a short spine (dorsal view); B, P5; C and D, ear-like lobes on inner margins of basipod 2 of P5 (arrows in B).
Bars: A 0.050 mm; B 0.100 mm; C, D 0.010 mm.


Species Heterorhabdus spinifrons - Plate 11 of morphological figuresIssued from: M.G. Mazzocchi, G. Zagami, A. Ianora, L. Guglielmo & J. Hure in Atlas of Marine Zooplankton Straits of Magellan. Copepods. L. Guglielmo & A. Ianora (Eds.), 1995. [p.38, Fig.3.3.2].
Female (SEM preparation): B, forehead (dorsal) with characteristic protuberance ending with a sharp spine; C, rostrum with 2 filaments; D, urosome (dorsal).
B 0.050 mm; C 0.005 mm; D 0.100 mm.


Species Heterorhabdus spinifrons - Plate 12 of morphological figuresIssued from : G.O. Sars in Résult. Camp. Scient. Prince Albert I, 69, pls.1-127 (1924). [Pl.LXII, figs.9-12].
Female: 9, habitus (dorsal); 10, forehead (lateral); 11, urosome (lateral left side).

Male: 12, right P5.


Species Heterorhabdus spinifrons - Plate 13 of morphological figuresissued from : R.B.S. Sewell in The John Murray Expedition, 1933-34, Scientific Reports, VIII (1), 1947. [p.166, Fig.44, L]. 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 Heterorhabdus spinifrons - Plate 14 of morphological figuresissued from : S. Ohtsuka, H.Y. Soh & S. Nishida in J. Crustacean Biol., 1997, 17 (4). [p.582, Fig.3, A, B].
Female (from S. Japan): A, right mandibular cutting edge; B, left cutting edge.
v = ventral teeth; c = central teeth.
Scale bars : 0.100 mm (A, B).


Species Heterorhabdus spinifrons - Plate 15 of morphological figuresissued from : S. Ohtsuka, H.Y. Soh & S. Nishida in J. Crustacean Biol., 1997, 17 (4). [p.583, Fig.4, F].
Female (S Japan): F, left side of labrum (posterior surface) with gland openings (small and large arrows).
Scale bar: 0.010 mm.


Species Heterorhabdus spinifrons - Plate 16 of morphological figuresissued from : C.O. Esterly in Univ. Calif. Publs. Zool., 1905, 2 (4). [p.183, Fig.37].
Female (from San Diego Region): a, forehead (lateral, left side); b, Mxp (b.1 = 1st basal).


Species Heterorhabdus spinifrons - Plate 17 of morphological figuresissued from : C.O. Esterly in Univ. Calif. Publs Zool., 1906, 3 (5). [Pl.14, Fig.93].
Male (from San Diego, California): 93, P5.


Species Heterorhabdus spinifrons - Plate 18 of morphological figuresissued from : D.F.R. Arcos in Gayana, Zool., 1975, 32. [Lam.VIII, Figs.68-70].
Female (from Bahia de Concepcion, Chile): 69, P5.

Male: 68, habitus (dorsal); 70, P5.


Species Heterorhabdus spinifrons - Plate 19 of morphological figuresissued from : T.E. Bowman in Smithson. Contr. Zool., 1971, 96. [p.47, Fig.43, d].
Female (from Florida): d, distal end of Mx2.


Species Heterorhabdus spinifrons - Plate 20 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. 51]. As Heterochäta spinifrons.
Male: 51, anal segment and caudal rami.


Species Heterorhabdus spinifrons - 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. 20 , Fig.11 ]. As Heterochäta spinifrons.
Male: 11, Md (mandibular palp).


Species Heterorhabdus spinifrons - Plate 22 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 , Fig.16]. As Heterochäta spinifrons.
Male: 16, Mx2 (posterior view).


Species Heterorhabdus spinifrons - Plate 23 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 , Fig.19 ]. As Heterochäta spinifrons.
Male: 19, basipod of Mxp (anterior view).


Species Heterorhabdus spinifrons - Plate 24 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 , Fig.31]. As Heterochäta spinifrons.
Male: 31, P5 (anterior view).
Ps = left P5; Pd = right P5; B1 = basipod 1 (= coxa); B2 = basipod 2 (= basis); Ri = endopod; Re = exopod.


Species Heterorhabdus spinifrons - Plate 25 of morphological figuresissued from : G.D. Grice in Fish. Bull. Fish and Wildl. Ser., 1962, 61. [p.221, Pl.24, Figs.10-11].
Female (from equatorial Pacific): 10, forehead (lateral); 11, Mx2.
Nota: One of the distal 3 spines of Mx2 is small and slender.


Species Heterorhabdus spinifrons - Plate 26 of morphological figuresissued from : R.-M. Barthélémy in These Doct. Univ. Provence (Aix-Marseille I), 1999. [Fig.18, B]. Female (Gulf of Marseille: France): B, external ventral genital double-somite.
go = genital operculum.
Scale bar: 0.100 mm.


Species Heterorhabdus spinifrons - Plate 27 of morphological figuresIssued from : G.S. Brady in Rep. Scient. Results Voy. Challenger, Zool., 1883, 8 (23). [Pl. XIII]. As Heterochaeta spinifrons.
Female: 1, habitus (dorsal); 2, A1; 4, A2; 5, Md; 6, Mx1; 7, Mx2; 8, Mxp; 9, exopod of P3; 11, P5; 12, urosome with spermatophore attached; 13, portion of spermarophore with spermatozoids (more highly magnified).

Male: 3, left A1; 10, P5.


Species Heterorhabdus spinifrons - Plate 28 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 , Fig.3 ]. As Heterochäta spinifrons.
Male: 3, A1 (segments 1 to 16)


Species Heterorhabdus spinifrons - Plate 29 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 , Fig.3 ]. As Heterochäta spinifrons.
Male: 3, A1 (segments 15 to 25).

Compl. Ref.:
Cleve, 1904 a (p.191); Wilson, 1942 a (p.190); Massuti Alzamora, 1942 (p.95, Rem.); Lysholm & al., 1945 (p.35); Sewell, 1948 (p.349, 503, 509, 516, 519, 521, 530, 534, 539, 547, 558, 567); C.B. Wilson, 1950 (p.240); King & Hida, 1955 (p.11); Fagetti, 1962 (p.30); Ganapati & Shanthakumari, 1962 (p.8, 15); V.N. Greze, 1963 a (tabl.2); Grice, 1963 a (p.496); Ahlstrom & Thrailkill, 1963 (p.57, Table 5, abundance); Björnberg, 1963 (p.53, Rem.); Unterüberbacher, 1964 (p.29); De Decker & Mombeck, 1964 (p.12); Grice & Hulsemann, 1965 (p.224); Pavlova, 1966 (p.44); Mazza, 1966 (p.71); 1967 (p.367); Fleminger, 1967 a (tabl.1); Grice & Hulsemann, 1967 (p.18); De Decker, 1968 (p.45); Morris, 1970 (p.2301); Park, 1970 (p.477); Deevey, 1971 (p.224); Roe, 1972 (p.277, tabl.1, tabl.2); Apostolopoulou, 1972 (p.328, 362); Björnberg, 1973 (p.345, 387); Guglielmo, 1973 (p.399); de Bovée, 1974 (p.109, 124); Vives & al., 1975 (p.49, tab.II, III); Deevey & Brooks, 1977 (p.256, tab.2, Station "S"); Carter, 1977 (1978) (p.36); Dessier, 1979 (p.206); Vaissière Séguin, 1980 (p.23, tab.1); Vives, 1982 (p.293); Kovalev & Shmeleva, 1982 (p.84); Guangshan & Honglin, 1984 (p.118, tab.); Tremblay & Anderson, 1984 (p.5: Rem.); De Decker, 1984 (p.317); Cummings, 1984 (p.163, Table 2); Scotto di Carlo & al., 1984 (1043); Regner, 1985 (p.11, Rem.: p.36); Brenning, 1985 a (p.28, Table 2); Madhupratap & Haridas, 1986 (p.105, tab.1); Lozano Soldevilla & al., 1988 (p.59); Heinrich, 1990 (p.18); Pancucci-Papadopoulou & al., 1990 (p.199); Suarez & al., 1990 (tab.2); Suarez & Gasca, 1991 (tab.2); Suarez, 1992 (App.1); Hattori, 1991 (tab.1, Appendix); Scotto di Carlo & al., 1991 (p.270); Kouwenberg, 1994 (tab.1); Shih & Young, 1995 (p.70); Errhif & al., 1997 (p.422); Hure & Krsinic, 1998 (p.66, 102); Suarez-Morales & Gasca, 1998 a (p.110); Lapernat, 1999 (p.20, 55); 2000 (tab.3); Razouls & al., 2000 (p.343, Appendix); Lopez-Salgado & al., 2000 (tab.1); d'Elbée, 2001(tabl. 1); Lapernat & Razouls, 2001 (p.123, tab.1); Holmes, 2001 (p.16); Vukanic, 2003 (139, tab.1); Hsiao & al., 2004 (p.326, tab.1); Lo & al., 2004 (p.89, tab.1); Dias & Araujo, 2006 (p.51, Rem., chart); Dur & al., 2007 (p.197, Table IV); Neumann-Leitao & al., 2008 (p.799: Tab.II, fig.6); Fernandes, 2008 (p.465, Tabl.2); Galbraith, 2009 (pers. comm.); Licandro & Icardi, 2009 (p.17, Table 4); Williamson & McGowan, 2010 (p.273, Table 3, Pacific central gyres: N and S); Schnack-Schiel & al., 2010 (p.2064, Table 2: E Atlantic subtropical/tropical); Hidalgo & al., 2010 (p.2089, Table 2); Mazzocchi & Di Capua, 2010 (p.425); Medellin-Mora & Navas S., 2010 (p.265, Tab. 2); Hsiao S.H. & al., 2011 (p.475, Appendix I)
NZ: 22

Distribution map of Heterorhabdus spinifrons by geographical zones
Species Heterorhabdus spinifrons - Distribution map 4issued from : C. de O. Dias & A.V. Araujo in Atlas Zoopl. reg. central da Zona Econ. exclus. brasileira, S.L. Costa Bonecker (Edit), 2006, Série Livros 21. [p.51].
Chart of occurrence in Brazilian waters (sampling between 22°-23° S)
Nota: sampling only 2 specimens.
Loc:
Antarct. (Indien), sub-Antarct. (Indien, Pacif. SE), Afr. S (E & W), Namibie, off Tristan da Cunha E, off I. Trinidade, off Ste Hélène S, Brésil, Congo, G. de Guinée, Is. du Cap Vert, off Is. du Cap Vert NW, off Mauritanie, Maroc-Mauritanie, Canaries, off Madère, off Portugal, off Amazone, Venezuela, Mer des Antilles, Caribbean Colombia, Caraïbes, G. du Mexique, Cuba, Floride, Mer des Sargasses, off Bermuda (Station "S"), G. du Maine, Woods Hole, Islande S, off Irlande SW, Açores, G. de Gascogne, off Cap Finisterre, Baie Ibéro-marocaine, Médit. (Mer d'Alboran, Castellon, Banyuls, G. du Lion, Ligurian Sea, Tyrrhénienne, Milazzo, Messine, off Malte, Adriatique, Ionienne, Mer Egée, bassin Levantin), G. d’Aden, Mer Arabe, Is. Laquedives, off Minicoy Is., Natal, Indien, Inde (Lawson's Bay), G. du Bengale, Indien S (convergence subtropicale), Indonésie-Malaisie, Philippines, mers de Chine (East China Sea, South China Sea), Taiwan (SW, E, N: Mienhua Canyon), Japon (Izu), off Japon S, off Sanriku, Nansei Is., Mer de Béring, Station "P", off British Columbia, Pacif. W (équatorial), Australie (Grande Barrière), Mer de Tasman N, Nouvelle-Zélande, off Tasmanie S, Pacif. (central sutropical N), off Hawaii S, off California, San Diego, W Baja California, Pacif. (équatorial), Pacific central gyres: N and S, Pâques NE, Galapagos SE, off Pérou, Pacif. SE (tropic.), Chili (N-S), Straits of Magellan (Pacific and Central area)
N: 130
Lg.:
(1) F: 3,3; (11) F: 3,433; (16) F: 3,4-2,8; M: 3,05-2,85; (17) F: 3; (22) F: 3,5-3; M: 3,4-2,95; (24) F: 4; (34) F: 2,14; M: 3,14; (35) F: 3,6; (36) F: 3,82-3,49; M: 3,59-3,43; (38) F: 3,12-2,96; M: 3,4-2,7; (45) F: 3,5-3; M: 3,3-2,95; (46) F: 3,15-3; (73) M: 3,04-2,77; (101) F: 3,57-2,1; M: 3,2-2,41; (121) F: 3,48; M: 3,19-2,95; (142) F: 3,4; (199) F: 3,5-2,74; M: 2,96-2,2; (205) M: 3,4-3,2; (207) F: 3,4-2,92; M: 3,2-2,76; (237) F: 2,5; M: 2,01; (260) F: 3,4-2,92; M: 3,16-2,88; (340) F: 3; (432) F: 3,8-3,3; (449) F: 3,4-3; M: 3,4-2,95; (808) F: 3,46-3,07; (824) F: 4-3,04; M: 3,84-3; (909) F: 3,4-3,9; M: 2,6-3,5; {F: 2,10-4,0; M: 2,01-3,84}
Rem.: "Spinifrons Group".
epi-meso-bathypelagic.
Sampling depth (Antarct., sub-Antarct.) : 0-600 m. Overall Depth Range in Sargasso Sea: 0-1500 m (Deevey & Brooks, 1977, Station "S");
After Björnberg (1963, p.53) this species is recorded for the first time off Brazil.
Confusions are possible between this species, previously described in the ancient literature and the new species described by Park (2000) from the Indo-Pacific.
After Lapernat & Razouls (2002, p.19) the Itoh's index value = 2751.6 (number of teeth: 4) on the right Md, and 3902 on the left Md (number of teeth: 3).
Last update : 13/05/2013
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