Species Card of Copepod
Calanoida ( Order )
    Calanoidea ( Superfamily )
        Calanidae ( Family )
            Calanus ( Genus )
Calanus hyperboreus  Kröyer, 1838   (F,M)
Syn.: Calanus magnus Lubbock,1854;
Calanus plumosus Lubbock,1854;
no C. hyperboreus : Wilson, 1942 a (p.173)
Ref.:
Kröyer, 1838 (in Damkaer & Damkaer, 1979, p.20); Giesbrecht, 1892 (p.91, 128, figs.F); Giesbrecht & Schmeil, 1898 (p.15); Sars, 1901 a (1903) (p.12, figs.F,M); Mrŕzek, 1902 (p.506); Farran, 1908 b (p.20); Lysholm, 1913 (p.5); With, 1915 (p.30, figs.F); Sars, 1925 (p.6); Rose, 1929 (p.6); 1933 a (p.57, figs.F,M); Jespersen, 1934 (p.34, Rem.); 1940 (p.7); Lysholm & al., 1945 (p.6); Brodsky, 1950 (1967) (p.86, figs.F,M); Farran & Vervoort, 1951 (n°32, p.3, fig.F); Wiborg, 1954 (p.103); 1955 (p.24); Tanaka, 1956 (p.258, Rem.); Conover, 1965 (p.153, figs.F,M, Rem. intersex.); 1965 a (p.308, figs.juv.5); Vinogradov, 1968 (1970) (p.45, 61, 63, 65, 86, 90, 95, 96, 112, 234, 262, 266); Shih & al., 1971 (p.36, 202); Vidal, 1971 a (p.11, 20, 113, figs.F,M); Marshall & Orr, 1972 (p.8); Brodsky, 1972 (1975) (p.9, 68, 84, 117, figs.F,M); Vyshkvartzeva, 1972 (1975) (p.188, figs.); Bradford & Jillett, 1974 (p.6); Vyshkvartzeva, 1976 (p.14); 1977 a (p.97, figs.); Brodsky & al., 1983 (p.174, figs.F,M, Rem.); McLaren & Marcogliese, 1983 (p.721, cell nucleus); Fleminger, 1985 (p.275, 285, Table 1, 4, fig.M), Rem.: A1); Bradford, 1988 (p.76, Rem.); Schnack, 1989 (p.137, fig.7: Md); Bucklin & al., 1995 (p.658); Harris, 1996 (p.95, 98); Melle & Skjoldal, 1998 (p.211, Rem.); Lindeque & al., 1999 (p.91, Biomol.); Bartthélémy, 1999 a (p.10, Fig.19); Hill & al., 2001 (p.279, fig.2: phylogeny); G. Harding, 2004 (p.6, figs.F,M); Dalpadado & al., 2008 (p.2266, Fig.2: Md, Table 2, 3)
Species Calanus hyperboreus - Plate 1 of morphological figuresissued from : R.J. Conover in Crustaceana, 1965, 8. [p.154, Fig.1].
Comparison of head appendages and fifth legs of normal male and female and the intersex (from Long Island, USA): a, P5 (normal female); b, idem (normal male; c, idem (intersex); d, mandible blade (normal female); e, idem (normal male); f, idem (intersex); g, precoxal gnathobase on Mx1 (male; h, idem (intersex); i, coxa and basipod of Mxp (male); j, idem (female); k, idem (intersex).

Nota: In normal animals, the inner margin of the coxae of P5 usually bears from 19 to 31 teeth which are not continuous to the distal end of the segment. There seems to be no sexual difference in the number of teeth, but the inner margin bearing them is straight or convex in the female and concave in the male. The exopod and endopod consist of 3 segments each in the adult of both sexes. The P5 male are slightly asymmetrical, particularly the most distal segment of the left exopod which is concave along the inner margin and lined with short, stiff bristles.
The P5 interxex were abnormal, with the exopod and endopod of left leg both 2-segmented, as was also the exopod of the right leg. The coxae of the intersex's P5 appear to be of the female type (compare figs. 1a and c), with coxal teeth counts of 19 and 20 on the left and right sides, respectively.
In normal adult, the mouth parts of C. hyperboreus are similar to those of C. finmarchicus, but because of the differences in feeding behavior between the sexes a more detailed study was made of those of C. hyperboreus: 1- A2 consists of an exopod 7-segmented and an endopod 2-segmented, each differing only slightly between sexes except in relative size; both rami are proportionally longer in males than in either stage V or adult females, suggesting allometric enlargement in the last molt. 2- Md blabe is of the typical Calanus type described by Beklemishev (1959), but there is a marked reduction in width of the chewing surface in the male, the entire blade in the male is relatively soft and the teeth lack a heavy sclerotic coat; also missing in the males are the short stiff bristles found in both stage V copepodids and females along the ventral edge of the blade near its narrowest dimension. 3- the anterior labrum is soft and has reduced musculature in the male. 4 - The length of Mx2 is relatively less than in males and the setation is somewhat reduced. 5 - Mxp was the only mouthpart measured taht failed to show allometric growth between the stages; the basal 2 segments (coxa and basipod) also show considerably reduced setation in the male as compared with the female.
In the intersex, the mouthparts were largely of the female type. On dissection, the genital system showed both male and female characteristics (see figure 2, below).


Species Calanus hyperboreus - Plate 2 of morphological figuresissued from : G.O. Sars in An Account of the Crustacea of Norway. Vol. IV. Copepoda Calanoida. Published by the Bergen Museum, 1903. [Pl. V].
Female & Male.


Species Calanus hyperboreus - Plate 3 of morphological figuresissued from K. Hulsemann in Invert. Taxon., 1994, 8. [p.1477, Fig.28, D].
Female: D, urosome (left: ventral); right: dorsal). Pore signature schematic by pooled samples (symbols are considerably larger than pores): Filled circle: 100 % presence; open circle: 95-99 % presence; triangle: 50-89 % presence. n = 15.


Species Calanus hyperboreus - Plate 4 of morphological figuresissued from : N.V. Vyshkvartzeva in Issled. Fauny Moreď, 1972, 12 (20). [p.166, Fig.5, i, a, b, g].
Femele Md (masticatory edge):1a, lateral aspect; 1b, left Md (bove view); 1g, right Md (above view)


Species Calanus hyperboreus - Plate 5 of morphological figuresissued from : R.-M. Barthélémy in These Doct. Univ. Provence (Aix-Marseille I), 1999. [Fig.19, B, G]. Female (from W coast of Greenland: Arctic): B, external ventral view genital double-somite; G, internal dorsal view genital area.
go = genital operculum; m1operculum muscle; m2 = muscles of egg-laying ducts; ed = egg-laying ducts; sr = seminal receptacles.
Scale bars: 0.200 mm (F); 0.100 mm. (G)


Species Calanus hyperboreus - Plate 6 of morphological figuresissued from : A. Fleminger in Mar. Biol., 1985, 88. [p.283, Fig.5 C]. As Calanus s.l. hyperboreus.
Male (from N Pacific, subarctic): Left A1 proximal segments (ventral view);
Nota: see remarks in Calanus s.l. pacificus californicus (Fleminger, 1985, p.275) concerning the dimorphism in the female A1.


Species Calanus hyperboreus - Plate 7 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, 19 , Atlas von 54 Tafeln. [Taf.6, Fig.2].
Female: 2, thoracic segment 5 and genital segment (lateral).


Species Calanus hyperboreus - Plate 8 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, 19 , Atlas von 54 Tafeln. [Taf.6, Fig.6].
Female (pro parte): 6, cephalosome and thoracic segment 1 (ventral).
Ce =cephalosome; Th1 = thoracic segment 1; Vord. Ant. = A1; Hint. Ant. = A2; Man = MdMax. = Mx1; Vord. Mxpd = Mx2;Hint. Mxpd. = Mxp.; 1-Schwimm-fuss = swimming leg 1 (P1).


Species Calanus hyperboreus - Plate 9 of morphological figuresissued from : S.B. Schnack in Crustacean Issue, 1989, 6. [p.144, Fig.7: 1].
1, Calanus hyperboreus (from Arctic): Cutting edge of Md.
V = ventral teeth; C = central teeth; D = distal teeth.

Nota: Schnack underlines the geographic differences in the development of the ventral teeth (see alsoin Beklemishev, 1959; Sullivan & al., 1975 and Vyshkvartseva, 1975. In polar regions, the 2nd ventral tooth is well developed, whereas it is small or absent in copepods of warmer regions, in relation wiith the size of diatoms, whereas in warmer regions, where small flagellates and coccolithophorids are more abundant, a 2nd well-developed ventral tooth would be unnecessary (see in the Md of Calanus carinatus).


Species Calanus hyperboreus - Plate 10 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, 19 , Atlas von 54 Tafeln. [Taf.7, Fig.12].
Female: 12, Md (anterior view).


Species Calanus hyperboreus - Plate 11 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, 19 , Atlas von 54 Tafeln. [Taf.8, Fig.10].
Female: 10, endopod of P1 (anterior view).


Species Calanus hyperboreus - Plate 12 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, 19 , Atlas von 54 Tafeln. [Taf.8, Fig.23].
Female: 23, P5 (inner margin of B1 for two individuals).
B1 = coxa.


Species Calanus hyperboreus - Plate 13 of morphological figuresissued from : H.-J. Hirche & B. Niehoff in Polar Biol., 1996, 16. [p.216, Fig.6].
Gonads of female Calanus hyperboreus. Examples of mature females from the field and from females collected in August 1992.
a: collected in June 1991; b: gonad with mature oocytes, which, however, were not deposited; c: Note the large oil sac in the anterior cephalothorax region in a and the thin remnants of the oil sac in b and c.


Species Calanus hyperboreus - Plate 14 of morphological figuresIssued from : R.J. Conover in Crustaceana, 1965, 8 (3). [p.313, Fig.1]
Calanus hyperboreus (from SE Cape Cod, Massachusetts): a, P5 of a normal stage V (sex undetermined); b, outer ramus of the left P5 of incomplete molt showing the formation of the male type P5 within the stage V structure.


Species Calanus hyperboreus - Plate 15 of morphological figuresIssued from : R.J. Conover in Crustaceana, 1965, 8 (3). [p.314, Fig.2]
Calanus hyperboreus (from SE Cape Cod, Massachusetts): Mandible blades of incomplete molts showing the adult structure whithin the stage V test. a, stage V to female, b, stage V to male.


Species Calanus hyperboreus - Plate 16 of morphological figuresIssued from : R.J. Conover in Crustaceana, 1965, 8 (3). [p.315, Fig.3]
Calanus hyperboreus (from SE Cape Cod, Massachusetts): a, ovary from stage V in medium stage of development; b, testis from stage V at medium stage of development; c, testis of recently molted male.

Nota : In his study, Conover points to a number of characters have been considered as possible criteria to distinguish the sex of living stage V copepodids.
In addition to urosome shape and the head length-body ratio, the appearance of the immature gonad as seen through the dorsal surface, the shape and color of the oil sac, and the pigmentation of the base of the first urosomal segment were used.
Dissection of preserved animals showed that the immature gonad was usually a reliable means of distinguishing sex during the fall and winter months. Early in its development, the immature gonad, probably in both sexes, cponsisted of a small cluster of spherical cells of uniform size (± 15µ) visible under high power of dissecting microscope.
As the female gonad developed, the oogonia could be seen to grade gradually into larger developing oocytes (a). The immature male gonad was distinguishable by its more dense, fine-grained appearance and its more nearly oval- or pear-shape (b, c).


Species Calanus hyperboreus - Plate 17 of morphological figuresIssued from : S. Falk-Petersen & al. in Deep-Sea Res. II, 2008, 55. [p.2282, Fig.9].
Arctic Ocean: the lipid-rich Calanus hyperboreus with a green gut and a well-developed lipid sac, collected at the Ice station 1 (82°N, 11°E) on 2 September


Species Calanus hyperboreus - Plate 18 of morphological figuresIssued from : B.J. Hansen, K. Degnes, I.B. Řverjordet, D. Altin & T.R. Střrseth in Polar Biol., 2013, 36. [p.1578, Fig.1, A].
Calanus hyperboreus Stage V from Kongsfjorden, Svalbard (79°N, 12°E).
Scale bar = 2 mm.

Compl. Ref.:
T. Scott, 1902 (p.450); Damas & Koefoed, 1907 (? part., p.352, tab.II, III); Sars, 1909 b (p.16); Sřmme, 1934 (p.1, production); Jespersen, 1939 (p.26, Rem., Table 25, 26, 27, 28, 29, 30); Barnes, 1949 (p.429, statistical variations); C.B. Wilson, 1950 (part., p.178, non stations Pacif.); Gundersen, 1953 (p.1, 15, seasonal abundance); Řstvedt, 1955 (p.14: Table 3, p.17, 35, 55); Lacroix, 1960 (p.11, 25); Conover, 1960 (p.399, Table I, respiratory rate); Raymont & Conover, 1961 (p.154, carbohydrate content); Grice, 1962 a (p.101, 102); Marshall & Orr, 1962 (tab.3); Conover, 1962 (p.190, metabolism-growth); Grainger, 1963 (p.66, fig.10, 12, chart, size); Grice, 1963 a (p.495); M.W. Johnson, 1963 (p.89, Table 1, 2); Mullin, 1963 (p.239, grazing rate vs. diet); Brodsky, 1964 (p.105, 107); Conover, 1964 (p.81, Table 2, fig.1, nutrition, assimilation); 1965 a (p.308, moulting cycle, sexual characters, sex ratio); Grice & Hulsemann, 1965 (p.223); Conover, 1966 (p.338, Table I, 2, assimilation, fecal pellets-bacteria); 1966 a (p.346, feeding, assimilation); 1966 b (p.187, feeding on large particles); Harding, 1966 (p.17, 65, 66, 71); Matthews, 1967 (p.159, Table 1, Rem.); Conover, 1967 (p.61, egg, development time, fecundity); Corner & Cowey, 1968 (p.393, Table 7, 8, respiration rate, assimilation); Conover & Corner, 1968 (p.49, 53, 65, respiration & nitrogen excretion); Dunbar & Harding, 1968 (p.319); Itoh, 1970 a (p.1, tab.2); Corkett, 1972 (p.171, eggs: development rate); Matthews & Sands, 1973 (p.19, Table 4); Beaudouin J., 1973 (p.69); Harding, 1974 (p.141, tab.2, gut contents); Lee R.F., 1974 a (p.313, lipids); Landry, 1975 a (p.434, Rem.: p.437, fig.3); Bousfield & al., 1975 (p.1, estuary); Sargent, 1976 (p.149, lipids); Skjoldal & Bamstedt, 1977 (p.197, adenosides); Deevey & Brooks, 1977 (p.256, Table 2, Station "S"); Matthews & al., 1978 (p.277, annual cycles); Dawson, 1978 (p.950, vertical distribution, generation length); Poulet & Marsot, 1980 (p.198, Table 1, feeding); Kosobokova, 1980 (p.84, caloric value); Bamstedt & Skjoldal, 1980 (p.304, weight-RNA); Hebert & Poulet, 1980-81 (p.121, feeding v.s. oil polluant); Huntley, 1981 (p.831, ingestion rate vs. food concentration); Gagnon & Lacroix, 1981 (p.401, Table 1, tidal effect); [Kovalev & Shmeleva, 1982 (p.82)]; Buchanan & Sekerak, 1982 (p.41, vertical distribution); Gagnon & Lacroix, 1982 (p.9, fig.4); 1983 (p.289, tidal estuary); Head & Conover, 1983 (p.219, digestive enzymes); Huntley & al., 1983 (p.143, Table 2, 3); McLaren & Marcogliese, 1983 (p.721, body size vs. nucleus counts); Bamstedt, 1983 (p.291, RNA variation); Tremblay & Anderson, 1984 (p.4); Sameoto, 1984 (p.213, Table 1, fig.3, 7, 8); 1984 a (p.767, vertical migration); Head & Harris, 1985 (p.99, feeding activities, digestive enzymes); Head & al., 1985 (p.281, gut pigment analysis); Smith & al., 1985 (p.693); Hargrave & al., 1985 (p.221, annual abundance); Bamstedt, 1985 (p.607, excretion rate); Bamstedt & Tande, 1985 (p.259, Table 2: literature data respiration & excretion); Groendahl & Hernroth, 1986 (tab.1); Head & al., 1986 (p.271, grazing); Conover & al., 1986 (p.878, Table1, filtration rate); Mikhailovsky, 1986 (p.83, Table 1, ecological modelling); Hirche, 1987 (p.347, activity, respiration v.s. temperature); 1987 a (p.431, spatial distribution, enzyme activities, nutrition); Falk-Petersen & al., 1987 (p.115, lipid composition); Huntley, 1988 (p.83, Table 1, feeding history); Conover & al., 1988 (p.267, 268); Head & al., 1988 (p.333, Table 1, 2, gut analysis, defecation rate); McLaren & al., 1988 (p.275, DNA content, development rate: egg-nauplius); S.L. Smith, 1988 (p.145, feeding, respiration, ammonium, excretion ice-edge effect); Bamstedt, 1988 (p.15, protein content); McLaren & al., 1989 (p.560, life history, annual production); Kattner & al., 1989 (p.473, Table 1, 2, 3, dry weight, lipids); Kosobokova, 1989 (p.27); Hirche, 1989 (p.431, spatial distribution, enzyme activity); Ikeda & Skjoldal, 1989 (p.173, oxygen consumption, N & P excretion, O:N vs. body weight); Citarella, 1989 (p.123, abundance); S.L. Smith, 1990 (p.59, egg production, lipid, gut content); Estep & al., 1990 (p.235, grazing); Hansen B. & al., 1990 (p.5, grazing); Fransz & al., 1991 (p.9); Conover & al., 1991 (p.177); Hirche, 1991 (p.351); Hirche & al., 1991 (p.477, Fig.7, 8, Table 2); Conover & Huntley, 1991 (p.1, fig.2, 3, 4, 5, Table 2, 3, 9, 10, 11, polar seas comparison); Head, 1992 (p.583, gut pigment destruction); Hirche & Mumm, 1992 (p.S485, geographic distribution); Herman, 1992 (p.395, fig.9 a, size distribution by OPC); Huntley & Lopez, 1992 (p.201, Table 1, A1, eggs, egg-adult weight, temperature-dependent production); Mumm, 1993 (tab.1, fig.2); Richter, 1994 (tab.4.1a); Vinogradov & al., 1994 (tab.1); Petryashov & al., 1995 (tab.1); Ashjian & al., 1995, p.4371, Fig.4, 5, 6, Table 2, 4); Hays, 1995 (p.301, fig.1, Table 1, vertical migration); Hirche & Niehoff, 1996 (p.209, vertical distribution, reproduction); DFO, 1996 (p.1, fig.6, interannual abundance); Albers & al., 1996 (p.347, lipids v.s. diet); Zauke & al., 1996 (p.141, Table 7, metal bioaccumulation); Hanssen, 1997 (tab.3.1); Hirche & Kwasniewski, 1997 (p.299, Table 1, 4, 5, Fig.4, 10, 12, 13); Falkenhaug & al., 1997 (p.449, spatio-temporal pattern); Daly, 1997 (p.319, Table 4, fecal pellet); Ashjian & al., 1997 (p.279, Table 1, 2, Figs. 2, 3, 4A,F); Weslawski & Legezynska, 1998 (p.1238); Kosobokova & al., 1998 (tab.2); Mauchline, 1998 (tab.21, 26, 33, 45, 46, 47, 48, 58, 63); Mumm & al., 1998 (p.189, Figs.3, 4); Niehoff, 1998 (p.53, gonad maturation); Melle & Skjoldal, 1998 (p.211, egg production, development); Sameoto & al., 1998 (p.1, 7, figs. 8, 9, spatial distribution, interannual variation); Conover & Gustavson, 1999 (p.41, tab.6); Halvorsen & Tande, 1999 (p.284, tab.2, 3, Rem.: p.281); B.W. Hansen & al., 1999 (p.233, seasonal abundance & biomass); Thibault & al., 1999 (p.1391); Kosobokova & Hirche, 2000 (p.2029, tab.2); Huggett & Richardson, 2000 (p.1843, tab.2); White & McLaren, 2000 (p.751, tab.1); Musaeva & Gagarin, 2000 (p.534, tab.1); Beare & al., 2000 (p.1545, Arctic index indicatot); DFO, 2000 a (p.1, Rem.: p.8, fig., interannual variations); Musaeva & Suntsov, 2001 (p.511); Lischka & al., 2001 (p.186); Sameoto, 2001 (p.749, Table 4, Rem.: decadal changes); Johns & al., 2001 (p.2121, Rem.: long-term series); Madsen & al., 2001 (p.75, development & production vs. annual); Fortier M. & al., 2001 (p.1263, fig.6, 7, diel vertical migration); Holmes, 2001 (p.37); Beaugrand & al., 2002 (p.1692); Beaugrand & al., 2002 (p.179, figs.5, 6); Auel & Hagen, 2002 (p.1013, tab. 2, 3); Pasternak & al., 2002 (p.147, Table 4, feeding activity vs. egg production, faecal pellets); Ringuette & al., 2002 (p.5081, Table 1, 2, Fig.6, population dynamic); Sameoto & al., 2002 (p.12); Astthorsson & Gislason, 2003 (p.843); Hirche & Kosobokova, 2003 (p.769, Fig.3, 9, Table 2, 3); Ashjian & al., 2003 (p.1235, figs.); Lindeque & al., 2004 (p.121, fig.2); Gislason & Astthorsson, 2004 (p.472, tab.1, fig.4); CPR, 2004 (p.50, fig.140); Veistheim & al., 2005 (p.382, tab.2, fig.1); Dmoch & Walczowski, 2005 (p.102 + poster); Thor & al., 2005 (p.341); Hirche & al., 2005 (p.310); Somoue & al., 2005 (Table I: p.66); Arnkvaern & al., 2005 (p.528, dynamic); Hopcroft & al., 2005 (p.198, table 2); Blachowiak-Samolyk & al., 2006 (p.101, tab.1); Lindeque & al., 2006 (p.221); Basedow & al., 2006 (p.1186: Table II); Hop & al., 2006 (p.182, Table 4, 5: inter-annual variability,); Olli & al., 2007 (p.84, Rem.: ice drift); Daase & al., 2007 (p.903, abundance/T°S); Deibel & Daly; 2007 (p.271, Table 1, 2, 3, 5, Rem.: Arctic polynyas); Blachowiak-Samolyk & al., 2007 (p.2716, Table 2); Falk-Petersen & al., 2007 (p.147, Table 9.1); Lane & al., 2008 (p.97, Tab.4, 6, fig.6); Sřreide & al., 2008 (p.2225, feeding strategy); Ota & al., 2008 (p.215, nauplii); Darnis & al., 2008 (p.994, Table 1, figs.8, 9); Tamelander & al., 2008 (p.2330, Table 1, organic matter); Gaard & al., 2008 (p.59, Table 1, N Mid-Atlantic Ridge); Jensen & al., 2008 (p.100); Pasternak & al., 2008 (p.2245, Table 1, 2, 3, grazing); Falk-Petersen & al., 2008 (p.2275, depth distribution); Blachowiak-Samolyk & al., 2008 (p.2210, Table 2, 3, 5, fig.4, biomass, composition vs climatic regimes); Walkusz & al., 2008 (p.1, Table 3, abundance); Pepin & al., 2008 (p.1, 9, figs. 21, 26, 31, 34, interannual variations); Dvoretsky & Dvoretsky, 2009 a (p.11, Table 2, abundance); Harvey & Devine, 2009 (p.5, interannual variation); Campbell & al., 2009 (p.1274, Table 2, 3, fig.3, grazing); DFO, 2009 (p.1, Rem. p.12, fig. 13, seasonal variability); Kosobokova & Hirche, 2009 (p.265, Table 4, fig.7: chart, biomass); Kosobokova & Hopcroft, 2010 (p.96, Table 1, fig.7); Head & Pepin, 2010 (p.1633, inter-decadal variability); Arendt & al., 2010 (p.49, Rem.: p.43); Bucklin & al., 2010 (p.40, Table 1, Biol mol.); Dünweber & al., 2010 (p.11, biomass, gut content); Dvoretsky & Dvoretsky, 2010 (p.991, Table 2); 2011 a (p.1231, Table 2: abundance, biomass); Templeman, 2010 (p.1, 15: fig.12, interannual variations); Kwasniewski & al., 2010 (p.72, Table 2, abundance vs hydrography); Dvoretsky V.G., 2011 (p.361, abundance, stage composition); Kosobokova & al, 2011 (p.29, Table 2, figs.4, 6, Rem.: Arctic basins); Tang & al., 2011 (p.77, composition & biomass); Pomerleau & al., 2011 (p.1779, Table III, IV, V, VI, VII); Swalethorp & al., 2011 (p.429, grazing, egg production and life strategies); Forest & al., 2011 (p.161, biomass, chemical composition); 2011 a (p.11418); 2012 (p.1301, figs.7, 8); Matsuno & al., 2012 (Table 1, 2, 3, fig.4, 7); Laakmann & al., 2012 (p.535, Table 1, fig.2, Rem.: mol. Biol.); Carstensen & al., 2012 (p.951, Fig.2, 10, biomass); Tammilehto & al., 2012 (p.165, nutition, algal toxicity); Dalpadado & al., 2012 (p.1, abundance vs. climate change); Johnson C & al., 2012 (p.1, 15, fig. 23a, 24a, interannual variarions); Davies & al., 2012 (p.614, energy content vs methods); Hansen B.H. & al., 2013 (p.1577, metabolism); Hsiao & Fang, 2013 (p.175, Table 2: Hg bioaccumulation); Questel & al., 2013 (p.23, Rem. p.31); Hirche, 2013 (p.2469, lefespan, reproduction)
NZ: 6

Distribution map of Calanus hyperboreus by geographical zones
Species Calanus hyperboreus - Distribution map 2
Chart of 1996
Species Calanus hyperboreus - Distribution map 3issued from : N. Mumm, H. Auel, H. Hanssen, W. Hagen, C. Richter & H.-J. Hirche in Polar Biol., 1998, 20. [p.192, Fig.1, p.194, Fig.3]
Fig.1 after Diepenbroek & al., 1997; Station map, the dark line connects stations of different expeditions to a transpolar transect (AB: Amundsen Basin, BS: Barents Sea; GL: Greenland; GS: Greenland Sea; LR: Lomonov Ridge; MB: Makarov Basin; MJP: Morris Jessup Plateau; NB: Nansen Basin; NG: Nansen-Gakkel Ridge; SB: Spitsbergen; WSC: West Spitsbergen Current; YP: Yermak Plateau

Fig.3: Biomass share (% of total mesozooplankton dry mass) of Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa and other taxa in 0- to 500 m depth of different Arctic regions (DM total: mean total dry mass).
Note the co-occuring of the three species, but the the different abundance according to the basins.
C. finmarchicus reached a maximum abundance in the WSC, this form was the dominant species in the West Spitsbergen Current and south of the central Nansen Basin.
Note C. hyperboreus is more important towards the north and less common in the West Spitsbergen Current.
Species Calanus hyperboreus - Distribution map 4issued from : E.H. Grainger in R. Soc. Canada, Spec. Publs., 1963, 5. [p.87, Fig.10].
Baffin Bay and Davis Strait. Circles indicate ocurrence of C. hyperboreus , darkened segments showing copepodite stages present.
Numbers of stations are shown.
Species Calanus hyperboreus - Distribution map 5issued from : G.C. Hays in Mar. Ecol. Progr. Ser., 1995, 127. [p.302, Fig. 1];
a: Areas (shaded) from which details of the temporal occurrence of copepodite stages V-VI Calanus hyperboreus in the CPR samples were examined.
b: Mean abundance of CV-VI (specimens per sample ± 1 SE) in different months.
Species Calanus hyperboreus - Distribution map 6issued from : H.-J. Hirche & B. Niehoff in Polar Biol., 1996, 16. [p.210, Fig.1].
Calanus hyperboreus: Multinet sampling locations in the Greenland Sea.
Species Calanus hyperboreus - Distribution map 7issued from : H.-J. Hirche & B. Niehoff in Polar Biol., 1996, 16. [p.212, Fig.2].
Vertical distribution of female and male Calanus hyperboreus in the Greenland Sea.
In November data from three stations were pooled (613, 616, 653), in February and March stations 118 and 130a data were pooled. No males or very few after March.
Species Calanus hyperboreus - Distribution map 8issued from : H.-J. Hirche & B. Niehoff in Polar Biol., 1996, 16. [p.215, Table 3].
Egg production of Calanus hyperboreus in the Greenland Sea (n number of females in experiments).

Nota: Surprisingly, spawning of C. hyperboreus in the central Greenland Sea seems to take place earlier than in both more southern and northern latitudes. The reproductive cycle may relate to the timing of food abundance. Thus the spring bloom in the ice-free parts of the Greenland Sea occurs 2-3 months earlier than in the ice-covered Northwest Paasge (see Conover & Siferd, 1993).
Species Calanus hyperboreus - Distribution map 9issued from : B.T. Hargrave, G.C. Harding, K.F. Drinkwater, T.C. Lambert & W.G. Harrison in Mar. Ecol. Prog. Ser., 1985, 20. [p.227, Fig.7].
Major species of zooplankton present at the central station in St. Georges Bay (45°45'N, 61°45'W) during 1977.
Nota: All zooplankton collections were made after sunset. The net towed obliquely throughout the water column (± 34 m in depth).
Species Calanus hyperboreus - Distribution map 10issued from : B.T. Hargrave, G.C. Harding, K.F. Drinkwater, T.C. Lambert & W.G. Harrison in Mar. Ecol. Prog. Ser., 1985, 20. [p.223, Fig.2].
Seasonal profiles of water temperature and salinity in St. Georges Bay (45°45'N, 61°45'W) near the central station during 1977.
Species Calanus hyperboreus - Distribution map 11Issued from : R.J. Conover in Crustaceana, 1967, 13 (1). [p.67, Fig.2].
Record of weekly egg laying by sixteen typical females of Calanus hyperboreus taken from the plankton in immature condition and maintened with food in the laboratory.
Letters and numbers in right corner of each individual graph represent the laboratory code name. Beneath is total egg production. Histograms indicate number eggs produced per week. Solid line represents percentage developing hatching.

C. hyperboreus was taken at depths greater than 200 m either in the Gulf of Maine or in the slope water to the east or southeast of Caope Cod (Massachusetts), and fed with the diatom Thalassiosira fluviatilis, in concentration of 5 to 10 x 10power 6 cells/l.

Nota: In the Gulf of Maine C. hyperboreus produces 1 generation a year. Molting to the adult stage is generally concentrated in the late fall and early winter, but gravid females have been found from September to May.
Species Calanus hyperboreus - Distribution map 12issued from : R.J. Conover in Some Contemporary Studies in Marine Science; Harold Barnes, Ed., 1966 b. [p.188, Fig.1].
Anterior and slightly ventral view of Calanus hyperboreus female (from NW Atlantic) showing limbs in feeding position.
Stippled margin surrounds the area within which contact with a large particle can result in a successful capture.
A: second antenna (Antenna: A2); B: first maxilla (Maxillule: Mx1); C: second maxilla (Maxilla: Mx2); D: maxilliped (Mxp). L: labrum; E: interdigitated setae originating from the second maxillae and maxillipeds; F: tips of swimming legs.

Nota: Food sources included the copepod’s own eggs (about 200 µ in diameter), the diatom Coscinodiscus sp. (about 330-355 µ indiameter, 100-150 µ high) and Artemia salina nauplii (500-1000 µ long).
The large food particles passed posteriorly over the labrum and were introduced into the ‘filter chamber’ under the tips of the lowered swimming feet. The Mxp prevented vertical and lateral escape of particles and brought cells into a position where they could be seized by Mx2. Some particles excaped dorso-laterally between the maxillary palps and the base of the Mxp, but most on entering the refion outlined in figure 1 were at least temporarily. Although Mx2 serve primarily as a passive filter when food particles are small, they could also be moved forward and backward. The baxckward motion increased the distance between the setae on the two opposing Mx2 allowing a larger particle to slip between them and be grasped. Forward movement closed the setae and drove the object toward the mouth (see fig.2). A sudden backward movement lifted and spread the setae, causing an unwanted object to be flicked away, often aided by a quick flip of the swimming feet.
Also important in handling larger food particles are a set of five shorter, robust setae, one on the inner margin of each of the proximal four segments of Mx2 and one on the distal segment (see fig.3).
Once a large particle was contacted, i twas manipulated by co-ordinated effort of Mxp, Mx2, and Mx1 so that the greatest dimension was parallel to the long dimension of the copepod’s body.
Movement of Mx1 caused ventrally directed setae on the endopod to oppose the forward motion imparted to the particle by the shorter inner setae on the Mx2 and Mxp until the particle was ‘juggled’ into proper orientation. It was then driven forwards and dorsally until it could be reached by the endites of Mx1, which pressed it against the Md blades until consumed.
Muscular movement of the anterior diverticulum of the midgut apparently sucked the cell fragments inside the body as they were shredded by the Md.
A Coscinodiscus cell could be captured and eaten in as little as 4 seconds.
In most instances the diatom test was entirely consumed, occasionally the copepod discarded or lost a partially eaten cell.
An actively feeding animal would ingest an empty diatom frustule almost as readily as intact cells. On the other hand, semi-liquid cell contents could be partially lost and feeding appendages sometimes became coated with sticky cell sap. Even when feeding on small cells a portion of the cell contents was probably lost in feeding. Fifteen percent of the particulate carbon removed from suspension by C. hyperboreus feeding on Thalassiosira fluviatilis (15-20 µ diameter) re-appeared as dissolved organic substances, having been leached from damaged cells and excreted by the copepods (Helleburst & Conover).
Even in animals which fed most successfully, feeding was not a continuous process. Vigorously feeding animals might consume Coscinodiscus cells at nearly 1 per minute for 30 to 40 minutes, but eventually they would start to reject everything offered. Other animals would take several to half a dozen cells in short succession and then rejects cells for 10 to 30 minutes before feeding again.).
Even in animals which fed most successfully, feeding was not a continuous process. Vigorously feeding animals might consume Coscinodiscus cells at nearly 1 per minute for 30 to 40 minutes, but eventually they would start to reject everything offered. Other animals would take several to half a dozen cells in short succession and then rejects cells for 10 to 30 minutes before feeding again.

The Artemia nauplii were handled by the copepods somewhat differently from Coscinodiscus. The presence of the moving nauplius was apparently sensed from few millimetres away and the feeding response, lowering of the tips of the swimming feet and vigourous movement of the Mxp, was instituted before actual contact. The nauplius was seized in much the same way as a large cell and was then oriented by Mxp and both Mxp2 so that either the head or anal end entered the moth first. The actual ingestion was relatively slow process taking from 30 seconds to several minutes.
The endites of Mx1 would push the nauplius forward; then all movement of the head appendages would cease except for that of the mandibular blades. After a few seconds, the maxillae would again push the nauplius forward and the mandibular ‘chewing’ would be repeated. This alternation of processes was continued, sometimes with a brief interruption, until the nauplius was entirely consumed.

Among the more than fifty female and stage V animals examined, there was considerable variation in the hability to handle different particles.
Animals that had been feeding heavily on a unialgal culture of small cells were particularly inclined to reject large particles.
Other food preferences were also exhibited. One female layed eggs prior to the start of the observations and had been feeding on them. When offered Coscinodiscus she consistently rejected the cell, but readily ate her eggs when presented. Several other copepods, which had been feeding on Coscinodiscus, rejected the eggs after first bringing them to the mouth. If a fecal pellet was encountered, i twas usually oriented so that one end was ‘tasted’ and then i twas flicked away ; examination of the pellet showed that the peritrophic membrane had been torn open and the contents exposed.

After a brief period of feeding, the anterior portion of the midgut contained an abundance of rather fluid, greenish cell debris which was pushed back and forth by opposing waves of peristalsis. At least 40 minutes elapsed from the start of feeding before the first fecal pellet was ejected. Subsequent pellets might be produced at more frequent intervals, but not less than 15 or 20 minutes apart.

C. hyperboreus might assimilate about 0.06 µg carbon from each Coscinodiscus cell eaten. The maximum respiratory rate of female during the fall and early winter (when these observations were made) was about 25 µl O2/copepod/day oxidized if fat was the primary energy substrate or 13 µg C if carbohydrate was metabolized. Somewhere between 150 and 220 Coscinodiscus cells would have to be eaten daily to satisfy this requirement.
Species Calanus hyperboreus - Distribution map 13issued from : R.J. Conover in Some Contemporary Studies in Marine Science; Harold Barnes, Ed., 1966 b. [p.189, Fig.2].
Diagrammatic representation of the range of movement shown by the feeding appendages when handling large particles.
A, Mx1; B, Mx2; C, Mxp. D, setae on inner margin of Mx2; E, setae on basal segment of Mxp; L, labrum.
Species Calanus hyperboreus - Distribution map 14issued from : R.J. Conover in Some Contemporary Studies in Marine Science; Harold Barnes, Ed., 1966 b. [p.190, Fig.3].
Idealized lateral view of the left Mx2 and left Mxp showing the interdigitated setae on both structures. A, Mxp; B, Mx2; L, labrum.
Species Calanus hyperboreus - Distribution map 15Issued from : J.E. Sřreide, S. Falk-Petersen, E.N. Hegseth, H. Hop, M.L. Carroll, K.A. Hobson & K. Blachowiak-Samolyk in Deep-Sea Res., II, 2008, 55. [p.2228, Fig.1].
Study sites in the Svalbard region. The biomass (dry-weight b/m2) of the population of Calanus hyperboreus, C. glacialis and C. finmarchicus (CI-adult) is shown for the main sampling locations (data from Stn. NK2 is missing) and the Atlantic and Arctic water masses are indicated by arrows (WCS: west Spitsbergen Current). The location of the ice edge (defined as 30% ice concentrations) is indicated for selected dates.Issued from : J.E. Sřreide, S. Falk-Petersen, E.N. Hegseth, H. Hop, M.L. Carroll, K.A. Hobson & K. Blachowiak-Samolyk in Deep-Sea Res., II, 2008, 55. [p.2228, Fig.1].
Study sites in the Svalbard region. The biomass (dry-weight b/m2) of the population of Calanus hyperboreus, C. glacialis and C. finmarchicus (CI-adult) is shown for the main sampling locations (data from Stn. NK2 is missing) and the Atlantic and Arctic water masses are indicated by arrows (WCS: west Spitsbergen Current). The location of the ice edge (defined as 30% ice concentrations) is indicated for selected dates.

Nota: Stable isotope and fatty acid trophic marker techniques were employed together to assess trophic level, carbon sources (phytoplankton vs. ice algae), and diet of the three Calanus species.
Patterns in absolute fatty acid and fatty alcohol composition revealed that diatoms were the most important food for C. hyperboreus and C. glacialis, followed by Phaeocystis, whereas diatoms, Phaeocystis and other small autotrophic flagellates were equally important for C. finmarchicus.
Species Calanus hyperboreus - Distribution map 16Issued from : S. Falk-Petersen & al. in Deep-Sea Res., 2008, 55. [p.2282, Table 5].
Arctic Ocean, Ice Stations 1 (82°N, 11°E) on 2 September 2004, and 2 (82°30'N, 21°E) on 4 September 2004: Depth distribution of mesozooplankton in the upper 1200 m.
Species Calanus hyperboreus - Distribution map 17Issued from : S. Falk-Petersen & al. in Deep-Sea Res., 2008, 55. [p.2281, Fig.8].
Arctic Ocean, Ice Stations 1 (82°N, 11°E) on 2 September 2004, and 2 (82°30'N, 21°E) on 4 September 2004: Temperature (black profile), relative fluorscence values (red line), salinity (dotted line).
Species Calanus hyperboreus - Distribution map 18Issued from : K.W. Tang, T.G. Nielsen, P. Munk, J. Mortensen, E.F. Mřller, K.E. Arendt, K. Tönnesson, T. Juul-Pedersen in Mar. Ecol. Prog. Ser., 2011, 434. [p.83, Fig.4]
Calanus hyperboreus (from the continental slope off Fyllas Bank to the inner part of Godthabsfjord, SW Greenland, corresponding to stations 0 to 20) in the summer (2008).
Contour plots of biomass (mg C/m3) of all developmental stages collected from 4 to 9 strata with a multinet samples (300 µm mesh aperture)
Dots are mid-points of sampling intervals. Numbers on top are stations. Hatched area = bottom topography.

Compare this distribution with the other dominant large zooplankton species Calanus glacialis, Calanus finmarchicus and Metridia longa for the same transect.
Species Calanus hyperboreus - Distribution map 19Issued from : K.W. Tang, T.G. Nielsen, P. Munk, J. Mortensen, E.F. Mřller, K.E. Arendt, K. Tönnesson, T. Juul-Pedersen in Mar. Ecol. Prog. Ser., 2011, 434. [p.79, Fig.1]
Station positions along Godthabsfjord in southwestern Greenland.
Species Calanus hyperboreus - Distribution map 20Issued from : K.W. Tang, T.G. Nielsen, P. Munk, J. Mortensen, E.F. Mřller, K.E. Arendt, K. Tönnesson, T. Juul-Pedersen in Mar. Ecol. Prog. Ser., 2011, 434. [p.81, Fig.2]
Contour plots of water temperature (°C), salinity, density (kg/m3) and chlorophyll a (mg/m3) along the transect of Godthabsfjord.
Distances were measured from Station o. Note the different contour line scales for different panels.
Hatched area in each panel represents bottom topography.
Species Calanus hyperboreus - Distribution map 21Issued from : DFO in DRO Sci. Stock Status Rep. G3-02 (2000). [p.8].
Abundance of C. hyperboreus in Emerald Basin (Nova Scotia) during the years 1982 to 1998.
Nota: Compare with Calanus finmarchicus and Calanus glacialis.
See in Sameoto & al., 1997.
Loc:
Arct. (all polar Basins), Arct. (Fletcher's Ice Is.), Resolute Passage, Devon Island, Nansen Basin, Pechora Sea, Kara Sea, Laptev Sea, Lomonosov Ridge, Chukchi Sea, SE Beaufort Sea, Canada Basin, Foxe Basin, Amundsen Gulf, Barrow Strait, Fram Strait, Jones Sound, Canadian abyssal plain, Barrow Strait, N Baffin Sea, Disko Bay, off shore Godthabsfjord, Labrador Sea, Davis Strait, Greenland Sea, Fram Strait, Kongsfjorden, Spitzbergen, Svalbard (Kongsfjorden), Iceland, Faroe, Bear Island, Norway Sea, W Norway (Korsfjorden, Malangen fjord), Barents Sea, Franz Josef Land, off W Ireland, Shetland Is., North Sea, NW Atlant., off Woods Hole, Gulf of Maine, Bay of Fundy, off SE Nova Scotia, Roseway Basin, Northumberland Strait, Browns Bank, Emerald Bank, St Georges Bay, Baie-des-Chaleurs, G. of St. Lawrence, upper St. Lawrence estuary, Sargasso Sea (in Grice & Hart, 1962; Deevey & Brooks, 1977: depth range 500-1000 m at Station "S": 32°10'N, 34°30'W); off E Cape Cod (42*N, 64°30'W) Harding, 1974, off coast of southern Morocco (in Somoue & al, 2005)
N: 242
Lg.:
(7) F: 9-7,5; (22) F: 10-7; M: 7-5; (47) F: 9,6-6,9; (65) F: 9; M: 6,5; (131) F: 10-7; M: 7-5; (328) F; ± 6,77; (790) F: 8,7-5,3; M: 6,77-5,55; (1099) F: 6,25-10,0; {F: 5,30-10,00; M: 5,55-7,00}
Rem.: epi-bathypelagic. Overall Depth Range in Sargasso Sea: 500-1500 m (Deevey & Brooks, 1977, Station "S': Rem: submergent species at Station "S"); 2000-1000 m (slope) Harding, 1974..
No doubt mistakenly reported in the Mediterranean Sea and in the Pacific.
Reported far south from its natural area, its abundance seems to have increased in the NW Atlantic by 39°N (Johns & al., 2001). Observed at 11 coastal stations in March from the southern Morocco.
After Hirche (2013, p.2469) females collected in October produced up to 1,000 eggs and had a maximum lifespan of 164 days without feeding, whereas fed females produced up to ca. 6,000 eggs and survived up to 806 days. These females are multiannual-iteroparous, i.e. capable to spawn in successive years, which would be unique for calanoid copepods. There was no difference in the timing of reproductive activity between females from the West Spitsbergen Current and the Greenland Sea Gyre. Fed and starved females collected in May and June began to spawn circa 2 and 4 months after collections, respectively, whereas females collected in August and October started spawning at the same time, in the middle of October. This indicates initiation of reproductive activity in the field in August, coincident with the descent into deep waters. The large size of females, robutness and combination of different types of diapause in their life cycle make C. hyperboreus a good model organism to study diapause controm mechanisms.
Last update : 10/07/2014
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