Species Card of Copepod
Calanoida ( Order )
    Calanoidea ( Superfamily )
        Calanidae ( Family )
            Calanus ( Genus )
Calanus helgolandicus  (Claus, 1863)   (F,M)
Syn.: Cetochilus helgolandicus Claus, 1863 (p.171, figs.F,M);
Calanus finmarchicus : Giesbrecht, 1892 (part., p.89, figs.F); Farran, 1908 b (p.20, Rem.); Farran, 1926 (? part., p.227); 1929 (part., p.207, 212); Brian, 1914 a (p.134); Pesta, 1920 (p.493); Marques, 1966 (p.2); Mageed, 2006 (p.168, Table 4); Raymont & Krishnaswamy, 1960 (p.239, carbohydrate content);
no C. helgolandicus : Mori, 1937 (1964) (p.14, figs.F,M); Sewell, 1948 (p.550, 551, 544, 555, 565); C.B. Wilson, 1950 (p.178); Shih & Young,1995 (p.68);
Calanus : Harris & al., 1986 (p.845, Table 2, comparison pump vs net)
Sars, 1901 a (1903) (p.11, figs.F,M); Thompson & Scott, 1903 (p.232, 241); With, 1915 (p.12, Rem.); Früchtl, 1923 a (p.138, figs.F, Rem.); Sars, 1925 (p.6); Rose, 1929 (p.5); Wilson, 1932 a (p.25, figs.F,M); Rose, 1933 a (p.59, figs.F,M); Rees, 1949 (p.219, figs.F,M, juv.4,5); Brodsky, 1950 (1967) (p.99, figs.F,M); Farran & Vervoort, 1951 (n°32, p.3, Rem.); Marshall & Orr, 1952 (p.527, Table I: egg sizes); 1953 (p.1, fig.1: egg); Woodhead & Riley, 1957 (p.47, Rem.V, figs.); Jaschnov, 1957 (p.191, figs.); Furnestin, 1960 (p.167); Jaschnov, 1961 a (p.1317, biogeography); Brodsky, 1961 (p.15, figs.F,M); Gaudy, 1962 (p.93, 99, Rem.: p.100, Pl.I: juv., Tableau I: development); Giron-Reguer, 1963 (p.24); Harding, 1963 (p.81, karyotype); Grice, 1963 a (p.497, fig.F, Rem.); Vilela, 1965 (p.5); Mazza, 1967 (p.59, 62: clé juv.,F,M); Manwell & al., 1967 (p.145, biochimie); Matthews, 1967 a (p.159, Rev.); Vilela, 1968 (p.7); Koga, 1968 (p.16, fig.: egg); Corral Estrada, 1970 (p.62, figs.F, Rem.); Jillett, 1971 (p.29, Rem.); Marshall & Orr, 1972 (p.8, figs.F); Brodsky, 1972 (1975) (p.9, 66, 81, 119, figs.); Vyshkvartzeva, 1972 (1975) (p.188, figs.); Williams, 1972 (p.53, figs.F, carte); Razouls, 1972 (p.91, 94, Tableau XXVI; Annexe: p.3, figs.F,M, Rem.); Bradford & Jillett, 1974 (p.6); Frost, 1974 (p.74, Rev.); Fleminger & Hülsemann, 1977 (p.233, figs.F,M, geographical range-taxonomic divergence); Arnaud & al., 1980 (p.213, gut structure); Brodsky & al., 1983 (p.160, figs.F,M); van der Spoel & Heyman, 1983 (p.62, fig.79); Roe, 1984 (p.356); Sazhina, 1985 (p.23, figs.N); Grigg & al., 1987 (p.253, Rem. st.V); Fleminger & Hulsemann, 1987 (p.43: geographical mrophometric variation); Bradford, 1988 (p.76, Rem.); Nishida, 1989 (p.173, table 1, 2, 3: dorsal hump); Schnack, 1989 (p.137, tab.1, fig.6: Md); Bradford-Grieve, 1994 (p.31); Kouwenberg, 1994 (tab.1); Bucklin & al., 1995 (p.658); Harris, 1996 (p.95, 99); Kaartvedt, 1996 (p.145); Runge & Plourde, 1996 (p.171); Hure & Krsinic, 1998 (p.13, 99, 112); Lapernat, 1999 (p.10, 55); Lindeque & al., 1999 (p.91, Biomol.); Barthélémy, 1999 a (p.10, Fig.19); Bucklin & al., 1999 (p.239, molecular systematic); Bucklin & al., 2000 (p.1237, Rem.: analyse génétique moléculaire); Hill & al., 2001 (p.279, fig.2: phylogeny); Buttino & al., 2003 (p.469, fig. egg); G. Harding, 2004 (p.7, figs.F,M); Papadopoulos & al., 2005 (p.1353, phylogeography, molecular biology); Conway, 2006 (p.7: copepodite stages 1-6, Rem.); Unal & al., 2006 (p.1961, genetic analysis, phylogeography); Avancini & al., 2006 (p.59, Pl. 27, figs.F,M, Rem.); Ferrari & Dahms, 2007 (p.32, 34, Rem. N, p.62: copepodites); Vives & Shmeleva, 2007 (p.895, figs.F,M, Rem.); Blanco-Bercial & al., 2011 (p.103, Table 1, mol. Biol., phylogeny); Laakmann & al., 2013 (p.862, figs.1, 2, 3, 5, Table 1, 2, 3, mol. Biol.)
Species Calanus helgolandicus - Plate 1 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. IIII].
Female & Male.

Nota: Forehead like a ribbed vault in lateral view.

Species Calanus helgolandicus - Plate 2 of morphological figuresissued from : S.M. Marshall & A.P. Orr in The Biology of a Marine Copepod, Springer-Verlag (Ed.), 1972. [Fig.2].
Above drawings: Calanus finmarchicus Female (from North Sea): Coxa of the left P5; Proximal teeth of the coxa of the right P5; Forehead (lateral view).

Middle drawings: Calanus finmarchicus Female (from Tromsö): Coxa of the left P5; Proximal teeth of the coxa of the right P5; Forehead (lateral view).

Below drawings: Calanus helgolandicus Female (from North Sea): Coxa of the left P5; Proximal teeth of the coxa of the right P5; Forehead (lateral view), front slightly pointed.

Species Calanus helgolandicus - Plate 3 of morphological figuresissued from K. Hulsemann in Invert. Taxon., 1994, 8. [p.1477, Fig.28, H].
Female: H, 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 = 400.

Species Calanus helgolandicus - Plate 4 of morphological figuresissued from : R. Williams in Bull. mar. Ecol., 1972, 8. [p.57, Fig.3].
Female (from N Atlantic): Lateral view (i) and ventral view (ii) of three urosomes showing the variation in shape of the spermathecae and their lobed appearance.

Species Calanus helgolandicus - Plate 5 of morphological figuresissued from : R. Williams in Bull. mar. Ecol., 1972, 8. [Plate XVII].
Female (from N Atlantic): lateral view of the urosome of the three species C. helgolandicus, C.finmarchicus and C. glacialis showing the differences in shape of their spemathecae. The edge of the operculum is easily seen in C. helgolandicus and C. finmarchicus.

Species Calanus helgolandicus - Plate 6 of morphological figuresissued from : R. Williams in Bull. mar. Ecol., 1972, 8. [Plate XVIII, XIX].
Female (from N Atlantic):
Above: Ventral view of the urosomes of the three species showing the obvious differences in shape of the spermathecae. The genital pore is in a more posterior position in C. glacialis than in the other two species.
Below: A dorsal view of the spermathecae still attached to the basal plate. The spermatophore sac secretion which precedes the extrusion of the spermatozoa, is clearly seen in the spermathecae of C. finmarchicus. The lobed appearance of the spermathecal sacs of C. helgolandicus is also shown.

Species Calanus helgolandicus - Plate 7 of morphological figuresissued from : N.V. Vyshkvartzeva in Issled. Fauny Moreï, 1972, 12 (20). [p.165, Fig.4, a, g].
Femele Md (masticatory edge, lateral): a, from North Sea; g, from Mediterranean Sea.

Species Calanus helgolandicus - Plate 8 of morphological figuresissued from : R.-M. Barthélémy in These Doct. Univ. Provence (Aix-Marseille I), 1999. [Fig.19, A]. Female (Gulf of Marseille: France): A, external ventral view genital double-somite.
go = genital operculum.
Scale bar: 0.100 mm.

Species Calanus helgolandicus - Plate 9 of morphological figuresissued from : C. Razouls in Th. Doc. Etat Fac. Sc. Paris VI, 1972, Annexe. [Fig.20].
Female (from Banyuls, G. of Lion): A, habitus (lateral).

Male: B, habitus (lateral).

Species Calanus helgolandicus - Plate 10 of morphological figuresissued from : C. Razouls in Th. Doc. Etat Fac. Sc. Paris VI, 1972, Annexe. [Fig.21].
Female: Forehead comparison between C. helgolandicus A, from Banyuls (Gulf of Lion) and C. finmarchicus B, from isle of Cumbrae, loaned by S.M. Marshall (Marine Station Millport, Scotland).
Nota; Forehead ogival in lateral view for C. helgolandicus (A) and perfectly rounded for C. finmarchicus (B).

Species Calanus helgolandicus - Plate 11 of morphological figuresissued from : S.M. Marshall & A.P. Orr in The Biology of a Marine Copepod, Springer-Verlag (Ed.), 1972. [p. 14, Fig.4].
Comparison of P5 male from C. finmarchicus (d) and C. helgolandicus (e).
Nota: Observe the lengths of endopodites and the inner edges of the coxa curvatures.

Species Calanus helgolandicus - Plate 12 of morphological figuresissued from : S.B. Schnack in Crustacean Issue, 1989,6. [p.143, 2].
2, Calanus helgolandicus (from off NW Africa, upwelling region): cutting edge of Md.

Species Calanus helgolandicus - 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, 19 , Atlas von 54 Tafeln. [Taf.7, Fig.8]. As Calanus finmarchicus.
Female: 8, masticatory edge of Md (anterior view).

Species Calanus helgolandicus - Plate 14 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, Figs.20, 21]. As Calanus finmarchicus.
Female: 20, P5 (anterior view); 21, inner margin of basipodite 1 (= coxa) of P5.

Species Calanus helgolandicus - Plate 15 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.13]. As Calanus finmarchicus.
Female: 13, Mx1 §anterior view).

Species Calanus helgolandicus - Plate 16 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.32]. As Calanus finmarchicus.
Female: 32, thoracic segment 5 (Th5) and genital segment (Ab 1-2 = genital double-somite).

Species Calanus helgolandicus - 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, 19 , Atlas von 54 Tafeln. [Taf. 8, Fig.15]. As Calanus finmarchicus.
Female: 15, exopodite 3 of P3 (anterior view).

Species Calanus helgolandicus - 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, 19 , Atlas von 54 Tafeln. [Taf. 8, Fig.3]. As Calanus finmarchicus.
Female: 3, A1, segments 1 to 17 (ventral view).

Species Calanus helgolandicus - Plate 19 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.3]. As Calanus finmarchicus.
Female: 3, A1, segments 17 to 25 (ventral view).

Species Calanus helgolandicus - Plate 20 of morphological figuresissued from : R. Gaudy in Rev. Trav. St. Mar. End., Bull. 27 (42). [p.125, Tableau I].
Identification of copepodids stages Calanus helgolandicus.
A: Stage; B: number of swimming legs; C: number of thoracic segments; D: number of abdominal segments; E, Size.

Species Calanus helgolandicus - Plate 21 of morphological figuresissued from : G. Trégouboff & M. Rose in Manuel de planctonologie méditerranéenne, 1957, CNRS, Paris. [Pl. 125].
Calanus helgolandicus female scematic ventral view (from Medirrranean Sea): Outer parasit Ellobiopsis chattoni fixed on copepod's appendages.

Species Calanus helgolandicus - Plate 22 of morphological figuresissued from : K.A. Brodsky in Zool. Zh., 1959, 38, 10. [p.1541, Fig.3].
Comparison of coxopodite inner edge of P5 female for Calanus glacialis (1), Calanus finmarchicus (2) and Calanus helgolandicus (3).

Calanus glacialis : Dentate plate on coxopodite has very short, blunt teeth and is sligh curved in central position. Teeth are close together, without spaces, numbering 30-34.
Calanus finmarchicus : Dentate plate on coxopodite has short, blunt teeth, with small spacings. Teeth-line not curved. Number of teeth 29-30.
Calanus helgolandicus : Dentate plate on coxopodite very characteristic; teeth have more or less parallel edges, are relatively small, strongly marked curve in middle of line; distal part of plate has closely set, elongated teeth; spaces between teeth only in central part of line, teeth here are rounded, not flat. Number of teeth 28 (according to Jaschnov most specimens from the North Sea had 28-33 teeth).

Species Calanus helgolandicus - Plate 23 of morphological figuresissued from : K.A. Brodsky in Zool. Zh., 1959, 38, 10. [p.1542, Fig.4].
Comparison of left leg of P5 for Calanus glacialis (1), Calanus finmarchicus (2) and Calanus helgolandicus (3).

Calanus glacialis : In segments of exopodite of left leg, the relation of width of 1st and 2nd segments to length of corresponding segments is 1 : 3. Left endopodite reaches almost half the length of the 2nd segment of the exopodite of the same leg.
Calanus finmarchicus : Relation of width to length of 1st and 2nd segments of exopodite of left leg is 1 : 2.5. Left endopodite extends beyond middle of 2nd segment of exopodite.
Calanus helgolandicus : Relation od width to length of 1st and 2nd segments of exopodite of left leg is 1 : 2.5. left endopodite reaches distal limit of first third of 2nd segment of exopodite of same leg.

Species Calanus helgolandicus - Plate 24 of morphological figuresIssued from : A. Fleminger & K. Hulsemann in Mar. Biol., 1977, 40. [p.243, Fig.6 a].
Pore signature patterns of female urosome (ventral view shown below, dorsal view of genital segment and segment 2 (above).
Specimens (n = 50) from North Atlantic localities.
Filled circles: integumental pore present in all specimens examined; open circles: pore present in from 90 to 99% of specimens examined. Symbols used to indicate pores are not proportionate to actual pore size (latter range from 1 to 3 µm in diameter).

Nota: The distribution of integumental organs on the urosome of adult females was determined in specimens selected at random from samples representing a variety of localities within the North Atlantic distribution.
Calanus helgolandicus is most distinctive in lacking the 2 pairs of pores found in C. finmarchicus and C. glacialis on the ventral side of urosomal segments 2 and 3 and in having only 3 pores on the dorsal side of the genital segment.

Species Calanus helgolandicus - Plate 25 of morphological figuresIssued from : A. Fleminger & K. Hulsemann in Mar. Biol., 1977, 40. [p.244, Fig.7].
Calanus helgolandicus Female: Integumental gland on ventral side of urosomal segments sites occupied in samples from (a) Western North Atlantic shelf and slope water (N = 19, MARMAP Cruise); (b) Mid-North Atlantic (N = 20); (c) Eastern North Atlantic off Europe (N = 20); (d) Eastern North Atlantic off Africa (N = 20).
Filled circles: pore present in all specimens of sample; open circles: pore present in 90 to 99% of specimens; triangles: pore present in 20 to 49% of specimens; dots: pore present in 1 to 19% of specimens.
Actual number of specimens with pore at indicated site is shown in arabic numerals when less than 100%.
Symbols used to indicate pores are not proportionate to actual pore size (latter range from 1 to 3 µm in diameter).

Nota: If the geographical variation we observed in the urosome pore signature of females proves to be real, it would demonstrate the lack of panmixis and the presence of two or more semi-independent populations in the North Atlantic Ocean.
It is not unlikely that this species was temporarily eliminated from the North Atlantic during Pleistocene glaciations. The population in the Mediterranean Sea probably served as the promary reservoir for C. helgolandicus replenishing the North Atlantic when conditions were favorable, while otherwise regularly contributing to the specie's North Atlantic gene pool.At present, the Mediterranean sub-surface outfall undoubtedly carries C. helgolandicus into the North Atlantic just as Atlantic surface waters provide a conduit for gene flow in the opposite direction.

Species Calanus helgolandicus - Plate 26 of morphological figuresIssued from : C. Lauritano, Y. Carotenuto, G. Procaccini, J.T. Turner & A. Ianora in Harmful Algae, 2013, 28. [p.27, Fig.3]
Egg images. Confocal microscope images of eggs spawned by females fed on the dinoflagellate Prorocentrum minimum (A) for one day or Karenia brevis for one (B) or three days (C).

Nota: After one day of feeding on Karenia brevis, eggs collected soon after spawning were morphologically similar to the control (A), even if the cytoplasm and nuclei were not clearly defined (B). Conversely, after 3 days of feeding, eggs exhibited altered membrane cell morphology and apoptotic features such as granulation and degeneration of the cytoplasm matrix (C), whereas control eggs were identical to the first day. Ingestion of Kerenia brevis causes adverse effects on copepods, included alterations in copepod swimming and photobehavior, reduction in egg production and egg viability, and in some cases, reduced survival. In the study, although none of the females died, expression patterns of selevted genes involved in stress responses, detoxification mechanisms and apoptosis regulation were significantly affected.

Species Calanus helgolandicus - Plate 27 of morphological figuresIssued from : P.M.J. Woodhead & J.D. Riley in J. Cons. int. Expl. Mer, 1957, 23. [p.48, Fig.1].
Side view of the urosome in potential male (a) and female (b) stage-V copepodites of Calanus helgolandicus from the western part of the English Channel in June 1950.
Nota: the two types are distinguished by the degree of curvature of the ventral surface of the first (proximal) urosome segment (indicated by arrows).

Species Calanus helgolandicus - Plate 28 of morphological figuresIssued from : P.M.J. Woodhead & J.D. Riley in J. Cons. int. Expl. Mer, 1957, 23. [p.50, Table 1].
The two size-groups in stage-V copepodites of Calanus helgolandicus separating the males from from females.

Compl. Ref.:
Pearson, 1906 (p.4, Rem.); Chatton, 1920 (p.16: Rem.; p.17); Rose, 1925 (p.151); Wilson, 1932 (p.21); 1942 a (part., p.172); Sewell, 1948 (p.347); C.B. Wilson, 1950 (part., p.178, non Pacif. stations); Duran, 1955 (p.52); Marshall & Orr, 1956 (p.587, feeding, young stages); Woodhead & Riley, 1957 (p.47; Jaschnov, 1958 (p.838, fig.2); Woodhead & Riley, 1959 (p.465, stage 5 M & F, sex ratio); Corner, 1961 (p.5, Table I: feeding, assimilation, filtering rate, oxygen consumption); Carlisle & Pitman, 1961 (p.827, neurosecretion -diapause); V.N. Greze, 1963 a (tabl.2); Zeiss, 1963 (p.110, respiration rates); Giron-Reguer, 1963 (p.24); Cowey & Corner, 1963 (p.495, amino acid composition, respiration); Rice, 1964 (p.163, hydrostatic pressure effects); Mazza, 1964 (p.293, weight); Grice & Hulsemann, 1965 (p.223, 225: Rem.); Linford, 1965 (p.16, Rem.: p.25, lipid content); Bodo & al., 1965 (p.219, annual cycle); Vucetic, 1965 (p.419, annual variations); 1965a (p.425, sex ratio, life history); 1965 b (p.431, annual variation length-weight); 1966 (p.1-91, life history); Mazza, 1966 (p.69); Ehrhardt, 1967 (p.737, geographic distribution), Rem.); Manwell & al., 1967 (p.145, electrophoresis-enzymes); Corner & Newell, 1967 (p.113, nitrogen excretion); Glover, 1967 (p.189, fig.5, annual abundance); Corner & Cowey, 1968 (p.393, Table 5, 8, aminoacid composition, assimilation); Robertson, 1968 (p.185, Text-Fig.5, abundance); Macdonald & al., 1972 (p.213, fig.4, hydrostatic pressure effect); Evans, 1968 (p.11); Vinogradov, 1968 (1970) (p.94, 268, 269); Keegan, 1969 (p.137, abundance); Mullin, 1969 (p.308, Table I: estimates of production); Champalbert, 1969 a (p.624); Richman & Rogers, 1969 (p.701, feeding v.s. growing diatom); Singarajah, 1969 (p.171, Table I, II, behaviour); Butler & al., 1969 (p.977, nutrition/metabolism); 1970 (p.525, Rem.: p.529); Kovalev, 1969 a (p.146); 1970 a (p.87, Tableau 1, 2, comparison hyponeustonic & planctonic forms); Jaschnov, 1970 (p.204, chart); Paffenhöfer, 1970 (p.346, cultivation); Paffenhöfer & Sreickland, 1970 (p.97, feeding); Shih & al., 1971 (p.36); Lee R.F. & al., 1971 (p.99, lipids); Lincoln, 1971 (p.677, fig.2, 3, hydrostatic pressure & illumination effects); Bainbridge & Forsyth, 1972 (p.21, Table I: indictor species); Roe, 1972 (p.277, tabl.1, 2); 1972 a (p.316); Corkett, 1972 (p.171, eggs: development rate); Corner & al., 1972 (p.847, feeding, excretion); Razouls S., 1972 (p.1, respiration); Nival & al., 1972 (p.63, respiration); Gaudy, 1972 (p.175, 218, figs.25-27, annual cycle, Rem.: p.223-4); Eriksson, 1973 (p.37, fig.6, 7, 9, annual cycle); 1973 b (p.113, 118); Guglielmo, 1973 (p.399); Champalbert & al., 1973 (p.529, CHN composition); P. Nival & S. Nival, 1973 (p.135, mouth parts, grazing); Nival & al., 1973 (p.123, respiration); Nival & al., 1974 (p.231, respiration & excretion); S. Razouls, 1974 (147, oxygen rate); Corner & al., 1974 (p.319, feeding, nauplii diet); Corral Estrada & Pereiro Muñoz, 1974 (tab.I); Vives & al., 1975 (p.35, tab.II, III, IV); Fernandez, 1975 (p.1, fig.5, 6, 7, 8, 9, 10, 22, metabolism/lux); Corner & al., 1976 (p.345, animal diet, metabolism); Corner & al., 1976 (p.121, hydrocarbon effects); Paffenhöfer, 1976 (p.49, feeding behaviour); Deevey & Brooks, 1977 (p.256, Table 2, Station "S"); Sargent & al., 1977 (p.525, grazing); Greve, 1977 (p.83, feeding); Harris & al., 1977 (p.187, polluant effect); Colebrook, 1978 (tab.1); Fernandez, 1978 (p.97, metabolism/food, Rem.: Table 19); Comaschi Scaramuzza, 1978 (p.16); O'Hara & al., 1979 (p.331, sex hormone); Spooner & Corkett, 1979 (p.197, feeding rates vs oil effects); Andronov & Maigret, 1980 (p.65, Table 1, 2, 3); Hallberg & Hirche, 1980 (p.283, gut ultrastructure, enzymes); Hirche, 1981 (p.174, enzymes); Collins & Williams, 1981 (p.273, monthly distribution-salinity); Brenning, 1981 (p.1, spatial distribution, T-S diagram, Rem.); Kovalev & Shmeleva, 1982 (p.82); Vives, 1982 (p.290); Williams & Robins, 1982 (p.271, preservation modes vs. length, weight, nitrogen & carbon); Borgmann, 1982 (p.668, size/production); Castel & Courties, 1982 (p.417, Table II, spatial distribution); Hirche, 1983 (p.281); Rivière, 1983 (p.19, enzymes); Tremblay & Anderson, 1984 (p.4); Bottrell & Robins, 1984 (p.259, size/weight/Carbon & Nitrogen); Scotto di Carlo & al., 1984 (p.1042); Alvarez-Marques, 1984 (p.110, annual abundance, figs.F,M); Petipa & Ostravskaya, 1984 (p.631, energy expenditure), Baars & Fransz, 1984 (p.120, Table 1, grazing); Baars & Oosterhuis, 1984 (p.97, diurnal feeding rhythms); Mayzaud O. & al., 1984 (p.15, feeding/enzyme); Prahl & al., 1984 (p.317, assimilation); Hirche, 1984 (p.233, seasonal distribution); Boucher, 1984 (p.469, spatial distribution/hydrological front); Regner, 1985 (p.11, Rem.: p.20); Brenning, 1985 a (p.3, Table 2, fig.9); Jansa, 1985 (p.108, Tabl.I, , II, III, IV, V); Williams & Collins, 1985 (p.28); Nott & al., 1985 (p.271, gut histology, fecal pellet); Neal & al., 1986 (p.1, feeding); Diel & Klein Breteler, 1986 (p.85, growth, development rate); Robinson & Hunt, 1986 (p.791, Table 1, 2, fig.2); Harris & Malej, 1986 (p.75, vertical migration v.s. ammonium excretion); Robinson & al., 1986 (p.201, seasonal & spatial distributions); Gill & Poulet, 1986 (p.193, appendage activity); Brylinski, 1986 (p.457, spatial variations); Quisthoudt & al., 1987 (p.995, spatial distribution); Comaschi Scaramuzza, 1987 (tab.1); Ibanez & Boucher, 1987 (p.205, Tableau, fig.5, 7, hydrological fronts); Boucher & al., 1987 (p.133, spatial distribution/physical structure); Williams R. & al., 1987 (p.247, vertical distribution); Gill & Harris, 1987 (p.785, feeding behaviour); Grigg & al., 1987 (p.253, biometry, development); Poulet & Gill, 1988 (p.259, feeding behaviour); McLaren & al., 1988 (p.275, DNA content, development rate: egg-nauplius); Gorsky & al., 1988 (p.133, Table I, C-N composition); Lozano Soldevilla & al., 1988 (p.57); Roy & al., 1989 (p.145, feeding); Giguère & al., 1989 (p.522, Table 1, formaldehyde preservation); Kattner & Krause, 1989 (p.261, Table 3, 4, 5, lipids); Poulet & al., 1989 (p.1325, Table 2, 3: vitamin content); Kouwenberg & Razouls, 1990 (p.23, climatic change); Krsinic, 1990 (p.337, Table I-II, vertical distribution); Fransz & al., 1991 (p.5 & suiv.); Hay & al., 1991 (p.1453, Table 2, 5); Scotto di Carlo & al., 1991 (p.271); Green & al., 1992 (p.1631, Nauplii-fecal pellets); Bautista & Harris, 1992 (p.41, ingestion rate, gut contents); Seguin & al., 1993 (p.23); Fromentin & al., 1993 (p.285, Ligurian transect abundance); Morales C.E. & al., 1993 (p.185); Kouwenberg, 1993 (p.215, fig.3, seasonal abundance); 1993 a (p.281, fig.3, 4, sex ratio); Guerin-Ancey & David, 1993 (p.119, table 1, biovolume, vertical distribution); Hays & al., 1994 (tab.1); Harris, 1994 (p.431, ingestion & egestion); Poulet & al., 1994 (p.79, eggs); 1995 (p.85, eggs, Nauplii); Krause & al., 1995 (p.81, Rem.: p.128); Laabir & al., 1995 (p.1125, egg production vs. algal concentrations, fecal pellets); Southward & al., 1995 (p.127, fig.2, 3, 8, 13, distribution-abundance vs time); Pond & al., 1995 (p.75,gut-pH); Fromentin & Planque, 1996 (p.111, distribution vs NAO); Stöhr & al., 1996 (p.457, abundance); Zauke & al., 1996 (p.141, Table 7, metal bioaccumulation); Siokou-Frangou, 1997 (tab.1); Gilabert & Moreno, 1998 (tab.1, 2); Reid & Hunt, 1998 (p.310, figs.2, 3, Rem.); Kouwenberg, 1998 (p.203, seasonal variation, fig.3: global change); Beare & McKenzie, 1999 (p.241, fig.6, 7); Seridji & Hafferssas, 2000 (tab.1); Kang H-K & Poulet, 2000 (p.241, diet vs reproduction); Kang H-K & al., 2000 (p.2171, diets vs reproduction); Ku Kang & al., 2000 (p.2171, reproduction vs diets); Moraitou-Apostolopoulou & al., 2000 (tab.I); d'Elbée, 2001 (tabl. 1); Lapernat & Razouls, 2001 (tab.1); Holmes, 2001 (p.36, Rem.); Fransz & Gonzalez, 2001 (p.255, tab.1); Weikert & al., 2001 (p.227, tab.4, fig.6, 8, 9, Rem.); Bressan & Moro, 2002 (tab.2); Zerouali & Melhaoui, 2002 (p.91, Tableau I); Beaugrand & al., 2002 (p.1692); Beaugrand & al., 2002 (p.179, figs.5, 6); Lindley & Reid, 2002 (p.153, abundance veriation); Reid & al., 2003 (p.260, figs.3,5); Titelman & Kiørboe, 2003 a (p.137, nauplius behaviour); Gaudy & al., 2003 (p.357, tab.1); Kane, 2003 (p.151); Morgado & al., 2003 (p.S205, abundance vs ebbs and floods in estuary); Vukanic, 2003 (p.139, tab.1); Bode & al., 2003 (p.85, Table 1, abundance); Vieira & al., 2003 (p.S163, Table 2, abundance); Wexels Riser & al., 2003 (p.301, grazing vs faecal pellets, toxins algae); Lindeque & al., 2004 (p.121, fig.2); Fernandez de Puelles & al., 2004 (p.654, fig.7); Vinogradov & Musaeva, 2004 (p.503, Rem: p.512); CPR, 2004 (p.50, fig.139); Lopez-Urrutia & al., 2004 (p.303, clearance rate, predation); Vallet & Dauvin, 2004 (p.539, tab.2); Veistheim & al., 2005 (p.382, tab.1, 2, fig.1); Souissi & al., 2005 (p.161); Siokou-Frangou & al., 2005 (p.160); Molinero & al., 2005 (p.156); Yebra & al., 2005 (p.1367, growth: C5-C6); Jonasdottir & al., 2005 (p.1239, egg production & hatching success); Bonnet & al., 2005 (pp.1-53, tab.2. overview in European waters); Queiroga & al., 2005 (p.195, table 1); Hays & al., 2005 (p.337, fig.1, climate change); Rawlinson & al., 2005 (p.205, tidal exchange); lindeque & al., 2006 (p.221); Albaina & Irigoien, 2006 (p.413, effect of the parasite Ellobiopsis); Isari & al., 2006 (p.241, tab.II); Marques & al., 2006 (p.297, tab.III); Zervoudaki & al., 2006 (p.149, Table I); Jansen & al., 2006 (p.102, nutrition); Varela & al., 2006 (p.272, Table 3, oil spill effects); Ceballos & Alvarez-Marqués, 2006 (p.1, dynamic of population); 2006a (p.189, reproduction); Koppelmann & Weikert, 2007 (p.266: tab.3); Kouwenberg & Lantoine, 2007 (p.239: UV-B effects); Cook & al., 2007 (p.757, Naupliar development times); Helaouët & Beaugrand, 2007 (p.147, climate change effects); Fernandez de Puelles & al., 2007 (p.338, 348); Albaina & Irigoien, 2007 (p.433, Tab.1); Valdés & al., 2007 (p.103: tab.1); Poulet & al., 2007 (p.415, reproduction); Busatto, 2007 (p.26, Tab.3); Ferrari & Dahms, 2007 (p.63, Rem.); Fileman & al., 2007 (p. 70, grazing); Khelifi-Touhami & al., 2007 (p.327, Table 1); Vincent & al., 2007 (p.295, Table 1, 3, 6, 7, fig.2, nitrogen transfert); Hirst & al., 2007 (p.189, seasonal dynamics, mortality rates); Cabal & al., 2008 (289, Table 1); Gaard & al., 2008 (p.59, Table 1, N Mid-Atlantic Ridge); Wichard & al., 2008 (p.30, reproduction); Molinero & al., 2008 (p.271, fig.4); Brylinski, 2009 (p.253, Tab.1); Labat & al., 2009 (p.1746, Table 2); Licandro & Icardi, 2009 (p.17, Table 4); Park & Ferrari, 2009 (p.143, fig.2, biogeography); Mackas & Beaugrand, 2010 (p.296, figs.11, 12); Bonnet & al., 2010 (p.725, Rem.: predation by Sagitta setosa); Fileman & al., 2010 (p.709, Rem.: feeding); Eloire & al., 2010 (p.657, Table II, temporal variability); Mazzocchi & Di Capua, 2010 (p.424); Mazzocchi & al., 2011 (p.1163, fig.6, long-term time-series 1984-2006); Fileman & al., 2011 (p.403, abundance vs bloom condition); Nowaczyk & al., 2011 (p.2159, Table 2); S.C. Marques & al., 2011 (p.59, Table 1); McGinty & al., 2011 (p.120, seasonal abundance vs N Atlantic zones); Jonasdottir & Koski, 2011 (p.85, vertical distribution, production); Bicknell & al., 2011 (p.1794, Table 1, 2, effects of formalin preservation); Isari & al., 2011 (p.51, Table 2, abundance vs distribution); Zervoudaki & al., 2011 (p.45, fig. 3 as Calanus spp, Table 3, abundance, egg production: Turkish Straits); Castellani & al., 2011 (p.104: Rem.); Van Ginderdeuren & al., 2012 (p.3, Table 1, Rem.: p.10); Lauritano & al., 2011a (p. 1, feeding vs toxicity, molecular expression); Lauritano & al., 2012 (p.22, Table 1, 2, gene/polluants); Zizah & al., 2012 (p.79, Tableau I, Rem.: p.86, 89); Miloslavic & al., 2012 (p.165, Table 2, transect distribution); Aubry & al., 2012 (p.125, table 3, fig. 6, 8 a, b, interannual variation); Salah S. & al., 2012 (p.155, Tableau 1, Rem.); Carotenuto & al., 2012 (p.46, population dynamics, egg production vs diets, cultivation); Lauritano & al., 2012 (p.1, genetic expression vs stress); Mayor & al., 2012 (p.258, fig.2, 3, hatching vs temperature & CO2 effects); McGinty & al., 2012 (p.122, time series abundance); Turner & al., 2012 (p.63, toxic dinoflagellate effects); Mazzocchi & al., 2012 (p.135, annual abundance 1984-2006); Bode & al., 2012 (p.108, spatial distribution vs time-series, % biomass); Alvarez-Fernandez & al., 2012 (p.21, Rem.: Table 1); Møller & al., 2012 (p.211, development time vs temperature & food); Gusmao & al., 2013 (p.279, Table 4, sex ratio); Belmonte & al., 2013 (p.222, Table 2, abundance vs stations); Helaouët & al., 2013 (p.1, long-term changes abundance; 1958-2008); Lauritano & al., 2013 (p.23, dinoflagellate toxic effect vs. genes); Sobrinho-Gonçalves & al., 2013 (p.713, Table 2, fig.8, seasonal abundance vs environmental conditions); Breckels & al., 2013 (p.2486, swimming behavior vs algal metabolites); Cole & al., 2013 (p.6646, microplastic ingestion); Peijnenburg & Goetze, 2013 (p.2765, genetic data); Kürten & al., 2013 (p.167, Table 1, C:N, fatty acid); Maar & al., 2013 (p.24, spatial distribution, long-term changes); Lidvanov & al., 2013 (p.290, Table 2, % composition); Fernandez de Puelles & al., 2014 (p.82, Table 3, seasonal abundance); Mazzocchi & al., 2014 (p.64, Table 3, 4, 5, spatial & seasonal composition %)
NZ: 6

Distribution map of Calanus helgolandicus by geographical zones
Species Calanus helgolandicus - Distribution map 3issued from : R. Williams in Bull. mar. Ecol., 1972, 8. [p.55, Fig.1].
Ditribution of stages V and VI in the North Atlantic from the Continuous Plankton Recorder.
The chart show the average abundance and distrubution derived from more than 43.000 samples taken a depth of 10 m during 1958 to 1968. The samples were assigned to rectangles of 1° lat. by 2° long. The boundary of the sampled area (defined as those rectangles sampled in more than 5 months) is shown by the straight lines in the chart; within this area the average abundance in each rectangle is shown by circular symbols; the presence of the species in the occasional samples outside this area is indicated by plus signs. The absence (in the sampled area) indicates that the species was not found in CPR.
large and small filled in circles and open circles, respectively, are used to indicate the following categories of abundance (average numbers per sample of 3.3 m3: >0.60 : 0.60-0.10 : <0.10
Species Calanus helgolandicus - Distribution map 4issued from : R. Williams in Mar. Biol., 1985, 86. [p.147, Fig.3].
Calanus helgolandicus. Annual mean distribution and abundance from sampling at 10 m depth by the Continuous Plankton Recorder. Data for all months sampled from 1960 to 1981. The boudary of the sampled area is shown by straight lines. Position of the 14°C isotherm in August is shown.
Species Calanus helgolandicus - Distribution map 5issued from : R. Williams in Mar. Biol., 1985, 86. [p.148, Fig.6].
Calanus finmarchicus (a) and C. helgolandicus (b) from Celtic Sea (53°30'N, 07°00'W). Vertical distribution of Copepodite V and VI. Numbers in each night (black) and day (white) hauls are plotted in 5 m depth intervals as percentagges of total numbers (n) present in the haul.
Temperature and Salinity profiles are shown for the day hauls. GMT: Greenwhich Mean Time.

Nota: The two congeneric copepods are morphologically very similar and show little phenotypic divergence. No hybrids are produced between the species which demonstrates that their isolating mechanisms are fully evolved. The only reason that C. finmarchicus, a northern species, is able to persist in the Celtic Sea, which has a sea-surface temperature of ca. 18° C, is because the sea becomes seasonally thermally stratified and a winter sea-temperature is retained below the thermocline.
Species Calanus helgolandicus - Distribution map 6issued from : U. Brenning in Wiss. Z. Wilhelm-Pieck-Univ. Rostock - 30. Jahrgang 1981. Mat.-nat. wiss. Reihe, 4/5. [p.3, Figs.4, 5].
Spatial distribution and T-S Diagram for Calanus helgolandicus from 8° S - 26° N; 16°-20° W, for different expeditions (V1: Dec. 1972-Jan. 1973; IV: Jun.-Jul. 1972; V: Feb.-Mar. 1973; VI: May 1974).
SO: Southern Surface Water (S °/oo: 34,50; T°C: 29,0); ND: Northern Water of the Surface Layer (S °/oo: 37,5; T°C: 21,0); SD: Southern Deep Water of the surface layer (S °/oo: 35,33; T°C: 13,4).
Water types basing on the works of Sverdrup & al. (1942), Wolf (1978) and Wolf & Kaiser (1978) distinguishing various quasi-permanent water types in the top 200 m of the water column in the NW Africa upwelling region. The water type SD, which usually forms the upper limit of the SACW (South Atlantic Central water: S °/oo: 34,35-35,33 & T°C: 5,15-13,4), can enter the surface layer as a result of upwelling, so that water types ND, SD and SO can be expected in this layer in upwelling regions. The boundary separating the NACW (North Atlantic Central Water: S °/oo: 35,10-37,35; T°C: 8,0-21,0) and SACW water masses is situated in the region between Bahia de Gorrei and Cap Blanc (Mauritania) throughout the whole year. The distribution of the water types SACW in the surface layer off North West Africa varies with the season and the meridional migration of the wind field. See in Brenning (1985 a, p.6).
Species Calanus helgolandicus - Distribution map 7issued from : W.A. Jaschnov in Int. Revue ges. Hydrobiol., 1970, 55 (2). [p.205, Fig.3]
Distribution of Calanus helgolandicus from literary sources and unpublished data.
Solid lines indicate the boundaries of Mediterranean water of definite salinity 36.0- 35.5 and 35.0 p.1000 (Defant, 1955. Arrows denote some of the currents. Dark circles indicate occurrence in reproduction areas, semi-dark circles in immigrated areas and white circles in expatriation areas (usual single findings).
Species Calanus helgolandicus - Distribution map 8issued from : R. Gaudy in Tethys, 1972, 4 (1). [p.237, Fig.36].
Schematic quantitative abundance of Calanus helgolandicus in the Gulf of Marseille (Mediterranean Sea) established from samples during the period between February 1961 to June 1967.

Gaudy (p.220) points to 3 generations per year in the neritic province from the winter to the spring, plus probably 2 generations off shore and deeper water (funding the mean development time determined graphically from nauplii to adults for the first three generations between 30 to 75 days); considering a mean development time of 45 days, interval separating two generations should be estimated 75 days; in consequence of the period August-December permitted a development of two genarations.
Species Calanus helgolandicus - Distribution map 9issued from : S. Eriksson in ZOON, 1973, 1. [p.46, Fig.9].
Size distribution of adult females of Calanus helgolandicus (offshore station H6:11°30'N, 57°40'.5, The Kattegatt) during 1969-70 in the main series.
Species Calanus helgolandicus - Distribution map 10issued from : S. Eriksson in Mar. Biol., 1974, 26. [p.320, Figs. 2-3]
Salinity and temperature curves for main series at offshore station H6 (11°30' N, 57°40'.5 E, The Kattegatt) from March 1968 to November 1970.
Species Calanus helgolandicus - Distribution map 11Issued from : E.D.S. Corner & S.C.M. O’Hara in The Biological chemistry of Marine Copepod, 1986. [p.305, Fig.6.10].
Diagrammatic representation of gross differences in the gut epithelium of (A) non feeding and (B) feeding Calanus helgolandicus. From Nott & al., 1985 ; copyright Springer-Verlag, 1985.

Nota : For the authors, gross cyrtological differences between cells of the digestive epithelium of feeding and non-feeding C. helgolandicus showed a ‘spent’ vacuolated cell disintegrating into the mid-gut lumen, its membranous contents clearly visible (B).
One interesting characteristic of the cells lining the gut, and one especially relevant to the presence of the animal’s own lipids in its faecal pellets, is the tendency of these epithelial cells to rupture during the digestive process and release cellular material into the lumen of the gut, where it would mix with undigested food. Such a mechanism may well account for lipids characteristic of the animal, sepecially fatty acids and cholesterol, becoming incorporated in faecal pellets, so providing the animal’s ‘signature’ on their lipid content. The re-absorption of cell contents of disrupted digestive epithelium by the R’ cells of the distal region (zone III) is likely to be an inefficient process.
In conclusion, the pelagic food webs constitute the link between the production and deposition of biolipids in the ocean. Copepods in particular have been shown to occupy a key position , as their faecal pellets contribute to sedimentary particulate material, especially in oceanic areas. Faecal pellets transport living material to the deep sea (see in Urrère & Knauer, 1981 ; Silver & Alldredge, 1981 ; Silver & Bruland, 1981 ; Fowler & Fisher, 1983). Such a mechanism has also been found to apply to the distribution of certain bacteria in the water column, these bacteria developing in the guts of animals and then being released in faecal material. Gowing & Silver (1983) regard these internal bacteria as facultative anaerobes, resistant to digestion and originating deep inside the pellets during their formation. Earlier, Silver & Alldredge (1981) observed both heterotrophic and autotrophic bacteria, including cyanobacteria, in faecal pellets and their fragments entrapped in fast-sinking marine ‘snow’.
Species Calanus helgolandicus - Distribution map 12Issued from : C.J. Corkett in J. exp. mar. Biol. Ecol., 1972, 10. [p.172, Table I].
Development time to hatching of eggs of Calanus helgolandicus.
Animals collceted off Plymouth (English Channel) on 24th May 1971.

Nota: The rate of development of eggs shows that the development time increases with decreasing temperature.
Corkett points to the larger eggs of C. hyperboreus (190 µ, after McLaren & al., 1969) and C. glacialis (178.6 ±2.5 µ, after McLaren, 1966) take longer to develop than the smaller eggs of C.finmarchicus (145 µ, after Marshall & Orr, 1953) and C. helgolandicus (163.9 ±0.866 µ at Plymouth).
A possible explanation for the linear relationship is that that development rate is determined by the surface : volume ratio which decreases with increasing diameter.
One could think that the metabolic exchange between the egg and its external environment is relatively slower for the larger eggs with their relatively smaller surface. Alternatively, one might assume that the development rate is determined by metabolic processes related to the size of the internal surfaces which in turn is proportional to the external surface.
Species Calanus helgolandicus - Distribution map 13issued from : N.R. Collins & R. Williams in Mar. Biol., 1981, 64. [p.282, Fig.11];
Calanus helgolandicus. Monthly distribution and abundance in Bristol Channel and Severn Estuary from November 1973 to February 1975, together with 33 p.1000 S isohaline.
Species Calanus helgolandicus - Distribution map 14issued from : N.R. Collins & R. Williams in Mar. Biol., 1981, 64. [p.283, Fig.12];
Calanus helgolandicus. Numerical abundance plotted against salinity for the four seasons in Bristol Channel and Severn Estuary from November 1973 to February 1975; 33 p.1000 S value is indicated.

Nota: This stenohaline species was found mostly in the southwestern area of the survey and was carried into the Channel by incursion of high salinity water from the Celtic Sea. Over most of the sampling period the approximate limit of penetration of the species into the Channel was represented by the position of the 33 p.1000 S isohaline, although in the spring it penetrated furyher up the Channel, close to the 30 p.1000 S isohaline.
Species Calanus helgolandicus - Distribution map 15Issued from : M. Laabir, S.A. Poulet, A. Ianora, A. Miralto & A. Cueff in Mar. Ecol. Prog. Ser., 1995, 129. [p.99, Fig.3].
Calanus helgolandicus (from Roscoff, France). Histograms of the relative proportions of diatoms, non-diatoms plus debris and bacteria estimated in the faecal pellets of wild females.
Each bar corresponds to the mean of at least 3 selected samples per month during 1993.
Species Calanus helgolandicus - Distribution map 16Issued from : G.A. Robinson, J. Aiken & H.G. Hunt in J. mar. biol. Ass. U.K., 1986, 66. [p.213, Fig.10].
Temperature and chlorophyll sections and plankton from a tow by Undulating Oceanographic Recorder (Aiken, 1981) on 23 August 1979 along the transect route Plymouth -Roscoff (west English Channel).
Scales for total copepods (x 10 power 4) on left and for C. helgolandicus on the right.
Species Calanus helgolandicus - Distribution map 17Issued from : A. Fleminger & K. Hulsemann in Mar. Biol., 1977, 40. [p.245, Table 4].
Length of prosome (mm) and number of teeth on coxopodite of P5 in adult females selected at random from samples used to determine urosome pore signature.
Species Calanus helgolandicus - Distribution map 18Issued from : A. Fleminger & K. Hulsemann in Mar. Biol., 1977, 40. [p.241, Fig.5 a].
Distribution of Calanus helgolandicus sensu stricto in North Atlantic and adjacent seas based on published litterature (squares: Edinburg Oceanographic Laboratory, 1973; open triangles: Jaschnov, 1970; present records: circles). Stippled areas approximate inhabited region beyond or at perimeter of North Atlantic Ocean.
Species Calanus helgolandicus - Distribution map 19Issued from : A. Lopez-Urrutia, R.P. Harris & T. Smith in Limnol. Oceanogr., 2004, 49 (1). [p.304, Fig.1].
Calanus helgolandicus feeding on Oikopleura dioica eggs, from off Plymouth (English Channel.
Lines represent the least square regression (slope = 0.490 ± 0.039; intercepts = - 1.16 ± 0.82 (µg Cby day) and 116 ± 82 (eggs by day); ± SE; r2 = 0.96; n = 8; P < 0.001 and 95% confidence limits for the parameter estimates (dashed lines).
Species Calanus helgolandicus - Distribution map 20Issued from : A. Lopez-Urrutia, R.P. Harris & T. Smith in Limnol. Oceanogr., 2004, 49 (1). [p.305, Fig.3].
Clearance rate of (A) Calanus helgolandicus from off Plymouth (English Channel), in relation to the size of Oikopleura dioica (Appendicularians).
The first x-axis data points represent the diameter of O. dioica eggs. Solid symbols represent the measured clearance rates and open symbols represent their mean. Vertical and horizontal error bars are the standard deviation of clearance rates and O. dioica size, respectively.
O. dioica cultures were maintened and experiments conducted in a constant temperature room at 15°C with a simulated 12:12 day: night cycle. Cohorts of cultured O. dioica , maintened according to Fenaux and Gorsky, 1985, were used to obtain eggs and juvenile appendicularians.
Species Calanus helgolandicus - Distribution map 21Issued from : M. Cole, P. Lindeque, E. Fileman, C. Halsband, R. Goodhead, J. Moger & T.S. Galloway in Environ. Sci. Technol., 2013, 47. [p.66,49, Fig.1, i].
Calanus helgolandicus containing 20.6 µm polystyrene beads (dorsal view), visualized using fluorescence microscopy;
Scale bar (gray line): 100 µm.
Species Calanus helgolandicus - Distribution map 22Issued from : E.F. Møller, M. Maar, S.H. Jonasdottir, T.G. Nielsen & K. Tönnesson in Limnol. Oceanogr., 2012, 57 (1). [p.214, Fig.1].
Experimental data and the fitted function on clearance rate (mL./ Copepode/day) of (a) Calanus helgolandicus (R2 = 0.57, p <0.001) and (b) Calanus finmarchicus (R2 = 0.45, p <0.001) as a function of temperature. Error bars are SD. (c) Clearance rate for both species normalized to fraction of its maximum.
For parameter values see Table 1.

Nota: The ingestion response to temperature was determined in the laboratory where the copepods were fed the diatom Thalassiosira weissflogii.
C. finmarchicus from Gullmar fjord (58°19.2'N, 11°32.8'E) on 06 April 2009; C. helgolandicus from off Plymouth (U.K.) in June 2009.
C. finmarchicus develops faster than C. helgolandicus below 11°C and slower above. The different relative development rates of the species are related to different temperature responses in ingestion rates.
A temperature increase of 1°C to 2°C may have consequences for the relative contribution of the two species to the copepod community, and both seasonal and spatial displacements of the Calanus populations can be expected under climate change.
Species Calanus helgolandicus - Distribution map 23Issued from : N. McGinty, A.M. Power & M.P. Johnson in J. Exp. Mar. Biol. Ecol., 2011, 400. [p.125, Fig. 2].
Locations of the ten clusters defined as ecosystem regions by K-means partitioning of satellite-derived Chl-a data. The locations of CPR samples throughout the study area are marked by black dots. Red contours indicate the underlying bathymetry at 200, 400, 800, 1600 and 3200 m intervals. Abbreviations and attributes that relate to each region are described in Table I.

Nota: Calanus finmarchicus is a sub-polar species whose presence in an area indicates a cold water influence while its congener, Calanus helgolandicus is seen as a more temperate oceanic species. These two species are used to test the inter-regional variation in the northern Atlantic. Only the adult stages (CV-CVI) are differentiated to species level.
Changes in phenology are implied when there is a significant month by time interaction. The centre-of-gravity date for seasonal zooplankton abundance was used as a measure of the timing of the seasonal peak, also known as the central tendency (T) is sensitive to monthly changes in abundance and can be viewed as a measure of the phenological change in the species annual cycle over time. T is estimated in decimal month units. See Beare & McKensie; 1999; Edwards & Richardson, 2004; Richardson & al., 2006.
Changes in sea surface temperature (SST) represent a strong environmental proxy for measuring climate related changes as this variable affects zooplankton at both the macro-ecological and the physiological level (SST data came from the HadISST1 dataset, provided by the Hadley Centre for Climate Prediction and Research Meterological Office, London, UK (Rayner & al., 2003).
Following the Chl-a based clustering, the long term SST pattern was compared among regions. The long term trend in SST was estimated from the slope of a linear regression of standardized SST against year. Variations in the link between SST and the dominant modes for the overall study area Atlantic Multidecadal Oscillation (AMO), the East Atlantic Pattern (EAP), and the North Atlantic Oscillation (NAO) index; (Cannaby & Husrevoglu, 2009) were examined using best fit regressions.
All the regions showed a warming trend over the time period considered:
- Warming was greatest in the Celtic Sea (CS) and Celtic mixed (CM) regions of shelf, at the Shelf Edge and in the more southern and central oceanic regions (WT, ABS and ABN; see Table 1).
- The influence of modes of variability affecting SST varied among regions. The AMO, itself a temperature signal, was present in all areas. The influence of climate indices varied, with the NAO a positive influence on SST in southern shelf and ocean regions and Irish Sea, but absent or with a negative association in central and northern off-shelf regions and Main Shelf (MS).
- The East Atlantic Pattern (EAP) was only selected as part of the best fit model in the Rockall Trough (RT) region.
The central tendency (T) for spatially averaged Chl-a in each region was calculated using values from March-October for each year. Off-Shelf there was a general northward progression in the timing of the seasonal peak.
- The earliest peak occurred in the Warm Temperate region (mean T of 5.76) with a gradual progression towards later seasonal peaks in the more northward regions. There was a difference of nearly a month between the most southern region, WT, and the two northern regions RT and RB, which had mean T of 6.61 and 6.64 months respectively.. On shelf there was less of a difference between the earliest and latest seasonal peak. The earliest T (SE) was 6.21 months while the latest (6.49 months) occurred in the Celtic Mixed (CM) region
Species Calanus helgolandicus - Distribution map 24Issued from : N. McGinty, A.M. Power & M.P. Johnson in J. Exp. Mar. Biol. Ecol., 2011, 400. [p.124, Table 1].
Ecosystem regions of the northen Atlantic studied by CPR. Data for all months between 1998 and 2008.

Nota: The measure of phenology, T, varied within and between regions over time.
For C. helgolandicus, the mean T was 6.4 months with an average range within of 1.2 months C. finmarchicus generally had the earliest central tendancy (5.7 months), and their average range was slightly larger (mean range 1.6 months).
In the majority of cases, increases in regional SST anomaly were negatively associated with the timing of the seasonal abundance peak (Fig.4, time series shown in Fig.5). This implies an earlier zooplankton ''bloom'' in warmer years. There were a few exceptions where this negative relationship with SST failed to hold. For C. helgolandicus, the region WT showed a relatively small correlation between ce,ntral tendaency and SST. For C. finmarchicus, the IS region had a weak positive correlation.
The annual average abundance of C. helgolandicus tended to be higher in warm years, with a positive correlation between abundance and SST in all areas except IS and SE. The strongest positive associations with temperature were in the northern regions of RT and MS. The pattern of mostly positive associations between annual copepod abundance and annual SST anomaly was reversed in the majority of cases in C. finmarchicus.
Species Calanus helgolandicus - Distribution map 25Issued from : N. McGinty, A.M. Power & M.P. Johnson in J. Exp. Mar. Biol. Ecol., 2011, 400. [p.126, Fig. 4].
Correlations betweej the timing of seasonal abundance peak (T) and standardized SST anomaly (first row), annual average abundance and standardized SST anomaly (second row) and annual average abundance and timing (last row) for each taxon. Gaps in the charts for the correlation between T and SST are where model fitting did not indicate an interaction between month and year.
Species Calanus helgolandicus - Distribution map 26Issued from : N. McGinty, A.M. Power & M.P. Johnson in J. Exp. Mar. Biol. Ecol., 2011, 400. [p.127, Fig. 5].
The central tendency (T) in decimal months, for Calanus helgolandicus and C. finmarchicus (black line) and local SST (red line) for each of the nine ecosystem regions.
Where zooplankton abundances varied between bimodal and unimodal peaks, only the unimodal pattern is shown. SST anomalies are inverted on the y axis. Abbreviations of each ecosystem regions are given in full in p.124, Table I.
Species Calanus helgolandicus - Distribution map 27Issued from : N. McGinty, A.M. Power & M.P. Johnson in J. Exp. Mar. Biol. Ecol., 2011, 400. [p.128, Fig. 6].
Mean accepted numbers (3 per m3) for Calanus helgolandicus in the nine ecosystem regions (three year running mean).
Note the scales differ.
Species Calanus helgolandicus - Distribution map 28Issued from : C. Wexels Riser, S. Jansen, U. Bathmann & P. wassmann in Mar. Ecol. Prog. Ser., 2003, 256. [p.303, Fig.2]
Scanningt electrom micrographs (SEM) of (a) Dinophysis norvegica, (b, c) Calanus helgolandicus faecal pellet containing D. norvegica; (d-f) light micrographs of C. helglandicus pellets containing D. norvegica.

Nota: Production rate of faecal pellets by individual and by day = 23.6 ± 1.6; faecal pellet volume produced (mm3 by ind. by day) = 0.026 ± 0.010; faecal pellet carbon produced (µg C by ind. by day) = 1.82 ± 0.75.
Faecal pellet length (µm) = 374 ±5.0; faecal pellet diameter (µm) = 65.9 ±0.4; faecal pellet volume (x 10000 µm3).
Species Calanus helgolandicus - Distribution map 29Issued from : H. Bi, K.A. Rose & M.C. Benfield in Mar. Ecol. Prog. Ser., 2011, 427. [p.157, Table 5].
Instantaneous mortality rates (individuals per day) from litterature.
Species Calanus helgolandicus - Distribution map 30Issued from S. Razouls in XXIII rd Congress of Athens, 3-11 November 1972. [p.2]. Oxygen consumed by individual (adult) in the Banyuls Bay.
(1) Hydrological season in the stability period: Eté = Summer: 18-20 °C; Hiver = Winter: 13-10°C.
Espèces = species; Saison = Season; Lg céph.= cephalothoracic length; an = individual.
Species Calanus helgolandicus - Distribution map 31Issued from : A.G. Hirst, D. Bonnet & R.P. Harris in Mar. Ecl. Prog. Ser., 2007, 340. [p.195, Fig.4].
Calanus helgolandicus egg production and surface water temperature and surface chlorophyll a concentration in the English Channel at Station L4 from March 2002 to March 2004
Congo, Morocco-Mauritania, Canary Is., off Madeira, Portugal, off W Cape Finisterre, Chesapeake Bay, New-York Bight, off Woods Hole, off Newfoundland, Sargasso Sea, Station "S" (32°10'N, 64°30'W), off SW Ireland, Galway Bay, Lough Hyne, Aran Islands, Ireland, Celtic Sea, Bristol Channel, Rnglish Channel, NE Scotland, Norway, North Sea, Fladen Ground area, Gullmarfjord, Kattegatt, Pas de Calais, English Channel, Plymouth, Roscoff, Bay of Biscay, Glenans Islands, Arcachon Bay, Portugal, Mondego estuary, Ria de Aveiro, off Coruña, Vigo, Santander, off Gijon, Cap Ghir, Ibero-moroccan Bay, Morocco, Medit. (Alboran Sea, Castellon, Baleares, off Barcelona, Banyuls, Marseille, Villefrance-s-Mer, Ligurian Sea, Tyrrhenian Sea, Naples, Milazzo, G. of Annaba, Strait of Messina, Gulf of Taranto, Malta, Adriatic Sea, Mljet Is., Venezia, Po delta, Aegean Sea, Thracian Sea, Lebanon Sea, Bardawill Lagoon, ? Black Sea)
N: 319
(22) F: 3,2-2,7; M: 2,8-2,5; (45) F: 3,25-2,75; M: 2,8-2,5; (65) F: 3; M: 2,8; (131) F: 3,5-2,7; M: 3,1-2,5; (180) F: 3,16-2,61; M: 2,58; 2,4; (199) F: 3,31-2,36; M: 3,19-2,36; (207) F: 3,09-2,82; M: 2,59-2,47; (327) F: 3,41-2,4; M: 2,98-2,35; (340) F: 2,8; (373) F: 2,6; (449) F: 5,4-2,7; M: 3,2-2,35; (920) F: 2,654-2,746; (927) F: 2,80-2,86 [winter]; 2,51-2,65 [Fall]; (1009) F: 2,75-3,25; (1037) F: 1,9-2,5; (1107) F: 2,5-3,13; M: 2,5-3,04; (1127) F: 2,6-3,3; M: 2,6-3,0; (1137) F: 2,26-2,57; M: 1,98-3,08; {F: 1,90-3,50; M: 2,35-3,20}
It does'nt take into account the female maximal value very high from Pesta (1920) in the Adriatic Sea.

The mean female size is 2.884 mm (n = 34; SD = 0.5716), and the mean male size is 2.681 mm (n = 23; SD = 0.3247). The size ratio (male : female) is 0.93.
Rem.: Overall Depth Range in Sargasso Sea: 0-2000 m (most numerous between 1000 and 1500 m) (Deevey & Brooks, 1977, Station"S").
This species has sometimes been confused with C. finmarchicus in the literature, the latter doubtful in the Mediterranean Sea.
In the Pacific it has been replaced by C. sinicus or C. australis, in the southern Atlantic and near South Africa by C. agulhensis.
A doubt subsists in considering C. euxinus as a valid species.
After Vucetic (1965) the number of teeth on coxopodite is for the females 24-26 (23-35), and 16-20 for males, and in C. helgolandicus var. ponticus (Yashnov, 1955) (= Calanus euxinus 24-29 (18-37) for females, and 11-21 for males.

For Itoh (1970 a, fig.2, from co-ordonates) the Itoh's index value from mandibular gnathobase = 370; for Schnack: 370, and for Lapernat & Razouls (2002, p.19) the Itoh's index value = 500.2 (number of teeth: 8).
After Corner & al. (1974) this species could survive the winter in the Clyde sea-area by feeding carnivorously on nauplians forms.

See in Mazzocchi M.G., 2006
Last update : 25/10/2016
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