Calanoida ( Order )
    Calanoidea ( Superfamily )
Calanidae Dana, 1846 ( Calanoidea )
Ref.: Giesbrecht, 1892 (p.41); Giesbrecht & Schmeil, 1898 (p.12); Sars, 1901 a (1903) (p.8); Esterly, 1905 (p.122); van Breemen, 1908 a (p.5, clé des G); Wilson, 1932 a (p.19); Rose, 1933 a (p.55); Mori, 1937 (1964) (p.2, clé des G); Brodsky, 1950 (1967) (p.81, 84); Vervoort, 1951 (p.55); Farran & Vervoort, 1951 (n°32, p.3); Chen & Zhang, 1965 (p.25); Björnberg, 1972 (p.24); Brosdsky, 1972 (1975) (p.1, 115, Rev.); Bradford & Jillett, 1974 (p.5, Rev.); Andronov, 1974 a (p.1005); Brodsky, 1976 (p.5); Vyshkvartzeva, 1976 (p.11); Björnberg & al., 1981 (p.616); Bowman & Abele, 1982 (p.10); Razouls, 1982 (p.57); Brodsky & al., 1983 (p.147); Sazhina, 1985 (p.107, 109, N); Fleminger, 1985 (p.273, 277, Table 2, Rem. A1); Mauchline, 1988 (p.719); Bradford, 1988 (p.73, Rev.); Zheng Zhong & al., 1984 (1989) (p.224, clé des G.); Nishida, 1989 (p.174, Rem.); Huys & Boxshall,1991 (p.461); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.22, Déf.); Chihara & Murano, 1997 (p.737); Barthélémy, 1999 a (p.30); Bradford-Grieve & al., 1999 (p.877, 902, 904, 905, 906, clé des G.); Ohtsuka & Huys, 2001 (p.445, 461); Boxshall & Halsey, 2004 (p.13; 49; 78: Def.; p.80: Genera Key); Mulyadi, 2004 (p.10, Genera Key for Indonesian Seas); Vives & Shmeleva, 2007 (p.889, Genera Key); Blanco-Bercial & al., 2011 (p.103, Table 1, Fig.2, 3, 4, Biol. mol, phylogeny)
Bradford-Grieve J.M., (2002 onwards). Key to calanoid copepod families. Version 1 : 2 oct 2002. http://www.crustacea.net/crustace/calanoida/index.htm
Rem.: Fleminger (1985) studies the number and the arrangement of the setae and aesthetascs on the A1 of 25 species in the Calanidae family. The author defines 4 types of setae and 3 forms of aesthetasks. Besides the sexual dimorphism of the A1, which occur always in the (M) by a fusion of the 2 first proximal articles, and, according to the genera, the fusion of certain other articles, one observes in certain (F) of numerous species, an arrangement of groups of setae and aesthetascs homologous to those in the (M) in weak proportion and variable in time. These (F) having a (M) phenotype present two problems: one of systematic and phylogenetic interest, the other concerns the hypothesis of a sex change in the course of development and of its determinism. For the author the views of Brodsky (1972) as well as those expressed by Bradford et Jillett (1974) are not satisfactory.
Thereby Neocalanus is limited to 2 species N. gracilis and N. robustior likewise Calanus, sensu lato, holds the taxa of which the generic status is still uncertain (C. cristatus , C. hyperboreus , C. plumchrus , C. propinquus , C. simillimus , C. tonsus ). May be considered as Calanus sensu stricto the species grouped in the complex ' Calanus finmarchicus ' ( C. chilensis , C. finmarchicus , C. glacialis , C. pacificus C. orientalis , C. sinicus , C. australis , C. helgolandicus , C. marshallae ). The genus Nannocalanus is maintained, as well as those proposed by Bradford & Jillett (1974): Calanoides, Canthocalanus, Cosmocalanus, Mesocalanus, Undinula.
Bradford (1988) contests the scheme established by Brodsky (1972). She refers to the morphological relationships that permit the species groups more strict and more neutral concerning the phylogenetical relationships between the genera.
In addition to only the morphological criteria, various aspects of their biology and genome can now be contemplated (Bucklin, Frost & Kocher, 1992, 1995, Bucklin & Lajeunesse, 1994; Bucklin, Thomas & Kocher, 1996; Bucklin, Lajeunesse, Curry, Wallinga & Garrison, 1996).
8 G.: Calanoides, Calanus, Canthocalanus, Cosmocalanus, Mesocalanus, Nannocalanus, Neocalanus, Undinula.
Family Calanidae - Plate 1Issued from : A. Fleminger in Mar. Biol., 1985, 88. [p.277].
Morphological types of setae and aesthetascs.
Nota: Four types of antennal setae may be recognized among the Calanidae:
Type 1: smooth-surfaced, acuminate filaments varying from shorter than the bearing segment to more than 10 times its length (D), and located anteriorly on the segment.
Type 2: short, coniform processes (E), always found on the anterior, distal end of Segment 8 and in many species on Segment 12.
Type 3: pseudo-annulated setae bearing spike-like setules (F), located on Segments 22, 23 and 24.
Type 4: short and with a more or less capitate apex, usually filled with lipid-like globules (G); they occur only in adults males on one or more of the following segments depending upon the species (fused Segment 1-2, Segment 7 and Segment 9. The capitate setae occupy the distal position that is filled by a Type 1 seta in the female and in the copepodite stage V.

Aesthetascs:appear in 3 forms:
Category 1: slender, vermiform processes (A) located on most segments of the female antenna.
Category 2: digitiform, irregularly swollen or pinched along their length, and rounded at the apex (B); they occur on Segments 1 thriugh 9 in males.
Category 3: occurring in males on Segments 10 to 23, vary structurally in fine detail among the genera, but all have a central cuticular keel articulating proximally with the antero-distal corner of the segment and extending to the succeeding antennal segment (C). A 'fleshy' layer covers the keel and extends as dorsal and ventral lamellae over the length of the Category 3 aesthetasc. These lamellae may be fused over the entire length of the aesthetasc or may separate at the distal end. The 'fleshy' layer is fragile and often is partially to totally eroded by capturing nets or during storage in formaldehyde.

Family Calanidae - Plate 2Issued from : A. Fleminger in Mar. Biol., 1985, 88. [p.278, Table 2].
Calanidae Number of setae (Set) and aesthetascs (Aes) per antennule segment in the Calanidae genera examined.
Individual segments with similar numbers of Set and Aes are grouped together.

Arrangement of setae and aesthetascs:
In all species studied, most or all of the adult females had a trithek arrangement (i.e. 1 proximal seta, 1 distal seta, and 1 distal aesthetasc) on each segment of A1 except Segments 1 and 20 to 25 (see Calanus pacificus californicus Fleminger, 1985). Segment 1 bears 3 setae and 1 aesthetasc. Segment 2, apparently a fusion of 3 segments, bears 3 such tritheks in positions 2a, 2b, and 2c. Segments 20 and 21 each lack 1 seta and the latter usually lacks the aesthetasc as well. Segments 22 and 23 have only 1 seta and 1 aesthetasc anteriorly and 1 posteriorly positioned, plumose, pseudoannulated seta (Type 3). Segment 24 has 2 setae similar to those of Segment 23, but no aesthetasc. Segment 25 bears 4 relatively short setae, i minute seta and 2 aesthetascs (see fig. A1 of calanus pacificus californicus and Calanoides philippinensis from Fleminger, 1985) ; it may be derived from the fusion of two or more segments.
The A1 of adult males exhibits alternation of trithek and quadrithek combinations, the latter consisting of 2 setae and 2 aesthetascs and occurring on Segments 2b, 3, 5, 7 and 9. Segments 2a, 2c, 4, 6, 8 and 10 through 19 bear tritheks (as in the female).Segment 21 bears 1 seta and 1 aesthetasc in the male. The remaining segments carry the same numbers and types of setae and aesthetascs as found in the female. All the setae and th escale-like male aesthetascs are articulated and rotate through about 90° from normal to the antenna surface to flat against it.

Family Calanidae - Plate 3Issued from : A. Fleminger in Mar. Biol., 1985, 88. [p.279, Table 2 (continued)].
Calanidae Number of setae (Set) and aesthetascs (Aes) per antennule segment in the Calanidae genera examined.
Bracketed segments are completely fused; Segments 8 and 9 are incompletely fused. Individual segments with similar numbers of Set and Aes are grouped together.

Family Calanidae - Plate 4Issued from : A. Fleminger in Mar. Biol., 1985, 88. [p.279, Table 3].
Calanus s.s. Antennule segmentation, setation and aesthetasc.
Individual segments with similar numbers of Set and Aes are grouped together.
Setae: Set = setiform; Spi = spiniform; Plu = plumose and pseudoannulated; Cap = capitate.
Aesthetascs: Ver = vermiform; Dig = digitiform; Sca = scale-like.
(1) Calanoides Brady, 1883
Syn.: Carinocalanus Brodsky, 1972 (1975) (p.112)
Ref.: Brady, 1883 (p.74); 1914 (p.4); A. Scott, 1909 (p.10); Sewell, 1929 (p.25); Vervoort, 1946 (p.29); 1951 (p.55, Rem.); Carvalho, 1952 a (p.137); Tanaka, 1956 (p.259); Chen & Zhang,1965 (p.28); Ramirez, 1966 (p.6); Brodsky, 1972 (1975) (p.113, 114, 116); Bradford & Jillett, 1974 (p.6, 9, 13, Rev.); Fleminger, 1985 (p.273, 275, 278, 285, Table 2, Rem.: A1); Mauchline, 1988 (p.719); Razouls, 1982 (p.74); Bradford, 1988 (p.73,76, Rev.); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.23, Déf.); Chihara & Murano, 1997 (p.741); Mauchline, 1998 (p.68); Bradford-Grieve & al., 1999 (p.907: clé spp.); Boxshall & Halsey, 2004 (p.80) ; Vives & Shmeleva, 2007 (p.890); Sabatini & al., 2007 (p.341, Rem.: Comparison of species)
Rem.: type: Calanoides patagoniensis Brady,1883, not C. carinatus Kröyer,1849.
5 spp. + 1 doubtful.
Genus Calanoides - Plate 1issued from : M. E. Sabatini, F. C. Ramirez & J. Bradford-Grieve in Invert. Syst., 2007, 21. [p.361, Table 4]. Comparison of Calanoides species: key characters to distinguish female specimens.

Genus Calanoides - Plate 2issued from : M. E. Sabatini, F. C. Ramirez & J. Bradford-Grieve in Invert. Syst., 2007, 21. [p.362, Table 5]. Comparison of Calanoides species: key characters to distinguish male specimens.
(2) Calanus Leach, 1816
Syn.: Monoculus Gunnerus,1770; Cyclops (part.) Müller,1776;
Cetochilus Roussel de Vauzème,1834;
Nannocalanus (part.) Sars,1925
Ref.: Kröyer, 1848 (in Damkaer & Damkaer, 1979, p.12); Brady, 1878 (p.37); Giesbrecht, 1892 (p.45, 88); Dahl, 1894 (p.74, clé spp.); Giesbrecht & Schmeil, 1898 (p.12, clé spp.); Wheeler, 1901 (p.164); Sars, 1901 a (1903) (p.8); Esterly, 1905 (p.123, clé spp.); van Breemen, 1908 a (p.6, clé spp.); A. Scott, 1909 (p.7); Wolfenden, 1911 (p.190); Esterly, 1924 (p.83); Sars, 1925 (p.5); Sewell, 1929 (p.19, Rem.); Wilson, 1932 a (p.19, clé spp.); Rose, 1933 a (p.55, clé spp.); Mori, 1937 (1964) (p.11, clé spp.); Vervoort, 1946 (p.22); Brodsky, 1950 (1967) (p.85, clé spp.); Tanaka, 1956 (p.251); Brodsky, 1959 a (p.1548, clé spp., Biogéo); 1961 (p.5, clé spp.); Chen & Zhang, 1965 (p.25); Ramirez, 1966 (p.5); Brodsky, 1972 (1975) (p.4, 112, 115, Rev., clé S/G & spp.); Vyshkvartzeva, 1972 (1975) (p.186); 1976 (p.11); Frost, 1974 (p.77); Bradford & Jillett, 1974 (p.5, Rev.); Kabata, 1979 (p.20); Razouls, 1982 (p.62); Gardner & Szabo, 1982 (p.135); Brodsky & al., 1983 (p.149, clé spp.); Van der Spoel & Heyman, 1983 (p.147); Zheng Zhong & al., 1984 (1989) (p.225, clé spp.); Fleminger, 1985 (p.277, Table 2, 3, 4, Rem.: A1); Mauchline,1987 (p.719); Bradford, 1988 ( p.73, Rev.); De Decker & al., 1991 (p.27); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.29, Déf.); Bucklin & al., 1995 (p.655); Chihara & Murano, 1997 (p.738); Mauchline, 1998 (p.67); Bradford-Grieve & al., 1999 (p.907: clé spp.); Boxshall & Halsey, 2004 (p.80); Vives & Shmeleva, 2007 (p.892, spp. Key)
Rem.: type: Monoculus finmarchicus Gunnerus,1770. 14 spp.
(3) Canthocalanus A. Scott, 1909
Ref.: A. Scott, 1909 (p.8); Vervoort, 1946 (p.36); Tanaka, 1956 (p.259); Chen & Zhang, 1965 (p.29); Brodsky, 1972 (1975) (p.114, 116, Rev.); Bradford, 1972 (p.14); Bradford & Jillett, 1974 (p.6, 10); Razouls, 1982 (p.76); Zheng Zhong & al., 1984 (1989) (p.227); Fleminger, 1985 (p.273, 278, Table 2, Rem.: A1); Mauchline, 1987 (p.719); Bradford, 1988 (p.73, 76, 79, Rem.); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.33, Déf.); Chihara & Murano, 1997 (p.739); Mauchline, 1998 (p.68); Boxshall & Halsey, 2004 (p.80)
Rem.: type: Canthocalanus pauper (Giesbrecht,1888). 1 sp.
(4) Cosmocalanus Bradford & Jillett, 1974
Ref.: Bradford & Jillett, 1974 (p.6, 12); Razouls, 1982 (p.83); Fleminger, 1985 (p.273, 278, Table 2, Rem.: A1); Mauchline, 1987 (p.719); Bradford,1988 (p.74, 75, 79); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.35, Déf.); Chihara & Murano, 1997 (p.737); Mauchline, 1998 (p.67: F, M); Barthélémy, 1999 a (p.26); Boxshall & Halsey, 2004 (p.80)
Rem.: type: Undina darwini Lubbock,1860. 2 spp.
(5) Mesocalanus Bradford & Jillett, 1974
Syn.: Neocalanus (part.): Brodsky & al., 1983 (p.184, clé spp.)
Ref.: Bradford & Jillett, 1974 (p.6, 12); Razouls, 1982 (p.79); Gardner & Szabo, 1982 (p.147); Fleminger, 1985 (p.273, 278, Table 2, Rem.: A1); Bradford, 1988 (p.74-76); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.38, Déf.); Chihara & Murano, 1997 (p.741); Mauchline, 1998 (p.68); Boxshall & Halsey, 2004 (p.80); Vives & Shmeleva, 2007 (p.897)
Rem.: For Brodsky (1980: pers. comm.) the creation of this genus appears not well guaranteed. Type : Calanus tenuicornis Dana,1849. 2 spp.
(6) Nannocalanus Sars, 1925
Syn.: Canthocalanus (part.) : Brodsky, 1972 (1975) (p.116); Calanus (part.) : Bradford & Jillett, 1974 (p.6, 9)
Ref.: Sars, 1925 (p.9); Vervoort, 1946 (p.25); Chen & Zhang, 1965 (p.30); Razouls, 1982 (p.79); Zheng Zhong & al., 1984 (1989) (p.227); Fleminger, 1985 (p.273, 278, Table 2, Rem.: A1); Bradford, 1988 (p.76, 79); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.39, Déf.); Chihara & Murano, 1997 (p.737); Barthélémy, 1999 a (p.26); Boxshall & Halsey, 2004 (p.80); Vives & Shmeleva, 2007 (p.899)
Rem.: type: Cetochilus minor Claus,1863. 2 spp.
(7) Neocalanus Sars, 1925
Syn.: Tropocalanus & Neocalanus (part.) Brodsky, 1972 (1975) (p.116); Brodsky & al., 1983 (p.184)
Ref.: Sars, 1925 (p.7); Sewell, 1929 (p.26, Rem.); Wilson, 1932 a (p.27); Vervoort, 1946 (p.38, Rem.); Oliveira, 1946 (p.454); Tanaka, 1956 (p.261); Chen & Zhang, 1965 (p.26); Bradford & Jillett, 1974 (p.10, Rev.); Razouls, 1982 (p.77); Gardner & Szabo, 1982 (p.141); Zheng Zhong & al., 1984 (1989) (p.227); Fleminger, 1985 (p.273, 278, Table 2, Rem.: A1); Mauchline, 1987 (p.719); Bradford, 1988 (p.74, 76, 79); Miller, 1988 (p.226, 270); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.41, Déf.); Chihara & Murano, 1997 (p.739); Mauchline, 1998 (p.68); Bradford-Grieve & al., 1999 (p.908: clé spp.); Miller, 2002 (p.139); Boxshall & Halsey, 2004 (p.80); Vives & Shmeleva, 2007 (p.901, spp. Key)
Rem.: type: Calanus gracilis Dana,1849. 6 spp.
Genus Neocalanus - Plate 1issued from : R.J. Machida, M.U. Miya, M. Nishida & S. Nishida in Mar. Biol., 2006, 148. [p.1077, Fig.4].
Relationship between the phylogeny and distribution pattern of the Neocalanus species.
Shaded portions of the maps represent distributions based on the litterature. Clades A-E correspond to those in Fig.3, p.1075.
By comparing the habitat localities and the phylogenetic relationship, it has been demonstrated that major linkeages of Neocalanus species diverged into different habitat localities. ''Groups'' classified by Bradford & Jillett (1974) is the first bifurcation among Neocalanus species (clade A), those separated by the temperature gradient between tropical to subtropical waters (''gracilis\" group: N.gracilis and N. robustior) and subarctic waters (''tonsus'' group: N. tonsus, N. cristatus, N.plumchrus, N. flemingeri). Second bifurcation is between N. tonsus and the rest of the species in the ''tonsus'' group (N. cristatus, N. flemingeri, N. plumchrus), those separated by the subantarctic and subarctic Pacific (clade B). Although these subantarctic and subarctic Pacific species constitute the sister clade, their distribution areas are widely separated. The occurrence of closely related animals to the north and south of the tropical zone but not within the tropics, known as bipolar or antitropical distribution, among zooplankton assemblage is not an exceptional phenomenon (Reid, 1978). Several mechanisms have been proposed for the development of antitropical distributions, divisible into two classes: dispersal and vicariance (Burridge, 2002). The dispersal mechanism assumes that a taxon was originally represented on one side of the tropics, and subsequently moved across the equator to colonize the opposite hemisphere. On the other hand, the vicariance mechanism assumes that taxa that once occurred in the tropics but were later expatriated, resulting in osolated Northern and Southern Hemisphere populations.
The four species that occur in subarctic Pacific and subantarctic clustered monophyletically and all of the species are known to perform ontogenetic vertical migration (Miller, 1988). Therefore, the dispersal mechanism is much more likely than the vicariance mechanism for the explanation of the antitropicality of thes four Neocalanus species.
Two explanations have been proposed for the transequatoril exchange of populations in the plankton community (Dunbar, 1979). One is through the deep water link between populations, a low-temperature route. Another explanation is transequatorial dispersal during glaciations, implicating cooler surface water temperature, only applicable to carnivorous or omnivorous zooplankton that complete its life cycle in mesopelagic environment, and not for Neocalanus species, which mainly depend on primary production.
(8) Undinula A. Scott, 1909
Syn.: Undina (part.) Dana,1847
Ref.: A. Scott, 1909 (p.16); Sars, 1925 (p.10); Wilson, 1932 a (p.29); Dakin & Colefax, 1940 (p.87, clé spp.); Vervoort, 1946 (p.72); Carvalho, 1952 a (p.138); Tanaka, 1956 (p.264); Chen & Zhang, 1965 (p.30); Brodsky, 1972 (1975) (p.114, 116, 117, 120); Bradford & Jillett, 1974 (p.11, Redef.); Razouls, 1982 (p.80); Zheng Zhong & al., 1984 (1989) (p.228); Fleminger, 1985 (p.273, 278, Rem.: A1); Mauchline, 1988 (p.719); Bradford, 1988 (p.74, 77); Razouls, 1993 (p.308); Bradford-Grieve, 1994 (p.44); Chihara & Murano, 1997 (p.739); Mauchline, 1998 (p.68); Boxshall & Halsey, 2004 (p.80); Vives & Shmeleva, 2007 (p.905)
Rem.: type: Undina vulgaris (Dana,1849). 1 sp.
Additional information for the Calanidae family ( back to the main information )

Remarks on sizes :
 
The minimum and maximum values of the adult females of Calanidae family sare presented in figure 1 (in ordinates: total length in mm; in abscissa : Calanoides from 1 to 5, Calanus from 6 to 19, Canthocalanus : 20, Cosmocalanus : 21 and 22, Mesocalanus : 23 and 24, Nannocalanus : 25; Neocalanus from 26 to 31; Undinula : 32
 

Figure 1



Table I: species, length, depth and location characteristics (E: epipelagic; M: meso-; B: bathy-).

species
Lg (mm)
 
 
Neocalanus cristatus
7,50 à 10,50
Arct. & sub-arct. (Pacif.)
E-B
Calanus hyperboreus
5,50 à 10
Arct. & sub-arct. (Atlant.)
 
Calanoides acutus
± 3,50 à ± 6
Antarct. & sub-antarct.
E-M
Calanus glacialis
-
Arct. & sub-arct.
 
Calanus propinquus
-
Antarct. & sub-antarct.
E-M
Neocalanus flemingeri
-
sub-arct. (Pacif.)
 
Neocalanus plumchrus
-
sub-arct. (Pacif.)
 
Calanus pacificus
2,15 à 5
sub-arct., cold temperate
 
Neocalanus robustior
-
cosmop, tropic., subarct.
E-M
Neocalanus tonsus
-
sub-antarct., tropic.
 
Calanus finmarchicus
-
cold temperate
E-M
Calanoides carinatus
± 2,50 à ± 4
cosmop, sub-antarct, tropic.
E-B
Calanoides macrocarinatus
-
sub-antarct.
E-B
Calanus australis
-
sub-antarct.
E-B
Calanus chilensis
-
cold temperate
 
Calanus euxinus
-
temperate
 
Calanus helgolandicus
-
temperate
E-M
Calanus simillimus
-
sub-antarct.
 
Calanus sinicus
-
tropic., cold temperate
 
Calanus marshallae
-
sub-arct.
 
Calanus jaschnovi
-
temperate
 
Neocalanus gracilis
-
cosmop., topic., cold temperate
 
Calanoides patagoniensis
± 2 à ± 3
sub-antarct.
E-?M
Calanoides philippinensis
-
tropic.
M
Calanus agulhensis
-
temperate
 
Cosmocalanus caroli
-
tropic.
 
Cosmocalanus darwini
-
cosmop., topic., cold temperate
E-?M
Mesocalanus lighti
-
sub-tropic.
 
Mesocalanus tenuicornis
-
cosmop., topic., temperate
E-B
Undinula vulgaris
-
cosmop., tropic.
 
Nannocalanus minor
-
cosmop., tropic., cold temperate
E-M
Canthocalanus pauper
-
tropic. (Indo-Pacif.)
E

 Any use of this site for a publication will be mentioned with the following reference :

Razouls C., de Bovée F., Kouwenberg J. et Desreumaux N., 2005-2012. - Diversity and Geographic Distribution of Marine Planktonic Copepods. Available at http://copepodes.obs-banyuls.fr/en 
[Accessed May 23, 2013]

© copyright 2005-2012 CNRS, UPMC

Webmaster
CNRS   Observatoire Océanologique de Banyuls sur Mer - Laboratoire Arago
UPMC - Paris Universitas

 

Version française
English version

 

Search

On the WEB of CNRS

 


Marine Planktonic Copepods

Marine Planktonic Copepods

 

Imprimer Contact Accueil Plan du site Accès restreint Retour Une du Labo Imprimer Contact Plan du site Crédits Téléchargez les Plug-Ins