Calanoida ( Order )
    Spinocalanoidea ( Superfamily )
Spinocalanidae Vervoort, 1951 ( Spinocalanoidea )
Syn.: no Arctokonstantinidae Markhaseva & Kosobokova, 2001 (p.319)
Ref.: Vervoort, 1951 (p.10, 72, Déf.); Farran & Vervoort, 1951 g (n°39, p.3); Mazza, 1967 (p.99); Grice, 1971 (p.273, 279, clé); Andronov, 1974 a (p.1005); Damkaer, 1975 (p.13, clé des G.); Bowman & Abele, 1982 (p.9); Razouls, 1982 (p.137); Fleminger, 1983 (p.605, 607, clé des G., Rem.); Brodsky & al., 1983 (p.144, 147, 242, clé des G.); Park, 1986 (p.192, 193, 194); Mauchline, 1988 (p.724, 740: pores cuticulaires.); Huys & Boxshall, 1991 (p.467); Razouls, 1991 (p.209, Rem.); Schulz, 1992 (p.176, clé des G.); Razouls, 1993 (p.311); Bradford-Grieve, 1994 (p.94); Schulz, 1996 (p.599, Rem., clé des G.); Bradford-Grieve & al., 1999 (p.878, 903, 904, 913: clé des G.); Markhaseva & Kosobokova, 2001 (p.319, 323, Rem.); Ohtsuka & Huys, 2001 (p.461); Boxshall & Halsey, 2004 (p.12, 15; 49; 195: Def.; p.198: Clé des Genres); Schulz, 2004 (p.211: Rem.); Vives & Shmeleva, 2007 (p.820, part. key G.); Markhaseva, 2008 (p.15, Rem.); 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.: According to Vervoort (1951, p.72), Farran, in his manuscripts of the 'Zooplankton Sheets', formed a new familly, Spinocalanidae, to comprise such genera as Spinocalanus Giesbrecht, 1888, Monacilla Sars, 1905 and Mimocalanus Farran, 1908. Although Farran gives no diagnosis of this new family it seems clear that he based it on the quite different arrangement of the setae on the swimming legs and the family has been defined on such lines in the revision of the 'Sheets'. 10 G.: Damkaeria, Isaacsicalanus, Kunihulsea, Methanocalanus, Mimocalanus, Monacilla, Mospicalanus, Rhinomaxillaris, Spinocalanus, Teneriforma. When estimating whether a particular characteristic is primitive or advanced, Markhaseva (2008, p.12 and see table) follows the criteria generally accepted among copepodologists : the evolution of copepods is characterized by oligomerization (V.A. Dogel’, 1954) and follows, primarily, the pathway of reductions, fusions, and losses (Boxshall, Ferrari & Tiemann, 1984), i.e, the condition of the greatest articulation and setation of appendages is considerd to be the most primitive one.
For Markhaseva (2008, p.12-13 and p.15: table) the genera Arctokonstantinus, Sognocalanus, Foxtonia and Foxtosognus belong to the new family Arctokonstantinidae.


After Markhaseva & Schulz (2008, p.287), concerning the ancestral mode of life of Spinocalanidae, some speculation are possible. Due to a more primitive morphology of pelagic representatives of spinocalanids, it may be assumed that they probably originated in the pelagic environment.
Typically, pelagic spinocalanid genera (Monacilla, Spinocalanus, Mimocalanus, Mospicalanus, Teneriforma) demostrate diversity of some primitive characters not observed in their benthopelagic relatives (Damkaeria, Kunihulsea, Methanocalanus, Isaacsicalanus). The following plesiomorphies are considered typical of pelagic spinocalanids :
1- 11 setae on Md endopod segment 2 in Spinocalanus, vs. 9-10 setae in other spinocalanids.
2- 15 setae on Mx1 praecoxal arthrite in Monacilla, vs. fewer in other spinocalanids.
3- 6 setae on Mx1 coxal endite in Spinocalanus, vs. fewer in other spinocalanids.
4- 6 setae on Mx1 distal basal endite in Teneriforma, vs. fewer in other spinocalanids.
5- 16 setae on endopod and 11 setae on exopod of Mx1 in Monacilla, Spinocalanus, Mimocalanus (but 23 on endopod in M. crassus) ; exopod setation of this type also in Mospicalanus, vs. fewer in other spinocalanids.
6- Mx2 with 9 setae (2+7) on endopod fused to distal basal endite in Monacilla, Spinocalanus, vs. fewer in other spinocalanids.
7- praecoxal endites of Mxp syncoxa with 1, 2, 3 setae in Monacilla, Spinocalanis, vs. fewer in other spinocalanids.
8- coxal endite of Mxp syncoxa with 4 setae in Monacilla, Mimocalanus, vs. fewer in other spinocalanids.
Typically, pelagic Spinocalanidae share some primitive characters with more derived benthopelagic spinocalanid genera :
1- the primitive type of A2 exopod setation 1, 1-1-1, 1, 1, 1, 1, 1 and 3 setae, is shared by pelagic Monacilla, Spinocalanus, Teneriforma and the benthopelagic genus Kunihulsea.
2- the most primitive condition in the armature of the proximal basal endite of Mx1 (4 setae) is found in the pelagic genera Monacilla, Spinocalanus , Mimocalanus, Mospicalanus as well as the benthopelagic Methanocalanus.
3- the most primitive armature of the proximal praecoxal endite of Mx2 (7 setae) is shared by the pelagic Monacilla and the benthopelagic Methanocalanus.
Family Spinocalanidae - Plate 1issued from : E.L. Markhaseva in Russian J. Mar. Biol., 2008, 34 (1). [p.15].
Details of armament of mouthparts and P1 in Spinocalanidae.
The table presents the most primitive condition of characters that has revealed for the genus.

Family Spinocalanidae - Plate 2issued from : E.L. Markhaseva & K. Schulz in Zootaxa, 2008, 1866. [p. 279].
Selected character states of Spinocalanoidea females (diagnostic characters of the family Arctokonstantinidae are in bold).
Refernces: Hulsemann & Grice, 1963; Fosshagen, 1967, 1983; Park, 1970; Damkaer, 1975; Brodsky & al., 1983; Schulz, 1989, 1996, 2004; Markhaseva & Kosobokova, 2001; Boxshall & Halsey, 2004; Ivanenko & al., 2007; Markhaseva & Schulz, 2008.

Nota: According to the authors (p.288), Arctokonstantinidae and Spinocalanidae belong to the Spinocalanoidea, the sister group of the Ryocalanoidea. Diagnostic characters considered to distinguis the superfamily Spinocalanoidea from Ryocalanoidea are not very strong.
Arctokonstantinus Markhaseva & Kosobokova, 2001
Ref.: Markhaseva & Kosobokova, 2001 (p.320); Boxshall & Halsey, 2004 (p.198); Markhaseva, 2008 (p.15, Rem.)
Rem.: Cf. Arctokonstantinidae
(1) Damkaeria Fosshagen, 1983
Ref.: Fosshagen, 1983 (p.260, 261); Schulz, 1989 (p.195, Rem.); Bradford-Grieve, 1994 (p.95, Déf.); Mauchline, 1998 (p.87, 89: F); Boxshall & Halsey, 2004 (p.198) ; Schulz, 2004 (p.198, Def.); Markhaseva, 2008 (p.15, Rem.)
Rem.: This genus, included previously among the Bathypontiidae, was transferred in this family by Bradford-Grieve (1994). 2 spp.
Foxtonia Hulsemann & Grice, 1963
Ref.: Hulsemann & Grice, 1963 (p.733); Razouls, 1982 (p.593); Fosshagen, 1983 (p.260); Razouls, 1993 (p.308); Mauchline, 1998 (p.66); Boxshall & Halsey, 2004 (p.198)
Rem.: Cf. Arctokonstantinidae
(2) Isaacsicalanus Fleminger, 1983
Ref.: Fleminger, 1983 (p.606, 607, 615); Schulz, 1989 (p.195, Rem.); Razouls, 1993 (p.311); Mauchline, 1998 (p.87: F); Boxshall & Halsey, 2004 (p.198); Markhaseva, 2008 (p.15, Rem.)
Rem.: 1 sp.
(3) Kunihulsea Schulz, 1992
Ref.: Schulz, 1992 (p.176); Mauchline, 1998 (p.87: F); Boxshall & Halsey, 2004 (p.198) ; Schulz, 2004 (p.206, Rem. : emend.); Markhaseva, 2008 (p.15, Rem.)
Rem.: 2 spp.:
(4) Methanocalanus Ivanenko, Defaye & Cuoc, 2007
Ref.: Ivanenko & al., 2007 (p.38); Markhaseva, 2008 (p.15, Rem.)
Rem.: 1 sp.
(5) Mimocalanus Farran, 1908
Ref.: Farran, 1908 b (p.22); Sars, 1925 (p.34); Rose, 1933 a (p.86); Vervoort, 1946 (p.156); Davis, 1949 (p.23); Farran & Vervoort, 1951 h (n°40, p.3); Damkaer, 1975 (p.67, clé spp.); Brodsky, 1950 (1967) (p.137); Tanaka, 1956 c (p.387); Fleminger, 1983 (p.607, 615); Brodsky & al., 1983 (p.303, clé spp.) ; Razouls, 1982 (p.147); Schulz, 1989 (p.195, Rem.); Razouls, 1993 (p.311); Bradford-Grieve, 1994 (p.98, Déf.); Mauchline, 1998 (p.87: F; p.90: M); Boxshall & Halsey, 2004 (p.198); Vives & Shmeleva, 2007 (p.821, spp. Key); Markhaseva, 2008 (p.15, Rem.)
Rem.: type: Mimocalanus. cultrifer . 10 spp.
Remarks on dimensions and sex ratio:
The mean female size is 2,082 mm (n= 10; S= 0,910; Cv= 0,437) and the mean male size is 1,770 mm (n= 6; S= 0,643; Cv= 0,369). The size ratio (M/F) is 0,902 ou 90,2 % (n= 6; S= 0,091; Cv= 0,101)
(6) Monacilla Sars, 1905
Syn.: Oxycalanus Farran,1908 b (p.25); A. Scott, 1909 (p.33); Wolfenden, 1911 (p.220); Hypsicalanus Wolfenden, 1911 (p.219)
Ref.: Sars, 1905 b (p.8); A. Scott, 1909 (p.34); Sars, 1925 (p.37); Sewell, 1929 (p.95); Rose, 1933 a (p.87); Vervoort, 1946 (p.157); Farran & Vervoort, 1951 h (n°40, p.3); Tanaka, 1956 c (p.395); Grice,1971 (p.274); Damkaer, 1975 (Déf., p.14, 62, clé spp.); Razouls, 1982 (p.149) Brodsky & al., 1983 (p.324, clé spp.); Fleminger, 1983 (p.607, 610, 612, 615); Mauchline,1988 (p.724); Schulz, 1989 (p.195, Rem.); Ferrari, 1992 (p.392, tab.3); Razouls, 1993 (p.311); Bradford-Grieve, 1994 (p.99, Déf.); Mauchline, 1998 (p.78: F; p.80: M); Bradford-Grieve & al., 1999 (p.914: clé spp.); Boxshall & Halsey, 2004 (p.198) ; Vives & Shmeleva, 2007 (p.828, spp. Key); Markhaseva, 2008 (p.15, Rem.)
Rem.: type: Monacilla typica . 3 spp. + 1 indet.
(7) Mospicalanus Schulz, 1996
Ref.: Schulz, 1996 (p.595); Boxshall & Halsey, 2004 (p.198); Markhaseva, 2008 (p.15, Rem.)
Rem.: 1 sp.
(8) Rhinomaxillaris Grice & Hulsemann, 1967
Ref.: Grice & Hulsemann, 1967 (p.39, 40, Rem.); Razouls, 1982 (p.593); Fosshagen, 1983 (p.260); Razouls, 1993 (p.309); Schulz, 1996 (p.599, Rem.); Markhaseva & Kosobokova, 2001 (p.319, 323, Rem.); Boxshall & Halsey, 2004 (p.198)
Rem.: Formerly included in the Bathypontiidae family. 1 sp.
Sognocalanus Fosshagen, 1967
Ref.: Fosshagen, 1967 (p.308, 313); Razouls, 1982 (p.594); Fosshagen, 1983 (p.260); Razouls, 1993 (p.309); Schulz, 1996 (p.599, Rem.); Mauchline, 1998 (p.87: F; p.90: M); Markhaseva & Kosobokova, 2001 (p.319, 323, Rem.); Boxshall & Halsey, 2004 (p.198)
Rem.: Cf. Arctokonstantinidae
(9) Spinocalanus Giesbrecht, 1888
Ref.: Giesbrecht, 1892 (p.52, 209); Giesbrecht & Schmeil, 1898 (p.31); Sars, 1900 (p.75); 1901 a (1903) (p.22); Wolfenden, 1906 (p.43); Esterly, 1906 a (p.55); van Breemen, 1908 a (p.28); Wolfenden, 1911 (p.215); Rose, 1933 a (p.84); Vervoort, 1946 (p.146); Davis,1949 (p.21); Brodsky, 1950 (1967) (p.122, clé spp.); Farran & Vervoort, 1951 g (n°39, p.3); Tanaka, 1956 c (p.389); Park, 1970 (p.482, clé spp.F); Grice, 1971 (p.275, 279, clé spp.); Damkaer, 1975 (p.15, clé spp., Rev.); Gardner & Szabo, 1982 (p.191); Razouls, 1982 (p.137); Brodsky & al., 1983 ( p.247, clé des spp.); Fleminger, 1983 (p.607, 615); Mauchline, 1988 (p.724, pores cuticulaires); Schulz, 1989 (p.195, Rem.); Razouls, 1993 (p.311); Bradford-Grieve, 1994 (p.100, Déf.); Mauchline, 1998 (p.78: F, p.80: M, figs.167, 169); Bradford-Grieve & al., 1999 (p.914: clé spp.); Boxshall & Halsey, 2004 (p.198); Vives & Shmeleva, 2007 (p.831, spp. Key); Markhaseva, 2008 (p.15, Rem.)
Rem.: type: S. abyssalis. 29 spp. + 3 unidentified.
Remarks on dimensions and sex ratio:
The mean female size is 1,954 mm (n= 25; S= 0,696; Cv= 0,356) and the mean male size is 2,032 mm (n= 19; S= 0,789; Cv= 0,388). The size ratio (M/F) is 0,919 or 91,9 % (n= 27; S= 0,097; Cv= 0,106). 0,953 or 95,3 % (n= 16; S= 0,15; Cv= 0,157). The sex-ratio (F/M) is 1,315. It is exceptional that the male size is superior to that of the females; the obtained mean size is from the sex ratio and is not confirmed in the ratio of the sizes (M/F) obtained from the paired female and male data.
(10) Teneriforma Grice & Hulsemann, 1967
Syn.: Tanyrhinus Farran, 1936 a (p.85)
Ref.: Grice & Hulsemann, 1967 (p.22); Grice, 1971 (p.274); Damkaer, 1975 (p.15, 82, Rev.); Fosshagen, 1983 (p.260, 261, Rem.); Brodsky & al., 1983 (p.331); Fleminger,1983 (p.607, 615); Razouls,1982 (p.152); Schulz, 1989 (p.195, Rem.); Razouls, 1993 (p.311); Bradford-Grieve, 1994 (p.105, Déf.); Mauchline, 1998 (p.87: F; p.90: M); Boxshall & Halsey, 2004 (p.198); Vives & Shmeleva, 2007 (p.846); Markhaseva, 2008 (p.15, Rem.)
Rem.: type: Tanyrhinus naso Farran,1936 . 3 spp.

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