Calanoida ( Order )
    Eucalanoidea ( Superfamily )
Eucalanidae Giesbrecht, 1892 ( Eucalanoidea )
Syn.: Eucalaninae Giesbrecht, 1892 (p.45); Esterly, 1905 (p.130)
Ref.: Sars, 1901 a (1903) (p.13); Gurney, 1931 a (p.84); Rose, 1933 a (p.66); Brodsky, 1950 (1967) (p.82, 99); Farran & Vervoort, 1951 b (n°34, p.3); Mazza, 1967 (p.63); Björnberg, 1972 (p.14, 24); Andronov, 1974 a (p.1005); Geletin, 1976 (p.75, Rev.); Björnberg & al., 1981 (p.619); Bowman & Abele, 1982 (p.10); Razouls, 1982 (p.90); Brodsky & al., 1983 (p.142, 146, 199); Sazhina, 1985; Mauchline, 1988 (p.723); Hiromi, 1987 (p.147); Zheng Zhong & al., 1984 (1989) (p.229, Genera Key); Huys & Boxshall, 1991 (p.463); Bradford-Grieve, 1994 (p.71, Def.); Madhupratap & al., 1996 (p.863, Table 5: %/copepods); Chihara & Murano, 1997 (p.787); Bradford-Grieve & al., 1999 (p.878, 901, 911, Genera Key); Ohtsuka & Huys, 2001 (p.445, 461); Goetze, 2003 (p.2321: phylogeny); Boxshall & Halsey, 2004 (p.15; 49; 115: Def.; p.116: Genera Key); Mulyadi, 2004 (p.105); Vives & Shmeleva, 2007 (p.869, Genera Key); Blanco-Bercial & al., 2011 (p.103, Table 1, Fig.2, 3, 4, Biol. mol, phylogeny); Laakmann & al., 2019 (p.330, fig. 2, 3, phylogenetic relationships); Hirai & al., 2020 (p.1, Fig.4: metabarcoding, Fig.8: OTUs distribution patterns, Fig.9: phylogenetic analysis)
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.: For Brodsky (1950 [1967, p.99]) the differences between the genera Eucalanus lato sensu and Rhincalanus are: absence of P5 from female, the absence of spines on thorax and abdomen, and pair of legs P5 uniramous in male for the first and P5 present in female, thorax and abdomen spinose in female, and one of the legs P5 of male biramous for the latter.
Geletin (1976) defines 3 sub-families: Eucalaninae (sensu stricto); Rhincalaninae , Subeucalaninae.
The genus Rhincalanus has become the type of the family Rhincalanidae.
3 G.: Eucalanus Dana, 1853; Pareucalanus Geletin, 1976; Subeucalanus Geletin, 1976.

Biology notes after Boxshall & Halsey (2004, p.116): Eucalanids are primary consumers in the epipelagic and mesopelagic plankton communities in both oceanic and neritic waters. They are primarily small particle feeders, utilising phyroplanktonic algal cells, and they exhibit a number of different feeding modes in the adults (Koehl & Strickler, 1981; Price, Paffenhöfer & Strickler, 1983; Price & Paffenhöfer, 1986a, b; Strickler, 1982; 1984) and naupliar stages (Paffenhöfer & Lewis, 1989).

Key to genera after Boxshall & Halsey (2004, p.116) :
1 - Female urosome 4-segmented .......... Eucalanus.
1' - Female urosome 3-segmented ......... 2.
2 - Male P5 uniramous on both sides, with small right leg not extending beyond distal margin of 2nd segment of long left leg; female Mx1 with defined coxal endite bearing 4 setae; A2 with 1st and 2nd exopodal segments separate ........... Pareucalanus.
2' - Male P5 uniramous on left side, right leg absent; Mx1 lecking defined coxal endite; A2 with 1st and 2nd exopodal segments fused .......... Subeucalanus.

Ohtsuka & Nishida (2017, p.571) point to that the taxonomy of coastal/oceanic family Eucalanidae is very difficult. Geotze & Ohman (2010) studied the molecular phylogeny and zoogeography of Eucalanidae, and revealed that intraspecific genetic variation is found even in oceanic taxa, raising interesting questions cobcerning the nature of any biological barriers to gene flow in oceanic waters. because calanids and eucalanids are considered not to have complex mating patterns, it is likely that sexual dimorphic features are less well developed.
Family Eucalanidae - Plate 1Issued from : G.A. Boxshall & S.H. Halsey in An Introduction to Copepod Diversity. The Ray Society, 2004, No 166, Part. I. [p.115].
Armature formula of swimming legs P1 to P4.

Nota: Female P5 absent.
- Male P5 asymmetrical; right leg in some species 3 or 4-segmented comprising unarmed coxa and basis and 1 or 2-segmented exopod. 1st exopodal segment unarmed or with outer spine, 2nd segment with apical seta. Right leg absent in many species. Left leg typically 4-segmented, longer than right leg when present, comprising unarmed coxa and basis plus 2-segmented exopod. 1st exopodal segment unarmed or with outer spine, 2nd segment often hirsute, armed with apical spine.
- Eggs release into water.

Family Eucalanidae - Plate 2Issued from : G.A. Boxshall & S.H. Halsey in An Introduction to Copepod Diversity. The Ray Society, 2004, No 166, Part. I. [p.117, Fig. 22].
Eucalanidae. A, Subeucalanus crassus (as Eucalanus crassus) habitus female; B, Pareucalanus attenuatus (as Eucalanus attenuatus) female Mx2; C, pareucalanus attenuatus 'as Eucalanus pseudattenuatus) female Md; D, Eucalanus elongatus female Mxp; E, male P5; F, Subeucalanus crassus (as Eucalanus crassus) male P5. [Giesbrecht, 1993a (1892): A-B and D-F; Sewell, 1947: C].
Eucalaninae Giesbrecht, 1892
Ref.: Giesbrecht, 1892 (p.45); Esterly, 1905 (p.130); Geletin, 1976 (p.92, Rev.)
Rem.: 2 G.: Eucalanus, Pareucalanus
Rhincalaninae Geletin, 1976
Ref.: Geletin, 1976 (p.92)
Rem.: Brodsky & al., 1983 (p.203) n'admettent pas la création de cette sous- famille. Boxshall & Halsey (2004, p.116) considèrent cette sous-famille comme une vraie famille à l'intérieur des Eucalanoida. Cf. Rhincalanidae
Subeucalaninae Geletin, 1976
Ref.: Geletin, 1976 (p.92)
Rem.: Cette sous-famille n'est pas retenue par Brodsky & al., 1983 (p.203).
1 G.: Subeucalanus
(1) Eucalanus Dana, 1852 ( Eucalaninae )
Ref.: Brady, 1883 (p.37); Giesbrecht, 1892 (p.46, 131); Giesbrecht & Schmeil, 1898 (p.19, spp. Key); Wheeler, 1901 (p.166); Esterly, 1905 (p.130, clé spp.); van Breemen, 1908 a (p.13); A. Scott, 1909 (p.19); Sewell, 1929 (p.46, Rem.); Wilson, 1932 a (p.30, clé spp.); Rose, 1933 a (p.66); Mori, 1937 (1964) (p.20, spp. Key); Dakin & Colefax, 1940 (p.95, spp. key); Vervoort, 1946 (p.83); Sewell, 1947 (p.36); Davis, 1949 (p.15); Brodsky, 1950 (1967) (p.99, spp. Key); Farran & Vervoort, 1951 b (n°34, p.3); Carvalho, 1952 a (p.139); Ramirez, 1966 (p.8); Fleminger, 1973 (p.966, Rev., spp. Key); Andronov, 1973 b (p.1720, Rem.); Geletin, 1976 (p.91, Rev.); Razouls, 1982 (p.90); Gardner & Szabo, 1982 (p.151); Brodsky & al., 1983 (p.202, clé spp., Rem.); Van der Spoel & Heyman, 1983 (p.147); Zheng Zhong & al., 1984 (1989) (p.229, clé spp.); Mauchline, 1988 (p.723); Hiromi, 1987 (p.150); Ferrari, 1992 (p.392, tab.3); Razouls, 1993 (p.309); Bradford-Grieve, 1994 (p.74, Def.); Chihara & Murano, 1997 (p.787, spp. Key); Mauchline, 1998 (p.77); Boxshall & Halsey, 2004 (p.116); Vives & Shmeleva, 2007 (p.870)
Rem.: Type: Calanus elongatus (Dana,1849).
Not all authors follow Geletin (1976) in his defenition of the genera Pareucalanus and Subeucalanus (Brodsky & al., 1983, p.203).
Fleminger (1973) splits up 17 spp. into 4 groups: 'subtenuis', 'pileatus', 'elongatus', 'attenuatus'.
17 spp. + 1 debated (E. pseudattenuatus ) + 2 to specify (E. muticus , E. peruanus ) + 3 very doubtful.
Bradford-Grieve (1994) rehabilitates the two genera Pareucalanus and Subeucalanus.
The genus Eucalanus comprises 6 spp. + 1 doubtful.

Diagnosis from Bradford-Grieve (1994, p.74) :
- As in the family definition.
- Urosome female 4-segmented, and posterior borders of pedigers 2-4 and urosome segment 1 without spines.
- Integumental organs located at Mx1 and Mx2 sites vary from 8-18 in number are not arranged in a rectangular pattern, posterior sets more lateral than the anterior sets. Mxp sites are in 4 or more pairs. Pediger segments 2 and 3 with 1 or 2 lateral sites ; pediger segment 4 lacking paired sites lying outside the dorsal quadrangle of hair sensilla ; abdominal segment 1-2 with at least 2 dorsal perforations.
- A2 endopod elongate, exopodal segments 1 and 2 separate.
- Md palp basis and exopod elongate ; endopod small, inserted just beyond midlength on basis which also bears 1 or 3 setae.
- Mx1 with 3 inner lobes bearing setae.
- P1 with a 2-segmented endopod and 3-segmented exopod.
P5 absent in female.
- P5 uniramous on both sides in male.
Remarks on dimensions and sex ratio:
The mean female size is 6.244 mm (n= 13; SD = 1.4723) and the mean male size is 4.732 mm (n = 12; SD = 0.9092). The size ratio (Male: Female) is 0.78 (n = 7; SD = 0.1174). The sex ratio (Female: Male) is 1.
(2) Pareucalanus Geletin, 1976 ( Eucalaninae )
Ref.: Geletin, 1976 (p.91); Razouls, 1982 (p.82, 98); 1993 (p.309); Bradford-Grieve, (p.80, Déf.); Bradford-Grieve & al., 1999 (p.912, pp. Key); Boxshall & Halsey, 2004 (p.116); Mulyadi, 2004 (p.106, Def.); Vives & Shmeleva, 2007 (p.872)
Rem.: type: Paracalanus attenuatus Dana,1849. Total: 6 spp. + 1 undetermined.
Brodsky & al., 1983 (p.203) do not acknowledge the creation of this genus. The genus is confirmed by Bradford-Grieve (1994)

Diagnosis from Bradford-Grieve (1994, p.80) :
- As in the family definition.
- Urosome female 3-segmented, and posterior borders of pedigers 2-4 and urosome segment 1 without spines.
- Integumental organs located at M11 and Mx2 with a total of 10 sites forming a rectangle and arranged anterior to posterior in 2 bilateral sets of 1 pair, 1 single and 1 pair of sites ; Mxp site has 6 pairs ; pedigers segments 2 and 3 in lateral view with 3 sites ; pediger segment 4 lacking paired sites lying outside of the dorsal quadrangle of hair sensilla ; urosome segment 4-5 with at least 3 dorsal sites.
- A2 endopod elongate, exopod segments 1 and 2 separate.
- Md palp basis and exopod elongate ; endopod small, inserted just at midlength on basis which also bears 2 or 4 setae.
- Mx1 with 3 inner lobes bearing setae.
- P1 with a 2-segmented endopod and 3-segmented exopod.
- P5 female absent.
- P5 male uniramous on both sides with right leg not extending beyond the distal part of left leg segment 2.
Remarks on dimensions and sex ratio:
The mean female size is 5.898 mm (n = 11; SD = 1.4656), and the mean male size 4.927 mm (n = 11; SD = 1.3540). The size ratio (Male: Female) is 0.835. The sex ratio is 1.
Rhincalanus Dana, 1852
Syn.: Calanus (part.) : Dana,1846
Ref.: Giesbrecht, 1892 (p.47, 152); Boxshall & Halsey, 2004 (p.116: Rem.)
Rem.: Cf. Rhincalanidae.
(3) Subeucalanus Geletin, 1976
Ref.: Geletin, 1976 (p.91); Razouls, 1982 (p.98); 1993 (p.309); Bradford-Grieve, 1994 (p.87); Bradford-Grieve & al., 1999 (p.912, spp. Key); Boxshall & Halsey, 2004 (p.116); Mulyadi, 2004 (p.106, Def.): Vives & Shmeleva, 2007 (p.874, spp. Key); Prusova & al., 2001 (p.51: Rem.)
Rem.: type: Eucalanus subtenuis Giesbrecht,1888. Total: 9 spp. + 3 undet.
Brodsky & al. (1983, p.203) do not acknowledge the creation of this genus. Bradford-Grieve (1994) affirms the validity of the genus.

Diagnosis from Bradford-Grieve (1994, p.69) :
- As in the family definition.
- Female urosome 3-segmented, and posterior borders of pedigers 2-4 and urosome segment 1 without spines.
- Integumental organs located at Mxp site in 2 pairs ; pediger segments 3 and 4 with a single mid-dorsal site lying between the anterior 2 hair sensilla ; pediger segment 4 with 1 pair of sites lateral to the quadrangle formed by 4 hair sensilla.
- A2 endopod elongate, exopod segments 1 and 2 fused.
- Md palp basis and exopod elongate ; endopod small, inserted on the terminal one third or quarter of basis which also bears 2 or 3 setae.
- Mx1 with 2 inner lobes bearing setae, 1 lobe absent.
- P1 with a 2-segmented endopod and 3-segmented exopod.
- P5 female absent.
- P5 male uniramous with right leg absent.
Remarks on dimensions and sex ratio:
The mean female size is 2.774 mm (n = 9; SD = 0.7719), and the mean size male 2.287 mm (n = 8; Sd = 0.6007). The size ratio (male: female) is 0.823; but if the species S. denudatus is excluded from the calcul because the great size difference between male and femelle (ratio male: female = 0.490), we obtain a ratio of 0.9498. The sex ratio is 1.125

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Razouls C., Desreumaux N., Kouwenberg J. and de Bovée F., 2005-2024. - Biodiversity of Marine Planktonic Copepods (morphology, geographical distribution and biological data). Sorbonne University, CNRS. Available at http://copepodes.obs-banyuls.fr/en [Accessed March 29, 2024]

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