Phaennidae Family - Marine Planktonic Copepods

Calanoida ( Order )	details
Clausocalanoidea ( Superfamily )	details

Abbreviations or symbols

Phaennidae Sars, 1902 ( Clausocalanoidea )

Ref.:

Sars, 1902 (1903) (p.42); Gurney,1931 a (p.84); Rose, 1933 a (p.125); Brodsky, 1950 (1967) (p.83, 225); Mazza, 1967 (p.173); Bradford, 1973 (p.137, Déf.); Andronov, 1974 a (p.1005); Bowman & Abele, 1982 (p.9); Razouls, 1982 (p.279); 1993 (p.310); Brodsky & al., 1983 (p.144,146); Park, 1983 a (p.317, Déf.); Bradford & al., 1983 (p.58); Mauchline, 1988 (p.735, 740 : cuticular pores); Huys & Boxshall, 1991 (p.356); Chihara & Murano, 1997 (p.852); Bradford-Grieve & al., 1999 (p.880, 902, 904, 927, clé des G.); Ohtsuka & Huys, 2001 (p.461); Ohtsuka & al., 2003 (p.53, 62: Rem.); Boxshall & Halsey, 2004 (p.15; 49; 158: Def.; p.160: key of genera); Ferrari & Markhaseva, 2005 (p.45, Rem.); Vives & Shmeleva, 2007 (p.678, part. Key G.); 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.:

8 G + 1 G.: Brachycalanus, Cephalophanes, Cornucalanus, Onchocalanus, Phaenna, Phaennocalanus, Talacalanus, Xanthocalanus.
Schulz (1998, p.42, 48), Vyshkvartzeva (2000, p.217) and Ohtsuka & al., 2003 (p.61, 62) regard the genus Xantharus as belonging to the Scolecitrichidae family. The belonging of this genus to this family is discussed by Schulz & Kwasniewski (2004, p.158).
Markhaseva & Semenova (2005) include provisionally the genus Kirnesius in this family.

The types of setae on the ramus of Mx2 include this family in the 'Bradfordian' grioup of families.

(1) Brachycalanus Farran, 1905

Species, varieties and details

Ref.:

Farran, 1905 (p.41); van Breemen, 1908 a (p.66); A. Scott, 1909 (p.81); Sewell, 1929 (p.176); Wilson, 1932 a (p.75); Rose, 1933 a (p.139); Tanaka, 1960 a (p.102); Bradford, 1973 (p.138); Campaner, 1978 a (p.976, Redef.); Razouls, 1982 (p.279); Bradford & al., 1983 (p.60, Def.); Othman & Greenwood, 1988 a (p.356, Rem.); Razouls, 1993 (p.310); Mauchline, 1998 (p.84: F); Ohtsuka & al., 2003 (p.62: Rem.); Bradford-Grieve, 2004 (p.287); Boxshall & Halsey, 2004 (p.161)

Rem.:

Redefinition after Campaner (1978 a, p.976) (Female).
Body robust and oval dorsally.
Cephalosome and thoracic segment 1 separated, thoracic segments 4 and 5 complete or incompletely separated or totally fused.
Rostrum short with 2 filaments.
Abdomen 4-segmented, densely covered with scale-like spines (anal segment extremely reduced).
A1 short, 24-segmented.
Exopod of A2 little greater than endopd.
Inner lobe 1 and 3 of Mx1 with respectively 2 and 4 setae.
Endopod of Mx2 with 1 vermiform seta and ( ?5) 7-8 brush-like setae.
The dorsal and dorsal surfaces of basis 1-2, and especially of exopod and endopod, from P1-P5 densely covered with scale-like spines.
P5 with 3 segments, last one with 2 lateral and 2 terminal aculeate spines.

Type-species : Wolfenden, 1904.
Campaner (1978 a, p.976) disagrees with Grice (1972), who placed B. minutus in this genus based mostly on the absence of rostral filaments.
Xanthocalanus ordinarius Grice (1972) is doubtless a new species of Brachycalanus.
Since B. gigas A. Scott, 1909, was definitely removed to the genus Lophothrix, the genus hitherto includes 3 species : B. atlanticus (Wolfenden, 1904), B. ordinarius (Grice, 1972), B. bjornbergae Campaner, 1978.
B. minutus Grice, 1972, is a species incertae sedis.

7 spp. (+1 doubtful) + 1 unident.

Remarks on dimensions and sex ratio:

The mean female size is 2,504 mm (n= 6; S= 0,906; Cv= 0,362). No male has been observed yet.

(2) Cephalophanes Sars, 1907

Species, varieties and details

Ref.:

Sars, 1907 a (p.15); Wolfenden, 1911 (p.283); Sars, 1925 (p.154); Steuer, 1926 b (p.214); Rose, 1933 a (p.141); Vervoort, 1957 (p.93, Rem.); Tanaka, 1960 a (p.124); Bradford, 1973 (p.138); Razouls, 1982 (p.280); Park, 1983 a (p.319, Déf.); Bradford & al., 1983 (p.60, Déf.); Mauchline, 1988 (p.735: pores cuticulaires); Razouls, 1993 (p.310); Chihara & Murano, 1997 (p.852); Boxshall & Halsey, 2004 (p.160); Vives & Shmeleva, 2007 (p.679, spp. Key)

Rem.:

Type: Cephalophanes refulgens Sars, 1907. 3 spp.

Remarks on dimensions and sex ratio:

The mean female size is 4,417 mm (n= 3; S= 0,255; Cv= 0,058) and the mean male size is 3,450 mm (n= 2; S= 0,534; Cv= 0,155). The size ratio (M/F) is 0,767 ou 76,7 % (n= 2; S= 0,126; Cv= 0,164)

(3) Cornucalanus Wolfenden, 1905

Species, varieties and details

Ref.:

Wolfenden, 1905 a (p.20); van Breemen, 1908 a (p.222); A. Scott, 1909 (p.83); Wolfenden, 1911 (p.281); Sars, 1925 (p.151); Sewell, 1929 (p.176); Wilson, 1932 a (p.74); Rose, 1933 a (p.138); Brodsky, 1950 (1967) (p.236); Vervoort, 1957 (p.90, Rem.); Tanaka, 1960 a (p.121); Razouls, 1982 (p.281); Bradford & al., 1983 (p.61, Déf.); Park, 1983 a (p.351, Déf.); Mauchline, 1988 (735: pores cuticulaires); Razouls, 1993 (p.310); Mauchline, 1998 (p.81: M; p.82: F); Bradford-Grieve & al., 1999 (p.929, clé spp.); Boxshall & Halsey, 2004 (p.160); Vives & Shmeleva, 2007 (p.682, spp. Key)

Rem.:

6 spp. + 1 unident.

Remarks on dimensions and sex ratio:

The mean female size is 6,755 mm (n= 6; S= 1,667; Cv= 0,247) and the mean male size is 6,727 mm (n= 3; S= 0,656; Cv= 0,098). The size ratio (M/F) is 0,838 ou 83,8 % (n= 3; S= 0,064; Cv= 0,076)

(4) Kirnesius Markhaseva & Semenova, 2005

Species, varieties and details

Ref.:

Markhaseva & Semenova, 2005 (p.40); Markhaseva & Schulz, 2009 (p.22, Rem.)

Rem.:

For Markhaseva & Schulz (2009, p.22) the family placement in Phaennidae is under review.

(5) Onchocalanus Sars, 1905

Species, varieties and details

Ref.:

Sars, 1905 b (p.19); van Breemen, 1908 a (p.65); A. Scott, 1909 (p.82); Wolfenden, 1911 (p.275); Sars, 1925 (p.144); Sewell, 1929 (p.176); Rose, 1933 a (p.136, clé spp.); Vervoort, 1950 a (p.3, clé spp. F); Brodsky, 1950 (1967) (p.234); Tanaka, 1960 a (p.115); Bradford, 1973 (p.136, 138); Razouls, 1982 (p.283); Gardner & Szabo, 1982 (p.269); Bradford & al., 1983 (p.61, Déf., Rem,.p.66); Park, 1983 a (p.327, Déf., clé des spp.); Mauchline, 1988 (p.735: pores cuticulaires); Razouls, 1993 (p.310); Chihara & Murano, 1997 (p.853); Mauchline, 1998 (p.81: M; p.82: F); Bradford-Grieve & al., 1999 (p.929, clé spp.); Boxshall & Halsey, 2004 (p.160, 161); Vives & Shmeleva, 2007 (p.689, spp. Key)

Rem.:

type: Onchocalanus trigoniceps Sars,1905. 10 spp.

Remarks on dimensions and sex ratio:

The mean female size is 6,558 mm (n= 9; S= 1,314; Cv= 0,200) and the mean male size is 5,943 mm (n= 8; S= 1,536; Cv= 0,258). The size ratio (M/F) is 0,888 or 88,8 % (n= 7; S= 0,089; Cv= 0,100). The sex-ratio (F/M) est de 1,12.

(6) Phaenna Claus, 1863

Species, varieties and details

Ref.:

Claus, 1863 (p.188); Giesbrecht, 1892 (p.57, 293); Giesbrecht & Schmeil, 1898 (p.50); van Breemen, 1908 a (p.55); A. Scott, 1909 (p.79); Rose, 1933 a (p.126); Mori, 1937 (1964) (p.54); Bradford, 1973 (p.138); Razouls, 1982 (p.286); Park, 1983 a (p.318, Déf.); Bradford & al., 1983 (p.67, Déf.); Mauchline, 1988 (p.734: pores cuticulaires); Razouls, 1993 (p.310); Mauchline, 1998 (p.78); Boxshall & Halsey, 2004 (p.160); Mulyadi, 2004, Def.): Vives & Shmeleva, 2007 (p.687); Markhaseva & Renz, 2011 (p.68, Rem.)

Rem.:

3 spp. (of which 2 doubtful)

(7) Phaennocalanus Markhaseva, 2002

Species, varieties and details

Ref.:

Markhaseva, 2002 (p.312); Boxshall & Halsey, 2004 (p.161)

Rem.:

1 sp.

(8) Talacalanus Wolfenden, 1911

Species, varieties and details

Ref.:

Wolfenden, 1911 (p.279); Campaner, 1978 a (p.976, Rev.); Razouls, 1982 (p.297); Park, 1983 a (p.327, Rem.); Bradford & al.,1983 (p.70, Rem.); Mauchline, 1988 (p.735, Rem.); Razouls, 1993 (p.310); Mauchline, 1998 (p.81: M; p.82: F); Bradford-Grieve & al., 1999 (p.928: Rem.); Boxshall & Halsey, 2004 (p.161)

Rem.:

type: Xanthocalanus greeni Farran,1905.
Genus maintained (Campaner, 1978 a; Bradford & al., 1983) or not according to the authors: Sars, 1925, p.134; Park, 1983 a et Tanaka & Omori, 1992 (p.268), very close to Xanthocalanus.
Redefinition after Campaner (1978, p.976) :
Body lengthened (about 6.0 mm or greater).
Cephalosome and thoracic segment 1 separated, thoracic segments 4 and 5 fused.
Anal segment strongly reduced.
A1 with 34 segments (with few and short setae), reaching about the body length.
Endopod and exopod of Md palp with nearly the same length, Basis 1 + 2 broad, with 3 setae.
Inner lobe of Mx1 very developed, with razor-like type setae in female and fleshy-type (flagellum-shaped from the middle to the tip) setae in male, inner lobe 3 with 3 setae.
Endopod of female Mx2 usually with 2 vermiform setae + 6brush-like setae (1 short and broad) ; endopod of male with brush-like setae forming a bud-like structure, and with ( ?) some vermiform setae.
Endopod of Mxp with razor-like type setae.
Endopod of P2-P4 with 1 or 2 groups of scale-like spines.
Female P5 with 3 segments, 3rd segment relatively short with 1 terminal plus 1 or 2 subterminal spines ; male P5 very asymmetrical in length, both with rudimentary endopod.

The genus Talacalanus differs from the genus Xanthocalanus mostly by :
The setation on Mx1 and endopod setation of Mxp, the relatively great body length.
Talacalanus specimen, an aspect which does not correspond to that figured by Wolfenden (1911, pl.31, fig.11).

Type-species : Xanthocalanus greeni Farran, 1905.
Other species :
Talacalanus calaminus Wolfenden, 1911. ; X. species 2 Grice, 1972 ; X. maximus Brodsky, 1950 (redescribed by Tanaka & Omori, 1967) certainly belong to this genus.

2 spp. + 1 doubtful + 1 unident.

Xantharus Andronov, 1981

Ref.:

Andronov, 1981 (p.1719); Bradford & al., 1983 (p.60, 104, Rem.); Schulz, 1998 (p.42: Rem.: Scolecithrichidae); Vyshkvartzeva, 1999 (2000) (p.217: Rem: Scolecitrichidae); Ohtsuka & al., 2003 (p.61, 62: Rem.)

Rem.:

Cf. Scolecitrichidae

(9) Xanthocalanus Giesbrecht, 1892

Species, varieties and details

Syn.:

Amallophora (part.) T. Scott, 1894 b (p.54); Tanaka, 1960 a (p.102, Rem.); ? Talacalanus Wolfenden, 1911 (p.279); Campaner, 1978 a (p.976); Bradford & al., 1983 (p.70); Mauchline, 1988 (p.735); Park, 1983 a (p.327) considère le genre Talacalanus comme un synonyme.

Ref.:

Giesbrecht, 1892 (p.57, 286); Giesbrecht & Schmeil, 1898 (p.49, clé spp.); Sars, 1900 (p.48); 1902 (1903) (p.45); Esterly, 1906 a (p.68); van Breemen, 1908 a (p.57); A. Scott, 1909 (p.80); T. Scott, 1909 (p.125, Rem.); Wolfenden, 1911 (p.272); Sars, 1919 (1921) (p.2); 1925 (p.125); Sewell, 1929 (p.176); Wilson, 1932 a (p.68, clé spp.); Rose, 1933 a (p.127, clé spp.); Brodsky, 1950 (1967) (p.226, clé spp.); Vervoort, 1957 (p.95, Rem.); Tanaka, 1960 a (p.86); Bradford, 1973 (p.133, 139, Redef.); Campaner, 1978 a (p.968, Rev.); Razouls, 1982 (p.287); Park, 1983 a (p.325, Déf.F.); Bradford & al., 1983 (p.70 & suiv., Déf.); Mauchline, 1988 (p.735); Ferrari, 1992 (p.392, tab.3); Razouls, 1993 (p.310); Chihara & Murano, 1997 (p.854); Mauchline, 1998 (p.81: M; p.82, 84, 85: F); Bradford-Grieve & al., 1999 (p.929, clé spp.); Vyshkvartzeva, 2003 (p.45: Rem.); Bradford-Grieve, 2004 (p.640, Rem.); Boxshall & Halsey, 2004 (p.161); Vives & Shmeleva, 2007 (p.695, spp. Key)

Rem.:

type : Xanthocalanus agilis & minor Giesbrecht,1892.
Campaner (1978 a, p.968) classifies the species into 2 groups of which the types for each group are: X. agilis et X. minor .
‘’Xanthocalanus agilis Group’’
Characters :
Cephalosome and 1st thoracic segment usually separated, 4th and 5th thoracic segments separated or fused.
Female A1 usually with 34 segments.
Md palp with endopod and exopod nearly aqual in length.
Mx1 : the large setae of the inner lobe of peculiar type (aculeate-straigth or slightly curved setae, bearing sharp spinules and/or setules) ; inner lobe 3 with 3 or 4 setae.
Endopod of Mx2 with 2 vermiform-type setae and 5 brush-like type setae, sometimes with an extra short brush-like seta.
Endopodite de P2-P4 usually with 1 or 2 groups of large spines.
P5 female with 3 segments, the 2 nd segment not much wider than the 1 st or 3 rd segment, this last either relatively narrow, long and with 1 or 2 terminal plus 2 lateral spines, or short, broad and with 2 or 3 spines : male P5 very or strongly asymmetical in length.
List of species :
Type-species : Xanthocalanus agilis Giesbrecht, 1892
? X. subagilis Wolfenden, 1904.
Species definitely in this group : X. fallax Sars, 1921 (= X. borealis Sars, 1903, part.) ; X. pinguis Farran, 1905 ; X. profundus Sars, 1907. Species possibly in this group : X. mixtus Sars, 1920 ; X. tenuiserratus Wolfenden, 1911.
Doubtful species in this group (most of them related with the genus Cephalophanes Sars, 1907, excepting by the absence of the frontal lenses) : X. antarcticus Wolfenden, 1908 ; X. squamatus Farran, 1905 ; X. agilis Wolfenden, 1911 ; X. kurilensis Brodsky, 1950 ; X. pavlosky Brodsky, 1955. .

‘’Xanthocalanus minor Group’’
Characters differing from the ‘’X. agilis Group’’:
Thoracic segments 4 and 5 fused.
Inner lobe 3 of Mx1 usually with 4 setae.
Endopod of Mx2 with 1 longer and 1 shorter vermifom-type setae, 5 brush-like type setae, usually with an extra short but wider brush-like seta ;
Female P5 with 2 or 3 segments, the last narrowed from the middle to the tip and with 1 or 2 terminal and 2 lateral spines ; male P5 slightly asymmetrical in length, with rudimentary endopod or none.
List of species :
Type-species : Xanthocalanus minor Giesbrecht, 1892.
Species definitely in this group : X. borealis Sars, 1900 ( = X. hirtipes van Höffen, 1897) non X. borealis Sars, 1903 (see Sars, 1921, pp.2-4 ; Sars, 1924, pl.35) ; X. medius Tanaka, 1937 ; X. penicillatus Tanaka, 1960 ; X. legatus Tanaka, 1960, X. marlyae Campaner, 1978.
Species probably in this group : X. fragilis Aurivillius, 1898 ; X. amabilis Tanaka, 1960 ; X. muticus Sars, 1905.

50 spp. (of which 1 doubtful) + 4 unident.

Remarks on dimensions and sex ratio:

The mean female size is 3,358 mm (n= 33; S= 1,304; Cv= 0,388) and the mean male size is 3,406 mm (n= 22; S= 1,244; Cv= 0,365).The size ratio (M/F) is 0,933 ou 93,3 % (n= 11; S= 0,139; Cv= 0,149). The sex ratio (F/M) is 1,5.