Calanoida ( Order )
    Diaptomoidea ( Superfamily )
Acartiidae Sars, 1903 ( Diaptomoidea )
Ref.: Sars, 1903 (p.147); Gurney, 1931 (p.85, 214); Rose, 1933 a (p.267); Brodsky, 1950 (1967) (p.83, 418); Gonzalez & Bowman, 1965 (p.255) ; Andronov, 1974 a (p.1005); Bradford, 1976 (p.162, Rev.); Björnberg & al.,1981 (p.661); Bowman & Abele, 1982 (p.9); Razouls, 1982 (p.596); Brodsky & al., 1983 (p.142,145); Zheng Zhong & al., 1984 (1989) (p.257); Sazhina, 1985 (p.119, N); Mauchline, 1988 (p.715, 740); Huys & Boxshall, 1991 (p.460); Yoo & al., 1991 (p.257, F,M Keys); Razouls, 1993 (p.307); Chihara & Murano, 1997 (p.669, G. & spp. Keys); Bradford-Grieve, 1999 (n°181, p.2); Barthélémy, 1999 a (p.56, 58, Rem.); Bradford-Grieve & al., 1999 (885, 901, 904, 961: clé des G.); Bradford-Grieve, 1999 b (p.213, Déf., Rem.); Ohtsuka & Huys, 2001 (p.461); Boxshall & Halsey, 2004 (p.12; 49; 50: Def.; p.52: Genera Key); Mulyadi, 2004 (p.136, Def.); Vives & Shmeleva, 2007 (p.401, Genera Key)
Bradford-Grieve J.M., (2002 onwards). Key to calanoid copepod families. Version 1 : 2 oct 2002.
Rem.: 5 G. : Acartia, Acartiella, Paracartia, Paralabidocera, Pteriacartia.
The genus Paracartia is not always accepted by all the authors.
For Barthélémy (1999 has, p. 59) the genital structures do not justify the creation of 6 sub-genera. The species of the various sub-genera are distributed in a variable way in 3 of 4 subgroups of species corresponding to four variants of the model acartiidae (cf. Fig.5, Plan 9). Account was already held by the absence of significant morphological criteria of distinction of the sub-genera, the characteristics of the genital area do not justify more to keep the sub-genera.
Family Acartiidae - Plate 1issued from : R.-M. Barthélémy in These Doc. Univ. Provence (Aix-Marseille I), 1999. [Fig.5]:
Female: Schéma of the genital area in modele Acartiidae.
A: external ventral view. gn = = genital slit; cp = copulatory pore; asterisk = cuticular shutter.
B: internal dorsal view. ap = apodemem; m2 = opercular muscle ; ed = egg-laying duct; gn = gonopore or gonoporal slit; gs = genital slit; sd = seminal ductsr = seminal receptacle; cp = copulatory pore.
C: Sectional drawings of genital structures. 1, 2, frontal cross-frontal; 3-4, cross-section
(1) Acartia Dana, 1846
Syn.: Dias Lilljeborg, 1853; Claus, 1863 (p.191); Brady, 1878 (p.50)
Ref.: Brady, 1883 (p.72); Giesbrecht, 1892 (p.75, 506); Giesbrecht & Schmeil, 1898 (p.150, clé spp.); Wheeler, 1901 (p.182); Sars,1903 (p.148); Esterly, 1905 (p.203); van Breemen,1908 a (p.155, clé spp.); A. Scott, 1909 (p.186); Steuer,1915 a (p.392, Rev.); 1923 (p.5); Esterly, 1924 (p.102); Früchtl, 1924 b (p.57); Steuer, 1929 (p.497); Gurney, 1931 a (p.216); Wilson, 1932 a (p.159, spp. Key); Sewell, 1932 (p.391); Rose, 1933 a (p.267, spp. Key); Mori, 1937 (1964) (p.100, spp. Key); Dakin & Colefax, 1940 (p.105, spp. Key); Farran, 1948 (n°12, p.3); Davis,1949 (p.64); Brodsky, 1950 (1967) (p.418, spp. Key); Carvalho, 1952 a (p.150); Tanaka, 1965 (p.386); Bowman, 1965 (p.149, Rem.: rostral filaments ); Gonzalez & Bowman, 1965 (p.255); Ramirez, 1966 (p.18); M.S. Wilson, 1966 (p.109, Rem.); Owre & Foyo, 1967 (p.100, clé spp.); Björnberg, 1972 (p.73-78: Nauplius); Bradford, 1976 (p.163 : Redef.); Razouls, 1982 (p.597); Gardner & Szabo, 1982 (p.413); Zheng Zhong & al. , 1984 (1989) (p.257, spp. Key); Mauchline, 1988 (p.715); Huys & Boxshall, 1991 (p.337, 340); Yoo & al., 1991 (Rev., p.255); Ferrari, 1992 (p.392, tab.3); Razouls, 1993 (p.307); Madhupratap & Haridas, 1994 (p.74, Rem. Subgenera); Chihara & Murano, 1997 (p.669, Subgenera & spp. Keys); Mauchline, 1998 (p.95); Bradford-Grieve, 1999 (n°181, p.3, spp. Key); Bradford-Grieve & al., 1999 (p.961, 962: spp. Key); Bradford-Grieve, 1999 b (p.214, Def., Rem.); Boxshall & Halsey, 2004 (p.52); Mulyadi, 2004 (p.137, Def., spp. Key in Indonesian waters); Vives & Shmeleva, 2007 (p.394, spp. Key)
Rem.: In a genus revision, Steuer in 1915 establishes two subdivisions based on the presence/absence of rostral filaments (Arostratae et Rostratae), but later Bowman (1965) when studying a population of A. liljeborgi demonstrates that the rostral filaments can be present or absent. The environmental conditions, notably the salinity, could be responsable for this fact.
It is probable that other morphological characters show a certain variability like the spinulation of the abdomen, the structure of the P5. Various anomalies have been shown by Brylinski (1984, p. 961)
The genus divides currently into 6 sub-genera: Acartia (Planktacartia ), Acanthacartia , Acartiura , Euacartia , Hypoacartia , Odontacartia, contested by Barthélémy (1999, p.868; 1999 a, p.59, scheme 9) after analysis of the genital structures. Among the majority of species, we can distinguish two main groups, based on the external morphology of the genital area on one hand, and the seminal duct morphology on the other. The first group includes ten species, viz., A. clausi, A. hudsonica, A. omorii, A. longiremis of the subgenus Acartiura, A. bifilosa, A. chilkaensis, A. italica, A. levequei, A. tonsa of the genus Acanthacartia, and A. lilljeborgi of the subgenus Odontacartia, exhibing a compact external area, with well-defined structure for each of them. All exhibit a very homogeneous configuration of their internal area with seminal ducts in characterisytic loop-like form. Accorfding to Steuer (1923), this loop is visible in ventral view after clearing the specimens. The duct seems less voluminous only in the subgenus Acanthacartia species. The similitude in the organization of the genital structures af all these species confirms the closerelationship between the subgenera Acartiura and Acanthacartia. Thus, the morphology of the genital complex does not justify the distinction of two subgenera. The second group comprises A. danae, A. negligens of the subgenus Acartia, A. amboinensis, A. japonica of the subgenus Odontacartia and A. southwelli of the subgenus Euacartia; these species offer a fragmented external area with ventrolateral genital slits, and seminal ducts, simple bend-shaped opening in the proximal zone of the egg-laying ducts, as in the precedent group. Therefore, the genital anatomy does not justify the distinction between the different subgenera.
56 spp. + 2 indet. + 10 doubtful.

Diagnosis after Bradford-Grieve (1999, p.3): As for the family, with the following additional characters.
Anal somite is without an anal operculum, as anus opens between last two urosomites into a dorsal grove on anal somite.
Caudal rami short, separated from anal somite.
A2 basis fused with endopodal segment 1, which is long and slender and bears 9 setae; exopod shorter than endopodal segment 1.
P1 exopodal segment 1 and 2 each with a long slender outer distal spine, and with 2 spines on exopodal segment 3.
Female P5 3-segmented , uniramous, with last segment modified into a long, slender spine.
Male P5 larger on right, exopodal segment 2 with a large inner lobe, and exopodal segment 3 in the form of a clasper.
Genus Acartia - Plate 1Issued from : H.Y. Soh, S.Y. Moon, E.O. Park & B.A.V. Maran in J. Crustacean Biol., 2013, 33 (5). [p.728, Fig.9].
Schematic illustration on the geographical distribution of the sibling species of subgenus Euacartia.
A. sarojus Madhupratap & Haridas, 1994 (black square), A. southwelli Sewell, 1914 (white square) and A. forticrusa Soh, Moon, Park & Maran, 2013 (black circle).
1- Gulf of Kutch, Gujarat: 2- Mandovi-Zurai estuaries, Goa; 3- Cochin Harbor, Kochi; 4- Tuticorin, Tamil Nadu; 5- Kilakarai, Tamil Nadu: 6- Porto Novo, Tamil Nadu; 7- Kakinada Bay, Andha Pradesh; 8- Chilka Lake, Orissa; 9, Sri Lanka; 10- Wenchang River, Hainan; 11- Pearl (Zhu) River, Gwantung; 12- Taiping Lake, Zhangtsui; 13- Tianjin estuary, Tianjin; 14- Beolgyo Sream, Jeollanamdo; 15- Seomjin River estuary, Jeollanamdo.
Number 1 is recorder from Mahdrupatap & Haridas (1994) and numbers 2-9 are recorded from Sewell (1914, 1924, 1932), Steuer (1915, 1923), Tranter & Abraham (1971), Madhupratap & Haridas (1994), and the prest study, respectively. Numbers 10-15 are cited from Shen & Song (1979) and the present study.
Acanthacartia Steuer, 1915
Ref.: Steuer, 1923 (p.21, 55); Gurney, 1931 a (p.217); Rose, 1933 a (p.268); Bradford, 1976 (p.164); Chihara & Murano, 1997 (p.670); Bradford-Grieve, 1999 (n° 181, p.5); Barthélémy, 1999 a (p.32); Bradford-Grieve, 1999 b (p.214, Def., Rem.)
Rem.: Subgenus of Acartia defined by Steuer, 1915 a (p.396)
Type: Acartia pietschmanni Pesta, 1911.

Diagnosis after Bradford-Grieve (1999, p.5): As for the genus, with the following additional characters.
Rostral filaments usually present.
Ovaries fused.
A1 segments mostly unspined.
Posterior prosomal borders rounded and either naked or armed with spines, sometimes large and acute.
Spermathecal canal appears to be looped only in ventral view.
P5 as in Acartiura although heavy spine on male left exopodal segments 2+3 is not simple spine found in Acartiura but may have one or more accessory spines arising from its base or be expanded .
Coastal species.
Acartia Dana, 1846 (part.)
Syn.: Planktacartia Steuer, 1915 a (p.397); Steuer, 1923 (p.34,55);
Gurney, 1931 a (p.217); Rose, 1933 a (p.268)
Ref.: M.S. Wilson, 1966 (p.109, Rem.); Bradford, 1976 (p.163); Chihara & Murano, 1997 (p.669); Bradford-Grieve, 1999 (n°181, p.6); Barthélémy, 1999 a (p.32); Bradford-Grieve, 1999 b (p.218, Déf., Rem.)
Rem.: Type species: Acartia negligens.

Diagnosis after Bradford-Grieve (1999, p.6): As for the genus, with the following additional characters.
Rostral filaments present.
Ovaries fused.
Posterior prosomal borders rounded or drawn out into a point.
Female P5 coxopodite with long plumose seta, terminal spine denticulate.
Right male P5 exopodal segment 1 with a distal appendage.
Indopacific and Atlantic species. Oceanic in all tropical and sudtropical seas.
Acartiura Steuer, 1915
Ref.: Steuer, 1923 (p.5); Gurney, 1931 a (p.217, 220); Rose, 1933 a (p.267); Bradford, 1976 (p.159, Rev., spp. Key); Chihara & Murano, 1997 (p.669, clé spp.); Bradford-Grieve, 1999 (n°181, p.3); Barthélémy, 1999 a (p.31); Bradford-Grieve, 1999 b (p.221, Def., Rem.)
Rem.: Subgenus of Acartia defined by Steuer, 1915 a (p.394); 1923 (p.5)
Type: Acartia clausii Giesbrecht,1889.

Diagnosis after Bradford-Grieve (1999, p.3): As for the genus, with the following additional characters.
Ovaries fused.
Last prosomite rounded but may bear spines.
Rostral filaments absent.
Caudal rami short and slightly asymmetrical, with right ramus longest.
Female genital double-somite with spermathecal canal looped when viewed both laterally and ventrally, and genital apertures close together on ventral surface.
Female P5 has a smooth terminal spine bearing some distal hairs on both sides and with an evenly bulbous base, endopod absent.
Male P5 has right basis with 2 inner ridges and an outer distal plumose seta, exopodal segment 1 (not in A. discaudata) has a distal inner lobe and a proximal inner spine, exopodal segment 2 has its inner lobe bearing 1-3 spines, exopodal segment 3 has a terminal spine, and inner edge spine and at least 3 transverse rows of outer spines. Left basis has an inner proximal ridge, postrior surface spines, and an outer distal plumose seta; exopodal segment 1 usually has some spinules; the exopodal segments 2+3 is shaped like a hand with 2 lateral distal opposing spines, anterior spine much heavier and thicker-walled than posterior spine, but both are simple; patches of spinules and hairs are proximal to anterior spine;
Mainly temperate to cold species in both hemispheres (Bradford, 1976).
Euacartia Steuer, 1915
Ref.: Gurney, 1931 a (p.217); Rose, 1933 a (p.268); Bradford, 1976 (p.176); Madhupratap & Haridas, 1994 (p.74); Barthélémy, 1999 a (p.33); Bradford-Grieve, 1999 b (p.226, Déf., Rem.: p.214); Soh & al., 2013 (p.718, Table 1, 2, fig.9: geographical distribution)
Rem.: Subgenus of Acartia defined by Steuer, 1915 a (p.394).
Type: Acartia southwelli Sewell, 1914. 3 spp.
Hypoacartia Steuer, 1915
Ref.: Steuer, 1923 (p.14,54); Gurney, 1931 a (p.217); Rose, 1933 a (p.268); Bradford, 1976 (p.163); Bradford-Grieve, 1999 (n°181, p.6); Bradford-Grieve, 1999 b (p.226, Déf.)
Rem.: Subgenus of Acartia defined by Steuer, 1915 a (p.396).
Type: Acartia macropus Cleve,1901.

Diagnosis after Bradford-Grieve (1999, p.6): As for the genus, with the following additional characters.
Rostral filaments present.
Ovaries paired.
Female posterior prosomal borders with asymmetrical lateral extensions.
Female P5 inner border with a broad, dentate terminal spine.
Male P5 superficially similar to that of A. discaudata.
Coastal species.
Odontacartia Steuer, 1915
Ref.: Steuer, 1923 (p.26, 55, 2 groupes: 'erythraea' & 'centrura'); Gurney, 1931 a (p.217); Rose, 1933 a (p.268); Bradford, 1976 (p.163); Ueda, 1986 (Rem., p.17,18); Chihara & Murano, 1997 (p.671); Barthélémy, 1999 a (p.33); Bradford-Grieve, 1999 b (p.226, Déf., Rem.)
Rem.: Subgenus of Acartia defined by Steuer, 1915 a (p.396).
Type : Acartia lilljeborgi Giesbrecht,1889.
Planktacartia Steuer, 1915
Rem.: Subgenus of Acartia according to Steuer, 1915 a (p.397). M. S. Wilson, 1966 (p.109) considers the maintenance of this subgenus to be incorrect. It should be denominated Acartia in conformity with the International Code of Nomenclature.
Type: Acartia negligens . Cf. Acartia
(2) Acartiella Sewell, 1914
Syn.: Acartiella Steuer, 1915 a (p.394); 1923 (p.12, 54); Rose, 1933 a (p.268); Wellershaus, 1969 (p.268, clé spp.)
Ref.: Sewell, 1914 a (p.245); 1919 (p.2, 18, clé spp.); Gurney, 1931 a (p.215, 216, 217); Sewell, 1932 (p.392); Bradford, 1976 a (p.162); Zheng Zhong & al., 1984 (1989) (p.259); Barthélémy, 1999 (p.868, Rem.); 1999 a (p.59, Rem.); Bradford-Grieve, 1999 b (p.227, Déf.); Boxshall & Halsey, 2004 (p.52)
Rem.: Genus defined by Sewell (1914) with for species type: Acartia kempi Sewell, 1914. Considered by various authors as a sub-genus or a new family.
For Barthélémy (1999 a, p. 59) this genus has to constitute to him only a family. The species present a genital area of diaptomoid type deprived by opercule and by receptacle seminal, which distinguishes them from all other species of the family of Acartiidae possessing a genital model of area characterized well by the constant presence of receptacles and pores copulators neighboring to gonopores. We can also observe the other morphological characters: the length of the branches of the caudal rami, the shape flattened by the antennula, the biramous P5. These characters differentiate all the species of the genus of the other Acartiidae, with the exception of Acartiella kempi, only species of the genus presenting an antennule similar to that of Acartia (Sewell, on 1918).
According to Barthélémy (1999, p.868) the morphology of the external genital area and the absence of seminal receptacles very clearly isolate species of the genus Acartiella Sewell (1914), the validity of which is thus confirmed. Bradford (1976) pointed out that Acartiella should remain in the Acartiidae family especially because the exopods of P2-P4 are devoid of articulated outer edge spines, as in the other Acartiidae.
11 spp.:
(3) Paracartia T. Scott, 1894
Syn.: Paracartia T. Scott, 1894 b (p.68); Giesbrecht, 1897 b (p.254); Steuer, 1915 a (p.394); 1923 (p.15, 54); Rose, 1933 a (p.268)
Ref.: Sars, 1904 b (p.3); 1919 (1921) (p.15); Gurney, 1931 a (p.217); Sewell, 1948 (p.378, Rem.); Gonzalez & Bowman, 1965 (p.255); Bradford, 1976 (p.162, 163); Dussart, 1982 (p.20); Razouls,1982 (p.619); Ferrari, 1992 (p.392, tab.3); Razouls, 1993 (p.307); Bradford-Grieve, 1999 (n°181, p.14); Barthélémy, 1999 (p.867, 869: Rem.); 1999 a (p.34, 59, Rem.); Bradford-Grieve, 1999 b (p.227, Déf.); Boxshall & Halsey, 2004 (p.52); Vives & Shmeleva, 2007 (p.433, spp. Key)
Rem.: Considered as a subgenus by T. Scott, Steuer, then later as a genus by Sars (1904b).
For Barthélémy (1999a, p. 59) the organization of the external genital area , separated, in very lateral side and posterior genital splits, could constitute an important characteristic of the genus.
Type: Acartia dubia T. Scott,1894.Total: 5 spp.:
(4) Paralabidocera Wolfenden, 1908
Ref.: Wolfenden, 1908 (p.26); Vervoort, 1951 (p.148, Rem.); Bradford, 1976 (p.163); Brodsky & Zvereva, 1976 a (p.233-34, Rem.); Razouls,1982 (p.620); 1993 (p.307); Mauchline, 1998 (p.95); Barthélémy, 1999 (p.869: Rem.); 1999 a (p.34); Bradford-Grieve, 1999 b (p.227, Def., Rem.); Boxshall & Halsey, 2004 (p.52)
Rem.: Type: Paracartia antarctica Thompson,1898. Total: 3 spp.
For Barthélémy (1999, p.869) the sole examined species presents a fragmented area with two lateral genital slits, but internally there is no seminal duct and the receptacle seems to open distally in the egg-laying duct. The presence of paired genital orifices situated on both side of the genital segment is a constant character of the genus. The three species of the genus therefore present genital structures clearly differing from those of the Pontellidae. Thus, in spite of the resemblance with the genus Labidocera (Pontellidae), the comparison with the genital structures in Pontellidae (Barthélémy & al., 1998) confirms that the genus Paralabidocera must be referred to Acartiidae.
(5) Pteriacartia Belmonte, 1998
Syn.: Acartia josephinae Crisafi, 1974 (p.6)
Ref.: Belmonte, 1998 (p.360); Bradford-Grieve, 1999 (n°181, p.15); Barthélémy, 1999 a (p.34); Boxshall & Halsey, 2004 (p.52); Vives & Shmeleva, 2007 (p.436)
Rem.: type: Acartia josephinae. 1 sp.:

 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., 2005-2015. - Diversity and Geographic Distribution of Marine Planktonic Copepods. Available at 
[Accessed January 27, 2015]

© copyright 2005-2015 CNRS, UPMC

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


Version française
English version



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