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List species and varieties by family |
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| Megacalanidae Sewell, 1947 ( Calanoidea ) | |
| | | (1) Bathycalanus Sars, 1905 | |
| | Rem.: | type: Bathycalanus richardi Sars,1905. 10 spp. + 1 uncertained.
Diagnosis after H.B. Michel (1994, p.188) :
1 - Exopod of P1 with 0, 0, 2 outer marginal setae on the three, occasionally two, segments: 2nd inner lobe of Mx1 lacks setae; setae on terminal part of Mx2 ribbon-like and densely covered with spinules.
Diagnosis from Bradford-Grieve (1994, p.18) :
- Rostrum composed of two stout sausage-shaped, not tapering processes, located on a conspicuously thickened base of the anterior head.
- Mx1 inner lobes 1, 2 and 3 with 11, 0, and 2 setae respectively; basipod with 2-3 setae; endopod segments 1, 2, and 3 with 1-2, 1 and 4-6 setae respectively; exopod with 11 setae; outer lobes 2 and 1 with 1 and 9 setae respectively.
- Mx2 with setae very long, curved at the tips, each provided with a dense row of hairs on the inner margin forming a ribbon-like lemella.
Mxp with the 4 terminal segments with reduced setation compared to the other two genera.
- Male P5 very similar to that of Megacalanus |
| | [1] Bathycalanus adornatus Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F) [Figs] | |
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| | [2] Bathycalanus bradyi (Wolfenden, 1905) (F,M) [Figs] | |
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| | [3] Bathycalanus bucklinae Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F) [Figs] | |
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| | [4] Bathycalanus dentatus Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F,M) [Figs] | |
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| | Bathycalanus eltaninae Björnberg, 1968 (F,M) | | Ref.: | Björnberg, 1968 (p.75, figs.F,M) | | Rem.: | Cf. Elacalanus eltaninae (Björnberg, 1968) |
| | [5] Bathycalanus eximius Markhaseva, 1983 (F) [Figs] | |
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| | Bathycalanus maximus Wolfenden, 1911 (F,M) | | Ref.: | Wolfenden, 1911(p.198, Figs.F,M); Vervoort, 1946 (p.63, 71) | | Rem.: | Cf. Bathycalanus bradyi |
| | Bathycalanus inflatus Björnberg, 1958 (F) | | Ref.: | Björnberg, 1968 (p.81, figs.F) | | Rem.: | Cf. Elacalanus inflatus (Björnberg, 1968) |
| | [6] Bathycalanus milleri Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F,M) [Figs] | |
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| | Bathycalanus princeps Brady, 1883 (F,M) | | Ref.: | Farran, 1939 (p.360, Rem.); Grice & Hulsemann, 1967 (p.13) | | Rem.: | Cf. Elacalanus princeps (Brady,1883) |
| | [7] Bathycalanus pustulosus Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F) [Figs] | |
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| | [8] Bathycalanus richardi Sars, 1905 (F,M) [Figs] | |
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| | Bathycalanus rigidus Sars, 1920 (F) | | Ref.: | Sars, 1920 c (p.2); 1925 (p.19, figs.F); Farran, 1939 (p.355, Rem.); Jespersen, 1934 (p.46); Wilson, 1942 a (p.172); Vervoort, 1946 (p.70) | | Rem.: | Cf. Bathycalanus princeps (Brady,1883) |
| | Bathycalanus sverdrupi Johnson, 1958 (F) | | Ref.: | M. W. Johnson, 1958 a (p.258, Descr.F, figs.F) | | Rem.: | Cf. Elacalanus sverdrupi (Johnson, 1958) |
| | [9] Bathycalanus tumidus Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F) [Figs] | |
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| | [10] Bathycalanus unicornis Björnberg, 1968 (F,M) [Figs] | |
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| | Bathycalanus sp. Sewell, 1947 (Juv.) | | Ref.: | Sewell, 1947 (p.34, figs. juv.5) | | Loc: | Mer Arabe | | Lg.: | (11) 7,9 | | Rem.: | ? Cf. Bathycalanus bradyi | | | (2) Bradycalanus A. Scott, 1909 | |
| | Rem.: | Type: Bradycalanus typicus A. Scott,1909. Type locality: 0°17.6' S, 129°14.5' E. Total: 4 spp. (of which 1 doubtful).
For Owre & Foyo (1964) this genus is synonymous with Megacalanus, a position that will be given up by the authors in 1967.
Diagnosis after H.B. Michel (1994, p.187) :
1 - Exopod of P1 with 1, 1, 2 outer marginal setae on the three segments; 2nd inner lobe of Mx1 with 2-5 setae.
2 - A1 not so long that in Megacalanus; endopod 2 of Mx1 with 2 setae, 1 extremely small; terminal setae of Mx2 scythe-like, with rows of short, stiff hairs; rostrum extends into acuminate prongs of varying length and breadth.
Diagnosis from Bradford-Grieve (1994, p.20) :
- The genus resembles Megacalanus but:
- A1 apparently not modified for grasping in the male.
- Setae on the terminal parts of Mx2 scythe-like and densely ciliated.
- Mx1 with inner lobes 1, 2, and 3 with 13, 4, and 2 setae respectively; basipod with 4 setae; endopod segments 1, 2 and 3 with 2, 1, and 5 setae respectively; exopod with 11 setae; outer lobes 2 and 1 with 1 and 9 setae respectively.
- P1 basipod segment 2 without a hooked spine.
- Male left P5 exopod segment 2 with 1 simple spine without hairs on the inner border.
Differential diagnosis after Bradford-Grieve & al. (2017, p.51):
- Anterior head rounded or crested.
- Rostral filaments slender and tapering to point.
- Female A1 ancestral segment XXIII with aesthetasc. Dorsal surface of ancestral segments I-V each with small hair sensillum without maculae cribrosae.
- Right male A1 ancestral segments XXI-XXIII fused, segments XIX, XX and XXI with gripping elements.
- A2 Exopod ancestral segments I-III each bearing relatively well-developed seta each longer than its segment, or shorter than its segment (Bradycalanus abyssicolus), seta on segment IV extending short of distal border of exopod. Bathycalanus); endopod segment 1 with 4 setae, endopod segment 2 with 9 large setae and 2 vestigial setae
- mx1 praecoxal arthrite with 4 posterior surface setae, coxal endite with 4 + 1 small or 2 setae (Bradycalanus abyssicolus), basal endites 1 and 2 with 2, 4 setae, respectively; endopod segments 1-3 usually with 2, 1 + 1 vestigial, 5 + 1 small posterior surface setae, respectively.
- Mx2 relatively enlarged compared with other megacalanidae; longest setae extend as far as mid labrum and only setae of praecoxal endites 1 and 2 and coxal endite 1 with auxilliary setules; proximal inner seta of endopod segment 2 vestigial, convex border of distal inner seta of endopod segment 2 naked.
- Mxp not enlarged compared with other megacalanids, longest setae extend to rostrum, endopod segments 3-5 with variable numbers of setae poorly developed;
- P1 basis without hooked process, exopod segments 1 and 2 each bearing distolateral articulated spine, segment 3 with 2 outer border spines. Maculae cribrosae absent.
- Male P5 specialised seta on left leg tapering evenly, bordered by very long setules. |
| | [1] Bradycalanus abyssicolus Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F) [Figs] | |
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| | [2] Bradycalanus enormis Björnberg, 1968 (F,M) [Figs] | |
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| | [3] Bradycalanus gigas Sewell, 1947 (F) [Figs] | |
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| | Bradycalanus pseudotypicus Björnberg, 1968 (F) | | Rem.: | Cf. Bradycalanus typicus |
| | Bradycalanus pseudotypicus enormis Björnberg, 1968 (F,M) | | Ref.: | Björnberg, 1968 (p.85, figs.F,M); Michel, 1994 (p.182, Rem.) | | Loc: | Chili (Valdivia) | | Lg.: | (406) F: 17,5; 17; M: 13; {F: 17,00-17,50; M: 13,00} | | Rem.: | Cf. Bradycalanus enormis |
| | Bradycalanus sarsi Farran, 1939 (F) | | Rem.: | Cf. Bradycalanus typicus |
| | [4] Bradycalanus typicus A. Scott, 1909 (F, M) [Figs] | |
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| | Bradycalanus sp. Sewell, 1947 | | Ref.: | Sewell, 1947 (p.30: Descr.) | | Loc: | G. d'Aden | | Lg.: | (11) 7,2 | | Rem.: | Cf. ? Bradycalanus typicus | | | (3) Elenacalanus Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 | |
| | Rem.: | Type species: Heterocalanus medius Wolfenden, 1906. Total: 5 spp.
Mainly bathypelagic.
For Bradford-Grieve & al, (2017, p.132) Heterocalanus medius Wolfenden, 1906 is clearly a junior synonym of Calanus princeps Brady, 1883 since H. medius has a head crested and P1 exopodal segment 3 with 2 outer spines. The authors propose the new genus name Elenacalanus as replacement name for Heterocalanus in the sense of Wolfenden (1906) and designate H. medius Wolfenden (1906) as the type species, because the genus Heterocalanus T. Scott, 1894 ( in reality Pseudodiaptomus Herrick, 1889) was preoccupied.
Diagnosis after Bradford-Grieve, Blanco-Bercial & Boxshall (2017, p/129) :
As for Megacalanidae plus following character states:
- Anterior head usually rounded and without anterior spine-like processes but may be crested (E. princeps).
- Rostral filaments bluntly tapering.
- Genital double-somite bulbous, widest in dorsal view at about midlength, usually as wide as long.
- Posterolateral corners of pedigerous 5 usually bluntly triangular or rounded in E. sverdrupi.
- Female A1 ancestral segment XXIII with aesthetasc.
- Female and male ancestral segments I-V with hair sensillum on dorsal surface and adjacent macula cribrosa except sometimes on segment IV.
Right male A1 ancestral segments XIV-XV and XXI-XXIII fused,ancestral segments XIX, XX and XXI with fused gripping elements.
- A2 exopod ancestral segments I-III without seta, on ancestral segment IV seta absent (E. princeps) or, more usually, extending short of the distal end of exopod.
- Md gnathobase with ventral tooth set at right angles to main plane of gnathobase therefore appearing tapering, similar to other teeth; endopod segment 1 with 2 setae, endopod segment 2 with 9 setae.
- Mx1 with 2 or 3 posterior surface setae on praecoxal arthrite, coxal endite without setae, basal endites 1 and 2 with 2, 1-3 setae, respectively, endopod segments 1 and 2 with 1 seta each at most, endopod segment 3 with 4+1 small setae (often 4 in male), basal exite with or without seta, coxal epipodite usually with 7+2 proximal reduced setae (sometimes absent).
- Mx2: endopod with curled setae into semicircle distally but not completely curled on themselves, with row of fine, long dense setules along distal half of concave surface; long setae extend anteriorly as far as rostrum, none of setae with auxiliary spinules; 1 inner seta of endopodal segment 2 vestigial.
- Mxp endopod segments 2-6 with 4, 1, 1, 1 (outer seta absent), 2 long + 0-2 small setae (sometimes outer setae absent).
- P1 basis without anterodistal hook-like process ; exopod segments 1 and 2 without outer border spines; expopd segment 3 with 2 outer border spines.
- Setae on some male mouthparts reduced in size.
Remaks from Bradford-Grieve & al. (2017, p.131): The morphology-based cladistic analysis and the molecular-based analysis clearly identify two separate monophyletic clades within what was previously called Bathycalanus. The name Bathycalanus is retained for the group of species, including the type species Bathycalanus richardi, with a maximal number of setae on the endopod of the Md, Mx1 and Mxp and 1 outer border spine on P1 exopodal segment 3. |
| | [1] Elenacalanus eltaninae (Björnberg, 1968) (F,M) [Figs] | |
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| | [2] Elenacalanus inflatus ((Björnberg, 1968)) (F ) [Figs] | |
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| | [3] Elenacalanus princeps (Brady, 1883) (F,M) [Figs] | |
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| | [4] Elenacalanus sverdrupi (Johnson, 1958) (F) [Figs] | |
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| | [5] Elenacalanus tageae Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F,M) [Figs] | |
| | | Heterocalanus Wolfenden, 1906 | |
| | Rem.: | Cf. Bathycalanus | | | | Heterocalanus medius Wolfenden, 1906 (F) | | Ref.: | Wolfenden, 1906 (p.27, Descr.F, figs.F); 1911 (p.201, figs.F) | | Rem.: | Cf. Bathycalanus princeps |
| | | Rem.: | Cf. Megacalanus | | | | Macrocalanus longicornis Sars, 1905 (F) | | Ref.: | Sars, 1905 b (p.7, Rem.F) | | Rem.: | Cf. Megacalanus princeps Wolfenden,1904, non Brady,1883 | | | | (4) Megacalanus Wolfenden, 1904 | |
| | Rem.: | Type species : Megacalanus princeps Wolfenden, 1904. For Vervoort (1946, p.48) Wolfenden's Megacalanus princeps represents the type of the genus Megacalanus and has priority over Macrocalanus longicornis Sars, 1905 (p.7). Total: 4 spp. + 1 variety (doubtful)..
Bradford-Grieve & al. (2017, p.29) note a great deal of confusion caused by the incomplete description of Calanus princeps by Brady (1883), . Also the generic diversity of the Megacalanidae was not recognised until A. Scott's (1909) description of Bradycalanus, a genus which is morphologically closer to Megacalanus than to Bathycalanus. Wolfenden (1906) recognised that the Brady's type Calanus princeps was not a Megacalanus (resemble Heterocalanus Wolfenden, 1906, as Heterocalanus medius. The authors cited agree with Wolfenden's (1906) concept of a separate genus to take C. princeps Brady (1883) in a new genus Elenacalanus.
Diagnosis after H.B. Michel (1994, p.187) :
1 - Exopod of P1 with 1, 1, 2 outer marginal setae on the three segments; 2nd inner lobe of Mx1 with 2-5 setae.
2 - A1 very long, nearly twice the body length, extending beyond the caudal rami by about 8 segments; endopod 2 of Mx1 with 4 setae; terminal setae of Mx2 armed with moderately spaced spinules; bifurcated rostrum extends into slender, tapering filaments.
Diagnosis after Bradford-Grieve (1994, p.17) :
- As for the family definition.
- Head and pedigerous segment 1 and pedigerous segments 4 and 5 separate.
- Rostrum composed of 2 sharp, slightly tapering spines.
- A1 twice as long as the metasome.
- Mx1 inner lobe 1 (praecoxal arthrite) with 14 marginal spines and setae including 4 on the posterior surface and 1 on the anterior surface; inner lobes 2 and 3 (coxal and basal endites) with 4 and 4 setae respectively; basis with 4 setae; endopod segments 1, 2 and 3 with 3, 4 and 7 setae respectively; exopod with 11 setae; outer lobe 2 with 1 seta; outer lobe 1 with 9 setae.
- Mx2 with setae of moderate length, sparsely ciliated.
- P1 with exopod segments 1, 2, and 3 each with 1, 1, and 2 outer edge spines and a reflexed hook on each basipod 2 (= basis) (absent in some varieties).
- Males have slight differences in rostrum and appendages up to P4.
- A1 male asymmetrical, similar to female on the left, but with larger aesthetascs, with segments 7-9 imperfectly separated; 24-segmented on the right, with segments 7-9, 12-13, and 17-18 fused, segments 17+18, 19, and 20 edge expanded slightly into a flange; segment 9 bearing a clavate seta.
- Male P5 almost symmetrical, distinguished from those of the female only by the reduction in inner edge setae on exopod segment 3 and in the hairy projection terminated in a fine whip on the inner edge of the left exopod segment 2.
Differencial diagnosis from Bradford-Grieve & al. (2017, p.26):
- Anterior head without spine-like processes, rostral filaments tapering to point.
- Female A1 ancestral segment XXIII without aesthetasc, and segments XIV-XVII have ventral surface tooth row.
- Female and male ancestral segments I-V each with very small hair sensillum on dorsal surface.
- Right male A1 ancsetral segments XIV-XV and XII-XXIII fused, at least segments XX and XXI with fused gripping elements.
- A2 exopod ancsetral segments I-IV bearing relatively well-developed setae on segments I-III each seta longer than its segment, on segment IV seta extends beyond distal border of exopod;.
- Md gnathobase with ventral tooth set at right angles to main plane of gnathobase therefore appears tapering andsimilar to other teeth; endopod segment 1 with 4 setae, endopod segment 2 with 9 large and 2 short proximal setae.
- Mx1 praecoxal arthrite with 4 posterior surface setae, coxal endite with 5 setae, basal endites 1 and 2 with 4 setaze each, endopod segment 2 with 4 setae.
- Mx2, modestely developed; longest setae extend as far as anterior labrum; most setae bear auxilliary setules.
Mxp mosestly developed, longest setae extending to rostrum, endopod segments 3-5 with proximal seta shortest.
- P1 basis usually with anterior inner surface bearing large hook-like process o,to which straight inner seta inserted, exopod srgments 1 and 2 each bear distolateral articulated spine , segment 3 with 2 outer border spines.
- Specialised seta on male left P5 with base greatly enlarged into outer border bulge then tapering into short terminal section bordered by long setules.
- Circular or oval distoanterior 'macula cribrosa' present between rostral points; at base of all antennular aesthetascs, associated with some dorsal surface hair sensilla on ancestral segments I-V depending on species, and distoposteriorly on ancsetral segment XXVIII; on Md gnathobase, proximally near insertion of basis; on Mx1 outer proximal surface of exopod; on Mxp inner surface of basis and endopod segment 5; on anterior surface of basis of legs 1-5 adjacent to insertion of exopod segment1; and P1 exopod segment 2. |
| | Megacalanus bradyi Wolfende, 1905 (F,M) | | Ref.: | Wolfenden, 1905 a (p.3, Descr.F,M, figs.F); Farran, 1939 (p.360), Vervoort, 1946 (p.71, Rem.) | | Rem.: | Cf. Bathycalanus bradyi |
| | [1] Megacalanus ericae Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F,M) [Figs] | |
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| | [2] Megacalanus frosti Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F,M) [Figs] | |
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| | Megacalanus longicornis Sars, 1905 (F,M) | | Rem.: | All the authors do not agree concerning the synonymy or not between M. longicornis and M. princeps (problem of rule of prority).
Cf. Megacalanus princeps Wolfenden, 1904 |
| | [3] Megacalanus ohmani Bradford-Grieve, Blanco-Bercial & Boxshall, 2017 (F,M) [Figs] | |
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| | [4] Megacalanus princeps Wolfenden, 1904 (F,M) [Figs] | |
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| | Megacalanus princeps Brady, 1883 (F) | | Syn.: | Calanus princeps Brady, 1883 (p.36, figs.F) | | Ref.: | Vervoort, 1946 (p.70) | | Rem.: | Cf. Bathycalanus princeps (Brady,1883) |
| | Megacalanus princeps inermis Sewell, 1947 (F) | | Ref.: | Sewell, 1947 (p.25, figs.F); Guérédrat, 1969 (p.65, Rem.); Michel, 1994 (p.179, Rem.); Miller, 2002 (p.140, Rem.) | | Loc: | Médit., Mer d'Oman, Pacif. tropical | | Lg.: | (11) F: 8,73; (85) F: 11; 10,5 | | Rem.: | Cf. Megacalanus longicornis |
| | Megacalanus princeps princeps Wolfenden, 1904 | | Syn.: | Megacalanus princeps Wolfenden,1904 (p.112) | | Ref.: | Guérédrat, 1969 (p.67) | | Loc: | Pacif. (équatorial W & tropical S) | | Rem.: | Cf. Megacalanus longicornis |
| | Megacalanus princeps varius Guérédrat, 1969 | | Ref.: | Guérédrat, 1969 (p.67) | | Loc: | Pacif. (équatorial W & tropical S) | | Rem.: | Cf. Megacalanus longicornis |
| | Megacalanus sarsi Farran, 1939 (F) | | Syn.: | Macrocalanus Sars,1905; Megacalanus princeps Sars,1925; Wilson, 1932 a; Rose,1933 a; Wilson,1936 | | Ref.: | Farran, 1939 (p.360); Vervoort, 1946 (p.62) | | Rem.: | Cf. Bradycalanus sarsi |
| | Megacalanus typicus A. Scott, 1909 (F) | | Syn.: | Bradycalanus typicus A. Scott,1909 (p.14) | | Ref.: | Vervoort, 1946 (p.58, Rem.); Owre & Foyo, 1964 (p.343) | | Rem.: | Cf. Bradycalanus typicus | |
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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 April 20, 2024] © copyright 2005-2024 Sorbonne University, CNRS
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