Fiche d'espèce de Copépode
Calanoida ( Ordre )
    Calanoidea ( Superfamille )
        Calanidae ( Famille )
            Calanoides ( Genre )
Calanoides natalis  Brady, 1914   (F,M)
Syn.: Calanoides carinatus : Vervoort, 1963 (p.81); De Decker, 1984 (p.329); Schnack, 1989 (p.142, figs.); Ceballos & al., 2004 (p.739); Blanco-Bercial & al., 2006 (p.802)
Calanoides c.f. carinatus: Prusova & al., 2011 (p.29, figs.)
Ref.:
Brady, 1914 (p. 4, figs.F, M);
Viñas & al., 2015 (p.97, phylogeography, molecular index), Rem.; Bradford-Grieve & al., 2017 (p.816, F,M, table 1, , fig.2, phylogeny)
Espèce Calanoides natalis - Planche 1 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14). [p.817, Fig. 5].
Female: a-b, habitus (dorsal and lateral, respectively); c, A1; d, genital double-somite (lateral); e, same (dorsal); f, genital operculum (go) and left seminal receptacle (sr); g, caudal eamus (ventral); h, forehead (ventral).
Scale bars: 1.0 mm (a-c); 0.1 mm (d-h).

Nota:
- Anterior margin of forehead triangular in dorsal and lateral views.
- Rostrum extends into 2 long posteriorly directed filaments.
- Urosome 4-segmented.
- Genital double-somite symmetrical in dorsal view, widest part at anterior 1/4, bulges anteroventrally. Pair of seminal receptacles (sr) in lateral view slant anteroventral to posterodorsal at 18° relative to posterior border' of genital double-somite.
- Caudal rami seta I absent, setae II and III lateral, setae IV and V terminal, seta VI inner distal (seta V longest neing longer than urosome including caudal rami), seta VII inserted immediately anterior to seta VI; setules decorate inner border'.


Espèce Calanoides natalis - Planche 2 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14). [p.818, Fig. 6].
Female: a, A1 (ancestral segments I-XIX), b, A1 ( ancestral segments XX-XXVIII; c, A1 (ancestral segments X-XI; d, A1 (ancestral segments XXVII-VVVIII; e, A1 (segments I-XI; f, A1 (ancestral segment XXIII); g, A2.
a - aesthetacs; pa - pseudoantennulate seta, va - vestigial aesthetasc.
Scale bars = 0.1 mm.

Nota:
- A1 34-segmented, extending to about posterior border of 2nd urosomite; ancestral segments II-IV, X-XI, XXVII-XXVIII fused. Most setae are naked and of modified type (ms, weatherby & al., 1994; Lernz & al., 1996), aesthetascs (a) or vestigial aesthetascs (va), which are found on quadrithek females; 2 ventral-most setae on ancestral segment I and 1 posterior seta on each of ancestral segments XXIV-XXVI plumose; long setae on segments XXV and XXVI pseudo annulate (pa);

- A2 with separate coxa and basis; coxa with 1 long inner plumose seta; basis with 2 long inner setae each with rows of short setules; exopod slightly longer than endopod; endopod 2-segmented (although line of fusion between segment 2 and 3 visible on posterior surface and marked by row of small spinules; segment 1 with 2 inner setae and distal outer patch of outer setules, terminal segment with 8 + 7 setae; exopod with segments I and II fused, segments III-IV fused- fusion line visible; ancestral segments I-VIII each with long plumose seta, terminal segment IX-X with 1 + 3 terminal setae.


Espèce Calanoides natalis - Planche 3 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14). [p.820, Fig. 7].
Female: a, Md; b, Mx1; c, Mx2; d, Mxp.
Scale bar = 0.1 mm.

Nota:
- Md coxal gnathobase damaged with opaline teeth missing, largest tooth ventral, separate from adjacent 4 teeth by large gap; 4 small teeth follow and a longer dorsal tooth; lash-like element broken; basis with 4 inner setae; endopod 2-segmented, segment 1 with large inner lobe, 4 distal inner setae and outer distal setules, segment 2 with 11 terminal setae, 2 of them shorter and row of outer setules; exopod 5-segmented with 1, 1, 1, 2 setae.
- Mx1 praecoxal arthrite with 15 setae (4 of them on posterior surface, 1 on anterior surface) and 3 or 4large wide spines, and 2 to 5 small spines on posterior surface located midprowimally; coxal endite with 4 terminal setae, epipodite with 9 setae; basal endites 1 and 2 with 4 and 4 setae, respectively; exite with 1 seta; basis and endopod separate; endopod 2-segmented with segments 1 and 2 fused - fusion line visible, with 3, 4, 7 setae and 5 spinules on anterior surface; exopod with 11 setae.
- Mx2 praecoxal endites 1 and 2 with 6 + 1 small and 3 setae, respectively; endites 2 and 3 on coxa with 3 and 3 setae, respectively, epipodite with 1 long plumose seta; endite 5 of basis longest with 4 setae (one of them heavier); endopodal segment 1 with well-developed love bearing 2 short + 1 large setae, endopodal segments 2-4 with 2, 2, 3 setae, respectively; inner surfaces of endites 2-5 each with 1 seta that is shorter and more densely lined with long spinules than other setae; endopodal segments 2 and 3 with inner setae shorter than outer setae.
- Mxp directed ventrally so that setae on syncoxa and baisi directed into animal's midline; syncoxa (= a double segment derived from fusion of praecoxa and coxa) with 1, 2, 4 setae; basis with 3 setae; endopodal segment 1 partly separate bearing 2 setae, endopodal segments 2-6 with 4, 3, 2, 2 +1, 3 +1 setae, respectively.


Espèce Calanoides natalis - Planche 4 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14). [p.821, Fig. 8].
Female: a, P1 coxa, basis and endopod; b, P1 exopod; c, P2 (anterior surface); d, P3 (posterior surface); e, P4 (posterior surface); f, P5 (anterior surface).
Scale bar = 0.1 mm.

Nota: P5 extends beyo,d posterior border of caudal rami, with outer distal corner of endopodal segment 1 pointed; pore openi,gs on anterior surface at base of outer border spines.


Espèce Calanoides natalis - Planche 5 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14). [p.820].
Female: Armature formula of the swimming legs P1 to P5.
Roman numerals = spines, Arabic numerals = setae; outer border setation listed to the left in each group separated by '';''


Espèce Calanoides natalis - Planche 6 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14). [p.822, Fig. 9].
Male: a-b, habitus (dorsal and lateral, respectively); c, A1; d, forehead (lateral); e, left caudal ramus (dorsal); f, P5 (posterior view; R = right leg, L = left leg); g, left leg P5 exopod segment 3.
Scales: 1.0 mm (a-d); 0.1 mm (e--g).

Nota:
- Rostrum extends into 2 long, ventroposteriorly directed, tapering filaments that extend as far as labrum.
- Cephalic dorsal hump anterior border at about 1/3 the length of cephalosome.
- Urosome 5 free somites.
- Genital somite short.
- Urosomite 2 as long as wide.
- Caudal rami extend laterally; seta I absent, setae II and III lateral, setae IV and V terminal (seta V longest), seta VI on inner border, seta VII inserted at inner distal corner.
- A1 22-segmented, extends as far as caudal rami, symmetrical, with ancestral segments I-IV, V-VI, XXVII-XXVIII fused, IX-X partly fused on ventral surface, X-XI partly fused on posterodorsal surface. Aesthetascs larger than in female, doubled on segments III, V, VII, IX, XI. Most setae of modified (ms) type or aesthetascs (a); long setae on ancestral segments V, XXV, XXVI are of a 'pseudoannnulate' type (pa) (see Weatherby & al., 1994). The distal-most, non aesthetasc setae of segments X and XI foreshortened and clavate, resoectively (fig.10 d).
- P1 to P4 as in female.
- P5 asymmetrical. Left leg longest, extends beyond caudal rami; coxa and basis longer than those of right leg: when P5 extended straight, border between exopodal segments 2 and 3 distal to distal border of right leg exopodal segment 2; exopodal segment 2 length/width (C'/c') = 2.55; length of terminal spine of exopodal segment 3 relative to length of its segment ( A\"/B\") = 0.81; exopodal segment 3 length/width (F-b\") = 2.50; exopodal segment 3 with 5 patches of fine setules on anterior surface. Right leg endopod extends slightly short of distal border of exopodal segment 2 (C+D)/E = 1.11; length of exopodal segment 3 relative to terminal spine (B/A) = 1.60. See Table 4 for variability in the proportions of the male P5..


Espèce Calanoides natalis - Planche 7 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14). [p.823, Fig. 10].
Male: &, A1 (ancestral segments I-XVII; b, A1 (ancestral segments XVIII-XXVIII); c, A1 (ancestral segments XXVII-)XXVIII, setal numbers refer to numbering system of Weatherby & al., 1994); d, A1 (ancestral segments XX and XI); e, A2; f, Md; g, Mx1; h, Mx2; i, Mxp; J-k, syncoxae from Benguela Current.
Scale bars = 0.1 mm.

Nota:
A1 with ancestral segments I-IV, V-VI, XXVII-XXVIII fused, IX-X partly fused on ventral surface, X-XI partly fused on posterodorsal surface. Aesthetascs larger than in female, doubled on segments III, V, VII, IX, XI. Most setae of modified (ms) type or aesthetascs (a); long setae on ancestral segments V, XXV, XXVI are of a 'pseudoannnulate' type (pa) (see Weatherby & al., 1994). The distal-most, non aesthetasc setae of segments X and XI foreshortened and clavate, resoectively (fig.10 d). The proximal seta of ancestral segment III differs from other short mixed modality setae of most segments in that it is longer and with bluntly terminating tip.
- A2, Md, Mx1, Mx2 and Mxp sexually dimorphic with reduced setation.
- A2 exopod slightly longer than endopod; endopod 2-segmented
- Md coxal gnathobase very reduced with teeth scarcely developed; basis with 1 medium inner seta plus 1 vestigial seta and small process; endopod 2-segmented, segment 1 with inner lobe and 2 distal inner setae, segment 2 with 8 terminal setae; exopod 5-segmented with 1, 1, 1, 1, 2 setae, respectively.
- Mx1 praecoxal arthrite reduced with about 6 vestigial spines; coxal endite with 2 or 3 vestigial terminal setae, epipodite with 7 long setae; basal endite 1 with 3 or 4 reduced setae, basal sendite 2 with 4 reduced setae; exite withoit seta; basis and endopod separate; endopod 3-segmented, with 3, 3, 7 reduced setae; exopod with 10 + 1 vestogial setae.


Espèce Calanoides natalis - Planche 8 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14). [p.825, Table 4].
Proportions of key features of P5 male of Calanoides natalis from the Arabian Sea and off Namibia.

Setal formula:
Left leg: Coxa 0-0. Basis 1-0. Exopod I-0; I-0; II, 1, 0. Endopod 0-0; 0-0; 0,I,0.
Right leg: Coxa 0-0. Basis 1-0. Exopod I-0; I,1; II, 1,0.Endopod 0-0; 0-0; 0, 2,2. (Roman numerals = spines, Arabic numerals = setae; outer border setation listed to the left in each group separate by ';'

Nota: The authors note that among specimens from a single locationh there is a large degree of variability in the proportions of the male P5.


Espèce Calanoides natalis - Planche 9 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14) [p.828, Table 5]. Character states distinguished known of Calanoides.
A1 = antennule, a = aesthetasc; approx. = approximately; CR = caudal rami; Gns = genital double-somite; Go = genital operculum; ms = modified seta; Pd5 = pedigerous somite 5; P5 = fifth leg; Re = exopod; Ri = endopod; SR = seminal receptacle; St = terminal spine.


Espèce Calanoides natalis - Planche 10 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14) [p.812, Fig. 3]. Calanoides male P5 indicating the measurements made, from which the ratios in Table 4 and 5 were calculated.


Espèce Calanoides natalis - Planche 11 de figures morphologiquesIssued from : J.M. Bradford-Grieve, L. Blanco-Bercial & I. Prusova in J. Nat. Hist., 2017, 51 (13-14) [p.827, Fig. 12].
Charactristic to discriminate between females of C. natalis, C. carinatus and C. brevicornis.


Espèce Calanoides natalis - Planche 12 de figures morphologiquesissued from : S.B. Schnack in Crustacean Issue, 1989, 6. [p.143, Fig.6: 3].
3, As Calanoides carinatus (from off NW Africa, upwelling region): Cutting edge of Md.


Espèce Calanoides natalis - Planche 13 de figures morphologiquesissued from : S.B. Schnack in Crustacean Issue, 1989, 6. [p.144, Fig.7: 4].
4, As Calanoides carinatus (from off NW Africa, upwelling region): Cutting edge of Md.


Espèce Calanoides natalis - Planche 14 de figures morphologiquesissued from : S.B. Schnack in Crustacean Issue, 1989, 6. [p.146, Fig.8]. As Calanoides carinatus.
Schematic diagram of Mx2.
Distance between setae and length of setule not considerd. (Herbivore mode))
Nota: The intersetule spacing on the Mx2 may be used as an index for the feeding habit. Compare with Mx2 of the omnivore Centropages chierchiae.


Espèce Calanoides natalis - Planche 15 de figures morphologiquesIssued from : M.D. Viñas, L. Blanco-Bercial, A. Bucklin, H. Verheye, J.G.F. Bersano & S. Ceballos in J. Exp. Mar. Biol. Ecol., 2015, 468. [p.101, Fig.2].
TCS Network diagram showing frequencies, relationships and source locations for the 12 haplotypes observed for Calanoides carinatus s.l.
The size of the pie indicates relative frequency; color of the slice indicates the collection area (see also Fig.1); samples in legend are ordereds from North to South.
Due to the low number of individuals, the SE Atlantic samples from 16° to 34°S (Benguela Current) were pooled.

Nota: C. carinatus s.l. collected at 13 sampling locations was analysed (Table 1: localities, , date, number of individuals, sequence identification and GenBank). The TCS network diagram indicated complete isolation between the SW and NE/SE Atlantic populations, with no shared haplotypes between them. In addition, TCS analysis indicated no differentiation between NE and SE populations, with 2 groups of haplotypes found in both regions. Several individuals from the NE Atlantic, at the northern edge of the species distribution, showed very divergent haplotypes.
C. carinatus s.s. (SW Atlantic) and C. carinatus s.l. (NE/SE Atlantic) are clearly genetically differentiated from both C. macrocarinatus and C. acutus. Accirdingly, Sabatini & al. (2007) found morphological differences between specimens of C. carinatus s.s. and C. macrocarinatus from new Zealand and concluded that they were different species. They also found that both species may be distinguished from C. acutus by several morphological characters of the female and male.


Espèce Calanoides natalis - Planche 16 de figures morphologiquesIssued from : F. Höring in Master Thesis, Alfred Wegener Inst., Universität Bremen (Germany). [p.16, Fig.6]
Haplotype network of COI data.
Circles present the haplotypes which are connected by mutational steps (blue points).
The size of the circle corresponds to the number of individuals that share the same haplotype. The pie charts represent the proportion of origin of the individuals.


Espèce Calanoides natalis - Planche 17 de figures morphologiquesIssued from : F. Höring in Master Thesis, Alfred Wegener Inst., Universität Bremen (Germany). [p.34, Fig.16]
Haplotype network of the 484 bp COI fragment.
Combined data set of this study and available sequences on GenBank.


Espèce Calanoides natalis - Planche 18 de figures morphologiquesIssued from : F. Höring in Master Thesis, Alfred Wegener Inst., Universität Bremen (Germany). [p.8, Fig.4]
Stations map for samples of Calanoides natalis from the southwest African coast, the northern and southern Benguela upwelling systems, off Senegal/Mauritania to southwest Spain.
The aim of this research is the study of potentiel cryptic species, the spatial patterns in the distribution of genetic variance and identify potential barrriers to gene flow. Analysis of motochondrial (cytochrome c oxydase subunit I; COI) and nuclear (citrate synthase; CS) marker genes revealed a genetically cohesive populayion of C. natais) with a prevalent shift in allele frequencies.
Genetic differenciation between the northern and southern hemisphere was significant, which may point to a potential, but permeable barrier close to the equator.
The genetic analysis shows that horizontal differentiation was more pronounced than vertical structuring..
Retention mechanisms and the oxygen minimum zone did not have a strong impact on genetic differentiation of C. natalis in the Benguela region.


Espèce Calanoides natalis - Planche 19 de figures morphologiquesIssued from : G.S. Brady in Ann. Durban Mus., 1914, Vol. I. [Pl. I, figs. 6-9].
Male (from Durban Bay, Natal): 6, habitus (from right side); 7, Abdomen; 8, A2 female; 6, Md female; 10, P5 male (right side); 11, P5 male (left side).

Nota female:
- Cephalothorax distinctly separated, and much stouter than the posterior abdominal portion, rounded off in front.
- Forehead bearing a pair of slender, curved, tentacular processes;
- Abdomen 4-segmented.
- Caudal stylets short, scarcely longer than the last abdominal segment, bearing 5 apical setae.
- A1 24-segmented, sparingly setiferous, reaching beyond the distal end of the abdomen.
- Branches of A2 nearly equal in size; inner branch 2-segmenred, outer indistinctly 4 or 5-segmented.
- Md of the usual calanoidd type.
All the swimming legs biramose, the branches 3-segmented.
Nota male:
- Abdomen very slender, 5-segmented.
- Biting plate of Md entirely wanting.
- P1 to P4 as in female.
- P5 very slender and prehensile, destitute of lateral setae; inner branch of the foot of the right side reduced to a very minute ovate, 1-segmented process, outer branch slender, flexile, 3-segmented, its last segment much the shortest; inner branch of the left foot 3-segmented, much reduced in size, reaching only to the middle of the 2nd segment of the outer branch.

NZ: 4 + 1 douteuse

Carte de distribution de Calanoides natalis par zones géographiques
Espèce Calanoides natalis - Carte de distribution 3issued from : C. Courties in Thèse Univ. Bordeaux I., 1978. [Fig.13]. As Calanoides carinatus.
Seasonal abundance along Congo and Gabon coast during cold season (August).
Number/20 cubic meters after cotation from Frontier (1969): 1 = 1-3 individuals; 2 = 4-17; 3 = 18-80; 4= 80-350; 5 = 350-1500; 6 = 1500-6500; 7 = 6500-30000.
Mesh aperture of the net: 0.330 mm.
Nota:The author pointsto a demographic bloom during the cold season.

Probably = Calanoides natalis.
Espèce Calanoides natalis - Carte de distribution 4issued from : C. Courties in Thèse Univ. Bordeaux I., 1978. [Fig.8]. As Calanoides carinatus
Seasonal abundance along Congo and Gabon coast during warm season (March).
Number/20 cubic meters after cotation from Frontier (1969): 1 = 1-3 individuals; 2 = 4-17; 3 = 18-80; 4= 80-350; 5 = 350-1500; 6 = 1500-6500; 7 = 6500-30000.
Mesh aperture of the net: 0.330 mm.

Probably = Calanoides natalis
Espèce Calanoides natalis - Carte de distribution 5Issued from : C. Courties in Thèse 3ème Cycle, N°1432, Univ. Bordeaux I, juin 1978. [Fig.38]. As Calanoides carinatus.
Seasonal variations of abundance. Number of generations from 1 to 6. at two stations a (120 milles) and b (160 milles) during the colder season 1974 off Pointe Noire (Congo)

Sampling by WP2 (mesh aperture 200 µm).
Station ''a'': 22°5 C and 35.86 p.1000 at 10 m; 18°6 C and 35.88 p.1000 at 30 m. Station ''b'': 22°9 C and 35.88 p.1000 at 10 m; 18°.8 C and 35.88 p.1000 at 30 m.
Histograms indicate the cephalothoracic measures.

Nota: During the colder season friom 1974 C. carinatus show 6 generations.

Probably Calanoides natalis
Espèce Calanoides natalis - Carte de distribution 6Issued from : C. Courties in Thèse 3ème Cycle, N°1432, Univ. Bordeaux I, juin 1978. [Fig.45]. As Calanoides natalisBiological cycle.
Pronably = Calanoides natalis.
Espèce Calanoides natalis - Carte de distribution 7issued from : J.-P. Ehrhardt in Cah. Oceanogr., 1967 (9). [Fig.10].
Occurrence of Calanoides carinatus (= C. natalis collected at 14 stations on 39 stations between the southern Tyrrhenian Sea, Tunisia-Sardinia Strait and Cap Bon-west Sicily.

Nota: The species represents 0.3 % of adult copepods in the waters from salinity between 37.3 and 37.8 p.1000 (more frequently between 37.59-37.72), and between 4.35-5.93 cm3/l oxygen (more frequently between 4.69-5.50 cm3/l), probably under influence of the atlantic superficial water current.

Probably C.alanoides natalis
Espèce Calanoides natalis - Carte de distribution 8issued from : R. Stephen in Pelagic Biogeography ICoPB II. Proc. 2nd Int. Conf. Final report of SCOR/IOC working group 93. 9-14 July 1995. Workshop Rep. No.142. UNESCO, 1998.
P.343, Fig.7: Distribution of Calanoides carinatus (= Calanoides natalis) in the Indian Ocean.
Perhaps other species in the eastern part of the Indian Ocean. [CR]
Espèce Calanoides natalis - Carte de distribution 9issued from : U. Brenning in Wiss. Z. Wilhelm-Pieck-Univ. Rostock - 30. Jahrgang 1981. Mat.-nat. wiss. Reihe, 4/5. [p.6, Fig.8]. As Calanoides natalis.
Spatial distribution for Calanoides carinatus from 8° S - 26° N; 16°- 20° E, for different expeditions (V1: Dec. 1972-Jan. 1973; V2: Feb./Mar. 1973; IV: Jun.-Jul. 1972; VI: May 1974.
Legend of circles in T-S diagram.

Nota: Taking into account the seasonal temperature variations, the number of generations to be expected in a given region when the duration of upwelling in the region concerned is known, we have: 7-8 generations at Nouachott (Mauritania), 5-6 at Cap Vert (Dakar) and 3 at Cap Roxo (Casamance).

Probably = Calanoides natalis.
Espèce Calanoides natalis - Carte de distribution 10issued from : U. Brenning in Wiss. Z. Wilhelm-Pieck-Univ. Rostock - 30. Jahrgang 1981. Mat.-nat. wiss. Reihe, 4/5. [p.7, Fig.9].
T-S Diagram for Calanoides carinatus (= calanoides natalis (all copepodid stages) from 8° S - 26° N; 16°- 20° E, and Namibia (SW Africa) for different expeditions (V1: Dec. 1972-Jan. 1973; V2: Feb./Mar. 1973; IV: Jun.-Jul. 1972; VI: May 1974; VIII: 21 Sept.-17 Dec. 1976).
SO: Southern Surface Water (S °/oo: 34,50; T°C: 29,0); ND: Northern Water of the Surface Layer (S °/oo: 37,5; T°C: 21,0); SD: Southern Deep Water of the surface layer (S °/oo: 35,33; T°C: 13,4).
Water types basing on the works of Sverdrup & al. (1942), Wolf (1978) and Wolf & Kaiser (1978) distinguishing various quasi-permanent water types in the top 200 m of the water column in the NW Africa upwelling region. The water type SD, which usually forms the upper limit of the SACW (South Atlantic Central water: S °/oo: 34,35-35,33 & T°C: 5,15-13,4), can enter the surface layer as a result of upwelling, so that water types ND, SD and SO can be expected in this layer in upwelling regions. The boundary separating the NACW (North Atlantic Central Water: S °/oo: 35,10-37,35; T°C: 8,0-21,0) and SACW water masses is situated in the region between Bahia de Gorrei and Cap Blanc (Mauritania) throughout the whole year. The distribution of the water types SACW in the surface layer off North West Africa varies with the season and the meridional migration of the wind field. See in Brenning (1985 a, p.6).

Probally = Calanoides natalis.
Espèce Calanoides natalis - Carte de distribution 11issued from : U. Brenning in Wiss. Z. Wilhelm-Pieck-Univ. Rostock - 30. Jahrgang 1981. Mat.-nat. wiss. Reihe, 4/5. [p.8, Fig.10, f-g].
T-S Diagram for Calanoides carinatus (= Calanoides natalis) females and males.

Probably = Calanoides natalis.
Espèce Calanoides natalis - Carte de distribution 12issued from : U. Brenning in Wiss. Z. Wilhelm-Pieck-Univ. Rostock - 30. Jahrgang 1981. Mat.-nat. wiss. Reihe, 4/5. [p.9, Fig.11].
Vertical distribution for Calanoides carinatus (= Calanoides natalis) from 8° S - 26° N; 16°- 20° E.
T: daylight; N; night; G: all copepodis stages; W; females; M; males.

Probably = C.alanoides natalis.
Espèce Calanoides natalis - Carte de distribution 13Issued from : F. Höring in Master Thesis, Alfred Wegener Inst., Universität Bremen (Germany). [p.5, Fig.3]
Life cycle of Calanoides carinatus (= C. natalis) associated to the upwelling system (drawing after H Auel).
After Bradfoed-Grieve & al. (2017, p.803) C. natalis is known as an 'upwelling specialist' (Ceballos & al., 2004). Peterson (1998) considers such species to be pre-adapted to wind-driven upwelling situations where water circulation ensures that surface water is taken off shore, that deeper water drifts onshore, whereas, over the continental shelf, the drift is usually equatorward at the surface a,d polewards over the continental slope. Peterson (1998) summarizes the reasons why species such as C. natalis are able to maintain their position in an upwelling region through differing behaviour at different life stages. That is, diapause stage copepodids (CV) are carried at depth (> 200 m) polewards and, during upwelling, are carried inshore onto the continental shelf. When CV reach productive surface waters they exit dapause, mature males and females mate, and eggs are laid. The developing nauplii and copepodid stages remain within surface waters, drifting further offshore as they develop and equatorward as they mature to CV. If upwelling continues, CV stages, by migrating deeper in the water column, are again carried shoreward and may undergo several generations depending on the length of the upwelling season. The timing of this cycle differs depending on geographic region, the timing of the upwelling season and its length.
Loc:
SW South Africa, Namibia, Senegal-Mauritania, SW Spain, ? W Medit. Type locality: SW Indian (Durban Bay)
N: ? because confusion with Calanoides carinatus
Lg.:
(1319) F: 2,2-2,6; M: 2,1-2,41; (1320) F: 2,55; M: ?; {F= 2,20-2,60; M: 2,10-2,41}
Rem.: Confounded by numerous authors with Calanoides carinatus before 2017.
After Brady (1914, p.4), Giesbrecht (1892) has figured the P5 male, a form which he identifies with Calanus brevicornis Lubbock, his figure agreeing very exactly with P5 male of Calanoides natalis; but this lomb is neither figured nor described by Lubbock, and C. brevicornis is in other respects quite unlike the African species here described. It is, possible that Giesbrecht's figure may apply to C. natalis , in which case the reference to C. brevicornis is certainly erroneous.
For Bradford-Grieve & al. (2017, p.827)Calanoides natalis females are distinguished morphologically from the western south Atlantic species Calanoides carinatus stricto sensu by the setation of ancestral antennular segments XV, XVII, XIX and XXI. C. natalis has the setation of all these segments with 1 ms + 1a, whereas in C. carinatus the setation is 2 ms + 1a (see in Table 5). In this respect C. natalis is the same as C. brevicornis (as macrocarinatus) (Bradford-Griece, 1994. It is confirmed that the anterior surface of the praecoxal arthrite of Mx1 of C. carinatus does not have a group of large spines, whereas such spines are present on C. natalis (Fig.7b) and C. brevicornis (as macrocarinatus (Vyshkvartzeva, 1977). In males, the difference between C. natalis and C. carinatus is in the left P5: C. natalis (from the Arabian Sea) had the ratio of the length of the outer distal spine of exopodal segment 2 relative to the length of the inner border of rxopodal segment 3 (G/B) had a mean of 0.54 (range 0.47-0.67) (Table 4) whereas, in C. carinatus this ratio has a mean of 0.96 (Sabatini & al., 2007). Similarly, the length of the inner border of exopodal segment 2 relative to the maximum width of this segment (C/c') has a mean of 2.85 (range 2.53-3.30) (Table 4), whereas, in C. carinatus this ratio has a mean of 4.06 (Sabatini & al., 2007).
It is more difficult to distinguish female C. natalis from C. brevicornis, as their antennumar setation is the same. Their distributions overlap off southern Africa and no males were available of the latter species for re-examination.
Measurements of the female genital double-somite maximum width and posterior width were plotted against one another (Fig.12): the measurements for each species did not overlap in two dimensional space. Comparisons of these measurements were all significantly different, but the two populations of C. natalis (Arabian Sea and off Namibia) did not differ significantly (p > 0.1). It is known that C. natalis and C. brevicornis are genetically distinct (Viñas & al., 2015), but there are further subtle differences in shape and proportions that are difficult to quantify. See in Remarks for Calanoides brevicornis.
Dernière mise à jour : 24/08/2019

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Razouls C., de Bovée F., Kouwenberg J. et Desreumaux N., 2005-2020. - Diversité et répartition géographique chez les Copépodes planctoniques marins. Sorbonne Université, CNRS. Disponible sur http://copepodes.obs-banyuls.fr [Accédé le 27 septembre 2020]

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