Bo1m Inst. oceanogr •• S Paulo, 35(1):7-16. 1987
DISTRIBUTION AND CO-OCCURRENCE OF
Ca~a~o~de~ ~ah~~atu~AND LARVAE OF
Sahd~~e~~a bha~~~~e~~~~
AND
E~ghau~~~ a~~ho~taOVER THE SOUTHERN
BRAZILIAN CONTINENTAL SHELF
Antonio Frederico CAMPANER
&
Sumiko HONDA*Instituto de Biociências da Universidade de são Paulo, Departamento de Zoologia, (Caixa Postal 20520, 01498 são Paulo,SP)
Abstract
The distribution of adults and juveniles of Caia~o~d~ ~~at~ with co-occurring
copepods along two continental shelf transects off são Sebastião-Ubatuba and
Para-naguá in the SE and S Brazil respectively~ during December to February and May from
1975 to 1977 were studied by anaZysing plankton samples from 5 stations coUected
along each transect from 5 m above the bottom to the surface with a 0.333 mm mesh
Bongo neto
C.
~~at~ is present over the shelf along both transects only duringlate spring and early summer (December to FebruaryJ. Its density is approximately
20% and 5% of the mean copepod density estimated for são Sebastião-Ubatuba and
paranaguá respectively. Distribution of
C.
~~~at~ coincides with both that ofE~gha~ a~cho~'s Zarvae and the spatial distribution of the South Atlantic
Central waters over the são Sebastião-Ubatuba shelf from December to February.
Both C. ~~at~ and larvae of SMM~illa bh~~e~~ are locaUy abundant from
December to February but do not co-occur.
Descriptors: Crustacea, Copepoda, Calano~d~ ~~~at~, Abundance, Nerictic province. Pisces, fish 1arvae, SMM~etla b~~e~~, E~gha~
an~ho~a, Geographica1 distribution. são Sebastião-Ubatuba,
Paranaguá, SW At1antic, Brazi1.
Descritores: Crustacea, Copepoda, Caiano~de~ ~~natU6, Abundância, Província nerítica. Pisces, larvas de peixe. SMM~etla bh~~e~~,
E~gha~ a~~ho~a. Distribuição geográfica, são
Sebastião--Ubatuba, Paranaguá, Atlântico Sul Ocidental, Brasil. Introduction
Calano~d~ ~~at~ is widespread in
the tropical neritic zones of the three large oceans and in the western
Mediterranean (Brodskii, 1972; Vervoort, 1949). ln the Atlantic Ocean, it is distributed from 47°N to 37°S in waters wit~ temperature ranging from 12°C in
the southern subtropical convergence to 25°C in the Gulf of Guinea (Courties, 1978; Vervoort, 1949). It is considered an indicator of waters from the Malvinas Current off Argentina (Ramirez, 1977) and of subtropical water upwelling off Brazil (Bjornberg, 1981; Valentin,
1984b). The copepod has been studied in upwelling areas off western Africa but also is present along the Congo-gabonese continental shelf even in the absence of
* With a grant of the "Fundação de
Amparo i Pesquisa do Estado de são Pau 1 0" (FAPESP) for undergraduate
student (Proc. 85/0563-3).
Conth. no.
658do
I~t. o~eanogh. da U~p.upwelling if the surface waters are cool and trophically rich (petit
&
Courties, 1976) .ln the SE African marine ecosystem,
C.
~~at~ has a similar role as thatof Caia~~ 6úurrM~~~ in the North
Atlantic and of Caia~~ a~u;t~ in the
southern Atlantic (Bainbridge, 1972). It is considered the most important trophic link between the phytoplankfon and the fishes in upwelling areas (Hirche, 1980). The distribution of
C.
~~atU6over the sh~lf has been correlated with that of the ichthyoplankton (Weikert, 1984) and adult fish species (Brenning, 1985), especia11y c1upeids (Bainbridge 1960, 1972; Binet, 1983; Binet & 8uisse de Sainte Claire, 1975).
8
bJzaú.1Á..eJu.{,6 and
EnglLa.uLL6
a.ncÁo.aa. havebeen studied by Matsuura (1979, 1983) and Nakatani (1982) based on material and data collected during ten
oceanographic cruises of the R/V "Prof. W. Besnard" sponsored by the FINEP/IOUSP
(Financiadora de Estudos e Projetos/Ins-tituto Oceanográfico da Universidade de são Paulo). ln this paper a
quantitative, seasonal survey of the occurrence of Cala.noid~ ea.nina.tuh in two distribution area·s of these two fish species' larvae is presented with a discussion of their possible trophic relationships.
Material and methods
Plankton samples were collected with a 0.333 mm mesh Bongo net towed obliquely from 5 m above the bottom to the surface at five stations of each transect I and II, respectively on the continental shelf off são Sebastião-Ubatuba and paranaguá (Fig. 1) during December'75, January'76,May'76, December'76, and Jan./Feb.'77 (R/V "Prof. W. Besnard" cruises I-III and V-VI). Copepods were
+
N
\
48-Bo1m Inst. oceanogr., S Paulo, 35(1), 1987
counted under the stereomicroscope in a Bogorov's counting chamber either from the whole sample or from one subsample drawn off with a Folsom splitter (Cf. Campaner, 1985). Adult males and females, and juveniles (nauplii and copepodites I-V) of Ca.ta.noid~
ea.n1natU6
were counted separately.
Results and d-iscussion
Table 1 shows the copepod densities and furnishes the water mass composition in the upper 150 m of the water column at each station. The predominant
relationship of the Coastal, Shelf, Tropical, and SA Central water masses
(vide Campaner, 1985) was calculated by the percentage value of each water mass in the water column where 10 means 100% of the water. Areas of unprecise
delimitation or where there is the
overlap of two or more water mas ses were referred to as mixing zones.
The water mas ses had a similar distribution pattern as obtained by Matsuura (1986) for the SE-S Brazilian
Ilha
45-o stat ion transect
44-
24-
25-2r
43·W
CAMPANER
&
HONDA: C. e~natU6 x fish larvaeneritic region. The coas tal waters were only recorded in the shallower station offparanaguá (Table 1: St. A, Transect II), probably due to the inf1uence of estuarine waters. Tropical waters occupied the são Sebastião area more extensively, especial1y in Dec.'76 than
off paranaguá, showing larger oceanic inf1uence in the first area. Although spatial extent differed, shelf waters were a1ways present and situated above
the SA Central waters. These were more shallower in Dec./Jan. and deeper, far from the coast in May.
Table I. Relationship of the water masses, density of
Calanoide6
e~natU6,the other copepod species, and the copepod total in each station of transects I (são Sebastião-Ubatuba) and II (Paranaguá) on the southern Brazilian continental shelf from Dec. 175 to Jan./Fev. 177. Abbreviations: C= Coastal waters, T= Tropical waters, S= Shelf waters, A= SA Central waters, M= mixing zones, ad.= adults,
Tran s ect Seaso n
Oec. '75
II
Ooc. ' 75
J a n . '76
II
Jan . '76
Hay'76
II
Hay' 76
juv.= junveniles (nauplii + copepodites l-V)
Station A A A A A A local depth ( m) 18 63 83 116 1.41 2 I 42 60 97 127 19 72 81 120 280 22 44 64 95 135 16 ]1 77 11 7 127 20 45 64 95
Predominant relation s h i p
of water mas ses
C o T o S : A : M
0 : 0 : 100 : 0 0 : 0 : 6 : 4 : 0
o : O : 2 : : O
O : O : O o : 2 : O :
NO
DATA
AVAILABLE
0 : 0 : 100:0
o : O : : O
O : 4: O : : O
O : O : 4 :
0 : 2: 0:6 : 2
O : 10 O: O
O : O : 10 O : O
O : O : 4: : O
O : O : : O
O : O : O
0 : 0 1 0 0 : 0 0:0 : 100 : 0
O : O : : O
O : O 4 : 6 : O
O : 4
10 : O O : O 0:0 : 100 : 0 0:0 : 100 : 0
: O : O
O : 4 : 6 : O
c o p e p o d d e n s i t e 5
Ca l an oi d eh c a~ i natuh other
ad . Q ad o d juv . Total spp
0.45
31 . 40
0 . 77
O. 1 I
0.03
0.23
2 . 40
0 . 36
14 . 08
O. 12
0 . 04 0 . 09
0 . 20 0 . 02 0 . 10
0.04
O. 10
O. O 3
0 . 64
6.25
0 . 64 145 . 84
6.73997.11 4 . 00 830 . 15865 . 552032.84
I . 00
0 . 08
0 . 39 .0.16
2.34
O. 18
O. O 1
0.04
65 . 09 66 . 86 321.61 41.03 41 . 14 171.87
0 . 49
3 . 1 I
4 . 06 6.79
1~39.49
0 . 52 264.43
3 . 42 187 . 20 6 . 85 152 . 00 7 . 31 68.64
362 . 75
13.05 29.47 679 . 62
22.79 23 . 09 69 . 85 2 . 22
1. 94
0.03 0.40
0 . 60
6.34
0.05
0.08 o. 18
0 . 02
2 . 26 135 . 46
2 . 04 98 . 89
0 . 03 185.56
0.64
0 . 62
6.44
0 , 05
e5. 13 87.95 77.50 53 . 56
310 . 60 235.64 0.12 139.80 0 . 28
0 . 02
84.09 27 . 41
179.52 64 . 00 54 . 46
70.76
36 . 15
(No • m-3 )
Copepod
Total
146 . 48
1003.84
2898 . 39 388 . 47 2 13 . O 1
1039 . 49 264 . 95
190 . 62
158 . 85 75 . 95
362.75 709 . 09 92 . 94 137.72 100 . 93
185 . 59 85 . 77 88 . 57 83 . 94 53.61
310.60 235.64 139 . 92 84 . 37
27 . 41
179 . 52 64 . 00 54 . 46 70 . ,8 36 . 15
9
10 Bo1m I nst. oceanogr. t S Paulo, 35(1) t 1987
Tab1e 1. Conto
Tr.nsect St.tio" local 'redo ... ln.nt relatlon'hlp C o p e p o cI cI e n
•
I t I e,
(No....
)Se _, on clepth of water .'I.e,
(
..
) C : T : A , MA 2~ O : O : 10 : O : O
7~ O : O : ~ : 6 : O SI D : 6 : O : 4 : O
Dec. '76 124 O : 6 O : 4 : O
I ~~ O : 8 : O : 2 : O
A 19 10 O : O : O : O
~~ O : O : 10 O : O
II 64 O : O : : O
Dec . ' 76 9~ O : ~ : 2 : 4 : O
127 O : 6 : O
A 20 O : O : 10 O : O
70 O : O : 2 : e : O 81 O : O I : : O
Jan . /Feb . '77 120 O : 2 : : O
136 O : 6 : O : 2 : 2
1\ 22 4 : O : 6 : O : O
I,~ O : O : 5 : O
II 65 O : O 3 : 5 : 2
Jan./Feb.' 77 96 O : O : 2 : : I
131 o : 2 : 2 : 4 : 2
According to Bjornberg (1963), the above water masses on the Brazi1ian she1f shou1d be r~nked as follows,
relative to the decreasing mean plankton volume: Coastal, SA Central (Deep
Shelf), Shelf (Surface Shelf), and Tropical waters. Consequently, their distribution should influence the qualitative and quantitative
copepod composition greatly, as can be deduced from some results (see Table 1); e.g., the copepod densities were
relative1y high in St. A off paranagua where coastal waters were presento On
the other hand, densities were generally low in stations where tropical waters predominated over SA Central waters.
Cai.a.no.-tdu c.aJÚnMU6 was present on
the shelf during late spring and early summer (Dec./Jan./Feb.) and practical1y absent in autumn (May) (Campaner, 1985; present data) clearly showing its
seasonal occurrence. The stocks from ear1y summer were predominantly
juveniles (Tab1e 1). The reduction of the species over the shelf during
autumn has also been observed in the SE Atlantic off Africa (Courties, 1978;
Caluo.é.du C"J1:""4.t",· olhe, Copepod
ecl. 9 ed. d Ju v . Tot.1 ,pp Tot.1
o. 19 2.o~ ~ . 8o 7 . 03 618 . 81 625.8~
0.84 I. 40 10.87 13. I I 1048.44 1061.55 0.05 o. 10 8.95 9. 10 245 . 34 254 . 44 0.09 3 . 50 3.59 97.14 1 00.73 O. 12 O. 12 58.67 58.79
565 . 29 565.29 534.84 53~.64
354.04 354.0. 0.06 0.06 O. 12 178 . 33 173 . 45 0 . 03 0 . 05 0 . 08 88.28 a8.36
0.52 O. 10 0.62 510 . 55 511 . 17 35 . 56 I. 82 10.30 ~~.68 1201.24 1;:~5 . 92
5.20 2 . 53 202.35 210.08 151.69 361.77 0 . 34 o. ~ I 10 . 82 I I. 57 38.46 I 10 . 03 0.09 O.O~ 6. 07 6 . 20 81 . 13 87.33
91~.28 914.28 o. 19 O ~ 19 691 . 4~ 691.63 0.80 0.48 7 . 55 8.83 623 . 83 632.66 0 . 19 0.09 I .27 I. 55 196.57 138 . ~2
0 . 07 O. I I 8 . 71 8.89 153 . 87 162 . 76
Petit
&
Courties, 1976) and attributed to the migration of the summercopepodite stock to deeper waters. The development growth of
C.
c.aJÚnMU6 and its quick reproduction are not only favoured by the coo1ing of the surface waters (Binet&
Suisse de Sainte Claire,1975) but a1so by the phytoplankton growth (Smith, 1982). The more
superficial occurrence of the SA Central waters on the Brazilian shelf from
December to February (Tab1e 1) should contribute with nutrients for the
proliferation of the phytoplankton. This water mass did not reach depths below 50 m in May. Contrary to observations from são Sebastião-Ubatuba, the SA Central waters seemed to have little inf1uence on the increase of copepod abundance in Paranaguâ, where the greatest number occurred in the St. A (see Table 1: Transect II, Dec.'75) in shallow coasta1 water. ln this station
C.
c.aJÚnMU6 did not occur, it onlyappeared significantly in January, when the SA Central waters were more
CAMPANER
&
HONDA:C.
QakinatUó
x fish larvae 11approximately 20% and 5% af the mean copepad density in são Sebastiãa-Ubatuba and paranaguâ, respectively.
The spawning seasan and the greatest abundance of SaJLMneU.a blUt6ilÃ..en6-<-6
larvae aver the shelf occur from late spring to late summer, whereas in autumn no eggs and a few larvae were recarded (Matsuura, 1983). This seasonal occurrence pattern is similar to that observed for
C.
~natUó, but the density distribution of these twospecies are nat caincident: cansiderable numbers of C.
cakinatUó
are faund in are as of absence ar low numbers ofsardine 1arvae during Dec./Jan. (Fig. 2). Nakatani (1982) faund that spawning
af Eng~ ancho~aoccurs in twa
seasons, viz. late spring-ear1y summer and late winter-early spring. The former episade is restricted to upwe11ing areas near the coast whi1e the latter is
scattered throughout the she1f. ln the former season greater concentratians of 1arvae are found in areas of Transect l where
C.
QakinatUó
aIs o occursabundantly, especial1y its young forms (Fig. 3, Tab1e I).
The main habitat af the sardine 1arvae and juveni1es in the sauthern Brazi1 are the she1f waters with
temperature ranging fram 22 to 26°C and sa1inity from 34.5 to 36.0 % 0
(Matsuura 1979, 1983); these are
practica11y the sarne characteristics of the habitat of SaJLcü.neU.a aunita off the Senega1ese coast (Boe1y, 1982). The habitat of the anchavy 1arvae are the SA Central waters with temperature below 20°C (Matsuura, 1983; Matsuura et
al.,
1985; Nakatani, 1982). ln fact, areas of higher concentrations af anchovy 1arvae(Fig. 3) are genera11y coincident with stations af Transect I where the
SA Central waters predominate over the other water masses in the water column
(Tab1e 1).
Accarding to Goiten (1983), adu1ts af
SaJLcU..neU.a blUt6ilÃ..en6-<-6 are
predaminant1y phytap1anktivarous in the Cananéia region and zoop1anktivorous . off Ubatuba and Santos, the copepads
representing respective1y about 80 and 60% af the stamach contents af the sardines caught in the two areas.
Calano..idu Qakina.:b.L6
was not faund among the copepad species identified, these being cammon1y recorded in coas tal and she1f waters. No infarmatian isavai1ab1e abaut feeding habits of larvae and juveniles.
The feeding habit of larvae, juveniles, and adu1ts of Eng~
anQho~a has been studied in the
Argentine waters (Ange1escu, 1982; Ciechomski, 1967; Ciechomski& Weiss, 1974), According to the~e authors, alI deve10pmental stages feed
exclusive1y on zooplankton, the larvae predominantly on eggs, nauplii, young copepodites, and adults of copepods, especially the Calanoida, The cryophi1e species of calanoid capepods, including
C,
QakinatUó,
constitute the main item of the diet of juveni1e and adu1t anchovies.Angelescu (1982: 38) considered the quality and abundance of the main food (calanoid copepods) in the area under influence of the Malvinas Current, together with the environmental temperature and the thermocline formation, as the eco1ogica1 factors important in influencing the
anchovy's feeding - and growth. Anchovies may encounter these conditions on the southern Brazilian shelf - the cool SA Central waters spread upwards over the shelf off são Sebastião/Ubatuba
-during spring and summer when
Calano..idu QaJr.inatU6,
one of . the dominant copepod species in this waters mass, is reproducing quickly, Consequently, large numbers of copepod eggs and larvae are available tothe anchovy 1arvae and juveniles, During May, the anchovy 1arvae are more abundant far from the coast
(Fig, 3) in areas where
C.
~natUóis practically absent; this copepod species is replaced in abundance by another herbivore,
EUQalanU6 p..iteatU6,
(Campaner, 1985) which cou1d be an important food item in the anchovy diet during this period.
To conc1ude, the distribution af
Calano..ide..6 Qakina,tUó
over the southern Braziliancontinenta1 she1f coincides wi th both tha t of the EngJtau.W anQho.u:a' s1arvae and the SA Central waters in the são Sebastião-Ubatuba area during late spring and ear 1y summe r, l-li th a s easonal occurrence similar ·to C. ~na,tUó,
SaJtd..ineU.a bJUt.6ilÃ..en6..[,6' 1arvae are
12
Fig. 2
.
•
•
1'/
+ / ..
-
..-
.... ,; ++ /
/
,)
Bo1m Inst • oceanogr • I S Paulo, 35(1)
Densities
ç.
carinatus (no .m )
-3+ 0.0
• 0.01-0 .10
• 0 . 11 - 1. 00
$. bras ili ensl'
~~
S0-.."'" 0.1-10. 0
~ ~ 10 .1-100.0
o 1.01 · -10. 0
• 10.01-100. 0
• 100.01-1000 .0
(no
. m
- 2)9&
100.1-1000 .0Abundance
di st' SE rlbutiona1 of Ca1.an 'd 0-<' e..6 c.CVÚn
-S Braz i 1 . areas of 1 atU/.:, in
data. lan contin enta 1 sh arvae of e 1 f based S aJtLU.ne.li
~~ansects
on Ma.
bJr.a.6: I~e.n6.i6
and o : I I p.1otted· . I natsuurals ( 1979, 1983) In the
, . 1987
23'
• naXU6 x f i 5 h
_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _
CAMPANER & HONDA' C
, c.aJÚ ___~la~rv~ae
____ __Jan./Feb:77
Oensiti es
ç.
carinatus ( nO.m ) -3+ 0.0
• 0.01-0 .10
• 0.11-1.00
o 1.01-10.0 • 10.01-100.0
• 100.01-1000.0
E.anchoita (
~"'-::
nO.m ) -20-"'- 0. 1-10.0
~
~ 10.1-100.0
$
100.1 -1000.0 44'WF' 19,
3.
Abunda d' 1st r i nce of bu t 'C_O_'A
'M ... u.noidU .
B razll ian • lona I a continent I reas of 1
CAJuna.t·
arvae of E U6 ln t ransects I_
_ _ _ _ _ _ _ _ _
a_sh_e_lf~bas=eud
o~~<l~
a atani's (1982)ln the SE-Sancho~
!
I plotted in data •14
Resumo
Estudou-se a distribuição de
Calanoideó
caninatuh adultos e jovens e dos de~ais
copepodos ao longo de duas transecçoes, com 5 estações cada, sobre a platafoEma continental brasileira ao largo de Sao Sebastião-Ubatuba e Paranaguá, durante dezembro a fevereiro e maio de 1975 a 1977. Foram analisadas amostras de plâncton coletadas de 5 m acima do fun-do ate a superfície, com uma rede Bongo de O 333 mm de abertura de malha.
,
. ..
C, Q~atUó esteve prese~te
s1gn1f1-cativamente, nas duas regioes estuda-das somente durante o fim da prima-
,
-vera e início do -verao (dezembro a
fevereiro), Sua densidade represen-tou cerca de 20% e 5% da densidade media de todos os copepodos
cole-tados ao largo de são Sebastião--Ubatuba e de Paranaguá, respecti-vamente, A distribuição de
C.
Qani-natUó
sobre a plataforma ao largo desão Sebastião-Ubatuba coincidiu com a de larvas de Eng~~
anQhoita,
assim como com a distribuição espacial da massa de água Central do Atlântico Sul, durante o perío-do de dezembro a fevereiro, C.
Qa-~natUó e larvas de
SMdine1.ta
blUt6ilie.n6-L6 apresentaram o mesmo padrão sazonal de ocorrência
(estiveram presentes sobre a plata-forma em dezembro ate fevereiro e praticamente ausentes em maio), porem distribuíram-se abundantemen-te em áreas diferenabundantemen-tes.
Acknowledgements
We are grateful to Dr Y. Matsuura and Miss K, Suzuki (Instituto Oceanográfico, USP) for kindly putting at our disposal material and oceanographic data from the FINEP/IOUSP plankton collection, and Drs Tagea K. S. Bjornberg
(Instituto de Biociências, USP) and Frank D, Ferrari (Smithsonian Institution, Washington D. C.) for reading critically and commenting on the MS. S. Ronda is also acknowledging the FAPESP (Fundação de Amparo
ã
Pesquisa do Estado de são Paulo) fordonating a grant (Proc. 85/0563-3).
Bo1m Inst. oceanogr., S Paulo, 35(1), 1987
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