PEDRO MIGUEL ALFAIA BARCIA RÉ
ICTIOPLÂNCTON DA REGIÃO CENTRAL
DA COSTA PORTUGUESA E DO ESTUÁRIO DO TEJO
ECOLOGIA DA POSTURA E DA FASE
PLANCTÓNICA DE SARDINA PILCHARDUS (WALBAUM, 1792) E DE ENGRAULIS
ENCRASICOLUS (LINNÉ, 1758)
LISBOA
1984
ABSTRACT
Very few studies have
been made in Portugal on marine fish eggs and larvae. The present work deals
with: (i) the diversity and abundance of the ichthyoplankton along the central
portion of the portuguese coast and within Tejo estuary; (ii) The ecology of
the spawning and of the planktonic phase of Sardina pilchardus (Walbaum,
1792); (iii) The ecology of the spawning and of the planktonic phase of
Engraulis encrasicolus (Linné, 1758), within Tejo estuary. Fish eggs and
larvae were sampled over a period of two years off the central part of the
Portuguese coast and over a period of four years within Tejo estuary. The
diversity and abundance of the ichthyoplankton was analysed and related with
upwelling conditions and plankton biomass. Ichthyoplankton diversity was
highest in spring and summer and lower in winter, along the central part of the
Portuguese coast. Off the coast of Sines (1981/1982) the diversity of the
ichthyoplankton was probably influenced by upwelling conditions. Within Tejo
estuary the diversity was, with few exceptions, highest at exterior stations
and lower at interior stations during the four year survey (1978/1981). A
significant correlation was found to exist between plankton biomass and the
number of larvae sampled per month at each station off the coast of Peniche
(1979/1980). Within Tejo estuary the ichthyoplankton was nearly always more
abundant at interior stations. Sardina pilchardus eggs and larval stages
were the most abundant forms sampled along the central part of the Portuguese
coast. The analyses of the Euclidean distance cladograms of collected fish
larval stages, showed that off the central part of the Portuguese coast the
stations sampled were not very dissimilar and that there was not a definite gradient
of ichthyoplankton abundance. Within Tejo estuary interior stations were
normally very dissimilar in relation to the lower estuary. It was included
under the designations of winter spawners, spring spawners, summer spawners and
autumn spawners the species whose eggs and larvae were captured along the
central part of the Portuguese coast during these seasons. Within Tejo estuary
it was possible to distinguish between the species that use the estuary as a
primary spawning area from those that do not. The eggs of Sardina pilchardus
were sampled throughout a period of twelve months along the inshore waters of
the central portion of the Portuguese coast. Peak periods were found in
December and April (1979/1980) off the coast of Peniche and in March and June
(1981/1982) off the coast of Sines. Spawning was temperature dependent. Eggs
were sampled in greatest numbers when water temperatures were not higher than
15‑16ºC. Off the coast of Sines the upwelling conditions influenced both
spawning activity and duration. The existence of two spawning peaks off the
coast of Peniche in 1979/1980 was related to the age structure of the
spawners. The size and relative mortality of pilchard eggs varied seasonally.
The time of spawning was deduced from the presence of early stage eggs in the
plankton. Sardina pilchardus seems to spawn during the first hours of
darkness. Pilchard larvae showed a clear diel vertical migration being found
predominantly at a daytime depht somewhere near 20 meters and at the surface
during the night. Larvae also showed a cycle of feeding activity, suggesting
that feeding took place mainly if not exclusively during daytime. Sardine
larvae were aged using daily growth increments in sagittal otoliths. Growth
and thickness of daily units were found to varie in relation to the time of
the year. This variation was related to differences in food availability and
different feeding conditions. No obvious relationship was found with
temperature. The hour of the formation of the two parts of each daily unit
(organic and inorganic) was established. The protein‑rich portion of the
daily increment is deposited during the first hours of darkness. The
initiation of a new daily growth increment begins around midnight. The number
and biomass of Sardina pilchardus spawners was estimated from
quantitative egg surveys: coast of Peniche (1979/1980) (6,70.103
Ton); coast of Sines (1981/1982) (1,33.103 Ton). The eggs and larval
stages of Engraulis encrasicolus were sampled troughout a period of six
months (February to July) during four years (1978/1981) within Tejo estuary.
Spawning activity was highest in April and took place with greater intensity at
the interior portions of the estuary. The fact that anchovy larvae were present
in greater concentrations in the upper portions of Tejo estuary, and that
larvae of higher length‑classes were always found at interior stations,
led us to believe that there must be some mechanism responsible for its
retention within the estuary. Egg mortality increased and its dimensions
decreased as the spawning season progressed. Anchovy showed a crepuscular
spawning; early stage eggs were found between 18:00 and 00:30h, local time.
Retention of anchovy larvae within Tejo estuary was also studied. This
retention was mainly associated with the location of the spawning grounds and
coupled with the average conditions of water circulation inside the estuary.
I‑
INTRODUÇÃO. 4
II- MATERIAL E MÉTODOS. 11
2.1‑ Métodos de colheita e engenhos
utilizados. 11
2.2‑ Tratamento laboratorial do material
colhido. 15
2.3‑ Diversidade; métodos de associação
entre estações. 18
2.4‑ Terminologia utilizada (estados
larvares dos peixes). 19
III‑ ALGUMAS CARACTERÍSTICAS DO MEIO. 19
A‑ METEOROLOGIA E HIDROLOGIA. 19
3.1‑ Costa de Peniche 20
3.1.1‑ Meteorologia 20
3.1.2‑
Temperatura das águas 20
3.1.4‑
Correntes 21
3.2‑ Costa de Sines 21
3.2.1‑
Meteorologia 21
3.2.2‑
Temperaturas das águas 21
3.2.3‑
Salinidade e parâmetros químicos 22
3.2.4‑
Correntes 23
3.3‑ Estuário do Tejo. 23
3.3.1‑
Fisiografia 23
3.3.2‑
Largura e profundidade 23
3.3.3‑
Regime de marés 24
3.3.4‑
Temperatura das águas 24
3.3.5‑
Regime de salinidades 24
3.3.6‑
Circulação das águas no estuário. 25
3.4‑ Afloramento costeiro
("upwelling") ao longo da costa portuguesa 25
B‑ CICLO ANUAL DAS POPULAÇÕES FITO‑
E ZOOPLANCTÓNICAS. 26
3.5‑ Costa de Peniche 26
3.5.1‑
Fitoplâncton. 26
3.5.2‑ Zooplâncton. 27
3.6‑ Costa de Sines 28
3.6.1‑
Fitoplâncton. 28
3.6.2‑
Zooplâncton. 28
3.7‑ Estuário do Tejo. 29
3.7.1‑
Fitoplâncton. 29
3.7.2‑
Zooplâncton. 30
IV‑ DIVERSIDADE E ABUNDANCIA DO ICTIOPLANCTON NA
REGIÃO CENTRAL DA COSTA PORTUGUESA E NO ESTUÁRIO DO TEJO. 31
4.1‑ Diversidade 31
4.2‑ Abundância 36
4.3‑ Similaridade entre as estações
amostradas 38
4.4‑ Ecologia da postura de algumas
espécies 40
V‑ ECOLOGIA DA POSTURA E DA FASE PLANCTÓNICA DE SARDINA
PILCHARDUS NA REGIÃO CENTRAL DA COSTA PORTUGUESA. 47
5.1‑ Introdução. 47
5.2‑ Ocorrência anual e distribuição
espacial dos ictioplanctontes.