F. Pereira et al., Flux of decapod larvae and juveniles at a station in the lower Canal de Mira (Ria de Aveiro, Portugal) during one lunar month, INVERTEBR R, 38(3), 2000, pp. 183-206
Emigration and immigration of decapod larvae from estuaries depend on timin
g of larvae occurrence in the water column relative to the tidal, tidal amp
litude and day cycles. The phase relation of these natural cycles varies wi
th tidal regime and geographically, resulting in different time-patterns of
hatching of first stage larvae and of presence of late stage larvae in the
water column. Vertical migration behaviour according to phase of tide also
controls transport inside estuaries. These mechanisms were investigated in
a field study conducted on the northwest coast of Portugal where neap ebb
tides occur during the night around the quarters of the moon. Flux of decap
od larvae through one sampling station was measured during one lunar month
at the Canal de Mira (Ria de Aveiro) in the spring of 1990. The sampling pr
ogramme was comprised of a set of 25-h fixed station studies, separated by
25-h intervals during which no sampling took place. Plankton samples were c
ollected with a pump every hour at three depths. Current velocity and direc
tion at the standard depths, as well as height of the water column, were al
so measured every hour. Hourly instantaneous flux of larvae through a 1-m-w
ide vertical section of the Canal de Mira was calculated for the most abund
ant forms. A total number of 13 combinations of species and larval stages w
ere analyzed, belonging to the families Atelecyclidae, Pirimelidae, Portuni
dae, Pilumnidae, Grapsidae, Palaemonidae, Crangonidae and Thalassinidae. Pa
tterns of net larval flux along the lunar month could be grouped into three
types. Type 1 includes first zoeas that were consistently exported to the
sea. Type 2 comprises late zoeas, megalops and juveniles that were consiste
ntly imported into the estuary. First zoeas that were imported during some
of the 25-h studies but were exported during the others were included in Ty
pe 3, in species of this type import periods appeared to alternate with exp
ort periods according to lunar phase. Flux of Type 1 larvae followed a semi
-lunar pattern. Release activity of Type 1 zoeas took place during the nigh
t and started during neap tides around the quarters of the moon, but maximu
m releases occurred 3-4 h after high tide of average amplitude tides, 3-4 d
ays after the quadratures. These observations agree with the hypothesis tha
t hatching is timed to occur on ebb tides of the largest possible amplitude
so that larvae are easily dispersed from areas with a high density of pred
ator fishes. However, based on other observations on the Portuguese coast,
it cannot be ruled out that hatching might depend on a minimum number of ho
urs of darkness experienced by the females. Larvae included in Type 2 compr
ise forms that may have been retained inside the estuary for the entire lar
val phase, as well as one form that was imported from shelf waters. No semi
-lunar pattern of import was detected in this last form. Fluctuations of ne
t flux observed in Type 3 larvae, as well in other forms that were not incl
uded in any of the types, were more difficult to explain. These larvae were
first zoeas of species belonging to different taxonomic, morphological and
ecological groups and may show a diversity of adaptations to the way of li
fe of the adults. Imports and exports of larvae depended not only on time-p
atterns of abundance, but also on time-patterns of larval vertical distribu
tion. As a general rule, larval stages showed patterns of depth distributio
n that were consistent with vertical migration rhythmic behaviours synchron
ized with the tidal cycle. Though the effect was not always statistically s
ignificant, first-stage larvae were closer to the surface during ebb, espec
ially during the night, enhancing seaward transport.
On the contrary, later zoeal stages, megalops and juveniles were usually cl
oser to the surface during flood, suggesting migration to the water column
during this phase of the tide and landward transport.