Ms. L. (Laure) Pecquerie
Introduction
Anchovy, Engraulis encrasicolus, is an important target of
coastal fisheries, and the present stocks are rapidly declining. The
species is also prey for a large number of marine predators. Little is
known about details of the dynamics of its food, food intake,
migration, and energetics. An increase of knowledge of its energetics
could help in developing responsible fisheries management for this
species. The purpose of the project is to contribute to this knowledge
by application of the DEB theory to anchovy data. More in particular
the aims are to
- connect anchovy dynamics to spatially explicit models for lower
trophic levels
- spatialise life cycle energetics
- predict location and timing of spawning
Spatial distribution of the population
During May or June surveys (spawning season), the population is mostly
distributed in the south of the Bay of Biscay, with the small and big
individuals distributed in- and offshore respectively. The age classes
are intermingled. The explanation is possibly because swimming speeds
depends on size, not on age, and food density has a spatial structure.
There is some evidence from the spatial distribution of the French
pelagic fishery catches that the population is distributed in the
north of the Bay of Biscay at the end of summer and in autumn. There
is one study of the size of the fishes in the catches in the North in
July-September 2000, and there is also a distribution of the fishes
according to their size, with the big ones being more in the north.
Autumn surveys (2003 and 2005) showed that big Age 0 individuals
(juveniles) are integrated in the schools of the adults.
Description of the growth data
Otoliths show opaque regions that indicate growth conditions and
hyaline ones that correspond to growth arrest. Otoliths from juvenile
even show a daily ring pattern. The location of these rings has
information about the growth of the individuals. Growth generally
ceases during the low food densities in winter times. During spawning
the otolith tends to grow without aragonit deposits and for some
individuals, a spawning ring is identifiable (false winter ring). In a
similar way as for estimating annual growth, it was possible to
estimate the growth before and after spawning on those individuals
that showed a spawning ring: growth before spawning was that measured
between the previous winter ring and the spawning ring and the growth
after spawning was that measured between the spawning ring and the
next winter ring. Large individuals suffer from intensive harvesting
by fisheries, and are rare. Otolith data indicate:
- All individuals have at least one annulus on their otoliths :
seasonal growth, age of the fishes older than on year
- There is a linear relationship between growth between the first
and second winter and the growth before the first winter
- Age 1: All individuals have an otolith with an opaque border
in May: growth has resumed before the catch of the individual
- Age 2: All individuals have an otolith with a hyalin border in
May: growth has not resumed
- Age 3: All individuals present an annual growth during age 2: as
Age 2 individuals have not resumed growth in May, they resume growth
after May but before the following winter because we observed a
third winter ring
- 8 and 50 % of age 2 individuals typically have a spawning
ring. Petitgas and Grellier (2003) mention individuals that have a
spawning ring between the first and the second winter ring. It
indicates that their growth rate decreased significantly sometimes
between two winters.
Understanding anchovy growth & reproduction pattern
It turns out to be possible to find a set of parameters of the
standard DEB model that reproduces the mean length at age of the
population. For this purpose available data for anchovy had to be
supplemented with energetics studies on other fish species, such as flatfish. With
appropriate rules for the handling of the reproduction buffer a
realistic reproduction (i.e. spawning) pattern can be found: Hatching
occurs during spring and summer in 10-20 batches. Juveniles grow until
the next spring. At the end of winter (they are Age 1 fishes), they
still allocate energy to growth.
The fact that equally sized Age 1 fish resume growth before
spawning, while Age 2 and 3 fish do not, is more difficult to
understand in the context of the DEB theory. One possibility is that
allocation to reproduction varies with size and/or age. Such a
tradeoff between growth and reproduction is, however, inconsistent
with the DEB theory, in which allocation to growth competes with that
to somatic maintenance, not with reproduction. Another possibility is
the variation of parameter values among individuals. The capacity to
grow reduces with the size of the individual relative to its maximum
size, and so with age. A scatter in maximum sizes might explain why
size ranges of first and second year individual overlap. Body size
scaling arguments link covariation of several parameters, which needs
to be studied on their applicability to anchovy. A third possibility
is that first and other year individuals experience different food
densities, for instance by choosing different food items, or being in
different locations.
In terms of newly developed methodology to analyse energetics data
in the context of DEB theory, we developed a model for the growth and
the colour of otoliths with the aim to translate otolith data into
trajectories of temperature and food densities as experienced by that
individual fish. To this end we separate out the various contributions
of assimilation, dissipation and body growth to otolith growth and
otolith chemical composition (i.e. colour). This methodology has a much
wider field of application, even in archeology and
paleontology. Secondly we took the changes in shape into account in
the early juvenile stage. By assuming that this stage behaves as a
V1-morph, we extended the standard DEB model with a single
parameter. This extended version can capture the sigmoid length-to-age
curves that are typical for a wider class of fish data.
The further analysis of the various possibilities to interpret size
differences and growth potentials and the necessity to provide
estimates for realistic food density and temperature trajectories
involves a link with hydrodynamics models that include site-specific
primary (and secondary) production. We use the Mars 3D-model for this
purpose and constructed environmental forcing scenarios that lead to
new interpretations of fish growth data in the context of DEB theory.
This is the symposium that closed my project
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