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The
European Research Training Network on Fisheries-induced Adaptive
Changes in Exploited Stocks (FishACE) is set up to investigate
the prevalence and consequences of fisheries-induced adaptive changes
in exploited aquatic systems in European waters. This objective
necessitates the development and application of novel methodological
tools for
investigating empirical data, together with the careful construction
of theoretical models suitable for complementing empirical analyses
and evaluating managerial options. At the same time, the network
will provide advance training for a new generation of scientists
who will be educated to tackle the challenges posed by evolutionary
changes in exploited resources.
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Today,
fishing is the dominant source of mortality in most commercially
exploited fish stocks. According to the United Nation’s
Food and Agricultural Organization, world capture fisheries
have reached
a ceiling, with three stocks out of four being maximally
exploited or overexploited. Since all fish species were genetically
adapted
to the environmental conditions experienced prior to intensive
exploitation, the current, drastically altered conditions
cannot possibly leave their life-history patterns unaffected.
In other
words, fishing not only decreases the abundance of fish,
but also changes their genetic composition. This evolutionary
dimension
of fisheries has been overlooked or downplayed for decades,
so that fisheries scientists and managers are just now awakening
to
the formidable risks posed by further unmanaged fisheries-induced
evolution.
This awakening appears to have been facilitated by two independent
developments. First, over the past decade the formerly narrow
focus in fisheries management
has been widened, from the traditional goal of avoiding over-fishing
and securing maximum yield, to an ecosystem perspective recognizing
a much wider range of potential threats to marine biodiversity
and its
constituents. Consequently, in compliance with the FAO Code of
Conduct for Responsible Fisheries, most major fishing nations
in the North Atlantic
have recently agreed to respect a precautionary approach to “conservation,
management and exploitation of living aquatic resources in order
to protect them and preserve the aquatic environment, taking account
of the best
scientific evidence available”.
As a second factor, a suit of scientific advances have suggested that
fisheries-induced evolution is ubiquitous and that, if unmanaged, such
processes may pose serious threats to exploited stocks and to their value
as resources for humankind:
- There is growing recognition that fishing causes severe changes
in the demographic properties of exploited stocks. This
applies, in particular,
to maturation: in most stocks, fish mature earlier today than
they used to only a few decades ago.
- At the same time it has been shown that earlier maturation
may have adverse implications for the reproductive potential
of fish stocks,
not only because large females produce more offspring per unit
of body weight than smaller ones, but also because the size of
females and the
quality of their offspring tend to be positively correlated.
- Furthermore, thanks to recently developed improved statistical
methods, it is becoming increasingly clear that most of the documented
changes
in maturation are indeed evolutionary responses, and not mere
effects of phenotypic plasticity.
- Corroborating theoretical expectations, it has recently been
demonstrated experimentally that size-selective fishing can cause
genetic reductions
in growth that result in a decline of harvestable biomass.
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The scientific agenda proposed for this Research
Training Network will be based on two major lines of analyses: analyses
of empirical data and analyses of theoretical models. The notion
of reaction norms provides the conceptual framework for most of the
empirical studies suggested in this proposal. Reaction norms describe
the different phenotypes produced by a single genotype under different
environmental conditions. Reaction norms themselves are genetically
determined traits that can evolve under natural selection. Estimation
of reaction norms thus provides means to overcome confounding effects
of environmental changes in phenotypic data.
The empirical analyses are based on three main
data sources:
- Individual-level
biological data. Extensive time series of measurements of length,
body weight, age, and maturity stage are available through
scientific surveys and sampling of commercial catches, designed to
give a representative picture on the structure of populations and
for the basis of their rational management.
- Population-level
biological data. Individual-level data are supplemented by data
aggregated to population level estimates. These are often
available from the reports of the expert groups working under the
auspices of the International Council for Exploration of the Sea
(ICES).
- Environmental data. In addition to data on fish populations in
focus here, various types of environmental data are also available,
including abundance estimates of other fish species, phytoplankton
and zooplankton biomasses, temperature, salinity, and movements and
inflows of water masses.
Empirical data will be analysed using an array of statistical
methods. These include the established statistical toolbox, comprising
both classic methods and modern recent developments such generalized
linear models, mixed models and geostatistics. These methods will
be complemented by more specialised, recently developed tools for
estimating reaction norms for age and size at maturation from various
types of often incomplete fisheries data, partly relying on computation-intensive
methods such as bootstrapping and randomisation. Analyses of artificial
data generated with simple population models will be used to validate
analyses tailored for particular data.
For modelling the evolution of reaction norms, three approaches will
be used, each emphasizing different aspects ecological and evolutionary
dynamics:
- Quantitative genetic models. This
time-honoured framework has been developed for the study of quantitative
traits influenced by many genes of small additive effects, and originates
from the context of animal breeding.
- Adaptive dynamics models of reaction norms
represented as function-valued traits. Similarly to certain advanced quantitative
genetics models, these models represent reaction norms as infinite-dimensional
functions, and thereby allow any shape to evolve. However, in
contrast to quantitative genetics models, a realistic description
of the ecological setting is emphasised. In order to achieve
this, genetic detail is sacrificed.
- Eco-genetic
models. This novel approach aims at combining a realistic description
of the ecological
setting as well as population structure with a description of
genetic detail at a level that still allows predicting the rate
of evolutionary change.
More specific evolutionary repercussions of exploitation
will be studied based on two further model classes:
- Evolutionary food web models. The type and strength
of interactions between species pairs, as well as the distribution
of interactions
in the food web as a whole influences the evolutionary effects
of fishing. Food web models are characterized by their topology
and distribution of interaction strengths among the nodes in the
web.
- Evolutionary energy allocation models. Energetic
trade-offs constrain evolution of all key life-history traits. Energy
allocation
models consider the ecological and evolutionary dynamics of energy
allocations to growth, reproduction, and maintenance. In addition,
behavioural sub-models for describing foraging behaviour can be
included.
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