Work Packages 1 • 2 • 3 • 4 • 5

Objectives

1. Use statistical methods to disentangle demographic, plastic (i.e., direct environmental effects, including compensatory growth) and genetic components of phenotypic change in time series of biological data, refining the methods to best serve the specific case studies when necessary.
2. Widen the empirical basis of evidence of fisheries-induced evolution to new stocks, covering different ecosystems, geographic areas, life-history types, and fisheries.
3. Analyze fisheries-induced adaptive changes in traits for which current empirical evidence of fisheries-induced evolution, or absence thereof, is scarce (reproductive effort) or lacking (growth, fecundity).
4. Synthesize the empirical results on fisheries-induced life-history changes to reveal commonalities in responses, depending on the characteristics of fish stocks and fisheries.
5. Interface with WP2 on genetic approaches, explicitly linking phenotypic and genetic data, with WP3 on modelling approaches, and with WP4 on providing the integrative analyses for the selected focal stocks, cod and sole.

Description of work

General background

The notion of fisheries-induced evolution derives from the simple fact that if there is some genetic variation in traits that allow individuals to increase their contribution to next generations – by timing their life cycle so as to reproduce as much as possible before the mortal contact with the fisheries, by sacrificing future reproductive output for current reproduction by allocating less energy to growth and maintenance, or by decreasing their likelihood of being captured – then those genotypes that are best in coping with fishing tend to increase in relative numbers. It is likely that, through such evolution, stock attributes directly relevant for management, such as sustainable yield and recruitment, are at stake.
WP1 aims at broadening our understanding of the extent of evolutionary changes in exploited aquatic systems. Although evolutionary changes, by definition, imply genetic change, the genetics of macroscopic traits in fish are still largely unknown – providing the rationale for one of the key topics to be pursued in WP2. Therefore, empirical detection of fisheries-induced adaptive changes largely has to be based on phenotypic data. Furthermore, it is the phenotypes that matter for stock dynamics. Changes in phenotypic measurements, however, may reflect changes in demographic composition, plastic responses to environmental changes, and genetic changes. Whenever phenotypic observations may be confounded by an uncontrolled environmental and demographic effect – which, in principle, is always the case for data that do not originate from controlled experiments (inclusive all fisheries data) – all plausible environmental effects should be accounted for.
In this context, the concept of reaction norms is helpful. 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 for overcoming confounding effects of environmental changes in phenotypic data. The explicit consideration of the fact that phenotypic evolution has to be investigated in the context of reaction norms has fuelled the recent rise of studies of fisheries-induced adaptive changes into a new and vigorous branch of life-history theory – with real applications of uncontested value. This has been made possible by the development of a new, revised concept of probabilistic reaction norms for age and size at maturation (Heino, Dieckmann, and Godø, 2002, Evolution 56: 669-678) and by a whole set of new estimation methods that are needed for coping with various kinds of real data. These and similar reaction-norm-based approaches will form a cornerstone of empirical investigations in WP1.
The new research under WP1 benefits from other ongoing research efforts; for details see Section 5.5. In particular, fisheries-induced evolution is the focus of the Research Training Network FishACE (Fisheries-induced adaptive changes in exploited stocks), through which both empirical and model-based approaches are developed and applied.

Task 1.1: Fisheries-induced changes in Atlantic cod in the Barents Sea

The Barents Sea stock of Atlantic cod, the so-called Northeast Arctic cod, currently supports the biggest remaining cod fishery. It is also one of the best-studied fish stocks, with a long history of research and data collection. The aim of this task is to utilize individual life-history measurements collected since 1932 to 1) quantify the changes in age and size at maturation and 2) understand the likely nature of changes in these traits. A crucial question with regard to the latter aim is whether earlier maturation in this stock could be explained by compensatory growth responses alone – i.e., by the fact that faster individual growth resulting in earlier maturation is expected when a stock’s biomass is reduced through fishing – or if genetic changes in this stock’s maturation tendency have occurred as well.
Analyses will be based on estimating probabilistic reaction norms for age and size at maturation. This method offers the best tool currently available for disentangling, based on phenotypic data, the effects of demographic structure, of growth-related phenotypic plasticity arising from, e.g., variations in feeding regime and temperature, and of genetic changes in maturation tendency. Initial analyses based on data comprising 74,261 individually measured fish from 1932-1998 have already indicated a strong downward displacement of maturation reaction norms, supporting the hypothesis that the changes in age and size at maturation are partially caused by fisheries-induced genetic changes. The task here is to consolidate these analyses, including data collected after 1998.
The empirical analyses conducted under this task will be linked with analyses of molecular genetic variation based on historic tissue material (Tasks 2.2 and 2.3) as well as with model-based efforts under WP3 (Task 3.6 and 3.8), including a stock-specific focus on the economic dimension of fisheries-induced evolution (Task 3.8). The results will also contribute to the integrated case study on Atlantic cod (Task 4.1).

Task 1.2: Fisheries-induced changes in Atlantic cod in the Northwest Atlantic

Cod stocks in the Northwest Atlantic are sadly famous for the dramatic reductions in abundance, including the collapse of northern cod to commercial extinction. While adverse environmental conditions contributed to these declines, excessive fishing pressure was undoubtedly the main culprit. Strong fisheries-induced changes are thus expected. Recent analyses have already shown that maturation schedules of cod stocks off Newfoundland-Labrador shifted significantly towards earlier maturation from the late 1970s to the early 1990s, when directed fisheries on these stocks were put under moratoria.
The task here is threefold: 1) Utilizing indices of individual body condition, we want to understand the role body reserves (condition) play in maturation of cod. This is important both for refining the picture of phenotypically plastic and genetic components in the documented trends in maturation, and for improving our understanding of maturation process in this important species. The cod stocks off Newfoundland-Labrador (northern, Southern Grand Bank and St. Pierre Bank stocks – NAFO divisions 2J3KL, 3NO, and 3Ps, respectively) are ideally suited for such analysis as a large dataset of measurements of liver weight, gutted weight, and total body weight are available from DFO surveys that were initiated in 1970s. 2) Changes in reproductive effort in Atlantic cod off Newfoundland-Labrador are documented. Such changes are predicted by life-history theory. Here we aim at utilizing measurements of gonad weights from cod from Southern Grand Bank and St. Pierre Bank stocks (these stocks were selected as they are surveyed in spring, during the spawning season), collected during the aforementioned surveys to measure time trends in size-specific reproductive effort, expressed as gonadosomatic index (gonad weight/total weight, or gonad weight/gutted weight). This is an important parameter for reproductive potential of stock, and documenting and understanding changes in it are crucial. Dedicated research effort for these two sub-tasks on Canadian cod is already secured through a PhD student working at IMR with FishACE funding; complementary funding is critically needed for the integration of DFO to the project. 3) We will also analyze fisheries-induced changes in maturation of Flemish cap cod (NAFO division 3M), surveyed by DFO (1978-85) and EU (1988 onwards, data provided by IIM-CSIC). These data are particularly valuable as observations on individual maturation stages are based on histological methods: these not only allow accurate separation of immature and mature individuals (routinely used macroscopic examination is prone to some errors), but also separation of newly-maturing individuals from those that have spawned during some earlier seasons. This allows characterization of maturation trends with unusual accuracy, and also to evaluate the statistical methods that are used in other stocks where newly-matured individuals cannot be distinguished from those which matured earlier.
The results will also contribute to the integrated case study on Atlantic cod (Task 4.1).

Task 1.3: Fisheries-induced changes in North Sea gadoids

Demersal stocks in the North Sea have been exposed to high levels of exploitation for decades, and several stocks are currently at very low levels and under severe fishing restrictions (cod, Norway pout), while several others are showing poor recruitment (e.g., haddock). Under these conditions, strong evolutionary responses are possible, which may also have direct implications for the capacity of these stocks to recover. The North Sea is also an area where sampling programmes both through surveys and sampling of landings have been routinely conducted for decades, making it a natural laboratory for studying fisheries-induced change.
The goal here is to assess changes in life-history traits in the major gadoids in the North Sea: cod, haddock, whiting, Norway pout, and saithe (the other major group of demersal fish, the flatfish, will be dealt under Task 1.4). The question is approached at two fronts: 1) The first approach involves utilizing the extensive database from the International Bottom Trawl Surveys (IBTS) to assess changes in maturation. Several partners here contribute to the survey (Ifremer, FRS, IMR, IMARES, DIFRES) and thus are in good position to use these data effectively while acknowledging the limitations. The data are considered to be good from 1983 onwards when coverage was made spatially homogeneous (it might be also possible to utilize earlier data). Several analyses have already indicated trends towards earlier maturation, although the nature of these changes remains unclear. Here we will provide an overview of changes in maturation in North Sea gadoids by using reaction norms for age and size maturation to disentangle phenotypic plasticity and genetically-determined maturation tendency. 2) Cod and haddock are also studied in more detail, benefiting from the experience FRS has developed on reproductive biology of these two stocks. In particular, the goal is to i) look at maturation at finer geographical and biological scales (inshore and offshore stock components) and ii) analyze changes in other reproductive traits, namely reproductive effort (measured as gonadosomatic index) and relative fecundity in relation to condition. These analyses will rely on other data sources (samples from commercial landings), including some material from the west coast of Scotland that FRS has collected. Together, these analyses will offer a comprehensive overview of life-history changes in North Sea gadoids across several species and several traits.
The results on cod will also contribute to the corresponding integrated case study (Task 4.1).


Task 1.4: Fisheries-induced changes in flatfish

Substantial changes in growth, maturation, and reproductive investment have been documented in sole and plaice in the North Sea and the eastern English Channel during the 20th century. Plaice became heavily exploited at the beginning of the 20th century, whereas sole only became heavily exploited in the early 1960s. In female plaice, the changes in maturation and reproductive investment were shown to be partly due to fisheries-induced adaptive change. In sole, the changes have so far been interpreted as being related to density-dependent effects and an increase in food availability, but the possibility of fisheries-induced adaptations has not been rigorously considered.
In this task, the objective is to elucidate life history changes in sole and plaice: 1) In sole, observed changes in growth, maturation and reproductive investment will be analyzed using the biological monitoring data collected monthly since 1958. Dedicated research effort for this sub-task is already secured through a PhD student working at IMARES with FishACE funding. 2) In plaice, understanding of maturation changes in male plaice is still lacking as male plaice mostly mature under the legal landing size whereas the most comprehensive plaice data come from landings sampling programme. The analyses have therefore to rely on survey data collected on male plaice during two breeding seasons in 1980s, together with new data to be collected in 2007-2008. In addition, data that are collected in discard studies (observers on commercial vessels) could provide further insights. A major step forward will be achieved by the analysis of the covariance between growth, maturation, and reproduction by estimating the growth history, the onset of sexual maturity, together with the reproductive investment of individual fish.
The results will also contribute to the integrated case study on sole (Task 4.2).

Task 1.5: Fisheries-induced changes in small pelagics

Most of the empirical research on fisheries induced-changes has until now focused on demersal fishes characterized by relatively late maturation (prior to fisheries-induced changes) and long potential life span (the exception is Norwegian spring spawning herring, which is a pelagic species with these life history characteristics). One would expect that such species are more vulnerable to fisheries-induced changes in maturation as increasing fishing mortality very quickly erodes the benefits of late maturation. Higher levels of natural mortality, and consequently earlier maturation and shorter potential life span, on the other hand, characterize many small pelagic species. This may make them less susceptible to fisheries-induced life-history evolution, although the ensuing short generation time would make selection responses occur faster than in “slower” species.
This hypothesis is tested here with two pelagic stocks, sardine in the Portuguese coast and Barents Sea capelin: 1) Sardine is a commercially important clupeoid distributed in the north-eastern Atlantic from the North Sea to Senegal and throughout most of the Mediterranean Sea. Along the Portuguese coast, sardine mature at age 1-2 years, with life span up to 9-10 years. Recent analyses have shown a trend toward maturation at smaller size. This could be a purely phenotypic response as condition has at the same time increased. However, the data have not yet been analyzed with methods that would allow disentangling environmental effects on maturation and underlying genetically determined maturation tendency. The data available include samples from the commercial catch since the early 1980's complemented with data from surveys conducted every year since 1996, covering the north-western and southern waters of the Portuguese coast. 2) Capelin are a key element in the food chain of the Barents Sea, being the main forage fish for both marine mammals and the economically most valuable stock in the area, Northeast arctic cod, and utilizing themselves the abundant zooplankton production of the area. Capelin in the Barents Sea mature at ages 3-5 years (depending on growth consitions) and then die after spawning. This life history makes the capelin stock susceptible to strong fluctuations depending on recruitment and natural and fishing mortality – the stock has collapsed thrice during the last 30 years. Here we would analyze if the changes in age at maturation can be fully explained with growth-related phenotypic plasticity, or if residual components that can be attributed to fisheries-induced evolution remain. The analyses will be based on data from surveys conducted every year in the Barents Sea by IMR since 1973.

Task 1.6: Fisheries-induced changes in Atlantic salmon

Salmon spawn in rivers, usually in late autumn, and only some 10% of eggs survive to become young fish the following summer. Depending on temperatures and food supply, it takes between 6 months and five years longer for the young fish (parr) to grow enough to become smolts. Smolts migrate to the sea, where they feed and grow for one or more years before returning to their home river. Breeding success is correlated to size (and thus to age) through fecundity as well as mate selection. Wild Atlantic salmon rarely spawn more than once.
Fisheries-induced evolutionary changes could be manifest at several stages. In freshwater, male parr may become precociously mature, achieving breeding success without going to sea; the age of smoltification may become younger, avoiding additional years of freshwater mortality. At sea increased fishing mortality could favour one sea-winter (grilse) life-cycle over multi-sea-winter (MSW) strategies. Size-selective and season-specific human harvesting can affect salmon both offshore and when the adults return to the coast and their natal rivers. In Ireland and Scotland the grilse and MSW adults return at times separated by some months.
Although the main offshore fishery at Greenland Bank has now been closed, harvesting in the Baltic Sea feeding areas still occurs. This difference creates contrasting predictions on fisheries-induced evolution within and between populations: increased freshwater mortality will favour more precocious male parr, faster parr growth, and earlier smoltification. Human harvesting in the marine feeding areas, together with the recent increases in overall marine mortality, favour earlier adult sexual maturation, especially grilse life-cycles. Human exploitation of returning adults may also be size and run-timing selective, as grilse and MSW fish, and within those classes fish of different sizes, do not return randomly with respect to the fishing seasons, and are thus differentially exploited. In Scotland and Ireland data exist across both the onset and cessation of such human fishery activities.
Trends in several of these parameters have been documented for several stocks, i.e., increased fraction of 1 sea-winter salmon returning to spawn. In addition, data show that adult return run-timing has changed. The task here is to utilize the reaction norm approach to investigate the nature of life history changes in salmon: 1) In the Baltic Sea, along the coast of the Gulf of Bothnia, there is a mixed stock fishery on Atlantic salmon in their spawning migration. FGFRI has collected tagging data from this fishery. Stock of origin is known from tagging of smolts released from hatcheries. For tagged individuals, also length at release and age and length at recapture are known. Information on the immature part of the stock in the feeding area can similarly be based on fishery returns of tags. 2) MI has long-term angling datasets (since the mid 1920s) on returning salmon from several Irish rivers (Blackwater, Owenduff, and Newport). These describe the size of the fish caught with a monthly resolution from February to September. The multi-modality of size distribution will be used to reconstruct age of caught individuals before studying trends in age and size at maturation, by applying the probabilistic maturation reaction norm approach. The latter will require reconstructing the immature part of the stock through indirect methods. 3) FRS hosts extensive datasets on salmon from Scottish rivers. In particular a national catch data-base for rod caught adult salmon from the 1950s is supplemented by more detailed population data, most notably from the Girnock Burn and the River North Esk, both including the immature stages, since the 1970s. These can reveal trends in early life history (parr growth, proportions of precocious male parr, age at smoltification) as well as in adults (size and age, phenology).
The empirical studies on salmon will be linked to the model-based approaches pursued under Task 3.6.

Task 1.7: Fisheries-induced changes in landlocked salmonids

Arctic charr and whitefish are the main targets of inland fisheries in high latitude lakes, providing an important resource for both recreational and subsistence fisheries. These populations are exposed to selective harvesting, typically culling large adults foraging near shore. However, salmonids have particularly plastic life histories, which make detecting the expected fishery induced evolution in fecundity, age at maturity and somatic growth difficult. To disentangle the environmental and genetic components of life history variation in our long-term series on harvested land-locked salmonids, the reaction norm approach can be used. The advantage of lakes is that, being relatively closed systems with modest spatial dimensions, collecting representative data on the environment and exploitation pressure is potentially facilitated.
Here the goal is to study life history changes in charr and whitefish in three lakes in northern Norway where long time series collected by UT provide unusual opportunities for detecting life history evolution. In particular, access to detailed environmental information, spanning over 25 years and including temperature, food availability, and fish abundance, diet and parasite load, will facilitate the task of detecting evolutionary changes in fish life histories: 1) For Arctic charr, we will compare the temporal changes in fecundity, age and size at first reproduction, and somatic growth in lakes Takvatn and Fjellfroskvatn. These two neighbouring lakes harbour charr populations of common origin, but exposed to different harvesting regimes for over three decades, providing a unique field experiment on fishery induced evolution. 2) For whitefish, we will analyze the long-term data from lake Stuorajavri, to detect evolutionary changes in life histories along the lines of the Arctic charr study. The extensive data on life history and environment are accompanied by quantitative information on harvesting regimes, monitored throughout the long-term studies.
Under Task 3.6, an eco-genetic model supporting the analyses of empirical data will be developed.

Task 1.8: Comparative analysis and synthesis

Theoretically, evolutionary responses are proportional to 1) heritability of traits under selection and 2) selection differentials (the difference between mean phenotype before and after selection). The latter is determined by characteristics of the fish stock in interaction with the selection pattern of the fishery and the overall level of fishing mortality imposed by the fishery. The goal here is to utilize the growing body of empirical case studies to seek for common patterns in evolutionary responses in relation to the characteristics of fish stocks and fisheries, guided by our theoretical understanding of fisheries-induced evolution and population ecology in general. The salient questions include the following:

• Are short-lived species naturally adapted to high mortality and therefore less susceptible to fisheries-induced evolutionary changes?
• Fishing primarily on spawning grounds (Norwegian spring-spawning herring, and the historic exploitation pattern for Northeast Arctic cod) is theoretically expected to favour delayed maturation and high reproductive investment thereafter. Can real data substantiate this prediction?
• Do stocks with semelparous life history (death after first reproduction: Barents Sea capelin, in practice also Atlantic salmon) differ from those with iteroparous life history (all other studied species) in their sensitivity to fisheries-induced evolution?
• Do responses to fisheries-induced selection differ between males and females?
• Does the past fishing history have some influence on the response to recent fisheries-induced selection pressures? Can stocks with very long histories of fishing be considered “domesticated”?
• Fisheries selectivity: fishery on mature fish only vs. fishery on immature and mature fish, size limits
• Does understanding of life history changes in areas with few dominant species with strong linkages between them (e.g., capelin and cod in the Barents Sea, herring, sprat and cod in the Baltic) call for taking inter-specific interactions into account?
• Can observed life history changes fully be accounted for by compensatory responses and fisheries-induced evolution, or do large-scale environmental changes also play a significant role?

Objectives

The overall objective is to elucidate fisheries-induced adaptive genetic changes; both at the molecular level and in terms of quantitative genetics, in life-history traits relevant for the demography and productivity of exploited fish populations
Specific objectives:

1. Investigate temporal changes in neutral genetic variation using contemporary and historical tissue samples (otoliths), thereby providing knowledge on fisheries-induced changes in effective population sizes and migration rates among populations.
2. Explore temporal changes in the genetic make-up at candidate genes for important life-history traits, such as growth and size and age at maturation, to provide direct evidence of ongoing selective changes at the DNA level in genes of adaptive importance.
3. Provide evidence on selective quantitative genetic changes from wild exploited populations
4. Compare neutral and adaptive genetic changes over time, thereby illuminating the relative roles of random changes in the genetic composition caused by low effective population size and directional genetic changes due to selection.
5. Link genotypic information from candidate genes with phenotypic and quantitative genetic information at the individual level, to identify direct links between DNA variation and phenotype.
6. Unequivocally establish fishing as the main selective agent responsible for the observed adaptive genetic changes in wild exploited populations by assessing the agreement of fisheries-induced selection gradient (computed from long-term population ecological data and known historical fishing patterns) and the direction of observed selective genetic changes through the link between genotype and phenotype.
7. Evaluate the role of alternative evolutionary drivers to fishing, such as global warming by making use of available long-term time series of environmental variables for wild exploited populations and by subjecting the model species to alternative drivers (fisheries, temperature) and investigating its quantitative and qualitative (candidate gene) response.

Description of work

General background

Recent studies have strongly indicated fisheries-induced evolution of important life-history traits, such as age and size at maturation, growth or reproductive effort in exploited fish populations. Most evidence is based on the analysis of phenotypic data showing changes in maturation reaction norms, i.e. an individual’s propensity to mature at a certain size and age, which for many species and populations have changed markedly over historical times corresponding to the intensity of fishing. However, no “smoking gun” of fisheries-induced evolution at the DNA level or in terms of quantitative genetic parameters has been provided. To directly study genetic changes induced by fisheries retrospective analysis of DNA from historical tissue collections such as scales and otoliths in wild fish populations subjected to harvesting can be applied. This approach allows for direct identification of molecular and/or quantitative genetic changes in commercially exploited populations, which have been subject to intense fishing in the wild.
In WP2, we aim to develop this approach. Using historical tissue collections, we will investigate temporal molecular and quantitative genetic changes in two commercially exploited fish species: Atlantic cod (Gadus morhua) and common sole (Solea solea). The two species have been chosen based on their high economic and ecosystem importance, their large combined geographical coverage from Arctic to Mediterranean waters, their “classical” marine life-history, their known intense long-term exploitation patterns, and the availability of historical biological data and tissue samples for genetic analysis together with long-term phenotypic and population ecological data to corroborate potential fisheries-induced selection at the genetic level. Samples will be collected from several populations spanning an array of physical and biological environments and exploitation regimes.

Task 2.1: Biological samples collection

Recent studies have shown that historical tissue collections from fish such as scales and otoliths can be used as a source of DNA to investigate temporal changes in the genetic make-up of populations. Here, we will collect historical otolith samples for cod and sole from a number of European fisheries institutions. To maximize the likelihood of detecting fisheries-induced genetic changes, we aim at collecting samples from populations which conform to two prerequisites: the existence of long-term phenotypic trends in age and size at maturation and/or growth, and the evidence of some concordant fisheries-induced selection gradient affecting these traits. From a practical point of view, this requires that we aim at employing samples which have been, or can be, used as a source of phenotypic data (otolith reading) for the case studies on cod and sole in WP1, thus forming the basis of the WP4 “oversight of integrated case studies”. At the same time, the aim is to focus on population samples with very detailed time series allowing for resolution of changes in exploitation pattern and/or other potential drivers of evolution. For cod, samples are already available for the North Sea (DIFRES, Denmark), the Baltic Sea (DIFRES, Denmark and BFAF, Germany), and Northeast Arctic (IMR, Norway) For sole, collections are available for the English Channel and the North Sea (IMARES, The Netherlands and Ifremer, France), and the Danish Belt Sea (DIFRES, Denmark). Identification of other relevant samples is ongoing. A database of available historical samples for cod and sole from European fisheries institutions will be produced under this task.

Task 2.2: Baseline neutral genetic variation

Temporal studies of non-coding DNA loci can be used to evaluate genetic changes in exploited fish populations caused by neutral processes, i.e. genetic drift and migration. Exploitation will inevitably lead to a reduction in the census population size and most likely also to a reduction in the genetically effective population size (Ne), which, in popular terms, can be expressed as the number of individuals successfully propagating their genes to the next generation. Such reduction will favor temporal changes in the genetic composition of exploited populations due to purely random processes, i.e. genetic drift. Likewise, the reduction in census size can lead to altered migration patterns, such as increased isolation among populations by narrowing their range, or increased migration caused by the invasion of available habitats by populations subject to less – or less sensitive to – exploitation. Accordingly, by studying historical cod and sole samples, changes in the level and distribution of neutral genetic variation over time can be evaluated and, then, related to fishing regime history. This knowledge of neutral processes is a prerequisite as a baseline against which changes observed at candidate gene loci and quantitative genetic changes for life-history traits (see task 2.5) can be evaluated to detect selective genetic changes. Non-coding genetic markers will be employed to establish a neutral background, i.e. microsatellites and neutral SNPs (Single Nucleotide Polymorphisms). The choice of markers depends critically on the available genomic resources for the species in question. For cod, we will employ both microsatellites (10-20) and neutral SNPs (20-40) available from the literature and from a database of recently developed markers (Dr. Madjid Delghandi, Norwegian Institute of Fisheries and Aquaculture Research, Tromsø, Norway). For sole, we will employ microsatellites (10-20) available from the literature. Additionally, putative neutral SNPs will be developed from the outcome of a genomic project on Solea senegalensis (Pleurogene), which will be available in the next year. For both cod and sole, we aim at analyzing 50 individuals from each temporal sample.


Task 2.3: Genetic variation in candidate genes

Candidate genes are genes of known function suspected to have a large influence on a given trait. They can be structural genes or genes involved in a physiological process. The working hypothesis is that a molecular polymorphism is related to phenotypic variation. In the case of fisheries-induced selection, size and age at maturation and growth are among the traits expected and observed to be mostly affected. Accordingly, we aim at studying the evolutionary trajectories of candidate genes for growth and maturation. Two classes of genes are expected to play a major role: genes from the Growth-axis and the Brain-Pituitary-Gonad axis, the latter being involved in puberty determinism. We aim at screening temporal genetic changes in terms of modifications of allele frequencies for a number of these candidate genes for both cod and sole including (i) for the Growth axis, genes involved in the production of the Growth Hormone (GH), the Insulin Growth Factor (IGF), and somatolactin, and, (ii) for the Brain-Pituitary-Gonad axis, genes related to the secretion of the Follicle-Stimulating Hormone (FSH), the Luteinizing Hormone (LH), and aromatase. Since the development of genetic markers and screening of temporal population samples are very time consuming, we aim at screening candidate genes in succession. Depending on the detection of selection, the molecular analysis will then be pursued in more detail. Therefore, the total number of candidate genes that will be finely screened actually depends on the occurrence of detectable selection. For cod, genomic sequences of target genes are/will be available before the project start and genomic sequences of four additional candidate genes for the Brain-Pituitary-Gonad axis are already available (Dr. Christian Mittelholzer, Institute of Marine Research, Bergen, Norway). For sole, the available sequences from cod and sequences from other flatfish species will be used as a starting point for cloning sole-specific genomic sequences.

Task 2.4: Quantitative genetic variation

Quantitative genetics offer an alternative to candidate genes for establishing temporal changes in the genetic composition of populations. One advantage lies in that the relationship between genetic polymorphism and phenotypic variation does not need to be established because it is implicit, but one disadvantage arises from the need to access the relatedness between individuals to carry out the analysis, which, for wild populations, proves much more difficult than accessing molecular data. Still, recent studies have shown that quantitative genetic differentiation between wild populations can be estimated by combining molecular data on neutral genetic differentiation and data on phenotypic variation. Some limitations, however, are that the methods developed so far apply to spatial, and not temporal, differentiation and cannot account for other factors than genetics, such as environmental features, that may be responsible for part or all of the observed phenotypic differentiation. In this task, we aim at studying temporal quantitative genetic changes in the wild exploited populations of cod and sole studied at the molecular level. Regarding wild populations, based on existing methods for spatial quantitative genetic differentiation, new methods will be developed for estimating temporal quantitative genetic changes by combining data on temporal neutral genetic differentiation and data on temporal phenotypic changes, while accounting for environmental covariates that may be responsible for phenotypic changes at the same time. In addition, some of these methods will have to be specifically tailored to the analysis of probabilistic maturation reaction norms (WP1) which are in essence non-metric traits contrary to traits classically considered in quantitative genetics analysis. These methods will then be applied to neutral molecular data obtained from Task 2.2 and phenotypic data available from WP1 to estimate temporal quantitative genetic changes in growth, age and size at maturation, and probabilistic maturation reaction norms for both sole and cod wild populations.
In addition to the tracking of temporal quantitative genetic changes that will parallel the screening of temporal changes in candidate genes, this task will develop further methods to estimate within-population or within-generation quantitative genetic parameters (additive genetic variance, breeding values) for probabilistic maturation reaction norms, these parameters being critically needed to calibrate operational eco-genetic models for predicting fisheries-induced evolution (WP3). Probabilistic maturation reaction norms have indeed two specificities that make classical quantitative genetic methods obsolete: they are not metric but probabilistic traits, and they are not scalar but function-valued traits, i.e. traits described as a function instead of a single value. Identification of relevant existing datasets for exploited fish species to which the methods developed could be applied is ongoing (aquaculture cod from the Norwegian Institute of Fisheries and Aquaculture Research, Dr. Madjid Delghandi).

Task 2.5: Comparative analysis of neutral and adaptive genetic variation

Both neutral and adaptive genetic variation is influenced by random genetic drift, which in turn, is determined by the size of the genetically effective population size. Accordingly, to quantify the selective effects of fisheries on adaptive genetic variation, it is necessary to separate out random changes due to genetic drift from adaptive changes due to selection by comparing temporal changes in supposedly neutral and adaptive genetic variation. The subject of detection of selection has attracted a lot of attention recently, and a number of methods and software are available for identification of outlier loci not conforming to neutral expectations, i.e. loci under selection, for molecular data or to assess significant quantitative genetic differentiation relative to neutral genetic differentiation for quantitative genetic data. These methods are particularly well suited for classical marine organisms, including fish, where large effective population sizes are expected and, consequently, the drift “noise” relatively small. Further, within this project (task 3.6) model-based expectations of the effect of fisheries-induced selection on neutral and adaptive genetic variation will be generated. A comparative temporal analysis of neutral (microsatellite, SNP) and adaptive genetic variation (candidate genes and quantitative genetic differentiation) will be conducted for the natural populations of cod and sole
From the comparative analysis under Task 2.5., several putative candidate genes influenced by selection and some selective quantitative genetic changes are likely to be identified. However, further progress to relate selection at the genetic level unequivocally to fishing as the selective agent would need to (i) establish links between individual genotypes and the observed phenotype (this applies to candidate genes only, since in quantitative genetic analysis the link with the phenotype is implicit), (ii) then assess the agreement of the fisheries-induced selection gradient and the direction of genetic changes and (iii) finally rule out other potential selective agents than fishing. The last two tasks of this work package aim at unravelling these relationships.

Task 2.6: Linking adaptive genetic variation and phenotypic variation

Establishing links between individual genotypes and the observed phenotype may prove rather difficult, especially for wild populations. This requires that the actual genomic site under selection is identified, which is not a trivial task. First of all selection is not necessarily targeting different protein products caused by point mutations (SNPs) in the actual candidate gene, but could also be at gene regulation in the promoter (or other) region(s) situated several hundred (thousand) bases away from the gene. Secondly, the traits investigated are quantitative traits, most likely affected by many genes, thereby making the likely contribution to the total phenotypic change from selection at each single locus small. We aim to correlate phenotypic trait and single gene variation for wild samples of cod and sole, although relationships may to some extent be obscured by environmental and genetic noise. However, one or a few of the genes investigated could be a “major gene” explaining a large part of the phenotypic variance. In addition for cod we will try to access family samples from aquaculture to firmly establish the link between qualitative genetic variation and the phenotypic trait considered.

Task 2.7: Causal analysis

The ultimate step of the genetic work package consists in assessing the agreement between fisheries-induced selection and the direction of molecular and/or quantitative genetic changes and ruling out other potential selective agent. For wild cod and sole populations, the fisheries-induced selection gradient acting at the phenotypic level will be computed using the known history of fishing patterns and long-term population ecological data available from European fisheries institutions. It will then, be compared to the direction of selective genetic changes either at the molecular or quantitative level, in order to assess their agreement making use of the previously determined links between genetic polymorphism and phenotypic variation. The issue of disentangling several potential selective forces, including fishing, is perhaps more difficult, especially in wild populations. Indeed, during the decades where the intensity of fishing has increased dramatically and is expected to have induced life-history evolution in exploited fish populations, a number of other environmental factors have also demonstrated substantial changes. Among the most well known is “global warming”, which has led to a general increase in sea temperatures. Whether higher temperatures have had a substantial effect on life-history evolution in fishes is not known, but has often been mentioned as an alternative driver of the observed evolutionary changes in size and age at maturation. For wild fish, which have been subjected to both drivers, the relative effects can be difficult to disentangle. However, neither fishing intensity nor temperature has been increasing linearly. For example, new major technical breakthroughs have led to sudden increases in fishing pressure and other events such as the Second World War have dramatically reduced fisheries. For temperature, cyclical patterns of warm and cold periods respectively have been observed within the general increase. In view of that, temporal adaptive genetic changes for cod and sole will be compared with known historical changes in fishing intensity and temperature to see how much of the genetic variance over time is explained by each of the evolutionary drivers.

Objectives

The overall objective of WP3 is to harness the power of a variety of existing and emerging modelling techniques for understanding and forecasting fisheries-induced adaptive changes in life-history traits relevant for the demography and productivity of exploited fish stocks.
Specific objectives:

1. Establish how the shapes of maturation reaction norms, which critically affect the demographic response of fish populations to altered fishing pressures and climatic factors, are evolutionarily determined by the environmental variation of growth and mortality rates.
2. Extend existing models of fisheries-induced evolution from the current separate and limited focus on single adaptive traits into a synthetic framework for uncovering the complex life-history syndromes caused by fishing in multiple concomitantly evolving adaptive traits, including those affecting growth, maturation, and reproduction.
3. Investigate the vulnerability of prototypical life histories to fisheries-induced evolution based on strategic evolutionary models, so as to establish transferable knowledge and generalized predictions about the expected life-history impacts of alternative fishing regimes on exploited fish stocks.
4. Extend existing models of fisheries-induced evolution from the currently prevalent unisex approach to an analytical framework that can account for sex-specific differences in life-history determinants, like size-dependent fertility and the cost of reproduction, in order to accurately predict sex-specific evolutionary responses to exploitation.
5. Integrate salient population genetic components into existing models of fisheries-induced evolution, to generate new testable predictions on the effects of fishing on neutral and adaptive genetic variation in exploited populations.
6. Develop several tactical models of fisheries-induced evolution, structurally geared to and carefully calibrated for the specific conditions known to characterize selected fish stocks.
7. Systematically explore the implications of fisheries-induced evolution for stock stability and recovery potential, with particular reference to and exchange with corresponding research carried out under the auspices of the ongoing FP6 SSP-STREP UNCOVER.
8. Integrate salient socio-economic components into existing models of fisheries-induced evolution in order to establish sound predictions of the economic consequences of fisheries-induced evolution and to strengthen the long-term accuracy of the eco-genetic models with regard to variations in fishing effort and catch regulations.
9. Synthesize existing and emerging knowledge about the causes and consequences of fisheries-induced evolution into recommendations for the evolutionarily enlightened management of exploited fish stocks, with particular reference to practical options for slowing, stopping, or even reversing the genetic effects of fishing.

Description of work

General background

While descriptive empirical studies are necessary for documenting the actual course of fisheries-induced evolution in wild populations, they obviously suffer from difficult challenges. First, in such studies it is mandatory to weed out the influences of uncontrolled confounding variables, and second, sufficient amounts of data have to be collected for the detection of life-history trends to become feasible. Experimental studies, on the other hand, can control for extraneous variables and allow for the systematic collection of data, but typically lack scale and usually are limited to examining species with short generation times under unnatural conditions. The need to overcome these systematic difficulties creates a strong drive towards harnessing the power of theoretical models for systematically addressing key issues regarding the time course and outcome of processes of fisheries-induced evolution.
Modelling fisheries-induced evolution offers exciting opportunities for meeting challenges that cannot equally be tackled through any other approaches. Particular questions to be addressed are as follows. What evolutionary responses are expected under given ecological and environmental conditions? What are the time scales on which fisheries-induced evolution is expected to unfold? Are there measures for influencing the pace of fisheries-induced evolution? Which kinds of life history are particularly vulnerable to fisheries-induced evolution? How does the interaction among multiple life-history traits and the sex-specific nature of life-history trade offs affect expectations of fisheries-induced evolution? Which exploitation regimes are particularly likely to cause fisheries-induced evolution, and why? How does the size selectivity of fishing gear affect the course and outcome of fisheries-induced evolution? What evolutionary effects are expected in response to the implementation of fishing moratoria? What are the options for slowing down, stopping, or even reversing the effects of fisheries-induced evolution? What are the long-term socio-economic consequences of fisheries-induced evolution and of measures aimed at mitigating such evolution? How can we optimize (or redress) the balance between the short-term and the long-term consequences of exploitation? In addition to helping to answer these and other questions, models also play an important role in guiding future empirical work. Most importantly, they can be used to proactively predict the evolutionary effects of fishing and to test specific management strategies, without having to instigate large-scale, typically infeasible or unaffordable, empirical work. While model-based studies carried out to date have already contributed appreciably to our present understanding of the mechanisms and processes underlying fisheries-induced evolution, a wide range of crucial insights remain to be established. These are the subject of WP3.
The majority of existing evolutionary models have not been designed to match the specific and rather complex needs of studies of fisheries-induced evolution. This is because predicting the ecological and evolutionary dynamics of exploited fish stocks requires respecting a range of demanding considerations. Specifically, to do justice to the complexities of fisheries-induced evolution, models have to account for (a) the physiological structure of fish populations in terms of age, size, and maturity status; (b) trade-offs between the fitness consequences of changes in salient life-history traits; (c) the considerable degree of phenotypic plasticity typically underlying the dynamics of fish populations; (d) frequency-dependent selection pressures resulting from density-dependent growth and recruitment; and (e) the amount and distribution of additive genetic variance harboured in fish populations, allowing them to respond to exploitation pressures. Earlier modelling approaches, based on life-history optimization methods or on quantitative genetics theory, have succeeded in reflecting features (a) and (b), but have rarely incorporated feature (c), and typically fall short of reflecting features (d) and/or (e). More recent models, based on adaptive dynamics theory, can take care of features (a) to (d), but fail to do justice to (e).
This situation sets the stage for the development of a new generation of models of fisheries-induced evolution that are capable of addressing issues (a) to (e) simultaneously. These new models are called ‘eco-genetic,’ to reflect that they are specifically geared to incorporate a sufficient amount of ecological detail, thus tackling features (a) to (d), in addition to a suitable rendering of genetic detail, thus tackling feature (e). The eco-genetic modelling approach underlies most work planned for WP3 and offers versatile tools for helping fishery scientists and managers to understand and possibly mitigate the consequences of fisheries-induced evolution.


Task 3.1: Evolutionary determination of maturation reaction norms

The shape of maturation reaction norms has important consequences for the dynamics of maturation in fish populations. For straight maturation reaction norms with a positive slope, fast-growing individuals mature young and small, while slow-growing individuals mature old and large. By contrast, for straight maturation reaction norms with a negative slope, fast-growing individuals mature young and large, while slow-growing individuals mature old and small. For dome-shaped maturation reaction norms, individuals with intermediate growth rates mature large and at an intermediate age, those with fast growth mature small and young, while those with slow growth mature small and old. Since the age at maturation affects the cumulative probability of individuals to survive until maturity, while the size at maturation influences the size and fecundity of adult individuals, maturation dynamics, in turn, affect the age and size structure of the immature and mature life stages in a population, with evident repercussions for the reproductive potential (recruitment) of the spawning stock. This cascade of effect tightly links the dynamics of maturation and the shapes of maturation reaction norms to a stock’s demographic responses to exploitation.
The shape of maturation reaction norms is ultimately determined by evolution. The purpose of this task will be to characterize the ecological conditions leading to the evolutionary emergence of different maturation reaction norms. Previous studies have suggested that the joint probability distribution characterizing environmental variation in terms of growth and mortality rates experienced by individuals is a primary determinant of such evolution. However, no thorough analysis of such evolutionary determinism has been undertaken so far. This pending analysis is especially important in the context of exploited fish populations, since fishing affects the joint probability distribution of individual growth and mortality rates by (i) increasing mortality rates, (ii) modifying growth rates through density dependence and compensatory responses, and by (iii) preferentially removing individuals characterized by a given range of growth rates. The latter can occur, for instance, by targeting geographic areas where large fish are found.
Exploitation is therefore expected to induce evolutionary changes in the shapes of maturation reaction norms, in addition to the commonly observed changes in the absolute position (in the age-size plane) of such reaction norms. As part of the evolutionary analysis, the demographic consequences of different maturation reaction norms will be characterized in terms of the resultant age and size structures, demographic dynamics, and reactions to exploitation.


Task 3.2: Fisheries-induced multi-trait evolution

What life-history patterns emerge when multiple traits evolve under fisheries-induced selection pressures? To date, this question has remained completely unaddressed. Most published theoretical work has focused on the evolution of single traits in terms of age or size at maturation.
The recent wave of new research on the evolution of maturation reaction norms already amounts to a significant departure from this more limited traditional approach, since concomitant evolutionary changes in two-life history observables can thus be analyzed. The simplest reaction norm models of maturation evolution have assumed linear and deterministic maturation reaction norms. Task 3.1 extends this body of work to nonlinear reaction norms and thus addresses the question how the (nonlinear) shape of the reaction norms is affected by ecological conditions, environmental factors, and fishing pressures. Another important extension concerns the probabilistic nature of maturation reaction norms, which is formally represented by a probabilistic maturation envelope around a maturation reaction norm’s midpoint. In the simplest case, this envelope is of constant width across ages and can thus be characterized by a single adaptive trait. Together with the two traits needed for characterizing the slope and intercept on linear midpoint curves, this results in three-trait models of maturation evolution.
These reaction norm models, however, are focusing on evolutionary changes in maturation schedules alone, and are thus not yet sufficient to identify the life-history syndromes caused by fisheries-induced evolution. Two other crucial life-history processes in particular need to be considered. First, fishing is bound to affect the growth rates of exploited individuals, especially when technical measures like mesh-size regulations are in effect. If small individuals encounter smaller fishing mortalities than larger individuals, the gene pool of exploited stocks may gradually become biased towards slow-growing fish. In the absence of quantitative models, this effect is impossible to predict, since the size dependence of energy acquisition rates, fecundities, and natural mortalities also have to be taken into account when determining selection pressures on growth rates. Second, fishing must be expected to affect the reproductive effort of exploited individuals. The obvious rationale here is that, under conditions of heavy exploitation, saving resources for growth until the next season or for reproduction during the next season will be counter-selected, if the probability of still being alive during the next reproductive season is low due to fishing.
Together, these considerations create an imperative for investigating multi-trait models of fisheries-induced evolution. The simplest starting point for this endeavour is given by five-trait models in which the adaptive state of individuals is characterized by their juvenile growth rates and reproductive effort, in addition to three traits needed for describing the shape of linear probabilistic maturation reaction norms. Time permitting, also more advanced models of multi-trait fisheries-induced life-history evolution will be analyzed under this task, in which the adaptive state of individuals may be characterized by function-valued traits for the age- and size-specific energy allocation schedules for growth and reproduction.

While empirical evidence of fisheries-induced evolution has been documented in several species, it is currently far from clear if and how the pattern of evolutionary response significantly varies among species with significantly different life histories. Such differences, however, are clearly expected based on existing theoretical insights. For example, results of evolutionary analyses corroborate that species with longer generation times are generally more vulnerable to the selection pressures induced by exploitation. Yet, the question of how other life-history attributes of a species influence the out-come of fisheries-induced evolution has remained largely unstudied. Relevant attributes include natural mortality levels, average lifespan, patterns of determinate or indeterminate growth, semelparous or iteroparous reproduction, as well as typical somatic growth rates and ages at maturation. While some of these attributes may readily respond to fisheries-induced selection pressures, others are constrained by the overall life-history strategy and bauplan of species and thus must be interpreted as setting boundary conditions for processes fisheries-induced evolution.
This task will deliver a suite of strategic eco-genetic models parameterized for particular life-history prototypes. Each of these prototypes will idealize species or groups of species with a typical combination of life-history attributes. At least five such general life-history prototypes will be investigated, to provide general insights into the fisheries-induced evolution of anchovy-like, cod-like, salmon-like, ray-like, and whale-like life histories. Establishing measures of evolutionary vulnerability, these results will establish transferable knowledge and generalized predictions about the expected life-history impacts of alternative fishing regimes on exploited fish stocks. Interactions with Tasks 3.1 and 3.2 will thus be important. The resultant insights will support fisheries scientists and managers in their responsibility of prioritizing measures of monitoring and protection among a multitude of exploited stocks.

Task 3.4: Sex-specific dimensions of fisheries-induced evolution

Many fish are sexually dimorphic. Such dimorphisms are neither coincidental nor idiosyncratic, but instead result from different selection pressures and disparate life-history trade-offs experienced by males and females. For example, typical sexual dimorphisms in size imply that males are smaller than females. Males of many species also mature earlier and at smaller sizes. These phenomena have been documented in commercially exploited fish like flatfishes and gadoids. Fisheries-induced evolutionary changes in life-history traits are thus bound to differ between the male and female components of the same stock. Whenever harvesting is effectively sex-specific, these differences are amplified. Also differences in life-history trade-offs between the two sexes have to be considered: for example, the microgametic sex, usually the males, for obvious reasons experience smaller per gamete costs of reproduction, which, in turn, is bound to affect the allocation of energy between growth and reproduction, as well as between potential reproductive seasons.
We currently have next to no theoretical understanding of how fishing causes sex-specific changes in life histories and of how such changes may affect stock properties and dynamics over longer time scales. Depending on the underlying genetics, mating systems, and mechanisms of sexual and natural selection, the remoulding of male and female life histories through harvesting might possibly lead to increased or decreased sexual dimorphisms. It may also seriously disrupt the balance between sexual and natural selection pressures achieved – in each of the sexes, as well as between them – over long periods of adaptation in the wild. By disrupting such natural balances, the resulting fisheries-induced evolutionary changes may be mostly detrimental, contributing to the further deterioration of stock conditions. Several mechanisms can play a role in such processes. First, fishing can lead to a disproportionately larger evolutionary response in the larger sex, and this may cascade down to the otherwise invulnerable sex through shared somatic genes. Second, the preferential fishing of large females might release sexual selection pressures on male body size, resulting in larger males. Third, fishing can lead to biased sex ratios in the spawning stock, affecting and disrupting previously established mating systems. Fourth, fishing can further disrupt mating systems if mating preferences are driven by absolute or relative individual sizes, and this process may enhance or cancel out the effect of biased sex ratio. Fifth, disrupted mating systems may lead to emergent depensation in stock-recruitment relationships, thereby creating additional risks of population collapse.
Advanced methods for addressing these issues have already been developed for populations without sex structure. Benefits from intermediate results obtained in Tasks 3.1 to 3.3 can thus be expected. Task 3.4 will generalize the existing methods to sex-structured populations, incorporate qualitative knowledge and quantitative data on sex-specific life histories and mating systems, and investigate the complex dynamical interplay between the sex-specific selection pressures and trade-offs as outlined above.

Task 3.5: Fisheries-induced evolution of neutral and selected genetic markers

Fishing mortalities may have two effects on the genetic composition of exploited species. (a) Neutral genetic variation may be diminished, reflecting a decrease in effective population size and thus favouring random genetic drift. Such drift, in turn, may lead to changes in the allelic frequency distributions of genes of adaptive significance through purely random stochastic processes, unrelated to selection pressures. (b) A different group of effects occurs through changes in the allelic frequency distributions of adaptive genes due to fisheries-induced selection pressures. Enhancing our understanding of these latter effects through modelling is the aim of all other tasks in WP3.
Task 3.5, by contrast, aims at a closer analysis of three unresolved issues related to the interplay between the effects (a) and (b), an aim that can be addressed almost only though modelling. Firstly, previous expectations regarding effects (a) are based on classical population genetic models, which, in most cases, do not include a detailed description of the ecological setting, including fishing pressures, responsible for genetic changes. Such detailed descriptions, however, will be required if operational tools are to be developed and salient relationships between genetic parameters and relevant ecological properties – such as population size, spawning stock biomass, recruitment, or yield – are to be identified. Secondly, the fact that changes in the adaptive genetic make-up of exploited fish populations may result from random genetic drift, as well as from fisheries-induced selection, complicates the analysis of empirical genetic data on adaptive genetic variation. To disentangle effects (a) and (b), changes in the adaptive genetic composition of a population have to be assessed against the baseline changes observed in its neutral genetic composition, see Task 2.5. New methodologies have already been developed to carry out such comparisons. The existing methods, however, suffer from being based on rather simplistic ecological models and from being designed to deal with spatial, rather than temporal, genetic differentiation. New methods specifically tailored to analyzing temporal genetic differentiation will be developed in Tasks 2.4 and 2.5, based on classical population models. An important component of Task 3.5 will be to test the robustness of these new methods against the ecologically more detailed and realistic models considered in WP3. Thirdly, the links between changes in the neutral genetic composition, the adaptive genetic composition, and the ecological characteristics of exploited fish populations presently remains very unclear. Due to the ease and cost-effectiveness of conducting molecular analysis on neutral markers, it is highly desirable to develop operational tools for inferring some relevant changes in the adaptive genetic composition of exploited stocks from observed changes in their neutral genetic composition. If this were feasible, neutral molecular markers could be used to establish early warning signals for significant and relevant fisheries-induced evolutionary changes in exploited stocks.
Task 3.5 will deliver, as a first step, a generic model for studying these questions and thus the interplay between effects (a) and (b). This model will be either deterministic or individual-based, depending on the complexities encountered during model’s development, it will include sufficiently detailed account of the underlying ecological setting, and it will be specifically tailored to study temporal genetic differentiation. In a second step, this generic model will be parameterized for the species (cod and sole) used for genetic analysis in WP2, in order to provide testable predictions against which to interpret the empirical genetic data.

Task 3.6: Fisheries-induced evolution of specific stocks

Much of the research foreseen for WP3 will establish general and transferable model-based insights into the causes and consequences of fisheries-induced evolution. Task 3.3 will be devoted to analyzing the evolutionary vulnerability of specific life-history prototypes. Task 3.6 goes one step further in this development and aims at establishing carefully calibrated quantitative models of fisheries-induced evolution for a few selected stocks. Altogether, such dedicated modelling efforts are proposed for four stocks: Northeast Arctic cod, sole in the English channel, Arctic charr in Norwegian lakes, and Atlantic salmon in Scotland.
Northeast Arctic cod & Sole in the English channel. These two stocks have been selected in reflection of the comprehensive empirical investigations of fisheries-induced evolution, both ecological and genetic, in cod and sole that will serve as integrative case studies across the different work packages of this entire STREP project; for further details please see the descriptions of Tasks 1.1 to 1.4, WP2, and WP4. Work on the stock-specific eco-genetic model of Northeast Arctic cod will benefit from substantial preparatory work that has already been carried out for this purpose at IIASA and IMR. The stock-specific eco-genetic model on sole in the English channel will be based on the expert knowledge and quantitative data available on this stock at Ifremer and IMARES.
Arctic charr in Norwegian lakes. The stock-specific eco-genetic model of Arctic charr will be constructed as part of an overarching effort carried out under Tasks 1.7 and 3.6, aiming at elucidating fisheries-induced evolution in two Norwegian lakes for which unusually detailed time series on Arctic charr have been collected. For the charr populations of lakes Takvatn and Fjellfroskvatn, extensive environmental and life-history data available at UT will allow estimation of the relevant parameters of the planned eco-genetic model. The utility of this data will be complemented by data on genetic variation and harvesting pressure, which will allow incorporating accurate assumptions about selective pressures and evolutionary rates. The parameterized models will be analyzed in order to derive quantitative expectations that can be validated against the trends in the available long-term data. Time permitting, an analogous modelling approach will be applied to the whitefish stock in lake Stuorajavri, thus extending the analysis to the main target species of fisheries in Northern European lakes. Present management practices for inland fisheries in Norway and elsewhere are derived from models that do not incorporate evolutionary changes in exploited stocks. The analysis of eco-genetic models developed for Arctic charr and whitefish will provide guidelines for the evolutionarily enlightened management of salmonid stocks. Variations in fishing pressures, size selectivities, and harvested habitats will be considered to move towards the sustainable exploitation of stocks exposed to rapid evolution in growth, maturation, and reproduction. The evolutionarily enlightened management recommendations obtained for Arctic charr and whitefish can easily be extended to other salmonid species with similar evolutionary ecologies living in Scandinavia, Russia, and Canada. Adding insights into the evolutionarily implications of fisheries to Norwegian research and training programs will contribute to mitigating fisheries-induced evolution in landlocked salmonids and other fish stocks.
Atlantic salmon in Scotland. The fourth stock-specific model, on Scottish salmon, has been selected because of the complementariness of salmon life history compared with the other two targeted marine species, and since excellent data, as well as preliminary model-based investigations, on this stock provide a first-rate basis for developing and calibrating the planned model. It is envisaged that this modelling effort will focus primarily on the North Esk river stocks, since here historical records are strongest. A second motivation for the planned focus on the North Esk stems from the large demographic and genetic sampling program currently being carried out there by FRS. Three classes of model, partly already established within FRS, will be parameterized using the insights gained from the empirical data analysis, Task 1.6, and will be utilized to examine fisheries-induced evolution in Atlantic salmon. First is a run-reconstruction model that is used by international fishery organizations like ICES to reconstruct Scottish Atlantic salmon prefishing abundance. It is envisaged that this model will be useful in gaining insights into the stock structures available before fishery impacts. It will provide estimates that can be compared and/or combined with output from the other two model classes below. Second is a structured population model of growth, development, and size-at-age currently being developed for salmon. It is envisaged that this model will provide a strategic overview of how different spatially structured populations interact and how fishing efforts could influence both the viability and adaptive traits of the resultant metapopulation. Third is an individual-based eco-genetic model developed for describing the dynamics of wild Atlantic salmon populations. The model follows individual Atlantic salmon from egg, through the various developmental stages, until death after spawning. As an individual moves between stages, its growth rates and probabilities of maturation and survival are calculated. These are based on a synthesis of population demographic parameters collected from the literature and reflect various individual trait values of a particular fish. While literature-based values are used as defaults, all model parameters can simply be altered, so that the model can characterize a particular river population of interest. Early versions of this model have already successfully been validated against empirical measurements of unimpacted salmon populations. Development of the model’s genetic component is in progress.

Task 3.7: Implications for stock stability and recovery potential

This task will focus on analyzing the effects of fisheries-induced evolution on the stability and recovery potential of exploited stocks. Since such evolution occurs in life-history traits affecting growth, survival, maturation, reproduction, and offspring quality, its demographic consequences are undeniable.
There are many reasons to expect that the demographic consequences caused by fisheries-induced evolution are mostly detrimental for the resilience of stocks. First, fisheries-induced evolution towards earlier reproduction at smaller sizes coerces individual fish into physiologically inefficient energy allocation patterns that result in poor reproductive efficiencies. The allometric dependence of fecundities on body size here conspires with the extra effects of body size on the quality and survival probabilities of eggs to create a very strong total effect. The evolutionary change also diminishes the copious contribution of large and old females to a stock’s recruitment, thus undermining the stock’s insurance against streaks of unfavourable environmental conditions. In species with physically demanding spawning migrations, the reduced age and size of mature fish may readily increase size-specific migration mortalities – in principle up to a point at which mature individuals become too feeble to reach their genetically imprinted spawning grounds reliably. Second, fisheries-induced evolution towards higher reproductive efforts may exhaust individual fish energetically during their reproductive seasons and thereby bring about elevated levels of mortality between reproductive seasons. Effects of diminishing return on reproductive investments, resulting from physiological or morphological constraints, will further exacerbate this effect, resulting in the wasteful allocation of energy to reproduction. The reduced number of reproductive seasons inevitably resulting from increased reproductive effort can lessen the average fecundity of individuals, since such a reduction undermines bethedging strategies against the perils of environmental fluctuations. Third, fisheries-induced evolution towards lower growth rates results in smaller body sizes, which in turn may expose individuals to higher levels of size-dependent natural mortality. Rates of energy acquisition can also be negatively impacted by reductions in body size. In addition to these three groups of effects operating through specific individual traits, fisheries-induced evolution may also damage mutually co-adapted trait complexes by overriding the fine-tuning of multi-trait life-history strategies afforded through long preceding periods of natural selection. The potential disruption of mating systems through fisheries-induced evolution adds a further dimension to this problem.
For all these reasons, fisheries-induced evolution may render exploited stocks less stable with regard to environmental fluctuations. In a similar vein, the potential of stocks to successfully recover from periods of overexploitation may become jeopardized. Accordingly, fisheries-induced evolution may adversely contribute to the collapse of fish stocks and may in addition impede their recovery once a crash has occurred. To mention just one particularly well known example, findings on fisheries-induced evolution preceding the infamous collapse, and essential lack of recovery ever since, of northern cod off the coast of Newfoundland and Labrador will have to be considered in this light. In carrying out the research agenda for this task, close connections will be sought with corresponding efforts unfolding under the auspices of the ongoing FP6 SSP-STREP UNCOVER.

Task 3.8: Economic models of fisheries-induced evolution

This task aims at integrating and strengthening the interface between ecological, evolutionary, and economic modelling, in order to contribute to an improved understanding of the long-term effects of fishing.
It has been demonstrated that high fishing pressures may cause evolution towards maturation at earlier ages and smaller sizes, bound to decrease the profit of the corresponding fishery. For example, the world’s largest stock of Atlantic cod, the Northeast Artic cod population of the Barents Sea, has experienced a considerable reduction in its mean age at maturation from 10 to around 6 to 7 years during the second half of the twentieth century. There is a price premium of about 70% on fish above 3 kg compared with fish below 1 kg, and this premium will in all likelihood increase when the abundance of large fish in the stock is diminished. Considering that the total value of Norwegian cod fisheries was 3 billion Norwegian Kroners (approximately 400 M€) in 2005, it is clear that fisheries-induced evolution can lead to considerable economic losses. Analyzing the economic effects of fisheries-induced evolution on a particular stock will benefit from the inclusion of climate effects on the marine ecosystem. Currently we know very little about how such climate effects will interact with harvesting effects. A warmer climate will probably lead to faster growth and potentially to larger-sized fish, which can be sold for a higher price. This benefit, however, is likely to be offset by the fisheries-induced evolution of the stock. It will therefore be worthwhile to investigate how concomitant changes in genetics and climate influence the economic optimal fishing patterns, as seen by different stakeholders. For this purpose, the disparate economic perspectives of individual fishermen and of society at large have to be taken into account.
The interplay between demography, evolution, climate, fishing behaviour, and socio-economic forces on the one hand, and alternative management strategies on the other is quite complex. To our knowledge, this important nexus has as yet not been investigated in a interdisciplinary framework. Based on the close collaboration between ecologists, evolutionary biologists, and economists, work on this task will thus analyze how the economic returns from fish stocks are affected by this interplay. Strengthening the interface between biological and economic models, this task will deliver a bio-economic model of Northeast Arctic cod that will be utilized to examine what harvest strategies are optimal, both economically and ecologically. The biological component of this approach will be based on an individual-based eco-genetic model for the Northeast Arctic cod, to be developed as part of Task 3.6. Pilot studies on this key component, carried out in collaboration between the University of Oslo (UO) and IIASA, are already in progress, and preliminary results indicate that this approach will indeed be successful. The required quantification of the behaviour of fishermen will be based on an ongoing effort at UO, dedicated to estimating the relevant parameters for assessing the cost structure of the Norwegian fisheries. This will include studying how the fleet structure will change in response to evolutionary and climatic changes. Once the bio-economic model is in place, different management scenarios will be combined with climate scenarios, to study their joint repercussions for the economics of fisheries-induced evolution in Northeast Arctic cod. The primary management objective to be analyzed will be to maximize the long-term profitability of the fisheries as measured by the net present value of the fisheries-generated cash flow. Making the economic and ecological costs of fisheries-induced evolution explicit in this way will serve as an important step forward in the development and implementation of improved management practices.

Task 3.9: Evolutionarily enlightened stock management

Eco-genetic models are designed to explore the ecological and evolutionary consequences of fishing. This type of comprehensive and flexible approach is needed as fisheries managers increasingly struggle to deal with multiple objectives and thus need a basis on which to make balanced, informed decisions.
Task 3.9 will synthesize the findings of Tasks 3.1 to 3.8 from the perspective of fisheries management. Two groups of questions will thus be addressed. (1) Which types of life histories are particularly vulnerable to fisheries-induced evolution? (2) Which types of fishing regimes are particularly likely to cause fisheries-induced evolution? The non-synthetic components of Task 3.9 will address two further groups of questions. (3) In particular, work on this task will carefully address the measures that can be taken to slow, stop, or reverse evolutionary responses to fishing. It has been suggested, for example, that fishing on a stock’s spawning grounds can elicit evolutionary responses that counter the negative evolutionary consequences of fishing on the same stock’s feeding grounds. More recently, model-based studies have demonstrated that reducing harvest mortality and implementing marine reserves can help to counter evolution towards maturation at young age and small size. Yet the effects of geographic or seasonal restrictions on fishing in spatiotemporally structured stocks remain to be analyzed systematically, and particular attention will be devoted to clarifying the expected effect of marine reserves and marine protected areas on the pace and direction of processes of fisheries-induced evolution (preparatory work on this question is already underway at IIASA). Also the evolutionary consequences of size-based restrictions on fishing will still have to be explored comprehensively, with particular regard to technical measures like mesh size regulations or regulations based on protective slot limits. In general, the effectiveness of all these potential management measures in controlling multi-trait evolution is as yet poorly understood. (4) A fourth group of questions to be investigated in this task will elucidate the conditions under which evolutionary responses are relatively easy or more difficult to reverse. A fundamental insight already highlighted by existing research is that fisheries-induced selection on key life-history traits typically is much stronger than natural selection acting in the opposite direction, once fishing is reduced or ceased. This suggests that there might usually be a long wait for evolutionary reversals, even under ideal conditions. While some reversal of phenotypic life-history traits thought to be affected by fisheries-induced evolution has been observed in natural populations after fishing pressures were relaxed, selection intensities were not estimated and the extent to which such reversals represented genetic changes currently remains unknown.
Answers to these four suites of questions will help fisheries managers to take suitable decisions regarding the implementations of catch regulations and will provide guidance for prioritizing limited resources for monitoring and intervention on the most imperilled and/or promising stocks, thus reaping maximal practical benefits from the advent of evolutionary fisheries science.

Objectives

The global objective of this work package is to coordinate research carried out in WP1, WP2, and WP3 for two cross-cutting case studies on cod and sole.
Specific objectives:

1. Insure the careful integration of the two case studies by a timely combination of research and results from WP1, WP2, and WP3 according to a pre-established stepwise progression.
2. Produce a final synthesis about fisheries-induced evolution in these two species utilizing the broadest possible knowledge base.

Description of work

General background

Establishing unequivocally fisheries-induced evolutionary changes requires a carefully integrated analysis based on a stepwise progression. The analysis necessitates different types of data - time series of phenotypic data, historical collections of tissue samples, and population ecological data - and expertise from rather diverse scientific fields - life-history theory, evolutionary ecology, population genetics, quantitative genetics, modelling, and fisheries sciences - which can hardly be found within a single department or within a single institute. In addition to the various specific insights that will be produced by work packages 1 to 3 (WP1 on case studies, WP2 on genetic analyses, and WP3 on eco-genetic models), this project offers a unique opportunity for gathering the various data and bringing together the diverse scientific expertise required to conduct such an analysis. This work package aim to (i) insure optimal integration for two specific case studies on cod and sole by coordinating the step-wise evolutionary analysis, and (ii) produce a synthesis about fisheries induced evolutionary changes for these two species.

Task 4.1: Coordination and synthesis of case study on cod
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Task 4.2: Coordination and synthesis of case study on sole

The overall progression of the work will be the same for the two case studies. Therefore, the two tasks are the subject of a common description below.
A fully integrated analysis of fisheries-induced evolution requires combining different types of data:

• Times series of phenotypic data on life-history traits, or closely related traits, expected to be under fisheries-induced selection, e.g., length- and/or weight-at-age (growth curves), reproductive effort or fecundity-at-age, age and size at maturation
• Historical collection of biological tissue (ideally, coming from the same individual as those measured for phenotypic traits) and allowing for DNA extraction and thereby molecular genetic data acquisition. Biological archives, like otoliths or scales, can provide both phenotypic (length- and weight-at age, age and size at maturation) and molecular genetic data at the same time.
• Population ecological data including
• Biological data, e.g., length-weight and weight-fecundity relationships, models of energy allocation between growth and reproduction, maturity ogives;
• Time series of catch statistics, e.g., total catch, age and size distribution in catch, catch sex-ratio, maturity status in catch;
• Time series of population dynamic data, e.g., abundance (census, biomass), age and size structure, sex-ratio, natural mortality, maturity ogives;
• Fishing regime history, e.g., time series of fishing mortality, fishing selectivity (gear type, selectivity ogives per gear type, minimum landing size…); and
• Environmental data, i.e. changes in other potential environmental drivers of adaptive changes, e.g., temperature, salinity, species composition.

This work package will insure that all the necessary data will be gathered for the case studies on cod and sole and will orchestrate the sharing of these data between the teams working on these two species in WP1, WP2, and WP3.
After gathering the set of data required, this work package will orchestrate the evolutionary analysis across WP1, WP2, and WP3 according to the following stepwise scheme:

• Step 1: Documenting temporal trends in life-history traits and removing plastic components (WP1). Phenotypic data will be used to document temporal trends in life-history traits. Part or all of the plastic component of these trends will be assessed and removed using time series of environmental factors susceptible to trigger plasticity, or based on specific methods such as probabilistic maturation reaction norms. Any remaining residual trend will be suggestive of evolutionary changes.
• Step 2: Estimating fisheries-induced selection gradients (WP1 and WP3). Fisheries-induced selection gradients affecting the focal life-history traits will be estimated using population ecological data. Their agreement with the residual temporal trends documented in Step 1 would strongly suggest fisheries-induced evolution. If the proper data are available, environmentally-induced selection gradients will also be estimated to evaluate alternative environmental drivers for evolutionary changes.

The next steps will consist of including genetic data in the analysis in order to assess whether evolutionary changes suggested by the phenotypic analysis are detectable at the genetic level:

• Step 3: Documenting significant temporal adaptive genetic changes (WP2). Two different, though complementary, methods will be implemented to detect adaptive genetic changes: (i) assessing significant temporal differentiation at the molecular level for some candidate genes and/or (ii) assessing significant temporal differentiation in terms of quantitative genetic parameters. In both cases, the principle of the analysis will be the same: temporal genetic differentiation at the molecular or quantitative level will be compared with the baseline temporal genetic differentiation at neutral markers. Any significantly higher differentiation at the molecular or quantitative level would establish that directional selection affects the adaptive genetic composition of the population. Yet, the link with fisheries-induced selection gradients will remain to be ascertained.
• Step 4: Linking temporal adaptive genetic changes and fisheries-induced selection (WP2). If directional selection acting on adaptive genetic variation is established, knowledge of the link between genotypes and observed phenotypes is required to clearly relate temporal adaptive genetic differentiation with fisheries- or environmentally-induced selection gradients documented in Step 2. For quantitative genetics, this link is implicit but for candidate genes, it will need to be investigated. Once the phenotypic effect of adaptive genetic variation is known the agreement between temporal adaptive genetic changes, fisheries-induced selection gradient and temporal trends in life-history traits will be evaluated, thus assessing whether adaptive genetic change in response to fisheries-induced selection could explain part of or all the temporal phenotypic trends documented in Step 1.
• Step 5: Predicting fisheries-induced evolutionary changes and their demographic consequences (WP3). Once fisheries-induced evolutionary changes are established empirically, the next steps towards more operational aspects are to be able to predict these changes and to evaluate their consequences on stocks demography and thereby fisheries yield. Some realistic eco-genetic models will be parameterized using the different types of data gathered for the two case studies to (i) assess whether it is possible to simulate the evolutionary changes empirically estimated, (ii) then predict future evolutionary changes and (iii) assess their effect of stocks’ demography with particular emphasis on stocks viability and yield.
• Step 6: Evaluating management practices to mitigate undesirable effects of fisheries-induced evolution (WP3). The eco-genetic models developed in Step 5 will be used to evaluate different management practices aiming at diminishing undesirable effects of fisheries-induced evolution and promote long-term viability and yield of fish stocks. These will include some indispensable economic ingredients advancing a more comprehensive understanding of the consequences of management practices on both the exploited stocks and the fisheries.

Objectives

The overall objective of WP5 is to ensure that all components of the FinE project will unfold and work together with maximal efficiency and output, so as to guarantee optimal coordination, integration, and dissemination of the undertaken research efforts.
Specific objectives:

1. Coordination, to ensure the effective organization and supervision of tasks throughout the project.
2. Integration, to exploit the synergies between tasks, promote the interfacing of results across tasks, and foster the successful liaison of FinE’s activities with related European activities.
3. Dissemination, to target audiences across a wide range of different groups, including the EC’s Commission Services, the scientific community, decision and policy makers, fisheries research agencies, other stakeholders, and the interested public.

Description of work

Task 5.1: Project web site

The project coordinator will establish and maintain the project’s web site. As a first purpose, this resource will contribute to the publicizing and wide-ranging dissemination of information about the FinE project, offering a one-stop access point to up-to-date information about the project’s participants, research plans, open positions, ongoing activities, and accomplished results. This will ensure that the appropriate information regarding the project will be widely accessible to facilitate the dialogue with society within Europe. Secondly, through its restricted-access internal part, this web site will serve as a clearing house for information emerging across the entire network of FinE participants. New developments, attained milestones, completed deliverables, and current research questions will be reflected. Documents and forms will be posted, and reprints and preprints will be deposited for retrieval across the network. Through these functions, the web site will provide a communicative backbone for the FinE network, strengthen the ties between all teams, and facilitate information exchange between remote teams. Email discussions initiated by the project coordinator and the participants will complement this web-based service.

Task 5.2: Project leaflet

A 2-4 page leaflet will be prepared by the coordinator. This will contain general information about the work programme, participants, expected results, and exploitation strategy. This leaflet will appear at least once, and will be broadly distributed (to the EU, all participants, selected targets in the fishing industry, relevant scientific meetings etc.). Optionally, the leaflet’s contents may be updated during the project’s lifetime.

Task 5.3: Annual consortium meetings

Annual consortium meetings will regularly bring together all FinE participants for several days during each year. Leaders and key staff of all partner teams are required to participate in these meetings. Organized by the team leaders at IMR, Ifremer, and DIFRES, these meetings will play a central role for the coordination of activities, the integration of team-specific efforts, and the dissemination of results. Collaborative efforts will be discussed and planned in detail, agreements on the division of labour between teams and individuals will be agreed on and reassessed as needed, and research plans will be updated in reflection of encountered obstacles and accomplished goals. Occasionally, external experts will be invited as required, to infuse the network’s collective scientific expertise with salient additional knowledge. The project’s kick-off meeting will take place within a short while after the start of the project, followed by two progress-oriented meetings after 12 and 24 months. The fourth network-wide consortium meeting will be integrated with the first international conference on fisheries-induced evolution, see Task 5.6. The meetings will be organized in a way that travel and subsistence costs are kept at a reasonable minimum. The European Commission will be informed about the meetings at least eight weeks in advance.

Task 5.4: Task monitoring and coordination

The team leaders at IIASA, IMR, Ifremer, and DIFRES will take the lead in the monitoring of and advising on the progress of research across all teams of the network. In this way it will be ensured that expectations and opportunities for research and collaboration are transparently communicated and well understood across the entire network, that responsibilities are clearly assigned and kept, that milestones and deliverables are achieved as required, that research plans and budgets are closely adhered to and adjusted as necessary, that problems and questions arising within or between teams are swiftly resolved, that the orchestration and synchronization across work packages and tasks proceeds with maximal efficiency, and that a synthetic perspective on the eventual integration of individual contributions prevails throughout the entire project. All other team leaders will naturally contribute to these overarching goals for organizing the project’s work.

Task 5.5: Liaison with related European initiatives

Special attention will be devoted to the liaison with other European initiatives working towards related objectives. The resulting interfacing efforts will be extended, in particular, to the Marie Curie Research Training Network FishACE (Fisheries-induced adaptive changes in exploited stocks); the Scientific Support for Policies initiatives UNCOVER (Understanding the mechanisms of stock recovery), EFIMAS (Evaluating scientific advice and decision-making processes in fisheries management systems), COMMIT (Committing to tailor-made long-term fishery management strategies), INDECO (Development of Indicators of Environmental Performance of the Common Fisheries Policy), and PROTECT (Marine protected areas as a tool for ecosystem conservation and fisheries management) as well as the Networks of Excellence MARBEF (Marine biodiversity and ecosystem functioning), EUROCEANS (European network of excellence for ocean ecosystems analysis), and MGE (Marine Genomics Europe). Important benefits will be reaped from gaining access to innovative insights generated within these initiatives as early on as is feasible. Likewise, it may be expected that a greater awareness of the scientific and practical issues surrounding processes of fisheries-induced evolution will helpfully contribute to the research agendas of these other projects. Exchange at the level of the initiative leaders will thus seek to foster mutually beneficial information flows.

Task 5.6: Progress reports

This task will ensure the efficient management of reporting activities through the lifecycle of the FinE project. Progress reports and salient intermediary communications will guarantee that the quality of exchange with the EC’s Commission Services is instigated and kept at the highest possible level. Specifically, the following sets of reports will be prepared:

• After 12 months, an ‘interim activity report’ giving project status and progress overview;
• At mid-term of the project, all reports as specified in Article II.7.2 of Annex II of the contract;
• At the end of the project, all reports as specified in Articles II.7.2, II.7.3, and II.7.4 of Annex II of the contract.

On suitable occasions, material prepared for the annual reports will amount to important preparatory steps towards publications in scientific journals, and vice versa. The involvement of the European Commission in this project will be demonstrated by adding the following sentence to each publication: “This study has been carried out with financial support from the European Commission, as part of the Specific Targeted Research Project Fisheries-induced Evolution (FinE, contract number SSP-2006-044276) under the Specific Support to Policies cross-cutting activities of the EC’s Sixth Framework Programme. It does not necessarily reflect the views of the European Commission and does not anticipate the Commission’s future policy in this area.”

Task 5.7: International conference on fisheries-induced evolution

The time will soon be ripe for organizing a first international conference on fisheries-induced evolution. While many, if not most, pioneers of this burgeoning field of research are European, fostering the international integration of efforts and insights is imperative, with particular reference to fisheries scientist and managers in countries like the USA, Canada, Japan, and Australia. Given the ubiquity of fisheries-induced evolution, also some capacity building in the developing world will have to be considered, before ignorance about the perils associated with unmanaged fisheries-induced evolution will have led to practically irreversible developments on their fishing grounds. To present and discuss the findings of the FinE project in a broad international forum and integrate project results into the international context, it is suggested that the organization of the important and ground-breaking first international conference on fisheries-induced evolution will be closely enmeshed with the activities and results of the FinE project. The planned conference is therefore envisaged for the final phase of the project, so as to provide a maximally resonant platform for displaying and disseminating the project’s overall findings. The conference will cover subjects under WP1 to WP3 with invited keynote contributions from outside and inside the project. While responsibility for the organization of this event will rest with the project coordinator, all four work package leaders will contribute to its planning and preparation. In this way, the FinE project will contribute to unprecedented momentum in the field, while at the same time showcasing the scientific and practical excellence of European fisheries research to a broader international audience.

Task 5.8: Booklet on fisheries-induced evolution

High-impact communication of the challenges resulting from fisheries-induced evolution to members of the wider scientific community, to decision and policy makers, fisheries research agencies, and other stakeholders will require documentation that significantly goes beyond the typical style of publications in scientific journals and the gray literature of fisheries research. The project therefore plans to produce a widely accessible booklet or brochure on fisheries-induced evolution based on the new findings obtained and the collective expertise held by all the participants of the FinE project. This will thus contribute to the effective dissemination of all the research carried out under the FinE project and thereby help building a greater awareness, especially among stakeholders, about what is at risk. By bringing together readily accessible and richly illustrated explanations of the key notions underlying fisheries-induced evolution with the latest results of empirical research and modelling, this comprehensive-yet-succinct digest will offer the finest available introduction into issues in the field. Foci on well-known and commercially important stocks, together with model-based illustrations of evolutionarily enlightened scenarios for their management, will round off this pioneering educational treatise.

Task 5.9: Final report and recommendations

Particular attention will be devoted to the comprehensive final reporting of all essential findings achieved under the FinE project. These synthetic efforts will culminate in the preparation of scientifically carefully evaluated recommendations, in the form of a Policy Implementation Plan, for updates in fishing and monitoring practices in European waters and beyond. In addition to coordinating the inputs of all teams into this product, fruitful synergies with the planned international conference (Task 5.7) and educational booklet (Task 5.8) on fisheries-induced evolution will be exploited. In particular, it will be ensured that the feedback provided by scientists from outside the FinE project, especially in connection with the planned international conference, on the project’s major and minor findings will be documented and addressed in the final report. Similar insights and feedback will also be used to further enhance the quality of the planned public-relations booklet. The project’s final report and the planned comprehensive booklet are among FinE’s most important deliverables. The project’s ambition is to offer the best summaries available anywhere in the world – comprehensive in the case of the final report, and inspiring in the case of the booklet – on documenting, understanding, and mitigating fisheries-induced evolution.

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Responsible for this page: Melanie Wenighofer
Last updated: 30 Jul 2009