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Knowledge of genetic structure at the finest level is essential for the conservation of genetic resources. Despite no visible barriers limiting gene flow, significant genetic structure has been shown in marine species. The common cockle (Cerastoderma edule) is a bivalve of great commercial and ecological value inhabiting the Northeast Atlantic Ocean. Previous population genomics studies demonstrated significant structure both across the Northeast Atlantic, but also within small geographic areas, highlighting the need to investigate fine-scale structuring. Here, we analysed two geographic areas that could represent opposite models of structure for the species: (1) the SW British Isles region, highly fragmented due to biogeographic barriers, and (2) Galicia (NW Spain), a putative homogeneous region. A total of 9250 SNPs genotyped by 2b-RAD on 599 individuals from 22 natural beds were used for the analysis. The entire SNP dataset mostly confirmed previous observations related to genetic diversity and differentiation; however, neutral and divergent SNP outlier datasets enabled disentangling physical barriers from abiotic environmental factors structuring both regions. While Galicia showed a homogeneous structure, the SW British Isles region was split into four reliable genetic regions related to oceanographic features and abiotic factors, such as sea surface salinity and temperature. The information gathered supports specific management policies of cockle resources in SW British and Galician regions also considering their particular socio-economic characteristics; further, these new data will be added to those recently reported in the Northeast Atlantic to define sustainable management actions across the whole distribution range of the species

A better understanding of growth drivers in shellfish populations including the common cockle Cerastoderma edule is essential, as their future is challenged by unsustainable fishing practices and climate change. In this study the spatial and temporal variabilities in common cockle growth across latitudes were assessed and compared with historical data. Six locations were examined at bimonthly intervals over 19 months; three Irish, two Welsh and one French, spanning the latitudes 54°N to 44°N. The results demonstrated that local abiotic and biotic factors have a larger impact on cockle growth than latitude. Cockles at similar latitudes grew at different rates and sizes, possibly due to factors such as density, fishing activity and interspecific competition. Cockles (0–3 years) impacted by low salinity and parasites (trematodes), exhibited reduced growth in later years. At the warmest, southernmost site growth was lowest in cockles >2 years. Previously, cockles at that site have been shown to spawn almost year-round, possibly diverting energy to gonad development rather than growth. The results opposed previously literature which demonstrated significantly greater growth at lower latitudes. These findings affirm that cockle growth and size is variable due to local abiotic (reduced salinity) and biotic (potentially trematode infection) drivers. Additionally, the synergistic relationship between these factors, i.e. warmer temperatures driving prolonged spawning, and the potential association between lower salinities and trematode prevalence, is concerning due to predicted climate related increases in temperature, precipitation and trematode prevalence/transmission, which may result in northern cockles reaching smaller maximum sizes.

Parasite species richness is influenced by a range of drivers including host related factors (e.g. host size) and environmental factors (e.g. seawater temperature). However, identification of modulators of parasite species richness remains one of the great unanswered questions in ecology. The common cockle Cerastoderma edule is renowned for its diversity and abundance of parasites, yet drivers of parasite species richness in cockles have not been examined to investigate the association of both macro and microparasite communities. Using cockles as a model species, some of the key drivers of parasite prevalence and parasite species richness were investigated. Objectives of this 19-month survey were to determine the influence of the environment, host-parasite dynamics and parasite associations on parasite species richness and prevalence at two different geographic latitudes, chosen based on environmental differences. The highest parasite species richness was recorded in the northern sites, and this was potentially influenced by a range of interactions between the host, the pathogens and the environment. Parasite prevalence increased with host size and age, and parasite species richness increased with reduced salinity. A number of interactions between parasites, and between parasites and pathologies may be influencing parasite infection dynamics. New and concerning information is also presented regarding interactions between parasites and their environment. A number of parasites and potential pathogens (bacteria, Trichodina ciliates, metacercariae, trematode sporocysts) may be advantaged under climate change conditions (warming seas, increased precipitation), increasing disease incidence, which may prove detrimental not just for cockles, but for other bivalve species in the future.

Parasite species richness is influenced by a range of drivers including host related factors (e.g. host size) and environmental factors (e.g. seawater temperature). However, identification of modulators of parasite species richness remains one of the great unanswered questions in ecology. The common cockle Cerastoderma edule is renowned for its diversity and abundance of parasites, yet drivers of parasite species richness in cockles have not been examined to investigate the association of both macro and microparasite communities. Using cockles as a model species, some of the key drivers of parasite prevalence and parasite species richness were investigated. Objectives of this 19-month survey were to determine the influence of the environment, host-parasite dynamics and parasite associations on parasite species richness and prevalence at two different geographic latitudes, chosen based on environmental differences. The highest parasite species richness was recorded in the northern sites, and this was potentially influenced by a range of interactions between the host, the pathogens and the environment. Parasite prevalence increased with host size and age, and parasite species richness increased with reduced salinity. A number of interactions between parasites, and between parasites and pathologies may be influencing parasite infection dynamics. New and concerning information is also presented regarding interactions between parasites and their environment. A number of parasites and potential pathogens (bacteria, Trichodina ciliates, metacercariae, trematode sporocysts) may be advantaged under climate change conditions (warming seas, increased precipitation), increasing disease incidence, which may prove detrimental not just for cockles, but for other bivalve species in the future.

A better understanding of growth drivers in shellfish populations including the common cockle Cerastoderma edule is essential, as their future is challenged by unsustainable fishing practices and climate change. In this study the spatial and temporal variabilities in common cockle growth across latitudes were assessed and compared with historical data. Six locations were examined at bimonthly intervals over 19 months; three Irish, two Welsh and one French, spanning the latitudes 54°N to 44°N. The results demonstrated that local abiotic and biotic factors have a larger impact on cockle growth than latitude. Cockles at similar latitudes grew at different rates and sizes, possibly due to factors such as density, fishing activity and interspecific competition. Cockles (0–3 years) impacted by low salinity and parasites (trematodes), exhibited reduced growth in later years. At the warmest, southernmost site growth was lowest in cockles >2 years. Previously, cockles at that site have been shown to spawn almost year-round, possibly diverting energy to gonad development rather than growth. The results opposed previously literature which demonstrated significantly greater growth at lower latitudes. These findings affirm that cockle growth and size is variable due to local abiotic (reduced salinity) and biotic (potentially trematode infection) drivers. Additionally, the synergistic relationship between these factors, i.e. warmer temperatures driving prolonged spawning, and the potential association between lower salinities and trematode prevalence, is concerning due to predicted climate related increases in temperature, precipitation and trematode prevalence/transmission, which may result in northern cockles reaching smaller maximum sizes.

1. Coastal systems provide many cultural ecosystem services (CES) to humans. Fewer studies have focused solely on CES, while those comparing CES across countries are even rarer. In the case of shellfish, considerable ecosystem services focus has been placed on nutrient remediation, with relatively little on the cultural services provided, despite strong historical, cultural, social and economic links between shellfish and coastal communities. The ecosystem services provided by the common cockle, Cerastoderma edule, have recently been described, yet the cultural benefits from cockles remain mostly unknown. 2. Here, we documented the CES provided by C. edule in five maritime countries along the Atlantic coast of western Europe, classifying evidenced examples of services into an a priori framework. The high-level classes, adapted from the Millennium Assessment and the Common International Classification of Ecosystem Services, were: inspirational, sense of place, spiritual & religious, aesthetic, recreation & ecotourism, cultural heritage and educational. A further 19 sub-classes were defined. We followed a narrative approach to draw out commonalities and differences among countries using a semi-quantitative analysis. 3. Examples of CES provided by cockles were found for all classes in most countries. Cockles supply important and diverse cultural benefits to humans across Atlantic Europe, making it an ideal model species to study CES in coastal areas. Most examples were in cultural heritage, highlighting the importance of this class in comparison with classes which typically receive more attention in the literature like recreation or aesthetics. We also found that the cultural associations with cockles differed among countries, including between neighbouring countries that share a strong maritime heritage. The extent to which cultural associations were linked with the present or past also differed among countries, with stronger association with the present in southern c

Reproduction and parasites have significant impacts on marine animal populations globally. This study aimed to investigate the associative effects of host reproduction and a host–parasite interplay on a marine bivalve, along a geographic gradient of latitude. Cockles Cerastoderma edule were sampled from five European sites (54°N to 40°N), between April 2018 and October 2019. A histological survey provided data on trematode (metacercaria and sporocyst life stages), prevalence, and cockle stage of gametogenesis to assess the influence of a latitudinal gradient on both interplays. Sex ratios at the northernmost sites were skewed toward females, and spawning size was reduced at the lower latitudes. Trematode infection did not follow a latitudinal gradient. Localized site‐related drivers, namely seawater temperature, varied spatially, having an impact on cockle–trematode interactions. Spawning was related to elevated temperatures at all sites. Prolonged spawning occurred at southern latitudes, where seawater temperatures were warmer. Trematode prevalence and the impact of trematodes on gametogenesis were found to be spatially variable, but not latitudinally. Therefore, it is not possible to determine the likelihood of boom and bust events in cockles, based on the latitudinal location of a population. In terms of sublethal impacts, it appeared that energy was allocated to reproduction rather than somatic growth in southern populations, with less energy allocated to reproduction in the larger, northern cockles. The demonstrated spatial trend of energy allocation indicates the potential of a temporal trend of reduced cockle growth at northern sites, as a result of warming sea temperatures. This awareness of the spatially varying drivers of populations is crucial considering the potential for these drivers/inhibitors to be exacerbated in a changing marine environment.

Reproduction and parasites have significant impacts on marine animal populations globally. This study aimed to investigate the associative effects of host reproduction and a host–parasite interplay on a marine bivalve, along a geographic gradient of latitude. Cockles Cerastoderma edule were sampled from five European sites (54°N to 40°N), between April 2018 and October 2019. A histological survey provided data on trematode (metacercaria and sporocyst life stages), prevalence, and cockle stage of gametogenesis to assess the influence of a latitudinal gradient on both interplays. Sex ratios at the northernmost sites were skewed toward females, and spawning size was reduced at the lower latitudes. Trematode infection did not follow a latitudinal gradient. Localized site‐related drivers, namely seawater temperature, varied spatially, having an impact on cockle–trematode interactions. Spawning was related to elevated temperatures at all sites. Prolonged spawning occurred at southern latitudes, where seawater temperatures were warmer. Trematode prevalence and the impact of trematodes on gametogenesis were found to be spatially variable, but not latitudinally. Therefore, it is not possible to determine the likelihood of boom and bust events in cockles, based on the latitudinal location of a population. In terms of sublethal impacts, it appeared that energy was allocated to reproduction rather than somatic growth in southern populations, with less energy allocated to reproduction in the larger, northern cockles. The demonstrated spatial trend of energy allocation indicates the potential of a temporal trend of reduced cockle growth at northern sites, as a result of warming sea temperatures. This awareness of the spatially varying drivers of populations is crucial considering the potential for these drivers/inhibitors to be exacerbated in a changing marine environment.

1. Coastal systems provide many cultural ecosystem services (CES) to humans. Fewer studies have focused solely on CES, while those comparing CES across countries are even rarer. In the case of shellfish, considerable ecosystem services focus has been placed on nutrient remediation, with relatively little on the cultural services provided, despite strong historical, cultural, social and economic links between shellfish and coastal communities. The ecosystem services provided by the common cockle, Cerastoderma edule, have recently been described, yet the cultural benefits from cockles remain mostly unknown. 2. Here, we documented the CES provided by C. edule in five maritime countries along the Atlantic coast of western Europe, classifying evidenced examples of services into an a priori framework. The high-level classes, adapted from the Millennium Assessment and the Common International Classification of Ecosystem Services, were: inspirational, sense of place, spiritual & religious, aesthetic, recreation & ecotourism, cultural heritage and educational. A further 19 sub-classes were defined. We followed a narrative approach to draw out commonalities and differences among countries using a semi-quantitative analysis. 3. Examples of CES provided by cockles were found for all classes in most countries. Cockles supply important and diverse cultural benefits to humans across Atlantic Europe, making it an ideal model species to study CES in coastal areas. Most examples were in cultural heritage, highlighting the importance of this class in comparison with classes which typically receive more attention in the literature like recreation or aesthetics. We also found that the cultural associations with cockles differed among countries, including between neighbouring countries that share a strong maritime heritage. The extent to which cultural associations were linked with the present or past also differed among countries, with stronger association with the present in southern c

Reproduction and parasites have significant impacts on marine animal populations globally. This study aimed to investigate the associative effects of host reproduction and a host–parasite interplay on a marine bivalve, along a geographic gradient of latitude. Cockles Cerastoderma edule were sampled from five European sites (54°N to 40°N), between April 2018 and October 2019. A histological survey provided data on trematode (metacercaria and sporocyst life stages), prevalence, and cockle stage of gametogenesis to assess the influence of a latitudinal gradient on both interplays. Sex ratios at the northernmost sites were skewed toward females, and spawning size was reduced at the lower latitudes. Trematode infection did not follow a latitudinal gradient. Localized site‐related drivers, namely seawater temperature, varied spatially, having an impact on cockle–trematode interactions. Spawning was related to elevated temperatures at all sites. Prolonged spawning occurred at southern latitudes, where seawater temperatures were warmer. Trematode prevalence and the impact of trematodes on gametogenesis were found to be spatially variable, but not latitudinally. Therefore, it is not possible to determine the likelihood of boom and bust events in cockles, based on the latitudinal location of a population. In terms of sublethal impacts, it appeared that energy was allocated to reproduction rather than somatic growth in southern populations, with less energy allocated to reproduction in the larger, northern cockles. The demonstrated spatial trend of energy allocation indicates the potential of a temporal trend of reduced cockle growth at northern sites, as a result of warming sea temperatures. This awareness of the spatially varying drivers of populations is crucial considering the potential for these drivers/inhibitors to be exacerbated in a changing marine environment.

Reproduction and parasites have significant impacts on marine animal populations globally. This study aimed to investigate the associative effects of host reproduction and a host–parasite interplay on a marine bivalve, along a geographic gradient of latitude. Cockles Cerastoderma edule were sampled from five European sites (54°N to 40°N), between April 2018 and October 2019. A histological survey provided data on trematode (metacercaria and sporocyst life stages), prevalence, and cockle stage of gametogenesis to assess the influence of a latitudinal gradient on both interplays. Sex ratios at the northernmost sites were skewed toward females, and spawning size was reduced at the lower latitudes. Trematode infection did not follow a latitudinal gradient. Localized site‐related drivers, namely seawater temperature, varied spatially, having an impact on cockle–trematode interactions. Spawning was related to elevated temperatures at all sites. Prolonged spawning occurred at southern latitudes, where seawater temperatures were warmer. Trematode prevalence and the impact of trematodes on gametogenesis were found to be spatially variable, but not latitudinally. Therefore, it is not possible to determine the likelihood of boom and bust events in cockles, based on the latitudinal location of a population. In terms of sublethal impacts, it appeared that energy was allocated to reproduction rather than somatic growth in southern populations, with less energy allocated to reproduction in the larger, northern cockles. The demonstrated spatial trend of energy allocation indicates the potential of a temporal trend of reduced cockle growth at northern sites, as a result of warming sea temperatures. This awareness of the spatially varying drivers of populations is crucial considering the potential for these drivers/inhibitors to be exacerbated in a changing marine environment.

Knowledge mobilisation is required to “bridge the gap” between research, policy and practice. This activity is dependent on the amount, richness and quality of the data published. To understand the impact of a changing climate on commercial species, stakeholder communities require better knowledge of their past and current situations. The common cockle (Cerastoderma edule) is an excellent model species for this type of analysis, as it is well-studied due to its cultural, commercial and ecological significance in west Europe. Recently, C. edule harvests have decreased, coinciding with frequent mass mortalities, due to factors such as a changing climate and diseases. In this study, macro and micro level marine historical ecology techniques were used to create datasets on topics including: cockle abundance, spawning duration and harvest levels, as well as the ecological factors impacting those cockle populations. These data were correlated with changing climate and the Atlantic Multidecadal Oscillation (AMO) index to assess if they are drivers of cockle abundance and harvesting. The analyses identified the key stakeholder communities involved in cockle research and data acquisition. It highlighted that data collection was sporadic and lacking in cross-national/stakeholder community coordination. A major finding was that local variability in cockle populations is influenced by biotic (parasites) and abiotic (temperature, legislation and harvesting) factors, and at a global scale by climate (AMO Index). This comprehensive study provided an insight into the European cockle fishery but also highlights the need to identify the type of data required, the importance of standardised monitoring, and dissemination efforts, taking into account the knowledge, source, and audience. These factors are key elements that will be highly beneficial not only to the cockle stakeholder communities but to other commercial species.

Coastal habitats provide many important ecosystem services. The substantial role of shellfish in delivering ecosystem services is increasingly recognised, usually with a focus on cultured species, but wild-harvested bivalve species have largely been ignored. This study aimed to collate evidence and data to demonstrate the substantial role played by Europe's main wild-harvested bivalve species, the common cockle Cerastoderma edule, and to assess the ecosystem services that cockles provide. Data and information are synthesised from five countries along the Atlantic European coast with a long history of cockle fisheries. The cockle helps to modify habitat and support biodiversity, and plays a key role in the supporting services on which many of the other services depend. As well as providing food for people, cockles remove nitrogen, phosphorus and carbon from the marine environment, and have a strong cultural influence in these countries along the Atlantic coast. Preliminary economic valuation of some of these services in a European context is provided, and key knowledge gaps identified. It is concluded that the cockle has the potential to become (i) an important focus of conservation and improved sustainable management practices in coastal areas and communities, and (ii) a suitable model species to study the integration of cultural ecosystem services within the broader application of ‘ecosystem services’.

Knowledge mobilisation is required to “bridge the gap” between research, policy and practice. This activity is dependent on the amount, richness and quality of the data published. To understand the impact of a changing climate on commercial species, stakeholder communities require better knowledge of their past and current situations. The common cockle (Cerastoderma edule) is an excellent model species for this type of analysis, as it is well-studied due to its cultural, commercial and ecological significance in west Europe. Recently, C. edule harvests have decreased, coinciding with frequent mass mortalities, due to factors such as a changing climate and diseases. In this study, macro and micro level marine historical ecology techniques were used to create datasets on topics including: cockle abundance, spawning duration and harvest levels, as well as the ecological factors impacting those cockle populations. These data were correlated with changing climate and the Atlantic Multidecadal Oscillation (AMO) index to assess if they are drivers of cockle abundance and harvesting. The analyses identified the key stakeholder communities involved in cockle research and data acquisition. It highlighted that data collection was sporadic and lacking in cross-national/stakeholder community coordination. A major finding was that local variability in cockle populations is influenced by biotic (parasites) and abiotic (temperature, legislation and harvesting) factors, and at a global scale by climate (AMO Index). This comprehensive study provided an insight into the European cockle fishery but also highlights the need to identify the type of data required, the importance of standardised monitoring, and dissemination efforts, taking into account the knowledge, source, and audience. These factors are key elements that will be highly beneficial not only to the cockle stakeholder communities but to other commercial species.

Coastal habitats provide many important ecosystem services. The substantial role of shellfish in delivering ecosystem services is increasingly recognised, usually with a focus on cultured species, but wild-harvested bivalve species have largely been ignored. This study aimed to collate evidence and data to demonstrate the substantial role played by Europe's main wild-harvested bivalve species, the common cockle Cerastoderma edule, and to assess the ecosystem services that cockles provide. Data and information are synthesised from five countries along the Atlantic European coast with a long history of cockle fisheries. The cockle helps to modify habitat and support biodiversity, and plays a key role in the supporting services on which many of the other services depend. As well as providing food for people, cockles remove nitrogen, phosphorus and carbon from the marine environment, and have a strong cultural influence in these countries along the Atlantic coast. Preliminary economic valuation of some of these services in a European context is provided, and key knowledge gaps identified. It is concluded that the cockle has the potential to become (i) an important focus of conservation and improved sustainable management practices in coastal areas and communities, and (ii) a suitable model species to study the integration of cultural ecosystem services within the broader application of ‘ecosystem services’.