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R. ter Hofstede

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Review (2026) - Zhiyuan Zhao, Tjisse van der Heide, Pauline Kamermans, Joop Coolen, Alex Oude Elferink, Christiaan van Sluis, Remment v. Hofstede, Eline van Onselen, Mark van Koningsveld, More Authors
Artificial reefs (ARs), a form of anthropogenic intervention in marine habitats, have a long history of deployment and continue to proliferate worldwide. Based on a comprehensive literature review and meta-analysis, we show that (i) ARs have evolved from socioeconomic-oriented tools into often-used components of active marine restoration, yet this transition conflicts with the continued use of eco-unfriendly materials, hindering upscaling; (ii) ARs positively impact marine organisms at community, population, and individual levels, but their contributions to organismal fitness remain limited compared to natural reefs. To address these limitations, we advocate a paradigm shift toward “rewilding ARs”─temporary structures designed to create opportunities for natural reef formation, enhance habitat quality, and gradually degrade to minimize human impact. These features support the transition from active intervention to spontaneous recovery, promoting sustainable biodiversity recovery by improving organism fitness and facilitating upscaling. Integrating insights from ecologists, engineers, legal experts, environmental consultants, and NGOs, we introduce six guiding principles for “rewilding ARs” to ensure effective, durable, no-regret, scalable, permit-friendly, and outcome-optimized implementation. Lastly, we present pioneering examples of innovative ARs progressing toward these principles, serving as references for future endeavors. ...
Journal article (2025) - Pascalle Jacobs, Tony Van Der Hiele, Jouke Heringa, Gabrielle P. Verbeeke, Linda Tonk, Remment Ter Hofstede
European oysters (Ostrea edulis) once covered large areas of the North Sea, but have disappeared due to a combination of overexploitation and the destruction of benthic habitats including hard settlement substrate. Offshore wind parks offer an opportunity for oyster restoration as fishing is banned inside these parks and scour protection provides hard settlement substrate. However, ecological restoration of marine systems is capital-intensive. The success of restoration projects is mainly determined by the choice of methods and techniques and consequently costs. Costs and cost-effectiveness information are therefore key in decision making processes concerning the selection of restoration efforts and techniques. So far, economic viability of marine ecosystem restoration have mainly focused on near-shore shallow habitats. The aim of this study was to provide insight into the most cost-effective deployment options to create a European flat oyster reef in an offshore wind farm in the North Sea. Within the current policy and legislation framework, several deployment scenarios were identified based on best practices, expert knowledge, and preliminary results of several pilots. The 9 scenarios included 'adults placed loose on the seafloor, 'adults glued on granite, 'spat settled on shells, 'spat settled on granite and a combined adult and spat scenario. Cost-effectiveness of the different scenarios was determined by modelling the expected reef biomass post-deployment both with and without the option to add additional settlement substrate post-deployment. The main conclusions from this exercise were that: 1. based on investment value, the scenarios adult loose on the seafloor, 'adults in cages and 'spat on shells had the highest revenues per Euro invested; 2. adding substrate in the years post-deployment increased cost-effectiveness in the model for all scenarios, and 3. the time post-deployment to reach a self-sustaining adult oyster population was, with 8-10 yr, shortest for the scenarios 'spat settled on shells' and the combined scenario of 'adults placed loose on the seafloor' and 'spat settled on shells'. ...
Journal article (2025) - Pauline Kamermans, Fleur Anteau, Abel Zempleni, Thomas E. Kodger, Tinka Murk, Karin Didderen, Remment ter Hofstede, Yanhua Zhao, Awen Le Graet, Diede Maas, Stephane Pouvreau, Sophie Valk, Tim Wijgerde
The flat oyster Ostrea edulis L., once common in the North Sea, declined rapidly due to intensive fisheries in the late 19th century and disease outbreaks at the beginning of the 20th century and is now listed as ‘threatened’ or ‘declining,’ with restoration of oyster beds now included in European directives and national plans. For oyster restoration, availability of suitable substrate is required to ensure successful settlement of oyster larvae. Off-shore windfarms are good candidates for restoration as bottom disturbance is not allowed and hard substrate is present in the form of so-called scour protection. This can provide settlement substrate for oyster larvae. In addition to the rock material that currently makes up the scour protection, studies focus on finding alternative and moldable materials that stimulate settlement. The aim of this study was to identify flat oyster larvae settlement preferences for different substrate materials. Oyster settlement on conventional scour protection rock (granite and eclogite), and currently used sandstone and concrete were compared to new types of scour protection rock (marble and limestone). In addition, three new substrates were included in the tests: a coating based on fine ground oyster shells (BESE-reef paste), substrate made of sandy dredged sediment (Geowall) and a bioinspired glue that binds crushed oyster shell fragments together (SeaCrete). Flat oyster larvae were exposed to the substrates in two hatchery experiments as well as under realistic, challenging field conditions. Flat oyster larvae settled on all substrates, with the lowest spat density on eclogite, granite and Geowall and the highest spat density on the two novel substrates SeaCrete and BESE-reef paste. These results promise to enhance native European oyster bed restoration with limited environmental impact as the novel substrates have low CO2 footprints and make use of wasted shells from the seafood industry. ...
Journal article (2024) - Remment ter Hofstede, Sterre Witte, Pauline Kamermans, Mark van Koningsveld, Linda Tonk
The native European flat oyster (Ostrea edulis) is an ecosystem engineer providing important ecosystem services, but became nearly extinct from the North Sea due to diseases and overfishing. There's a growing interest to restore these oyster reefs for their valuable contribution in re-establishing a rich ecosystem in the North Sea. In order to reintroduce the flat oyster population, the availability of hard substrate is crucial for initial settlement and reef development. Such substrate is offered by the infrastructure in offshore wind farms, by means of quarried rock placed at the base of the wind turbine foundations and on top of cable crossings to prevent scouring of the seabed. Further anthropogenic disturbances of the seabed are largely restricted, making wind farm areas promising sites for oyster reef restoration. For successful oyster reef initiation, offering a suitable type of substrate for larvae settlement is important. Here, we assess the settlement preference of flat oysters on 9 different types of substrate, by comparing total settlement, spat densities and spat survival. Oyster larvae settlement preference based on the total number of spat per surface area of the substrate was the highest for granite, a rock type conventionally used as scour protection in offshore wind farms. The lowest settlement preference was observed for steel and the biodegradable polymer BESE. The experiments were performed in a spatting pond and in a natural bay to be able to compare spat collection under controlled and natural conditions. Settlement rates in the spatting pond were much higher than in the natural environment, though survival rates were lower. Our results provide insight in the settlement preference of the European flat oyster for different types of substrate under controlled and natural conditions. Knowing these favorable substrates and conditions for oyster larvae settlement allows for the selection of pro-active measures that contribute to flat oyster reef restoration in the North Sea. ...
Doctoral thesis (2024) - R. ter Hofstede, M. van Koningsveld, Tjeerd J Bouma
Rapid changes in the marine environment are taking place worldwide, for which infrastructural development is one of the most extreme anthropogenic drivers. It comes in many forms and covers functionalities for multiple usages, such as coastal defence, transportation, and energy production. Marine infrastructure modifies seascapes by replacing natural habitats and changing environmental conditions critical to habitat persistence, potentially leading to its degradation and biodiversity loss. Although primarily built to meet functional criteria, their designs can incorporate nature-inclusive elements that benefit ecosystem components, i.e. species, habitats or ecosystem processes.

The implementation of nature-inclusive marine infrastructure is increasingly encouraged, but currently fails to achieve impact at scale due to the fragmented nature of individual measures. Without shared objectives, parallel efforts to enhance targeted ecosystem components might not lead to the desired effect, and could even interfere with each other. A jointly established strategy is required to design and implement nature-inclusive marine infrastructure that meets the wanted impact. Such as strategy is based upon overarching objectives for promoting selected ecosystem components at system-scale, i.e. the seascape dimension required to achieve the desired effect. It is furthermore essential to determine and develop design measures that would induce impact and to define the scale needed for these interventions. It is recognized that marine construction works first serve human needs, not nature goals, but nature-inclusive marine infrastructure does provide an opportunity to benefit ecological values at system-scale. Marine construction works can be synergized with the functioning of the ecosystem in which they are build much better than is currently practiced, and one should always strive for nature-inclusive features in their designs.

This dissertation provides insight into the process to identify, select and implement measures for nature-inclusive marine infrastructure to make a desired impact at system-scale, i.e. the seascape dimension required to achieve that impact. First, a stepwise approach is presented to define clear objectives for improving targeted ecosystem components, in which ruling polices, environmental conditions and the potential of using marine infrastructure are aligned. Stakeholders jointly select the most effective design measures for nature-inclusive marine infrastructure to reach shared targets for ecological impact. Next, it is key to define the scale of these interventions needed to achieve significant impact. A method is developed to select appropriate measures to benefit ecosystem components at a range of scales, from micro-scale (materials used) to mega-scale (connectivity between systems), and to assess their potential effects quantitatively. And finally, it is emphasized that nature-inclusive marine infrastructure can only make impact at system-scale if scientific knowledge about ecosystem functioning is paired with industry-based approaches used for infrastructural development. Five basic principles are provided for establishing this alignment, in order to effectively implement nature-inclusive design measures.

The approaches for engineering nature-inclusive marine infrastructure are demonstrated by defining a strategy to develop European flat oyster (Ostrea edulis) reefs in offshore wind farms in the Southern North Sea. The huge roll out of offshore wind farms aimed at renewable energy production in the North Sea is currently one of the most prominent marine infrastructural developments globally. Its potential for promoting targeted ecosystem components is recognized, as offshore wind farms provide an undisturbed seabed as well as hard substrate infrastructure, which both provide suitable habitat for a wide range of marine organisms. The results of a dedicated monitoring survey in existing offshore wind farms show that their presence indeed contributes to an increase in marine epibenthic biodiversity. Using the offshore wind farm areas specifically for the development of flat oyster reefs has gained particular interest. This species went near to extinct in the 20th century due to overfishing and diseases, and restoring flat oyster reefs in the North Sea meets international policy agreements. Offshore wind farms can be designed to include elements that benefit the restoration of this flat oyster population, such as using a type of hard substrate as scour protection that is favourable for oyster larvae settlement.

In conclusion, this dissertation provides guidance for defining management strategies for implementing nature-inclusive marine infrastructure to achieve impact at system-scale, with an emphasis on flat oyster reef development in offshore wind farms in the Southern North Sea. Application of the presented methods and outcomes of the studies could lead to the realisation of truly effective nature-inclusive marine infrastructure, seizing the opportunity offered by infrastructural developments to have a positive impact on the marine environment.
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Journal article (2024) - Remment ter Hofstede, Mark van Koningsveld
The marine environment faces continuous anthropogenic pressures, including infrastructural developments at a global scale. Integration of nature-inclusive measures in the design of infrastructural development is increasingly encouraged, but a lack of coordination results in fragmentation of project-based measures, failing to meet the desired overall effects. To realize impact at system-scale, i.e. the seascape dimension required to achieve the set objective for a selected ecosystem component, overarching policies with shared targets towards effective nature-inclusive marine infrastructure are needed. We present a stepwise approach to work towards operational objectives for promoting selected ecosystem components that can be species, habitats or ecosystem processes, in which ruling policies, environmental conditions and the use of infrastructural development are aligned, and agreement on achievable ambitions is reached. Having clear targets will provide guidance to project developers in designing the infrastructure nature-inclusive, and in setting up relevant monitoring programs to evaluate the measures taken. We demonstrate how this stepwise approach could be applied to derive operational objectives for the design of nature-inclusive marine infrastructure in the context of offshore windfarm development in the North Sea, currently one of the most prominent infrastructure developments that changes the marine environment drastically. The European flat oyster Ostrea edulis has been selected as target species in the case study, as its once abundant population is now nearly extinct from the North Sea due to human disturbances, and there’s growing interest to restore its reefs. The application of the stepwise approach indicates the potential for oyster reef restoration in the area, based upon a clear match between ruling policy, environmental conditions, and habitat suitability within offshore wind farms. An agreement between the main stakeholders on achievable ambitions can likely be established and would translate into the operational objective to actively introduce oysters to reach an initial critical mass and optimize settlement habitat in all future offshore wind farms in an area with suitable habitat characteristics. Such an agreement on overarching objectives is crucial to align separate initiatives to promote targeted ecosystem components and to jointly become most effective, which is ultimately in the best interest of the larger community using the system. ...

Nature-inclusive design of scour protection in offshore wind farms enhances benthic diversity

Journal article (2024) - Enzo M. Kingma, Remment ter Hofstede, Edwin Kardinaal, Rebecca Bakker, Oliver Bittner, Babeth van der Weide, Joop W.P. Coolen
In the past, a large part of the seabed of the southern North Sea was covered by hard substrates, including oyster beds, coarse peat banks, and glacial erratics. Human activities, particularly bottom trawl fisheries, led to the disappearance of most of these hard substrates, resulting in the loss of its associated diverse benthic life as well. However, the introduction of human-made structures such as offshore wind farms in the North Sea, offers a chance to provide habitat of similar functionality as the former hard substrates. The offshore wind farm infrastructure generally contains layers of rock material deployed at the base of the wind turbine foundations and cable crossings, so-called scour protection, aiming to prevent seabed erosion. The scour protection offers a unique habitat for rock-dwelling benthic organisms in an otherwise mostly soft-bottom environment. By designing the scour protection to be more nature-inclusive, the biodiversity of benthic life can be increased. In this study we examined the effect of substrate material and grading of the scour protection on the epibenthic biodiversity in situ. This was done by deploying research cages containing crates (n = 15) with different types of substrates (concrete, granite, and marble) on the scour protection within an offshore wind farm in the Dutch North Sea. The study revealed a significant (p < 0.05) positive relation between available substrate surface (pebble size) and taxonomic richness. Furthermore, a biological trait assessment of living habits (Tube dwelling, Burrowing, Free living, Crevice dwelling, Epi/endobiotic, and Attached) revealed variations in habit modes across substrate types, with marble and concrete samples showing greatest divergence. Marble samples contained a higher prevalence of tube dwelling organisms, whereas concrete samples contained a relatively higher prevalence of free living, epi/endobiotic and crevice dwelling organisms. The findings support the value of nature-inclusive scour protection designs, emphasizing that both taxonomic and functional diversity can be enhanced by increasing the available surface area of the scour protection and incorporating a variety of substrate types. By adopting these nature-inclusive design components, the coexistence of renewable energy production and a diverse marine benthic community can be further optimized. ...
Review (2024) - Maria L. Vozzo, Christina A. Buelow, Michael Sievers, M. Fernanda Adame, Paul Branson, Maddison Brown, Joseph R. Crosswell, Remment ter Hofstede, Mark van Koningsveld, More Authors...
Cross-habitat facilitative processes can enhance seascape restoration outcomes but there is uncertainty around the spatial dependencies of these processes across habitats. We synthesised the influence of environmental parameters on six processes underpinning cross-habitat facilitation and identified the linear distances over which they operate between habitats. All six process types occur at distances commonly used in seascape restoration demonstrating how harnessing facilitation can scale-up restoration to meet national and international goals. ...
Journal article (2023) - M. L. Vozzo, C. Doropoulos, B. R. Silliman, A. Steven, S. E. Reeves, R. Ter Hofstede, M. van Koningsveld, J. van de Koppel, T. McPherson, More authors...
Journal article (2023) - Remment Ter Hofstede, Gregory Williams, Mark Van Koningsveld
Incorporation of ecology and ecosystem services into marine infrastructural developments has gained interest over the last decades. Growing attention is given to combine the massive roll-out of offshore wind farms in the North Sea with reinstating the once rich but nowadays nearly extinct European flat oyster (Ostrea edulis). However, the practical upscaling of these pilots is hindered by the absence of clear management objectives and the lack of quantitative knowledge on the effect of technical interventions that could stimulate oyster reef development. Consequently, it is unclear what scale of intervention would actually be required to achieve overall management objectives. This paper presents a stepwise procedure designed in particular to guide the selection of appropriate measures and their required scale for pro-actively facilitating flat oyster reef development in offshore wind farms, in order to reach a desired state for oyster reef inclusive wind farms. The stepwise procedure addresses the historical and current situation of the physical system and social environment, provides options for intervention that stimulate oyster reef development at a range of scales, from micro-scale (materials used) to mega-scale (connectivity between wind farms), and quantitatively assesses the potential effect of applying these interventions. Assumptions have been made in quantifying the effort required for developing oyster reefs in offshore wind farms, and refinement is obviously needed. However, this is a first attempt to make such estimates. The outcomes provide direction in identifying research needs to fill knowledge gaps, as well as in decision-making during the design process for inducing oyster reef development in offshore wind farms. Herewith, application of the stepwise procedure supports authorities in restoration management for the successful reinstatement of flat oyster reefs in the southern North Sea. ...
Journal article (2023) - Miriam Schutter, Remment ter Hofstede, Joost Bloemberg, Jesper Elzinga, Mark van Koningsveld, Ronald Osinga
Acropora palmata is one of the major reef-building coral species in the Caribbean. The species has suffered drastic declines in abundance and sexual recruitment over the past decades. One method for active rehabilitation of A. palmata reefs is by assisting the production of sexual recruits under controlled lab conditions. Within this study, the effect of different aquaculture regimes and culturing periods on the survival rates of these recruits was investigated. In August 2016, coral spawn was collected on a reef nearshore New Providence, Bahamas, cross-fertilized, and reared in mobile laboratory facilities from Van Oord Dredging and Marine Contractors. Larvae were settled on pre-conditioned aragonite plugs. Sexual recruits were cultured under four different aquaculture conditions: ambient vs. high Total Alkalinity (TA) (∼2.8 mEq L−1 vs. 4.8 mEq L−1) and with vs. without feeding Artemia nauplii. Recruit size was monitored by tracking living tissue area and the number of polyps of a subset of recruits. Plates with recruits were outplanted to a nursery on the reef after 4, 9 and 14 weeks of aquaculture. Survival was determined during the aquaculture phase (at 4, 9 and 14 weeks after settlement), and after outplanting (at 27 and 44 weeks after settlement). During the aquaculture phase, survival was significantly lower in seawater with increased TA compared to ambient seawater conditions. The average number of polyps per recruit was significantly higher in the treatments with feeding. After outplanting to the reef, both survival and recruit size were highest in the feeding treatments. The most successful aquaculture treatment in this study was a combination of increased TA and feeding during 9 weeks of aquaculture, which resulted in a doubling of survival and recruit size at 10 months after settlement compared to ambient conditions. Ambient conditions did not enhance survivorship nor recruit size at 10 months after settlement, as compared to the other aquaculture treatments. Nevertheless, the success of ambient aquaculture conditions exceeded natural conditions, as no natural recruitment of A. palmata was observed in this study. We conclude that feeding during and ex-situ culture period enhances ex-situ growth rates and in situ recruit survival of A. palmata juveniles. No positive effects of the aquaculture treatment with only increased TA were found. Building on these results, recommendations are provided for future reef rehabilitation efforts using ex-situ rearing of sexually reproduced A. palmata recruits. ...
Journal article (2023) - Remment ter Hofstede, Tjeerd J. Bouma, Mark van Koningsveld
Marine reef ecosystems have degraded massively worldwide, and restoration efforts have as yet not managed to realize the scale required to reverse continued degradation. To achieve effective scales, scientific insights in restoration methods should be paired with industry-based approaches used for infrastructural development. We illustrate by five principles how long-standing experience of marine contractors with executing large-scale projects, can support reef restoration: i) utilizing industrial techniques to achieve positive impact at scale, ii) landscaping infrastructure to optimize habitat for targeted species, iii) inducing life to overcome connectivity bottle-necks and steer community composition, iv) designing nature development efforts to be self-sustainable, and v) ensuring continuity beyond project boundaries by early stakeholder engagement. Consciously connecting scientific knowledge to industry-based activities increases the likelihood that marine infrastructure development and ecosystem rehabilitation can be aligned. We plead that synergizing practices by science and industry is needed to upscale restoration efforts and truly improve marine reef ecosystems. ...
Journal article (2022) - R. ter Hofstede, F. M.F. Driessen, P. J. Elzinga, M. Van Koningsveld, M. Schutter
The North Sea was once abundantly covered with hard substrates such as oyster beds, coarse peat banks and glacial erratics, providing habitat to a rich community of marine species. Most of these habitats were destroyed by bottom-trawl fisheries over the past century, and today, the seabed hosts a relatively poor species community. Emerging offshore windfarms include the re-introduction of hard substrate by means of scour protection around the foundation of wind turbines. It is assumed that the new habitat will contribute to marine biodiversity, and this study aims to demonstrate that. Video data were collected using a Remotely Operated Vehicle in four wind farms in the southern North Sea. A quantitative assessment was made to determine the effect of scour protection on community structure. The assessment revealed distinct community clusters for geographic location and seabed type. Windfarms closely located to each other had a more similar epibenthic community compared to those further away. The epibenthic community at the rocky armour layer of the scour protection had a different species composition and a higher species abundance than the one at the sandy seabed surrounding it. Species diversity by means of richness, evenness and the Shannon diversity index was not consistently higher or lower for the communities at the different seabed types. This study shows that marine life inhabits scour protection in offshore wind farms and that it is different from the community living at the surrounding seabed. Knowing the potential epibenthic community structure at and around a scour protection supports the development of new wind farms that include components to enhance their ecological value. Herewith, our study contributes to efforts to restore biodiversity in the North Sea. ...

Review of concepts, policies, tools, and critical issues

Review (2011) - Stelios Katsanevakis, Vanessa Stelzenmüller, Giovanni D'Anna, Mike Duijn, Tatiana Filatova, Fabio Fiorentino, Helena Hulsman, Kate Johnson, Aristomenis P. Karageorgis, Ingrid Kröncke, Simone Mirto, Carlo Pipitone, Andy South, Susan Portelli, Wanfei Qiu, Henning Reiss, Dimitris Sakellariou, Maria Salomidi, Luc van Hoof, Vassiliki Vassilopoulou, Tomás Vega Fernández, Sandra Vöge, Anke Weber, Thomas Kirk Sørensen, Argyro Zenetos, Remment ter Hofstede, Peter J.S. Jones, Sandy Kerr, Fabio Badalamenti, Christos Anagnostou, Patricia Breen, Guillem Chust
Conventional sectoral management and piecemeal governance are considered less and less appropriate in pursuit of sustainable development. Ecosystem based marine spatial management (EB-MSM) is an approach that recognizes the full array of interactions within an ecosystem, including human uses, rather than considering single issues, species, or ecosystem services in isolation. Marine spatial planning and ocean zoning are emerging concepts that can support EB-MSM. EB-MSM is driven by high-level goals that managers aim to achieve through the implementation of measures. High-level goals and objectives need to be translated into more operational objectives before specific targets, limits and measures can be elaborated.Monitoring, evaluation and adaptation are necessary to ensure that marine management measures are both effective and efficient. Solid monitoring frameworks are the foundation of adaptive management, as they provide the necessary information to evaluate performance and the effectiveness of management actions. Marine protected areas (MPAs) - possibly set up in networks - constitute a key component in EB-MSM policies and practises and have been applied as a cornerstone in conservation of marine biodiversity, management of fish populations, development of coastal tourism, etc. Moreover, MPA experiences have provided methods and concepts (such as zoning) to a wider EB-MSM context. The assignment of values to biophysical features of the marine environment allows the direct assessment of related management choices and may assist EB-MSM.A range of monetary valuation techniques have been proposed to reduce attributes of goods and services to a single metric. However, in the marine environment such an approach is often over simplistic, and thus less reductive techniques may be necessary. Rather than producing a single metric, the results of non-monetary assessments guide policy allowing weight to be given as necessary to potential areas of conflict and consensus.Strategies to take into account climate change effects and geohazard risks in EB-MSM have been applied or proposed worldwide. EB-MSM regimes must be alert to such risks and flexible to account for changes. ...