COASTFRAG - Impact of habitat fragmentation and loss on coastal ecosystems and implications for sustainable management under climate change

Abstract

This is the project description for the COASTFRAG project, 2021-2024

Full text

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COASTFRAG – Impact of habitat fragmentation and loss on coastal ecosystems:
implications for sustainable management under climate change
1. Excellence
1.1 State of the art, knowledge needs and project objectives
Worldwide, the environment is undergoing rapid changes due to activities of the growing human
population, with climate change being a major concern. As more and more people settle along the coast,
human-induced pressures also increase (Halpern et al. 2015), causing coastal habitat loss and
fragmentation. These pressures lead to a dramatic decline in biodiversity and changes to ecosystem
functioning worldwide (Hennige & Roberts 2019, IPBES 2019). This in turn limits the potential for Blue
Growth (i.e. the economic growth based on marine and aquatic resources) and hampers the ambitions of a
doubling of the ocean economy in 2030 compared to 2010 (OECD 2016). This negative trend also
undermines the progress towards several of the assessed targets of the Sustainable Development Goals
(SDGs), related to poverty, hunger, health, climate and ocean life (SDGs 1, 2, 3, 13 and 14, respectively).
Loss of biodiversity is therefore considered to be not only an environmental issue, but also a
developmental, economic, security, social and moral issue.
Blue forests, such as seaweeds, kelp forests, seagrass meadows, salt marshes and mangroves, are coastal
vegetated habitats that cover huge areas around the globe, providing many different types of ecosystem
services (e.g. Barbier et al. 2011, Smale et al. 2013, Rebours et al. 2014). Littoral seaweeds, covering the
rocky shores everywhere, form the basis for many of the SDG targets by providing raw material for humans
(including food, animal feed and fertilizers), food and shelter for a variety of species (Spruzen et al. 2008,
Clayden et al. 2014) and by regulating global climate through carbon storage and sequestration (Krause-
Jensen & Duarte 2016). These ecosystems are experiencing multiple pressures caused by climate change
and human activities, but the effect of this is largely unexplored.
The overall aim of COASTFRAG is to contribute with the knowledge base needed to
safeguard the Blue forests’ ecosystem services and ensure a long-term sustainable Blue
growth (as stated in the EU Blue Growth strategy and the Norwegian Government's Updated
Ocean Strategy). To do so, we will study how pressures, individually and together, impact
seaweed communities across Europe (Figure 1). The findings will be communicated to a
wider audience, including to support regional and habitat specific management strategies.
Figure 1. COASTFRAG will
perform fragmentation field
studies (black dots) in four
European regions selected
to cover different
temperature (A) and water
quality (B) regimes: the
Norwegian Sea, the Baltic
Sea (Estonia), the Scottish
Atlantic Ocean (UK) and the
Mediterranean Sea (Spain).
Mesocosm experiments will
be carried out in Norway.
Note that the turbidity
model (B) is lacking data for
the Baltic Sea.
A
B
Water quality (light
attenuation, KdPAR)
High
Low
Seabed temperature
High
Low

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Littoral seaweed communities generally experience high variation in environmental conditions, such as
temperature (due to the tidal cycles) and wave exposure, which give rise to a variety of species and
habitats, different levels of productivity and differences in ecological function (Macreadie et al. 2017). The
climate and human activity related pressures that act on these ecosystems (Halpern et al. 2015,
Mieszkowska et al. 2019) and the subsequent loss and fragmentation of habitats (Tscharntke et al. 2012)
can result in small patches and the isolation of populations. This can in turn have profound consequences
on species diversity and ecosystem functioning (Fahrig 2003, Lindenmayer & Fischer 2007), including the
potential for regime shifts (i.e. abrupt and persistent shifts in ecosystem structure and function). Regime
shifts have been observed in kelp forests worldwide (Ling et al. 2015, Filbee-Dexter & Wernberg, 2018), but
have rarely been studied in intertidal seaweed communities (Petraitis & Latham 1999, Dudgeon & Petraitis
2001).
Habitat fragmentation, climate change and environmental variability affect species differently (Fischer &
Lindenmayer 2007), and the effect of multiple disturbances will therefore differ between habitats. Habitat
fragmentation has been thoroughly studied in terrestrial environments (Debinski & Holt 1999, Fischer &
Lindenmayer 2007), but few studies have addressed this for vegetated marine ecosystems (Airoldi et al.
2008), with seagrass habitats as the exception (Eggleston et al. 1998, Hovel & Lipcius 2001, 2002, but see
Caley et al. 2001 and Deza & Anderson 2010 on kelp).
COASTFRAG’s primary objective is to resolve how the pressure of habitat fragmentation and loss impacts
the species composition, structure and ecological function of intertidal seaweed communities while also
experiencing multiple pressures caused by climate change (increased temperature, reduced water quality,
higher rate of storm events, migration of species) and human activities (pollution). The data will feed into
models to assess seaweed communities, today and under future climate, providing robust predictions to
support decision makers and other stakeholders.
1.2 Research questions and hypotheses, theoretical approach and methodology
To answer COASTFRAG’s primary objective we will focus on four research questions (Q1-Q4):
Q1. Fragmentation, global stressors and scale – How does habitat fragmentation impact seaweed
communities under different climate regimes and at different spatial scales?
Q2. Fragmentation, environmental conditions and local human pressures – How does environmental
conditions, local human pressures and habitat fragmentation interact to impact coastal seaweed
communities and the potential for regime shifts?
Q3. Fragmentation and species interactions – How do different predatory and grazing pressures modify
the effect that habitat fragmentation has on seaweed communities?
Q4. Predicting the future – What changes will we see in seaweed communities facing climate change
COASTFRAG is structured into 6 work packages (WPs). WP1 will be responsible for compilation of data and
data management, coordination of Master and Postdoc activities and the sampling design. WP2-4 will be
responsible for the fragmentation and clearing experiments and the analyses across Europe. WP5 will
perform modelling and climate change projections and predict the future of seaweed communities. WP6
will extract the knowledge from all WPs and communicate this to a wider audience, including management
strategies to decision makers and stakeholders. Section 3.2 describes the organisation of the WPs in detail.
We expect that the response of the seaweed communities will be impacted by variation in temperature
and water quality (such as nutrient level). Some of the species already live close to their temperature
tolerance limit (Ugarte et al. 2010) and species abundances vary with nutrient levels (Bellgrove et al. 2010,
Macready et al. 2017). COASTFRAG will therefore be carried out across four regions in Europe to cover the
European range of temperature and water quality (Figure 1). The Norwegian Sea studies will take place in
Møre & Romsdal, on the West coast of Norway, a relatively pristine area with clear and cold waters, and a
wide range of wave exposure. Here, seaweeds are harvested commercially (incl. Ascophyllum nodosum,
which is one of the focal species in this project). Mesocosm experiments will be carried out at the NIVA
Marine Research Station Solbergstrand in the outer Oslofjord. The Baltic Sea study will take place in the
Gulf of Riga and the Baltic Proper, a relatively cold and turbid region. In the Baltic Proper the nutrient level
is moderate, but the Gulf of Riga has high levels due to the heavy riverine nutrient loads and limited water

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exchange (Kotta et al. 2008, 2020). The region is experiencing increased levels of habitat fragmentation due
to shifting patterns in salinity and storms, but also direct human-induced disturbance resulting in
eutrophication and increased turbidity. The Baltic Sea differs from the other regions in this project by
having brackish water and limited tides. The Scottish Atlantic Ocean studies will take place on the rocky
shores of West Scotland (UK), an area with moderately warm and relatively turbid waters and a wide range
of wave exposure. The area also has local nutrient enrichment, with higher levels in Loch Fyne (a fjord, sea
loch, connected to the catchment area of Glasgow and surrounding towns) that at the Firth of Lorn (an area
with few towns and little agriculture). A. nodosum is harvested commercially in this part of the Scottish
coast. The Mediterranean Sea studies will take place in the Catalan Coast and the Balearic Islands (NW
Mediterranean, Spain), an area with warm and relative clear waters. The area has different levels of
nutrients, with increased levels close to big cities and touristic places compared to more remote areas. The
Mediterranean Sea has highly fragmented habitats and is considered to be very sensitive to disturbances
(Giorgi 2006).
The accessibility to study sites (no need for diving or deep-water equipment in the littoral zone) and the
possibility to have relatively good control of the pressures acting on them, make intertidal shores ideal
systems for experimental manipulations. The habitats selected for this study (i.e. focal seaweeds)
comprise littoral seaweed beds of the species Fucus vesiculosus, Ascophyllum nodosum and the genus
Cystoseira. Despite being under pressure and sometimes heavily degraded, these habitats are still
dominating the coastal zone in the region in which they will be studied (as described under the WPs in
section 3.2) and are good representatives of European seaweed ecosystems. The relatively limited dispersal
ability of these seaweeds (Serrão et al. 1997, Dudgeon et al. 2001, Capdevila et al. 2018) makes them
practically suitable study objects for fragmentation experiments without having to work in too large areas.
F. vesiculosus beds are the most dominating littoral habitat forming macroalgae in moderately wave
exposed areas, found in three of the four study regions (the Norwegian, Baltic and Scottish Sea), which
makes this species very suitable for cross-region analyses. A. nodosum beds are dominating the more wave
sheltered parts. Being commercially harvested in both Norway and Scotland, the A. nodosum studies will
provide valuable knowledge for resource
management authorities and the harvesting
industry in these two countries. In the
Mediterranean Sea, seaweed habitats are
represented by Cystoseira beds, which show
the highest level of Mediterranean seaweed
complexity (Gianni et al. 2013, Gorman et al.
2013) and exhibit functional properties similar
to the F. vesiculosus and A. nodosum
communities in the other regions.
The fragmentation manipulation experiments
will be designed to separate the effect of
habitat fragmentation per se (sensu Fahrig
2003, i.e. change in habitat configuration with
constant area) from area loss (i.e. reduction of
area irrespective of arrangement, Figure 2).
The species and community properties to be
studied are the focal seaweed size, growth
and weight, habitat area and coverage, as well
as fertility indicators (i.e. the number of
receptacles, reproductive thalli and juveniles).
We will also measure coverage of other
macroalgae, and the biomass, abundance,
species composition, structure and functional
group diversity of seaweed associated species
(flora and fauna). Functional diversity will be
used estimate the functional redundancy, i.e.
Single large
Several small
Lots of tiny
Increasing area
Increasing fragmentation
Figure 2. Conceptual diagram of the nine types of experimental
plots (green=seaweed communities), designed to separate the
effect of fragmentation (i.e. changes in habitat configuration
with constant area, i.e. the effect of “single”, “several” and “lots
of…”) from that of habitat loss (i.e. reduction of area irrespective
of configuration, i.e. the effect of “large”, “small” and “tiny”).
The diagram is based on Loke et al. (2019).

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presence of several species having the same function (de Bello et al. 2007). We will use remote sensing
(satellites, aerial hyperspectral imagery and/or aerial photographs) dating back > 3 years for F. vesiculosus
and > 10 years for A. nodosum and the Cystoseira species (Ballesteros et al. 2009, Marine Scotland 2016,
Verdura et al. 2018) to ensure that our focal seaweed communities are successional mature (all WPs) and
to find areas that have had the relevant size and level of fragmentation for a sufficiently long period of time
(for the WP2 observational studies). Focal seaweed properties will be identified in situ, while properties of
the associated species will be identified at the lab.
Possible risks associated with this project are largely related to the field work under the WPs (described in
section 3.2). In WP2, data sampling will be taking place in areas with three types of fragmentation (Figure
2). Finding all three can be difficult, but the risk is heavily reduced by preparatory work with using remote
sensing to find suitable areas. WP2&3: To account for the possibility that stations in different regions don’t
have comparable wave exposure levels, wave exposure values from the Burrows (2020) model will be
integrated in the analyses. In WP4, cod and labrids will be added to the mesocosm basins. Cod is protected
in the Oslofjord throughout the year (fiskeridir.no/English/Fishing-in-Norway/Sea-angling-in-Norway), and
we will apply for a dispensation for fishing cod for scientific purposes. However, as the abundance of cod
has been drastically reduced in the Oslofjord, obtaining small fish for the mesocosm experiment can be
difficult. If we do not get the dispensation or we are not able to catch cod in the Oslofjord, we will get fish
from a region further south, permissions and routines that NIVA has already established for ongoing
experiments. To make sure that the fish are alive and healthy in the mesocosm basins throughout the
experiments, the basins will be looked after daily by NIVA staff on site.
In general, the fragmentation experiments impact relatively small areas. In the regime shift experiments
(WP3), where larger areas are cleared, only Fucus vesiculosus will be included, as this species recovers
relatively quickly (<3 years, Marine Scotland 2016), while this can take more than 10 years for Ascophyllum
nodosum (Marine Scotland 2016) and Cystoseira (Ballesteros et al. 2009, Verdura et al. 2018).
Consequently, we expect no significant negative environmental impact from this project. Where species
are added or removed, this will be done in accordance with Norwegian and European Research Ethics
Committees guidelines for “Protection of animals used in research”. COASTFRAG will follow the ethical
guidelines of both Norwegian and European Research Ethics Committees (www.etikkom.no/en,
www.eurecnet.org). All participating institutes strongly support gender equality. This project has more
women than men WP and Task leaders. Also, the project manager (Dr T. Bekkby) is female. Gender equality
issues will be considered when selecting additional projects participants, Master students and Postdocs for
exchange.
1.3 Novelty and ambition
COASTFRAG will fill a current knowledge gap regarding how pressures, alone and together, impact
biodiversity, structure and ecosystem functioning of coastal ecosystems. Such information is crucial to
understand present and predict future changes, and essential to support knowledge-based sustainable
management of European coastal communities. Until now, scientific understanding of this issue has mainly
been limited to terrestrial studies. COASTFRAG will provide scientific knowledge and conclusions that apply
across Europe in terms of geographical extent and environmental gradients. Further, COASTFRAG will
produce methodological standards and protocols for fragmentation studies across Europe. By developing
novel models that integrate theoretical, experimental and observational knowledge, we will move beyond
the common approach of making projections based on non-causal statistical relationships of environmental
and distributional data. This will improve our understanding of changes in ecosystems and our ability to
make better predictions of changes under future climate, also beyond the habitats studied by COASTFRAG.
2. Impact
2.1 Potential for academic impact of the research project
The Norwegian long-term plan for higher education (“Langtidsplanen for høyere utdanning” 2015-2024)
aims to develop outstanding academic communities, with strong ties to international research activities.
COASTFRAG involves collaboration between leading national and international experts, bringing together

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researchers at different stages of their careers, including Master students, Postdocs and junior research
assistants and researchers (co-leading WP1, 3 and 5). This will ensure training of Early Stage Researchers
and thereby invest in the education, training and recruitment of a new generation of marine scientists.
COASTFRAG funding will cover the exchange of two Postdoc students so that COASTFRAG data can be
included in their ongoing work. The Postdoc students will each work 50% for one year on the project (2022-
2023). The two students will be selected among already existing postdocs at the different universities
involved in the project, in tight dialogue with the members of the International Scientific Advisory Board
(ISAB), presented in section 3.1. By doing comprehensive communication and dissemination, to both the
scientific community and a wider audience, and making data and research findings openly accessible, we
aim to draw international attention to Norwegian research and to national and internationally relevant
research questions.
2.2 Potential for societal impact of the research project
COASTFRAG supports the aim of the United Nations Decade of Ocean Science for Sustainable
Development (www.oceandecade.org) by facilitating collaboration between scientists, policy makers,
managers and other stakeholder and ensure scientific knowledge delivery to the society. Climate change
and the loss of biodiversity are two of the most crucial environmental challenges, both in Norway and
globally. Evidence of the ecological impacts of habitat fragmentation is well established for terrestrial
ecosystems, and targets to improve connectivity are a fundamental part of both international and
European biodiversity legislation and policy (e.g. Article 3 Birds Directive, Article 10 Habitats Directive).
COASTFRAG will provide decision makers and other stakeholders (such as seaweed harvesters) with
knowledge on the situation in coastal ecosystems, to support decisions made on the use of the coastal zone
in a sustainable and climate adapted way, both now and in the future. COASTFRAG aligns with the targets
of the UN Sustainable Development Goal SDG 14 (ocean life) by increasing the scientific knowledge
needed to protect coastal ecosystems from significant adverse impacts, including strengthening their
resilience, and enhance conservation and the sustainable use of ocean-based resources, for the benefit of
all, now and in the future. As pointed out by the Blue papers of the High level panels (oceanpanel.org/blue-
papers), mitigating loss of biodiversity and habitats is crucial to combat poverty and hunger, safeguard
human health and fight climate change (though the protection, sustainable use and restoration of carbon
storing and sequestrating marine vegetation). COASTFRAG will thereby also support SDG 1, 2, 3 and 13.
Under the Norwegian long-term plan for higher education 2015-2024 (Meld. St. 7 (2014-2015) the
prioritised research needs of the Ministry of Climate and Environment (2016-2021) is to obtain a better
management of marine ecosystems and resources. To ensure sustainable and climate-adapted
development, harvesting and other activities in the coastal zone, the knowledge gained from COASTFRAG,
on how pressures act together to impact coastal ecosystems, is needed. COASTFRAG will support the
outspoken research needs of the Norwegian Environment Agency (for 2016-2021) on acquiring knowledge
on marine biodiversity and ecological processes. Knowledge on the effect of land use changes, such as
construction, and the effect of the total pressure along the coast is particularly lacking.
2.3 Measures for communication and exploitation
COASTFRAG has a simple yet strategic communication and exploitation plan centred around selected
target audiences (see WP6 in section 3.2). Communication to both specialised and non-specialised
audiences will occur through the project website and social media. Schools will also be directly engaged,
including with hands-on practices. Dissemination will primarily occur via scientific papers targeting
researchers and the specialised audience, as detailed in the online form for the Scientific Dissemination
Plan. Exploitation of the content of such papers, by decision makers and other stakeholders, will be
facilitated through periodic infographics and a webinar.

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3. Implementation
3.1 Project manager and project group
COASTFRAG involves collaboration between leading scientists from Norwegian and international institutes
at universities. We also involve international universities in an International Scientific Advisory Board
(ISAB). A National User Advisory Board (NUAB) will be invited. NIVA (Norway) will coordinate the project.
Dr Trine Bekkby will be the project manager, and will, with support from WP1, coordinate all WPs, ensure
a timely progress, communication among participants, project meetings, budget management and
reporting to the Research Council of Norway. NIVA will also, together with the University of Oslo,
coordinate Master student activities and Postdoc exchanges, in tight collaboration with both the
COASTFRAG research partners and the members of the ISAB. Dr Bekkby is a senior scientist (professor level)
with a strong track record of project management. She has been leading national projects since 2002 and
has been principal investigator, WP leader or having leading roles in international projects since 2007. Dr
Bekkby has extensive expertise in field work, including as survey leader using a variety of tools, with almost
400 days at sea. The NIVA project group has extensive knowledge of biodiversity, ecology, taxonomy,
structure and function of rocky seabeds, spatial statistics, distribution modelling and GIS. Prof. Stein
Fredriksen at the Department of Biosciences, University of Oslo (UniOslo, Norway) has collaborated with
NIVA researchers for decades on seaweed biology, ecology and taxonomy. He will be the University of Oslo
co-supervisor of all NIVA students. Dr Jonne Kotta from the Estonian Marine Institute, University of Tartu
(UniTartu, Estonia) has long-term experience working on patterns, dynamic and processes of coastal
ecosystems, including invasive species, macrophyte and invertebrate communities, trophic networks, and
scale-dependent relationships between environmental forcing and biotic patterns. Assoc. Prof. Emma
Cebrian and Dr Jordi Boada, from the Agencia Estatal Consejo Superior de Investigaciones Científicas;
Centre for Advanced Studies of Blanes (CEAB-CSIC, Spain), are leading scientists in the field of marine
community ecology, conservation and restoration of Mediterranean marine ecosystems, including on
biodiversity patterns, species interactions, habitat-species linkages and quantifying the cumulative impacts
of multiple stressors. Prof. Michael Burrows from the Scottish Association for Marine Science (SAMS, UK)
has extensive experience with ecology of individuals, populations, communities and whole ecosystems,
mainly in the littoral, studying ecosystem-scale patterns in biodiversity and the link to environmental
variables. Dr Martina Milanese, Managing Director and co-founder of the Studio Associato GAIA (Italy), has
a long experience in communicating science to different audiences, including to promote environmental
awareness, with strong expertise in large research projects such as H2020 consortia. H.B. Borchgrevink will
be co-leading WP5 together with Dr Milanese. He is the NIVAs Manager of Media Relations and
Dissemination, with a degree in Media and Communications and Political Science.
World-leading scientists have agreed to participate in the International Scientific Advisory Board (ISAB), to
provide guidance and critical advice on the work programme and methods, greatly enhancing the quality of
the project and the internationalisation of results. The ISAB will also contribute with setting the partners in
contact with Postdoc students that can potentially work on the COASTFRAG project. Prof Stephen Hawkins
(University of Southampton, UK) has agreed to Chair the ISAB. He is an expert on community ecology,
ecological connectivity and multiple drivers of change in ecosystems (including climate change); Assoc.
Prof. Isabel Sousa Pinto (University of Porto, Portugal) works with life-history traits, structural complexity
in communities and EU biodiversity policy; Prof. Dorte Krause-Jensen (Aarhus University, Denmark) works
with seaweed distribution and growth in relation to physical and chemical parameters, focusing on general
patterns in regulation of growth of macrophytes; Prof. Craig Johnson (University of Tasmania, Australia) is a
specialist in marine community ecology, incl. the drivers of dynamics, and how to interpret change,
including human-induced pressures, regime shifts, fragmentation and restoration. COASTFRAG will also
invite stakeholders and potential knowledge users (from the different counties) to participate in a National
User Advisory Board (NUAB). The aim of this board is to provide guidance on the user relevance of
COASTFRAG results, advice on the management support and communication to a wider audience. In
Norway, participants in the NUAB will include relevant County Councils, the Directorate of Fisheries, the
Norwegian Environment Agency (NEA) and members of the seaweed harvesting industry (e.g. DuPont
Norge). NEA has confirmed their participation (ref.nr. 2020/88). ISAB and NUAB will be invited to the kick-

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off meeting, annual online meetings, the final project meeting and to the webinar (described in WP6), with
sessions specifically tailored for NUAB members.
3.2 Project organisation and management
A Gantt chart of the WPs and Tasks is
shown in Table 1. Month for each
deliverable is shown in the WP
description. The institutions involved
in COASTFRAG have all the resources
and equipment needed to successfully
achieve the proposed objectives. This
includes field work equipment, lab
facilities, taxonomists and the licenses
and software needed for data
analyses, modelling and visualisation.
WP1 – ADMINISTRATIVE
SUPPORT AND SAMPLING
PROTOCOL
Lead: Dr H. Gundersen. Co-lead: M.S.
Brkljacic (NIVA, Norway)
The objective of this work package is to support the COASTFRAG project manager Dr T. Bekkby (NIVA) with
managing the project. WP1 will compile and manage all data, organise a field sampling training workshop
and develop a protocol for field sampling to ensure the ability for cross-region analyses. WP1 will also
coordinate the Master students and Postdoc exchanges from the different universities. Task 1.1 – Data
management and integration (Task lead: Dr H. Gundersen, NIVA): Develop a data management plan (DMP)
to clarify how to describe, store and share data, in accordance with the requirements of the Research
Council of Norway. See online form for more details on data availability. This task will also compile and
integrate (in GIS) all existing data and models, for use in fieldwork planning and the cross-region analyses.
Task 1.2 – Sampling protocol (Task lead: Dr K.Ø. Kvile, NIVA): This task will organise a training workshop for
all project participants and define the agreed-on protocol for the observational studies (WP2), the
fragmentation manipulation, and the clearing experiments (WP3 and WP4), with the aim to make the
analyses across Europe and the projections into the future (WP5) possible. It is particularly important to
discuss and agree on the size of the fragmentation plots (i.e. what is “single large”, “several small” and “lot
of tiny” in Figure 2), the minimal sampling area (MSA) for each habitat, how to measure properties (also
non-destructively), the areas to be cleared for the regime shift experiments (as discussed in Petraitis &
Latham 1999), the definition of functional groups and the level to which species are to be identified. The
European wave exposure model developed by Burrows (2020) will be used to standardise the stations
according to wave exposure. The EU Water Framework Directive classification for water quality levels will
be used to intercalibrate the nutrient levels. The sampling protocol will be developed in accordance with
the guidelines of the Research Ethics Committees (www.etikkom.no/en). Deliverable 1.1 (M3): Project
plan, a first version of a data management plan, platform for internal communication; Deliverable 1.2 (M3):
Invitation for Master student participations submitted to universities and Postdoc calls; Deliverable 1.3
(M4): Online GIS map for data and models; Deliverable 1.4 (M6): Training workshop on field design,
protocol for field studies; Deliverable 1.5 (M12): Technical paper on sampling protocol.
WP2 – FRAGMENTATION, GLOBAL STRESSORS AND SCALE
Lead: Dr T. Bekkby. Co-lead: Dr E. Rinde (NIVA, Norway).
Can we draw general conclusions on the impacts of fragmentation on littoral seaweed communities across
Europe? Is the effect of fragmentation different if it happens in an area that is already fragmented than if it
happens in an area with large and continuous habitat coverage? The objective of WP2 is to answers these
questions by addressing research question Q1 on how habitat fragmentation impacts seaweed
communities under different climate regimes and at different spatial scales. We will do observation studies
and data sampling, comparing communities associated with seaweed beds as they occur in the field. For
Table 1. Gantt chart of the WPs and Task in from 2021 to 2024.
Q1 Q2 Q3 Q4 Q1 Q2 Q3 Q4 Q1 Q2 Q3 Q4 Q1 Q2 Q3 Q4
Meetings
WP1 Task 1.1
Task 1.2
WP2 Task 2.1
Task 2.2
WP3 Task 3.1
Task 3.2
Task 3.3
Task 3.4
WP4 Task 4.1
Task 4.2
WP5 Task 5.1
Task 5.2
WP6 Task 6.1
Task 6.2
Task 6.3
2021
2022
2023
2024

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each of the four regions, we will find three “single large”, three “several small” and three “lot of tiny”
seaweed areas (Figure 2). We will use remote sensing tools (satellites, hyperspectral imagery and/or aerial
photographs) to identify potential suitable study areas (see more details in section 1.2). Within each of the
nine plots, species (flora and fauna) associated with the focal seaweed will be collected within a minimal
sampling area (MSA, standardised and agreed-on in WP1). Species and community properties (as listed in
section 1.2) will be estimated and data analysed across Europe. Before collecting the species in the
seaweed patches, some properties will be measured non-destructively (Cebrian & Ballesteros 2004): focal
seaweed habitat area and coverage and the coverage of different macroalgae species (as agreed on in
WP1); five replicate focal seaweed individuals within each patch will be size measured, and fertility
indicators (i.e. number of receptacles, reproductive thalli and juveniles) will be recorded. To be able to
assess possible “edge effects” (i.e. environmental changes that occur at the habitat boundary), we will put
out fauna traps (as described by Christie et al. 2007) in the middle and close to the edge of each patch.
“Edge traps” will be placed in the direction of and away from the water line, respectively. Task 2.1 –
Fragmentation under different global stressors (Task lead: Prof. M. Burrows, SAMS): To study the impact
of fragmentation across different global stressors, we will do observation studies in the four regions,
representing different climate (i.e. temperature and water quality) regimes (Figure 1). Fucus vesiculosus
beds are present, and will be the focal seaweed, in the Norwegian Sea, Baltic Sea and in Scottish Atlantic
Ocean; Ascophyllum nodosum in Norway and Scotland; Cystoseria beds in the Mediterranean Sea. The
studies will be performed under as similar as possible environmental conditions (such as wave exposure, by
the use of the model of Burrows 2020), and we will ensure minimum freshwater runoff and human
disturbances. Dr T. T. Bekkby will be the regional coordinator for the Norwegian Sea studies, Dr J. Kotta for
the Baltic Sea, Prof. M. Burrows for Scotland and Assoc. Prof. E. Cebrian for the Mediterranean Sea. Task
2.2 – Fragmentation at different spatial scales (Task lead: Dr T. Bekkby, NIVA): The effect of fragmentation
can wary with the extent of the habitat (Deza & Anderson 2010). We will therefore sample data for both F.
vesiculosus and A. nodosum, in two different areas in Norway, one representing an apparent lack of
previous fragmentation, i.e. a large extent of the habitat (i.e. at the magnitude of 500-1000 m2) and one
with smaller patches (i.e. ~100 m2). Deliverable 2.1 (M24): Paper on effects of fragmentation and global
pressures; Deliverable 2.2 (M24): Paper on effects of fragmentation at different scales.
WP3 – FRAGMENTATION, ENVIRONMENTAL CONDITIONS AND LOCAL HUMAN PRESSURES
Lead: Dr C.W. Fagerli. Co-lead: S.R. Moy (NIVA, Norway)
Will the impacts of fragmentation be the same regardless of the environmental conditions found in
different areas? And how does local human pressure impact the effects of fragmentation? The objective of
WP3 is to answer these questions by addressing research question Q2 on how variation in environmental
conditions, local human pressures and habitat fragmentation interact to impact coastal seaweed
communities and the potential for regime shifts. We will do mesocosm experiments and field habitat
fragmentation and clearing experiments in the four regions to study the effect of habitat fragmentation on
seaweed communities at different levels of wave exposure and human pressures (nutrient loads). At the
beginning of the experiments and during the revisits (to monitor and maintain the plots), some properties
will be measured non-destructively, as described in WP2. Also, five replicate focal seaweed individuals
within each plot will be size measured and identity marked so that they can be recovered and remeasured
for growth assessment. The sites will be revisited every spring and autumn (monthly for the mesocosm),
species and community properties will be measured non-destructively, the identity marked seaweeds will
be remeasured, and the fragmentation patters will be maintained. After 2 years (1 year for the mesocosm),
the non-destructive properties will be recorded one last time before all species (flora and fauna) associated
with the focal seaweed will be collected within a minimal sampling area (MSA). Species and community
properties will be estimated (as presented in section 1.2). Task 3.1 – Fragmentation and wave exposure
(Task lead: Dr C.W. Fagerli, NIVA): The mesocosm study will be carried out in Norway, in 12 outdoor
mesocosm basins, each with a wave machine and constantly receiving water from the fjord. Boulders with
F. vesiculosus and associated species will be collected in the fjord outside and placed in the basins. Six
basins will have a single large patch of seaweed (upper left plot of Figure 2), six basins will have several
small patches (upper mid plot of Figure 2), both groups with three replicates for each wave exposure level.
Task 3.2 – Fragmentation and local human pressures: (Task lead: Dr J. Kotta, UniTartu): Fragmentation
manipulation will be carried out in the Baltic Sea (Gulf of Riga and Baltic Proper, Estonia) and in the NW

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Mediterranean Sea (Catalan coast and Balearic Islands, Spain) as shown in Figure 2. In Estonia, the
fragmentation manipulation will be carried out in F. vesiculosus beds in three areas of high nutrient loading
(Gulf of Riga) and three areas with moderate nutrient loading (the Baltic Proper), under otherwise as similar
wave exposure levels as possible. In the Mediterranean Sea, the fragmentation manipulation will be carried
out in Cystoseira beds, with three replicate stations identified at two nutrient levels, keeping wave
exposure as similar as possible and freshwater runoff and human activity at a minimum. Dr J. Kotta will
coordinate the Baltic Sea studies and E. Cebrian the Mediterranean studies. Task 3.3 – Fragmentation,
wave exposure and local human pressures combined: (Task lead: Prof. M. Burrows, SAMS): Fragmentation
manipulation will be carried out in F. vesiculosus beds on the rocky shores of the Scottish Atlantic Ocean
(UK), under two different levels of wave exposure and two levels of nutrient loading (low, Firth of Lorn;
medium-high, Loch Fyne), keeping the freshwater runoff and human activity at a minimum. Task 3.4 –
Fragmentation, wave exposure and regime shifts: (Task lead: Dr T. Bekkby, NIVA): The potential for regime
shifts is impacted the number of species in a community that have the same function (Naeem 1998, Pillar et
al. 2013), so called functional redundancy (e.g. de Bello et al. 2007), which in is turn related to species
traits, such as morphology, physiology, phenology and behaviour of individual organisms (Christie et al.
2007, Diaz et al. 2013, Teixidó et al. 2018). Petraitis & Latham (1999) and Dudgeon & Petraitis (2001) have
suggested that the extent of the habitat (the patch size) has a great influence on the potential for regime
shifts. The clearing experiment will therefore be performed at different scales, i.e. by clearing patches of
different sizes. The experiments, i.e. clearing of F. vesiculosus beds with all associated species, will take
place in the Norwegian Sea and in the Baltic Sea. In Norway, the clearing will take place at two different
wave exposure levels, in the Baltic Sea in areas with high (Gulf of Riga) and moderate (the Baltic Proper)
nutrient loading. Within each wave exposure and nutrient class, three small, three medium and three large
patches will be cleared (in the magnitude of 5, 15 and 30 m, respectively, but the precise size of plots will
be agreed on as part of the protocol developed in WP1). Dr T. Bekkby will coordinate the Norwegian Sea
field study and Dr J. Kotta the Baltic Sea studies. Deliverable 3.1 (M33): Paper on effects of fragmentation
and wave exposure; Deliverable 3.2 (M37): Paper on effects of fragmentation and human impacts;
Deliverable 3.3 (M37): Paper on effects of fragmentation and the interaction between wave exposure and
human impact. Deliverable 3.3 (M33): Paper on the drivers of regime shifts (after clearing);
WP4 – FRAGMENTATION AND SPECIES INTERACTIONS
Lead: Assoc. Prof. E. Cebrian. Co-lead: Dr J. Boada (CEAB-CSIC, Spain)
Changes in grazers and grazing pressure are shaping macroalgae habitats and seascapes all over the world.
How do the coastal ecosystems respond to habitat fragmentation when new species enter or are lost from
the system? The objective of WP4 is to answer these questions by addressing research question Q3 on how
different predatory and grazing pressures modify the effect that habitat fragmentation has on seaweed
communities. The habitat fragmentation and species manipulation experiments will be performed in
Norway (Task 4.1) and Spain (Task 4.2). Task 4.1 – Top-down control by mesopredatory fish (Task lead: J.K.
Gitmark, NIVA): Norway has experienced decreased abundances of coastal cod (Gadus morhua) and an
increase in the number of labrids, such as the goldsinny wrasse (Ctenolabrus rupestris). Based on the
concept of mesopredator-release, the hypothesis is that this change has reduced the abundance of small
grazers (“cleaners”), resulting in changed communities, including increased levels of turf algae (Moksnes et
al. 2008, Kraufvelin et al. 2020). After the manipulation experiment of WP3 is terminated, new boulders
with F. vesiculosus will be placed in the Norwegian mesocosm basins. Six basins will have a single large
patch of seaweed (upper left plot of Figure 2), six basins will have several small patches (upper mid plot of
Figure 2). Small (10-15 cm) cods and labrids will be collected outside the station using beach seines and/or
fyke nets and placed in the basins. We will test the interactive effect of fragmentation and different top-
predators by having three replicates of each of the combination of the two different species of fish and the
two levels of fragmentation. At the beginning of the experiment, five replicate F. vesiculosus individuals
within each plot will be size measured and identity marked so that it can be recovered and remeasured for
growth assessment. During monthly visits, the fragmentation pattern will be maintained. At the beginning
of the experiment and during the revisits, some properties will be measured non-destructively, as
described in WP2. At the end of the experiment, after 6 months, the non-destructive properties will be
recorded one last time before the seaweeds and all associated species (flora and fauna) are collected.
Species and community properties (as described in section 1.2) will be estimated. Task 4.2 – Grazing by fish

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and sea urchins (Task lead: Assoc. Prof. E. Cebrian, CEAB-CSIC): A field experiment manipulating habitats of
Cystoseira displaying different species traits (in terms of complexity and structure) will allow for identifying
differential vulnerability and resilience to grazers. The experiment will be performed in the Balearic Islands,
where Cystoseira beds with different traits will be submitted to different grazing intensity by excluding the
two main grazers of the Mediterranean, the sea urchin Paracentrotus lividus and the sea bream Sarpa
salpa. Three replicates will exclude the sea urchins, three replicates will exclude the sea bream (using
excluding cages) and three replicates will serve as open controls. Initial algal coverage and community
structure will be measured non-destructively at the beginning of the experiment (photograph and image
analysis), and the effect of the different grazing pressure will be monitored monthly using photographic
surveys, comparing algal assemblages in the excluding cages and the controls. Deliverable 4.1 (M36): Paper
on the effect of mesopredatory fish grazing and fragmentation; Deliverable 4.2 (M42): Paper on the effects
of fish and sea urchin grazing.
WP5 – PREDICTING THE FUTURE
Lead: Dr G.S. Andersen. Co-lead: Dr K.Ø. Kvile (NIVA, Norway)
What will the coastal communities look like in the future? Who will be the "winners" and who will be the
"losers" in the face of global changes? The objective of WP5 is to answer these questions by addressing
research question Q4 on what changes we will see in future seaweed communities under different
environmental conditions, species interactions and human pressures facing climate change. This WP will
apply different approaches and model techniques that integrate theoretical, experimental and
observational knowledge to assess changes in the biodiversity, structure and ecosystem functioning of
littoral seaweed habitats. The framework enables projections into future climate scenarios that not only
consider the relationship between a given species and the changing environment, but also includes a
complex web of changing biotic interactions. Using characteristics derived from biological theory along with
empirical knowledge from survey and experimental data, the physical tolerance limits of species as well as
the inherent natural variability in regional conditions are jointly integrated. Specifically, the experimental
evidence serves to identify important cause-effect relationships, to better forecast changes in ecosystem
structure and functioning under future climate conditions. Task 5.1 – Space-for-time analyses (Task lead:
Dr G.S. Andersen, NIVA): We will analyse the patterns of spatial variation in the studied seaweed
communities (from all WPs), and use these results to predict the plausible future of littoral ecosystems (i.e.
change in time), applying both univariate and multivariate techniques. We will use a combination of well-
established methodology such as Maxent (Merow et al. 2013) or Boosted Regression Trees (Elith & Hastie
2008) and a range of ordination techniques. Although these models and analytical tools assume no change
over time in the niche-space preference function, they are very powerful to quantify non-linear
relationships and pinpoint plausible shifts in biotic patterns due to climate change. Species and community
level parameters (such as biotic interactions and functional characteristics) will be linked to different
environmental data, including climate change related predictor variables and human pressures. We will
then use these models and the future projections from ocean models, to predict the impact on the studied
seaweed communities in all studied regions. Projections like the regionally downscaled ocean
biogeochemical model for the European Seas (at 0.1° resolution), run for the 21st century under the RCP 4.5
and RCP 8.5 greenhouse gas concentration scenarios (A20 ROMS-ERSEM, Palmer et al. 2019), will be
utilised. Task 5.2 – Causality and interaction in predictions (Task lead: Dr J. Kotta, UniTartu): We will
develop a set of novel semi-parametric Joint Species Distribution Models (based on e.g. Vanhatalo et al.
2020), combining field observations and experimental evidence from the current project with previous
knowledge of the biological feedbacks (including species interactions) and tolerances to pressures
(including climate change). These models will specifically focus on (a) exploring causality in relationships
between predictor and response variables and thereby improve scenario predictions, (b) developing and
applying a novel modelling framework capable of targeting causality, nonlinearity, non-stationarity and
neighbourhood effects and (c) assessing uncertainty of model predictions related to the selection of
variables involved, the temporal variability, and known historical baselines. By combining results on the
specific sensitivity and vulnerability of seaweed communities to both global and local pressures Task 5.1
and 5.2 will together identify areas that will be the possible "winners" (i.e. are more resistant) and "losers"
(i.e. are more sensitive) facing global changes. Deliverable 5.1 (M45): Paper on future changes.

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WP6 – COMMUNICATION AND DISSEMINATION
Lead: Dr M. Milanese (GAIA, Italy). Co-lead: H.B. Borchgrevink (NIVA, Norway)
The knowledge gained from MARINFORSK will be highly relevant and usable by a range of audiences,
spanning academia, management, industry and the general public. In tight collaboration with all WP
leaders and the advisory boards (presented in section 3.1), WP6 will therefore ensure communication to
the identified target audiences. The plan for scientific dissemination is presented in the online form. Task
6.1 – Digital identity (Task lead: Dr M. Milanese, GAIA): The project website (with visuals and posts on
progress) will be the entry point to a broad audience. We will use social media to report on findings, field
work and other activities, lining to the website. Task 6.2 – School involvement (Task lead: H.B.
Borchgrevink, NIVA): At the beginning of the project we will contact nysgjerrigper.no (the Norwegian
Research Council's service for children and young people) and schools to: (a) organise talks and discussions
on marine ecosystems, climate change, scientific methods etc., and (b) invite schools to join field work and
enter the Nysgjerrigper (“Curious George”) competition with COASTFRAG issues. Task 6.3 – Dissemination
and exploitation (Task lead: Dr M. Milanese, GAIA): We will deliver infographics and a webinar to facilitate
the exploitation of COASTRAG results by decision makers and other stakeholders. A presentation will
specifically target the harvesting industry. One infographic will specifically aim to advise regional and
habitat specific management strategies. The project will link up to the Norwegian Blue Forest Network
(nbfn.no, in which NIVA is a member) to communicate results through their channels. Deliverable 6.1 (M3):
Web page and social media platform; Deliverable 6.2: Invitation to (M3) and report on (M45)
children/young people participation in field work and contribution to the Nysgjerrigper (“Curious George”)
competition; Deliverable 6.3 (M3-M45): Talks and discussions at schools; Deliverable 6.4 (M36+M45):
Article in the Norwegian Research Council's magazine service for children and young people; Deliverable
6.5 (M33+M45): Presentation at the Oslo Science Fairs (“Forskningstorget”); Deliverable 6.6 (M45):
Infographics; Deliverable 6.7: Webinar (M45).
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