Connectivity in coastal waters is described as the ability of animals, plants, sediments and organic material to spread freely between the open sea and coast, along the coastline and between coastal areas and inland waters. Since connectivity has an important and pivotal role regarding the biological, physicochemical, and hydromorphological conditions in aquatic systems, it is included as a quality element when estimating ecological status of surface waters in the coastal zone. The description of connectivity and how it should be evaluated is deficient in the regulations provided by the Swedish Agency for Marine and Water Management, SwAM (HVMFS 2019:25). As such, the Department of Aquatic Resources at the Swedish University of Agricultural Sciences (SLU Aqua) was assigned to give suggestions on what information on the biological connectivity concerning plants and animals that could be included in a revised version of these regulations. The assignment also included to provide a knowledge base on connectivity and physical disturbance in the coastal zone. In addition, the assignment included analysing physical disturbance on the connectivity of fish in the coastal areas of the Skagerrak, Kattegat and the Baltic Sea.
When revising the regulations, it was noted that information on the biological aspects of connectivity, and at what scales it operates, are missing. The current regulations do not account for connectivity being either active, through migrations of young and adult organisms, or passive, through transport of eggs, larvae, spores, seeds or fragments by currents (Chapter 2.1.2). This is important to keep in mind, since physical disturbance caused by jetties, piers, noise, boat traffic, etc. has different effects depending on what type of connectivity is in focus. The different dispersal types also occur over different time scales, where active migrations often occur on a seasonal basis and between different life stages, while passive dispersal generally occurs over days or weeks as larvae and spores develop in the free water column before settling on suitable habitat. There are also species that live their entire lives in open waters, such as phytoplankton, which is also included as a biological quality element to be assessed. Within the assessment process, it is essential to distinguish between typical home ranges and maximum migration and dispersal distances, as home ranges are relevant for population dynamics, while maximum migration and dispersal distances have a greater impact on genetic variation between populations. It is also important to consider how connectivity may be affected by a changing climate. Fish play a central role in marine ecosystems, and therefore connectivity of fishes – both between the coast and sea, within coastal areas and between the coast and freshwater – become crucial determinants of ecological status, both in freshwater and in coastal waters. As it is mainly fish that undertake migration between coastal and freshwater areas and the coast and sea, it is difficult to classify parameter 8.3 Connectivity between coastal waters and transitional waters and coastal land areas in the regulations (HVMFS 2019:25) without including fish. Fish should therefore also be included as a biological quality element in the regulations for coastal waters (Chapter 2.1.3). Fish are included in the assessment of ecological status in the Marine Strategy Framework Directive (MSFD), which geographically overlaps with the Water Framework Directive (WFD) in the coastal zone. On the other hand, the MSFD does not include connectivity as a criterion in ecological status assessments. This means that connectivity in the coastal zone, especially for fish, is currently overlooked and coordination between the directives is therefore required to address this. As connectivity is primarily a biological function linked to fish and other organisms, it is also worth considering whether connectivity should be included among the biological quality elements instead of the hydromorphological ones.
Summary
To our knowledge, Sweden seems to be one of the countries in the EU that has come furthest in terms of assessment of connectivity within the Water Framework Directive. The other EU countries seem to focus on the other two hydromorphological quality elements hydrographic condition and morphological condition in their assessments. This may be due to the fact that connectivity is not included in the Water Framework Directive, but that Sweden has chosen to include it as a separate quality factor in the regulations on status assessment in Swedish coastal waters. Connectivity is mentioned in some contexts in other EU countries, but is then most often linked to freshwater systems with a focus on migratory fish and barriers in the form of hydropower and other physical structures that interfere with the flow of water. Interest in developing the hydromorphological quality element, including connectivity, has grown in the EU and a number of reports and scientific articles are available on the subject (Chapter 2.1.4).
Connectivity can be measured in different ways and at different spatial scales. In Chapter 3, we summarise this information in a knowledge base for assessing connectivity in the coastal zone. Measuring and analysing the distribution of species in coastal and marine areas is a challenging task, and therefore knowledge in this area is still limited. Spatial analyses based on habitat maps and information on the dispersal distance of species have been used to analyse and assess active migration, especially in the Baltic Sea and the Skagerrak, as well as tagging studies and combinations of methods. To investigate passive dispersal, a combination of empirical data and hydrodynamic models have mainly been used to investigate dispersal routes and nursery habitats of different marine organisms and to identify key areas of connectivity such as sources and sinks. Chapter 3 also summarises the connectivity patterns of some biologically important groups of organisms and information on natural barriers such as salinity, depth and temperature that affect the dispersal of organisms and organic matter.
Few studies have examined the effects of physical disturbance on connectivity, both within Europe and other parts of the world. Chapter 4.2 describes the studies that have been carried out in Swedish waters, regarding the effects of physical disturbance on passive dispersal and active migration, and Chapter 4.3 summarises information on physical disturbance and connectivity on some key habitats. Chapter 5 describes the results of new analyses of physical disturbance on fish along the Swedish west coast that are part of the current assignment. The results of these studies show that physical disturbance can have a significant impact on connectivity, especially for those species that depend on shallow and wave-sheltered areas for their reproduction. This type of habitat is particularly sensitive to physical impacts, with a noticeable reduction in connectivity. These areas are also the most affected by loss of connectivity due to their high degree of coastal development, including jetties, buoys and marinas. In addition, the results from the modelling studies showed that macroalgae and macrophytes are strongly affected by physical changes. These findings underscore the importance of carefully considering and managing physical disturbance on species and habitats in the coastal zone in order to conserve and protect vulnerable marine ecosystems and the services they provide. The results are important for decision-makers and planners working on conservation measures and management of these environments.
There are significant knowledge gaps on connectivity and physical disturbance in coastal waters. Chapter 6 lists these knowledge gaps, including the need for high-resolution spatial data on the distribution of both organisms and pressures, as well as the relationships between these, both for the biological quality elements benthic fauna, macroalgae, marine plants and phytoplankton and for fish, which are not included as biological quality elements in the coastal zone. There is also a need for increasing the knowledge base on the effects of a changing climate and of various types of restoration measures on connectivity.