WEMo (Wave Exposure Model): Formulation, Procedures and Validation

This report describes the working of National Centers for Coastal Ocean Service (NCCOS) Wave Exposure Model (WEMo) capable of predicting the exposure of a site in estuarine and closed water to local wind generated waves. WEMo works in two different modes: the Representative Wave Energy (RWE) mode calculates the exposure using physical parameters like wave energy and wave height, while the Relative Exposure Index (REI) empirically calculates exposure as a unitless index. Detailed working of the model in both modes and their procedures are described along with a few sample runs. WEMo model output in RWE mode (wave height and wave energy) is compared against data collected from wave sensors near Harkers Island, North Carolina for validation purposes. Computed results agreed well with the wave sensors data indicating that WEMo can be an effective tool in predicting local wave energy in closed estuarine environments. (PDF contains 31 pages)

Does wave exposure determine the interactive effects of losing key grazers and ecosystem engineers?

The consequences of biodiversity loss in the face of environmental change remain difficult to predict, given the complexity of interactions among species and the context-dependency of their functional roles within ecosystems. Predictions may be enhanced by studies testing how the interactive effects of species loss from different functional groups vary with important environmental drivers. On rocky shores, limpets and barnacles are recognised as key grazers and ecosystem engineers, respectively. Despite the large body of research examining the combined effects of limpet and barnacle removal, it is unclear how their relative importance varies according to wave exposure, which is a dominant force structuring intertidal communities. We tested the responses of algal communities to the removal of limpets and barnacles on three sheltered and three wave-exposed rocky shores on the north coast of Ireland. Limpet removal resulted in a relative increase in microalgal biomass on a single sheltered shore only, but led to the enhanced accumulation of ephemeral macroalgae on two sheltered shores and one exposed shore. On average, independently of wave exposure or shore, ephemeral macroalgae increased in response to limpet removal, but only when barnacles were removed. On two sheltered shores and one exposed shore, however, barnacles facilitated the establishment of fucoid macroalgae following limpet removal. Therefore, at the scale of this study, variability among individual shores was more important than wave exposure per se in determining the effect of limpet removal and its interaction with that of barnacles. Overall, these findings demonstrate that the interactive effects of losing key species from different functional groups may not vary predictably according to dominant environmental factors.

Abundance of sedentary consumers and sessile organisms along the wave exposure gradient of subtropical rocky shores of the south-west Atlantic

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Macroalgae contribute to the diet of Patella vulgata from contrasting conditions of latitude and wave exposure in the UK

Analysis of gut contents and stable isotope composition of intertidal limpets (Patella vulgata) showed a major contribution of macroalgae to their diet, along with microalgae and invertebrates. Specimens were collected in areas with limited access to attached macroalgae, suggesting a major dietary component of drift algae. Gut contents of 480 animals from 2 moderately wave-exposed and 2 sheltered rocky shores in each of 2 regions (western Scotland, 55-56°N; and southwest England, 50°N), were analysed in 2 years (n = 30 site-1 yr-1). The abundance of microalgae, macroalgae and invertebrates within the guts was quantified using categorical abundance scales. Gut content composition was compared among regions and wave exposure conditions, showing that the diet of P. vulgata changes with both wave exposure and latitude. Microalgae were most abundant in limpet gut contents in animals from southwestern sites, whilst leathery/corticated macroalgae were more prevalent and abundant in limpets from sheltered and northern sites. P. vulgata appears to have a more flexible diet than previously appreciated, and these keystone grazers consume not only microalgae, but also large quantities of macroalgae and small invertebrates. To date, limpet grazing studies have focussed on their role in controlling recruitment of macroalgae by feeding on microscopic propagules and germlings. Consumption of adult algae suggests that P. vulgata may also directly control the biomass of attached macroalgae on the shore, whilst consumption of drift algae indicates that the species may play important roles in coupling subtidal and intertidal production.

Macroalgae contribute to the diet of Patella vulgata from contrasting conditions of latitude and wave exposure in the UK

Analysis of gut contents and stable isotope composition of intertidal limpets (Patella vulgata) showed a major contribution of macroalgae to their diet, along with microalgae and invertebrates. Specimens were collected in areas with limited access to attached macroalgae, suggesting a major dietary component of drift algae. Gut contents of 480 animals from 2 moderately wave-exposed and 2 sheltered rocky shores in each of 2 regions (western Scotland, 55-56°N; and southwest England, 50°N), were analysed in 2 years (n = 30 site-1 yr-1). The abundance of microalgae, macroalgae and invertebrates within the guts was quantified using categorical abundance scales. Gut content composition was compared among regions and wave exposure conditions, showing that the diet of P. vulgata changes with both wave exposure and latitude. Microalgae were most abundant in limpet gut contents in animals from southwestern sites, whilst leathery/corticated macroalgae were more prevalent and abundant in limpets from sheltered and northern sites. P. vulgata appears to have a more flexible diet than previously appreciated, and these keystone grazers consume not only microalgae, but also large quantities of macroalgae and small invertebrates. To date, limpet grazing studies have focussed on their role in controlling recruitment of macroalgae by feeding on microscopic propagules and germlings. Consumption of adult algae suggests that P. vulgata may also directly control the biomass of attached macroalgae on the shore, whilst consumption of drift algae indicates that the species may play important roles in coupling subtidal and intertidal production.

Predicting Pollicipes pollicipes (Crustacea: Cirripedia) abundance on intertidal rocky shores of SW Portugal: a multi-scale approach based on a simple fetch-based wave exposure index

Understanding and predicting patterns of distribution and abundance of marine resources is important for con- servation and management purposes in small-scale artisanal fisheries and industrial fisheries worldwide. The goose barnacle (Pollicipes pollicipes) is an important shellfish resource and its distribution is closely related to wave exposure at different spatial scales. We modelled the abundance (percent coverage) of P. pollicipes as a function of a simple wave exposure index based on fetch estimates from digitized coastlines at different spatial scales. The model accounted for 47.5% of the explained deviance and indicated that barnacle abundance increases non-linearly with wave exposure at both the smallest (metres) and largest (kilometres) spatial scales considered in this study. Distribution maps were predicted for the study region in SW Portugal. Our study suggests that the relationship between fetch-based exposure indices and P. pollicipes percent cover may be used as a simple tool for providing stakeholders with information on barnacle distribution patterns. This information may improve assessment of harvesting grounds and the dimension of exploitable areas, aiding management plans and support- ing decision making on conservation, harvesting pressure and surveillance strategies for this highly appreciated and socio- economically important marine resource.

Abundance of sedentary consumers and sessile organisms along the wave exposure gradient of subtropical rocky shores of the south-west Atlantic

Sedentary consumers play an important role on populations of prey and, hence, their patterns of abundance, distribution and coexistence on shores are important to evaluate their potential influence on ecosystem dynamics. Here, we aimed to describe their spatio-temporal distribution and abundance in relation to wave exposure in the intertidal rocky shores of the south-west Atlantic to provide a basis for further understanding of ecological processes in this system. The abundance and composition of the functional groups of sessile organisms and sedentary consumers were taken by sampling the intertidal of sheltered and moderately exposed shores during a period of one year. The sublittoral fringe of sheltered areas was dominated by macroalgae, while the low midlittoral was dominated by bare rock and barnacles. In contrast, filter-feeding animals prevailed at exposed shores, probably explaining the higher abundance of the predator Stramonita haemastoma at these locations. Limpets were more abundant at the midlittoral zone of all shores while sea urchins were exclusively found at the sublittoral fringe of moderately exposed shores, therefore, adding grazing pressure on these areas. The results showed patterns of coexistence, distribution and abundance of those organisms in this subtropical area, presumably as a result of wave action, competition and prey availability. It also brought insights on the influence of top-down and bottom-up processes in this area.

Wave forecasts associated with hurricane landfalls in the vicinity of Cape Lookout, North Carolina

Hurricanes can cause extensive damage to the coastline and coastal communities due to wind-generated waves and storm surge. While extensive modeling efforts have been conducted regarding storm surge, there is far less information about the effects of waves on these communities and ecosystems as storms make landfall. This report describes a preliminary use of NCCOS’ WEMo (Wave Exposure Model; Fonseca and Malhotra 2010) to compute the wind wave exposure within an area of approximately 25 miles radius from Beaufort, North Carolina for estuarine waters encompassing Bogue Sound, Back Sound and Core Sound during three hurricane landfall scenarios. The wind wave heights and energy of a site was a computation based on wind speed, direction, fetch and local bathymetry. We used our local area (Beaufort, North Carolina) as a test bed for this product because it is frequently impacted by hurricanes and we had confidence in the bathymetry data. Our test bed conditions were based on two recent Hurricanes that strongly affected this area. First, we used hurricane Isabel which made landfall near Beaufort in September 2003. Two hurricane simulations were run first by passing hurricane Isabel along its actual path (east of Beaufort) and second by passing the same storm to the west of Beaufort to show the potential effect of the reversed wind field. We then simulated impacts by a hurricane (Ophelia) with a different landfall track, which occurred in September of 2005. The simulations produced a geographic description of wave heights revealing the changing wind and wave exposure of the region as a consequence of landfall location and storm intensity. This highly conservative simulation (water levels were that of low tide) revealed that many inhabited and developed shorelines would receive wind waves for prolonged periods of time at heights far above that found during even the top few percent of non-hurricane events. The simulations also provided a sense for how rapidly conditions could transition from moderate to highly threatening; wave heights were shown to far exceed normal conditions often long before the main body of the storm arrived and importantly, at many locations that could impede and endanger late-fleeing vessels seeking safe harbor. When joined with other factors, such as storm surge and event duration, we anticipate that the WEMo forecasting tool will have significant use by local emergency agencies and the public to anticipate the relative exposure of their property arising as a function of storm location and may also be used by resource managers to examine the effects of storms in a quantitative fashion on local living marine resources.

Wave-induced changes in seaweed toughness entail plastic modifications in snail traits maintaining consumption efficacy, link to supplementary material

1. Environmental stress can influence species traits and performance considerably. Using a seaweed-snail system from NW (Nova Scotia) and NE (Helgoland) Atlantic rocky shores, we examined how physical stress (wave exposure) modulates traits in the seaweed Fucus vesiculosus and indirectly in its main consumer, the periwinkle Littorina obtusata. 2. In both regions, algal tissue toughness increased with wave exposure. Reciprocal-transplant experiments showed that tissue toughness adjusts plastically to the prevailing level of wave exposure. 3. Choice experiments tested the feeding preference of snails from sheltered, exposed, and very exposed habitats for algae from such wave exposures. Snails from exposed and very exposed habitats consumed algal tissues at similar rates irrespective of the exposure of origin of the algae. However, snails from sheltered habitats consumed less algal tissues from very exposed habitats than tissues from sheltered and exposed habitats. Choice assays using reconstituted algal food (triturated during preparation) identified high thallus toughness as the explanation for the low preference of snails from sheltered habitats for algae from very exposed habitats. 4. Ultrastructural analyses of radulae indicated that rachidian teeth were longest and the number of cusps in lateral teeth (grazing-relevant traits) was highest in snails from very exposed habitats, suggesting that radulae are best suited to rupture tough algal tissues in such snails. 5. No-choice feeding experiments revealed that these radular traits are also phenotypically plastic, as they adjust to the toughness of the algal food. 6. Synthesis. This study indicates that the observed plasticity in the feeding ability of snails is mediated by wave exposure through phenotypic plasticity in the tissue toughness of algae. Thus, plasticity in consumers and their resource species may reduce the potential effects of physical stress on their interaction.

Georeferenced standardized apparent wave power along the coastline of Cabrera National Park for analysis of essential habitats of littoral species

Wind- induced exposure is one of the major forces shaping the geomorphology and biota in coastal areas. The effect of wave exposure on littoral biota is well known in marine environments (Ekebon et al., 2003; Burrows et al., 2008). In the Cabrera Archipelago National Park wave exposure has demostrated to have an effect on the spatial distribution of different stages of E.marginatus (Alvarez et al., 2010). Standarized average wave exposures during 2008 along the Cabrera Archipelago National park coast line were calculated to be applied in studies of littoral species distribution within the archipelago. Average wave exposure (or apparent wave power) was calculated for points located 50 m equidistant on the coastline following the EXA methodology (EXposure estimates for fragmented Archipelagos) (Ekebon et al., 2003). The average wave exposures were standardized from 1 to 100 (minimum and maximum in the area), showing coastal areas with different levels of mea wave exposure during the year. Input wind data (direction and intensity) from 2008 was registered at the Cabrera mooring located north of Cabrera Archipelago. Data were provided by IMEDEA (CSIC-UIB, TMMOS http://www.imedea.uib-csic.es/tmoos/boyas/). This cartography has been developed under the framework of the project EPIMHAR, funded by the National Park's Network (Spanish Ministry of Environment, Maritime and Rural Affairs, reference: 012/2007 ). Part of this work has been developed under the research programs funded by "Fons de Garantia Agrària i Pesquera de les Illes Balears (FOGAIBA)".

Population dynamics of blue mussels in a variable environment at the edge of their range

Spatial and temporal variation in the abundance of species can often be ascribed to spatial and temporal variation in the surrounding environment. Knowledge of how biotic and abiotic factors operate over different spatial and temporal scales in determining distribution, abundance, and structure of populations lies at the heart of ecology. The major part of the current ecological theory stems from studies carried out in central parts of the distributional range of species, whereas knowledge of how marginal populations function is inadequate. Understanding how marginal populations, living at the edge of their range, function is however in a key position to advance ecology and evolutionary biology as scientific disciplines. My thesis focuses on the factors affecting dynamics of marginal populations of blue mussels (Mytilus edulis) living close to their tolerance limits with regard to salinity. The thesis aims to highlight the dynamics at the edge of the range and contrast these with dynamics in more central parts of the range in order to understand the potential interplay between the central and the marginal part in the focal system. The objectives of the thesis are approached by studies on: (1) factors affecting regional patterns of the species, (2) long-term temporal dynamics of the focal species spaced along a regional salinity gradient, (3) selective predation by increasing populations of roach (Rutilus rutilus) when feeding on their main food item, the blue mussel, (4) the primary and secondary effects of local wave exposure gradients and (5) the role of small-scale habitat heterogeneity as determinants of large-scale pattern. The thesis shows that populations of blue mussels are largely determined by large scale changes in sea water salinity, affecting mainly recruitment success and longevity of local populations. In opposite to the traditional view, the thesis strongly indicate that vertebrate predators strongly affect abundance and size structure of blue mussel populations, and that the role of these predators increases towards the margin where populations are increasingly top-down controlled. The thesis also indicates that the positive role of biogenic habitat modifiers increases towards the marginal areas, where populations of blue mussels are largely recruitment limited. Finally, the thesis shows that local blue mussel populations are strongly dependent on high water turbulence, and therefore, dense populations are constrained to offshore habitats. Finally, the thesis suggests that ongoing sedimentation of rocky shores is detrimental for the species, affecting recruitment success and post-recruit survival, pushing stable mussel beds towards offshore areas. Ongoing large scale changes in the Baltic Sea, especially dilution processes with attendant effects, are predicted to substantially contract the distributional range of the mussel, but also affect more central populations. The thesis shows that in order to understand the functioning of marginal populations, research should (1) strive for multi-scale approaches in order to link ecosystem patterns with ecosystem processes, and (2) challenge the prevailing tenets that origin from research carried out in central areas that may not be valid at the edge.

Physical Constraints to Aquatic Plant Growth in New Zealand Lakes

The nature of aquatic plant communities often defines benthic habitat within oligotrophic and mesotrophic lakes and lake management increasingly recognizes the importance of maintaining plant diversity in order to sustain biological diversity and capacity within lakes. We have developed simple statistical relationships between key physical and vegetation variables that define the habitat requirements, or “habitat-templates”, of key vegetation types to facilitate management of plant communities in New Zealand lakes. Statistical relationships were derived from two datasets. The first was a multi-lake dataset to determine the effects of water level fluctuation and water clarity. The second dataset was from a comprehensive shoreline survey of Lake Wanaka, which allowed us to examine within-lake variables such as beach slope and wave action. Sufficient statistical relationships were established to develop a habitat template for each of the major species or assemblages. The relationships suggested that the extent and diversity of shallow-growing species was related to a combination of the extent of water level fluctuation and wave exposure. (PDF contains 9 pages.)

Preliminary comparison of natural versus model-predicted recovery of vessel-generated seagrass injuries in Florida Keys National Marine Sanctuary

Each year, more than 500 motorized vessel groundings cause widespread damage to seagrasses in Florida Keys National Marine Sanctuary (FKNMS). Under Section 312 of the National Marine Sanctuaries Act (NMSA), any party responsible for the loss, injury, or destruction of any Sanctuary resource, including seagrass, is liable to the United States for response costs and resulting damages. As part of the damage assessment process, a cellular automata model is utilized to forecast seagrass recovery rates. Field validation of these forecasts was accomplished by comparing model-predicted percent recovery to that which was observed to be occurring naturally for 30 documented vessel grounding sites. Model recovery forecasts for both Thalassia testudinum and Syringodium filiforme exceeded natural recovery estimates for 93.1% and 89.5% of the sites, respectively. For Halodule wrightii, the number of over- and under-predictions by the model was similar. However, where under-estimation occurred, it was often severe, reflecting the well-known extraordinary growth potential of this opportunistic species. These preliminary findings indicate that the recovery model is consistently generous to Responsible Parties in that the model forecasts a much faster recovery than was observed to occur naturally, particularly for T. testudinum, the dominant seagrass species in the region and the species most often affected. Environmental setting (i.e., location, wave exposure) influences local seagrass landscape pattern and may also play a role in the recovery dynamics for a particular injury site. An examination of the relationship between selected environmental factors and injury recovery dynamics is currently underway. (PDF file contains 20 pages.)

Conceptual Ecological Modelling of Sublittoral Rock Habitats to Inform Indicator Selection

The purpose of this study is to produce a series of Conceptual Ecological Models (CEMs) that represent sublittoral rock habitats in the UK. CEMs are diagrammatic representations of the influences and processes that occur within an ecosystem. They can be used to identify critical aspects of an ecosystem that may be studied further, or serve as the basis for the selection of indicators for environmental monitoring purposes. The models produced by this project are control diagrams, representing the unimpacted state of the environment free from anthropogenic pressures. It is intended that the models produced by this project will be used to guide indicator selection for the monitoring of this habitat in UK waters. CEMs may eventually be produced for a range of habitat types defined under the UK Marine Biodiversity Monitoring R&D Programme (UKMBMP), which, along with stressor models, are designed to show the interactions within impacted habitats, would form the basis of a robust method for indicator selection. This project builds on the work to develop CEMs for shallow sublittoral coarse sediment habitats (Alexander et al 2014). The project scope included those habitats defined as ‘sublittoral rock’. This definition includes those habitats that fall into the EUNIS Level 3 classifications A3.1 Atlantic and Mediterranean high energy infralittoral rock, A3.2 Atlantic and Mediterranean moderate energy infralittoral rock, A3.3 Atlantic and Mediterranean low energy infralittoral rock, A4.1 Atlantic and Mediterranean high energy circalittoral rock, A4.2 Atlantic and Mediterranean moderate energy circalittoral rock, and A4.3 Atlantic and Mediterranean low energy circalittoral rock as well as the constituent Level 4 and 5 biotopes that are relevant to UK waters. A species list of characterising fauna to be included within the scope of the models was identified using an iterative process to refine the full list of species found within the relevant Level 5 biotopes. A literature review was conducted using a pragmatic and iterative approach to gather evidence regarding species traits and information that would be used to inform the models and characterise the interactions that occur within the sublittoral rock habitat. All information gathered during the literature review was entered into a data logging pro-forma spreadsheet that accompanies this report. Wherever possible, attempts were made to collect information from UK-specific peer-reviewed studies, although other sources were used where necessary. All data gathered was subject to a detailed confidence assessment. Expert judgement by the project team was utilised to provide information for aspects of the models for which references could not be sourced within the project timeframe. A multivariate analysis approach was adopted to assess ecologically similar groups (based on ecological and life history traits) of fauna from the identified species to form the basis of the models. A model hierarchy was developed based on these ecological groups. One general control model was produced that indicated the high-level drivers, inputs, biological assemblages, ecosystem processes and outputs that occur in sublittoral rock habitats. In addition to this, seven detailed sub-models were produced, which each focussed on a particular ecological group of fauna within the habitat: ‘macroalgae’, ‘temporarily or permanently attached active filter feeders’, ‘temporarily or permanently attached passive filter feeders’, ‘bivalves, brachiopods and other encrusting filter feeders’, ‘tube building fauna’, ‘scavengers and predatory fauna’, and ‘non-predatory mobile fauna’. Each sub-model is accompanied by an associated confidence model that presents confidence in the links between each model component. The models are split into seven levels and take spatial and temporal scale into account through their design, as well as magnitude and direction of influence. The seven levels include regional to global drivers, water column processes, local inputs/processes at the seabed, habitat and biological assemblage, output processes, local ecosystem functions, and regional to global ecosystem functions. The models indicate that whilst the high level drivers that affect each ecological group are largely similar, the output processes performed by the biota and the resulting ecosystem functions vary both in number and importance between groups. Confidence within the models as a whole is generally high, reflecting the level of information gathered during the literature review. Physical drivers which influence the ecosystem were found to be of high importance for the sublittoral rock habitat, with factors such as wave exposure, water depth and water currents noted to be crucial in defining the biological assemblages. Other important factors such as recruitment/propagule supply, and those which affect primary production, such as suspended sediments, light attenuation and water chemistry and temperature, were also noted to be key and act to influence the food sources consumed by the biological assemblages of the habitat, and the biological assemblages themselves. Output processes performed by the biological assemblages are variable between ecological groups depending on the specific flora and fauna present and the role they perform within the ecosystem. Of particular importance are the outputs performed by the macroalgae group, which are diverse in nature and exert influence over other ecological groups in the habitat. Important output processes from the habitat as a whole include primary and secondary production, bioengineering, biodeposition (in mixed sediment habitats) and the supply of propagules; these in turn influence ecosystem functions at the local scale such as nutrient and biogeochemical cycling, supply of food resources, sediment stability (in mixed sediment habitats), habitat provision and population and algae control. The export of biodiversity and organic matter, biodiversity enhancement and biotope stability are the resulting ecosystem functions that occur at the regional to global scale. Features within the models that are most useful for monitoring habitat status and change due to natural variation have been identified, as have those that may be useful for monitoring to identify anthropogenic causes of change within the ecosystem. Biological, physical and chemical features of the ecosystem have been identified as potential indicators to monitor natural variation, whereas biological factors and those physical /chemical factors most likely to affect primary production have predominantly been identified as most likely to indicate change due to anthropogenic pressures.