Ecology

Conducting an ecosystem survey during a pandemicCancellation of the survey aboard its primary National Oceanic and Atmospheric Administration (NOAA) survey vessel was overcome through acquisition of a charter for a commercial fishing vessel, following all COVID-19 guidelines (Supplementary Figs. 1 and 2). Initial plans were for 15 days at sea, rather than the 45 typically conducted. This lower effort, along with adverse weather and vessel constraints, resulted in only 25% of the average number of mid-water trawls being collected in the long-term core survey area (Fig. 1 and Supplementary Fig. 1). Despite the data reduction, this effort was one of the only fisheries independent surveys to occur on the US West Coast after the first lockdown in March 2020, furthering the need to evaluate impacts of reduced sampling and provide a robust synthesis of survey results for fishery management. Here we provide updated indices for a selection of ecologically and commercially important species that are critical for assessing ecosystem status.The 2020 sampling was spatially biased towards inshore (shallow) stations (Fig. 1) and thus the previously used method for calculating abundance indices (averaging log-transformed catch-per-unit-effort (CPUE), across all sampled stations) was expected to result in biased indices, in particular for species with strong nearshore (e.g., market squid Dorytheuthis opalescens, anchovy) or offshore (YOY Pacific hake Merluccius productus, myctophids Myctophidae, octopus Octopoda, krill) habitat associations (Supplementary Fig. 3). We confirmed that this bias does indeed occur by recomputing indices for the past 30 years, but using only 1 trawl from each of the 15 stations that were sampled in 2020, and comparing these indices to those using all available trawls (Fig. 2 and Supplementary Fig. 4). In contrast, model-based indices computed from equivalently subsampled past data did not show systematic bias due to the incorporation of spatial covariates (Fig. 2). Thus, although the average log CPUEs were well correlated with model-based indices for well-sampled years (1990–2019), average log CPUEs were determined to be inappropriate for 2020 reporting, and the model-based results were used to develop indices for all taxa for years 1990–2020.Fig. 2: A model for uncertainty and unavoidable effort reduction.a SE of log index vs. number of hauls for a given year from the delta-GLM model. Each point is a year, with 2020 indicated in red. Lines are predicted relationship between SE and sample size for each year, color indicating the mean log index for that year, scaled within taxa. b Relative bias in the index point estimate using 15 hauls from the 2020 stations vs. all hauls from all stations sampled in a given year, computed as (x2020 − xall)/xall. Boxplots show spread of results across all years, 1990–2019 (n = 30 independent years, center: median, box: first and third quartiles, whiskers: smallest and largest values no further than 1.5× IQR from the first and third quartiles; IQR, interquartile range). In the left panel, the index was computed by averaging values of log(CPUE + 1) from all available hauls in a given year. In the right panel, the index was computed from the maximum likelihood estimate (MLE) of a delta-GLM model with spatial covariates, as log(MLE + 1). For the model-based index, the x2020 estimate excludes hauls from the focal year but includes complete data from all other years. CPUE, catch-per-unit-effort; GLM, generalized linear model.Full size imageThe 2020 model-based indices for total rockfish and sanddab (Citharichthys spp.) were the second lowest on record and continued a decline from record high abundance levels observed during the 2014–2016 marine heatwave (Fig. 1)22,23. Pacific hake, myctophids, and octopus were also below average. In contrast, the 2020 index for adult northern anchovy continued a multi-year period of persistently high abundance (Fig. 1). Market squid indices were below average, following a mostly positive trend over the past 7 years. Following the steep decline in 2019, the krill index in 2020 was lower than average (Fig. 1); however, as discussed below, uncertainty may be underestimated for this highly patchy taxonomic group. As a consequence of the low sample sizes, a more rigorous evaluation of the trade-off between sample size (trawls) and uncertainty was conducted, as well as further evaluation of trends through application of existing ecosystem science tools.Quantifying uncertainty by resampling the pastFor most taxa, the uncertainty associated with the 2020 relative abundance estimate was the greatest in the time series, an intuitive result of the sparse sampling for that year (Figs. 1b and 2). The SE was estimated to be over three times the long-term average SE for rockfish and Pacific hake, myctophids, and octopus, and the largest (but less than double the long-term mean) for sanddabs and krill (Fig. 2a). By contrast, the uncertainty associated with the adult anchovy index was lower than the long-term average, due to the great abundance and high frequency of occurrence of anchovy in 2020, compared to years in past decades. This reflects the general trend of uncertainty (on the log scale) being greater for a given taxon when abundance is lower, which generally held for all taxa except krill in our explorations (Fig. 2 and Supplementary Fig. 5). Through time, the relative bias of the subset of stations (2020) vs. the full sample size is also consistently lower for the model-based solution compared to using the average estimate (Fig. 2 and Supplementary Fig. 6). There is also a strong relationship between the number of trawls conducted and the resulting error for each point estimate, with the error essentially doubling when the number of trawls is reduced from the long-term average of 62 to the 15 that were conducted in 2020 (Fig. 2a). By contrast, reducing the total number of trawls from 62 to 40 increases the relative error by just under 25%, while increasing the number of trawls from 62 to 90 only decreases the relative error by 16%. The extent to which the mean relative abundance scales that error up or down, regardless of sample size, is taxon specific. There is an approximate doubling of the error at lowest abundance levels relative to the highest levels for rockfish, sanddabs, hake, and market squid, an increase of more than fourfold over the same range for anchovies and octopus, and relatively modest scaling of the error for myctophids and krill (Fig. 2). This trade-off between survey effort and the error of the ecosystem indices provides critical guidance for future survey planning with respect to the complex trade-off between effort and uncertainty in the face of highly variable interannual catch rates.A seabird’s perspectiveThe Farallon Islands (National Wildlife Refuge) are located in the center of the survey region and host the largest breeding colony of common murre (Uria aalge) in the region (Fig. 1). Interannual variability of Farallon Island seabird population dynamics, reproduction, and foraging ecology are well understood and also track RREAS observations6,17. In particular, patterns such as alternating cycles of forage species occurrence and subsequent reproductive output are known to be linked to regional ocean and climate conditions17,20. Long-term observations of seabird diets in the Farallon Islands were fortunately not impacted by the pandemic. As common murre feed their chicks predominantly either juvenile rockfish or northern anchovy (Supplementary Fig. 7), and common murre prey selection is known to covary with prey abundance in the surrounding ecosystem17,20, these observations provide a critical data stream for evaluating 2020 rockfish and anchovy abundance index estimates from the limited trawl sampling. We updated regression models relating the proportion of rockfish and anchovy in murre diets, respectively, to model-based abundance indices for rockfish and anchovy using past data (Fig. 3). Linear models provided the best fit for YOY rockfish and anchovy, (r2 = 0.70; r2 = 0.58, respectively, both p  More

Alpha-pinene, the most abundant volatile found in all propolis, was also the most abundant volatile found in propolis produced in the Adriatic Sea coast of Italy, and its likely botanical origin was suggested to be native conifer trees from that region13. α-Pinene together with β-pinene were previously found as the two most abundant volatiles in propolis produced in the Rio Grande do Sul14 and Paraná3 states, in a not specified Brazilian propolis sample15, and propolis from South Africa16 and Uruguay1. It is noteworthy that South Africa and Uruguay are in similar latitudes to southern Brazil, indicating a characteristic profile related to that propolis location.Furfural, which was found in almost all samples, is a product of sugars dehydration commonly found in agricultural byproducts and was identified by SHS-GCxGC-TOF-MS in South African propolis, in which volatiles was extracted by heating at 45 °C/5 min, at concentrations ranging from trace to 11.3%16.β-Eudesmol was found as the most abundant volatile in propolis produced in France, Hungary, Bulgaria, and Northern Italy and was also the most abundant volatile in the distilled essential oil of Populus nigra buds, which likely is its primary botanical origin1. However, in our study, β-eudesmol was also found in the resins of AA as a minor volatile (from 0.7 to 2.4%). Additionally, AA resins were the only ones containing sabinene, α-thujene and α-bisabolol. Thereby, AA may be plant sources of these volatiles for brown propolis from southern Brazil. α-Bisabolol was also found as a major volatile in propolis produced in temperate zones of China and Turkish1.It is noteworthy that the temperature used to extract the volatiles, 180 °C, was higher than those commonly used for volatile profile characterization of 50–75 °C3,15,17. The degradation rate of pure monoterpenes at 120 °C varied greatly, depending on the compound, as it was 100% after 4 h for α-terpinene, 50% after 24 h for limonene, and 38% after 72 h for camphene18. The thermal degradation led to p-cymene, eucarvone and 1,2-epoxyde derivatives from limonene; thymol, ketoaldehydes, and eucalyptol from α-terpinene; and camphenilone, verbenone, and aromatic compounds from camphene18.Although verbenone was found in samples of our study (up to 1.3%), it was also tentatively identified in brown propolis extracted at 75 °C/30 min3, while p-cymene and verbenone were tentatively identified in Mediterranean propolis extracted at 60 °C/45 min17.Furthermore, McGraw et al.18 quoted Punsuvon, who reported the degradation at 90–130 °C (not specifying the time length) of pure α-pinene (23–37%), forming β-pinene, α-pinene oxide, α-campholenal, verbenol, pinocamphone, myrtenol and verbenone, and of pure β-pinene (22%), forming mainly myrtenol. From those thermal degradation products of α-pinene, some are reportedly relevant in propolis and conifer tissues, such as β-pinene, α-campholenal, myrtenol and verbenol1,15,19. Hence, the terpene diversity in natural products seems to result from naturally occurring chemical reactions catalyzed by microorganisms or enzyme systems20. At the same time, induced heating is a non-natural way to get it, and it is not simple to differentiate whether the terpene diversity is natural or induced by extraction conditions.The increase in temperature increased peak intensities up to 180 °C, and the number of peaks also increased, which likely indicates volatiles release from the propolis’s complex resinous/waxy matrix (Supplementary Fig. S1). However, the formation of low percentages of the tentatively identified carvone oxide (Supplementary Table S1), which likely had the added internal control l-carvone as a precursor, is an indication of oxidation. Nevertheless, l-carvone was pierced outside the samples within the vials. Thereby, it was more exposed to O2 and more prone to oxidation than the other volatiles present in the propolis/resins matrices.Concerning the multivariate analysis, the PCA showed that the SHS-GCMS method was sensible to discriminate propolis samples produced in different municipalities, even when the distance between the apiaries was 72 km (from ‘Beira do mato’ to ‘Vila Zulmira sede’). Moreover, the PCA indicates that A. angustifolia may be more attractive than Pinus species for bee foragers as a plant resin source to produce propolis.It is noteworthy that the possibility of Araucaria sp. resins be used as a botanical source for bees to produce brown propolis in southern Brazil was previously suggested by5, based on the identification of a single non-volatile compound, which is typical in some Araucaria species, in propolis samples from Paraná state. AA is a dominant species in subtropical and temperate rainforests in southern Brazil and adjacent areas. These areas were intensively explored over the nineteenth century. Nowadays, it is legally protected in permanent preservation areas since AA is endangered10. Therefore, the likely presence of AA resins in OP1 reinforces the need for sustainable preservation of natural environments since it may be related to OP1’s outstanding antioxidant activity4.From the tentatively identified volatiles in the hierarchical clustering heatmap, α-campholenal, α-phellandrene, β-bourbonene and trans-verbenol were found in essential oils of Pinus species19. This finding may indicate PT as another plant resin source for propolis production in those areas. To our knowledge, p-mentha-1,8-dien-7-ol was tentatively identified in plants from the Araucariceae family for the first time. p-Mentha-1,8-dien-7-ol, also known as perilla alcohol, is found in many plants’ essential oils, such as lavendin, peppermint, spearmint, and cherries21. Therefore, further studies should be conducted with authentic standards to confirm the identified volatiles in brown propolis and conifer resins from southern Brazil to be further used as phytochemical markers.In conclusion, there are indications that the resin from native Araucaria angustifolia is more attractive for bees to produce propolis in southern Brazil, although there is also an indication that non-native Pinus elliott and Pinus taeda are plant resin sources as well. However, the singularities on the chromatograms of propolis from each apiary/municipality illustrated in the heatmaps and the not complete overlap of the propolis and the conifer resins in the PCA may indicate that there are other botanical sources for bees to produce propolis within the permanent preservation areas of southern Brazil, which remain unknown. More

Coastline segmentsFor reasons of data availability and socioeconomic relevance, the analysis was limited to latitudes between 66° N and −60° S. In this area of interest, the world was divided in 1 arcmin (~2 km) grid cells. To define a logical position for the establishment of an efficient levee, the coastline location was derived from the OpenStreetMap68, moved 100 m land inward and smoothed. For every cell containing a coastline segment, coastline length and a coast-normal transect were derived at the center of segments resulting in 495.361 transects that are on average 1.1 km apart. Bootstrapping revealed that transect distances up to 2 km give very similar results. All transects stretch 4 km seaward and 4 km inland to fully capture most foreshores.Elevation dataA global intertidal bathymetry/elevation dataset from high-resolution EO data (USGS Landsat and Copernicus Sentinel-2), the Foreshore Assessment using Space Technology (FAST) intertidal elevation map69, was produced to compliment commonly used global data products with low resolution and higher inaccuracy in intertidal zones. Global coastlines were divided over 25000 tiles of each 40 × 40 km2. For these tiles, all available images were collected for the period between 1997 and 2017. Surface water was identified, using normalized difference spectral indices (NDSI, here SWIR1 and Green band) for all images (median of 317 images per tile) covering various tidal conditions, and the per pixel mean calculated to derive time-ensemble average (TEA) NDSI images. We developed a new technique to transform TEA images to intertidal elevation independently of in situ calibration data. TEA-NDSI images were normalized by the spatially averaged NDSI values of regions identified (using global elevation datasets) as land and water, respectively. This resulted in a single image per tile that represented the inundation probability for each pixel in the intertidal zone. The inundation probability represents the long-term average tidal inundation, because it was derived from a collection of images that span a time period similar to the tidal epoch (period of 19 years). Pixels having a probability of 1 represent permanent water, and have elevations less than or equal to the lowest astronomical tide (LAT), whereas land (p = 0) represents elevations higher than or equal to the highest astronomical tide (HAT). By deduction, p = 0.5 is equivalent to local mean sea level (LMSL). Tidal statistics from the global tide model FES2012 were used to couple the derived inundation probability to an elevation. The main source of bed level data originates from this map and has a 20 m horizontal resolution and typically a 30–50 cm vertical accuracy (RMSE = 0.52 m, MAE 0.42 m, as assessed at a number of sites with high quality elevation data (Supplementary Fig. 7)). Bathymetry data (GEBCO35; 30 arc-second horizontally, tens of metres vertically) and topography data (MERIT36; 3 arc-seconds, 2 m vertically) were merged to create a continuous bathymetry-elevation map by changing the vertical datum of MERIT from EGM96 to MSL by assuming 0 m +MSL at the OSM coastline. Global bathymetry datasets (e.g. GEBCO) and elevation datasets (e.g. SRTM and MERIT) lack accuracy (especially nearshore), but are commonly used17,18,23,34. The final bed level was constructed using FAST intertidal data where sufficient valid data points were available, complemented by the merged GEBCO-MERIT data where these points were lacking.Vegetation extentThe FAST coastal vegetation map69 was based on Landsat-8 and Sentinel-2 satellite images collected between 2013 and 2017. The map provides actual vegetation presence at 10 m resolution. Vegetation presence was obtained by applying an individual NDVI threshold per tile, with a total of 25,000 tiles, based on the yearly NDVI average and NDVI amplitude. The FAST coastal vegetation map is validated based on NDVI comparison with local measurements taken at Zuidgors, The Netherlands (R2 = 0.92) (Supplementary Fig. 8). If vegetation was present, the vegetation type was determined by global salt marsh32 and mangrove14 maps, complemented with Corine Land Cover30 (CLC, Europe only) and GlobCover v2.231 maps when there is no coverage. Determining global coastal vegetation extent is difficult and affected by eutrophication in coastal environments. This behaviour is observed on the coast along the Persian Gulf and the Red Sea. To improve accuracy only vegetated transects identified by the global salt marsh32 and mangrove14 map and confirmed by the FAST coastal vegetation map are included for these areas. Moreover, vegetated transects with a green belt width smaller than 250 m identified by GlobCover are excluded from the study for accuracy reasons (Supplementary Fig. 8). To avoid mixed vegetation types, the vegetation type was determined by the most dominant type. The vegetation width constituted of the sum of vegetated grid cells between the start and the end of the vegetated zone.Water level and wave dataThe design water levels were based on a combination of tide and storm surge for the selected probability of occurrence (return periods 2, 5, 10, 25, 50, 100 default, 250, 500, 1000 years) and came from the GTSR dataset34. SLR and subsidence were not taken into account because this study focuses on the present situation. Moreover, quantifying the future role of vegetated foreshores would not only require SLR scenarios but also an insight in the development of wetlands over time, which is strongly determined by local conditions such as sediment supply56,57,60. Offshore wave conditions were obtained from ERA-Interim33 re-analysis, based on data from 1979 till 2017 and reprojected to Dynamic Interactive Vulnerability Assessment (DIVA)70 points. Next, the Peak Over Threshold method was applied to construct representative values for the significant offshore wave height, Hs and the peak wave period Tp for all the return periods. The nearshore wave height was limited by the local water depth at the start of the (vegetated) foreshore using a breaker criterion (gamma = 0.55). This is a fairly low value considering the range of values cited in literature71 leading to conservative wave attenuation by vegetation results. Wave-bottom interactions in the sub-tidal zone and processes such as refraction and diffraction are not explicitly simulated. The conservative breaker criterion is chosen to implicitly account for these processes in a conservative manner. The wave period remained unchanged and the wave direction was assumed coast normal and wave growth along the transect due to wind effects was excluded. However, for the current study a more sophisticated approach to account for longshore wave variability based on topography was considered infeasible at the global scale and considered to yield limited outcome looking at the uncertainty in socioeconomic factors. The average Hs,offshore = 4.6 m (std = 2.0 m) and the average Hs,startforeshore = 0.7 m (std = 0.7 m).Profile constructionThe 8 kilometre coast-normal transects consisted of 321 gridpoints, thus a horizontal grid resolution of 25 m. We used four different methods: Foreshore method 1 (based on the FAST intertidal elevation map), Foreshore method 2–4 (based on MERIT-GEBCO). The properties of the FAST intertidal elevation map, MERIT and GEBCO are described under the header ‘Elevation data’. Foreshore method 1 produced the most accurate profiles and foreshore method 4 the least accurate profiles. The profile construction steps are described hereafter. Validity checks were performed to identify false indications of intertidal area in the FAST intertidal elevation map. Individual data points were marked invalid and removed in case: (1) MERIT points were situated above the surge level with a return period of 2 years, while data from the intertidal map indicated a lower elevation. (2) Data from the FAST intertidal map was situated at open sea. (3) Data from the FAST intertidal map along the transect dropped below a minimum range threshold of 10 cm. A fourth check was performed based on the continuity of the data. Data from the FAST intertidal map contain discontinuities along the profile. These continuities exist on pixel level due to the use of the modified normalized difference water index and in some instances cloud coverage was preventing full coverage. Lastly, discontinuities arise due to the presence of (high elevated) tidal flats and banks in coastal areas. (4) Data length was defined as the length of continuous data points along the transect. If the data length of a patch decreased below a threshold of 100 m, the points were marked invalid. Gaps between valid data patches were filled using linear interpolation if the gap was smaller than 250 m. Eventually, one, none or multiple valid data patches were found along the transects. See Supplementary Fig. 2 for example transects.Global coastline shapes range from straight sandy coastal stretches to complex coastlines often found in estuaries. With a transect length of 8 km, the start and the end of the transects could both be situated on land, hampering an unambiguous identification of the foreshore of interest. We designed the algorithm such that the last foreshore was selected. For profiles using data from the FAST intertidal map (foreshore method 1, 50.9% of populated susceptible coastlines), the last valid patch corresponds to the last foreshore. The inclusion of tidal flats as part of the foreshore was determined based on the gap length. In case no (sufficient, thus not satisfying the minimum data length criterion of 100 m) valid data was available from the FAST intertidal map based on the four described checks, the profile was based on a merged GEBCO-MERIT set (methods 2, 3 and 4), respectively, 46.1%, 3.0% and 0.01%. For the second method, data points were selected between a minimum threshold of −2 m MSL and a maximum threshold equal to the surge level with a return period of 2 years. Next, for the selected points the direction of the slope was determined by comparing elevation between the data point concerned and the next data point. This resulted in patches of upward sloping sets of data points between the minimum and maximum threshold. Similar to foreshore method 1, the validity of the patches was checked using data length, gap length and the corresponding thresholds of 100 m and 250 m. The start and the end of the foreshore were determined by the first and last valid point of the last patch. Foreshore method 3 was used if not sufficient foreshore data were available to satisfy the minimum data length threshold (100 m). In these cases, the start of the foreshore was defined as the first upcrossing intersection with −2 m MSL along the transect. The end of the foreshore corresponded to the intersection between the elevation profile and the governing surge level with a return period of 2 years. Foreshore method 4 was used if no start and or end of the foreshore could be found. In this case the start and/or end point of the foreshore corresponded to the first and last data point, respectively.In some cases, elevation for the end of the foreshore was missing due to several reasons. First, the upper part of the intertidal zone was sometimes missing from the FAST intertidal map, due to low frequency of inundation of the upper intertidal zone or cloud cover. Second, bed elevation in mangrove belts was hard to define based on satellite imagery, as the canopy is detected as the earth surface. These uncertainties were counteracted by consulting the mangrove and salt marsh maps. If vegetation was present in one of these maps, the derived foreshore was extended until the end of the vegetated zone. An elevation equal to the surge level with a return period of 2 years was chosen as elevation for extended foreshore points with an elevation exceeding this surge level.Vegetation parametersAs deducting the type and size of mangrove trees and salt marshes from EO data at global scale is not possible (yet), the current modelling approach relies on field and literature observations. For the scope of this research the properties of the mangrove trees occurring at the seaward side of the mangrove belt are the most relevant. To avoid overestimation of wave attenuation in young mangrove forests, the mangrove dimensions are chosen such to be representative for young fringing pioneering mangroves up to a height of 3 m that are practically vertically uniform compared to mature trees. The modelling approach uses four parameters to represent vegetation: height, diameter, number of stems and drag coefficient. The exact characteristics are based on observations in literature8,9,72,73,74,75,76 (N = 30 m−2, d = 35 mm, h = 3.0 m).High quality observations on wave attenuation by mangroves under storm conditions do not exist. For the drag coefficient the theoretical value, 1, of a rigid cylinder is chosen, because mangrove trunks can be considered rigid. For salt marshes a winter state representative as found in NW Europe is chosen. The values are defined based on FAST field tests (Romania, UK, Spain and the Netherlands) and literature10,24,77,78 (N = 1225 m−2, d = 1.25 mm, h = 0.30 m). A drag coefficient (CD) of 0.19 is chosen, which is the lower limit found during large-scale flume tests10. The drag coefficient depends on biophysical characters as well hydrodynamics. The drag coefficient represents drag due to skin friction and pressure differences, but also effects like swaying motion of stems24. The 1D modelling approach takes into account gaps in vegetation cover, e.g. due to the presence of channels. Zonation of vegetation types is not implemented, because this level of detail is insignificant in relation to the inaccuracies induced by the use of global datasets.Wave attenuation modelTo determine wave attenuation along the foreshore transects and the resulting significant wave heights relevant for the flood defence on a transect, we used a lookup-table approach. The lookup table was generated by combining 668,304 model output values for different combinations of foreshore slopes, vegetation covers and hydrodynamic conditions. The table contained wave heights modelled by XBeach79 in surfbeat mode (a nearshore numerical wave model that accounts for the presence of vegetation) at regular intervals along a steady slope, both with and without vegetation. XBeach uses for wave-vegetation interaction the rigid cylinder80 approach and includes an energy sink term to the wave energy balance to implement wave dampening81. We used conservative vegetation characteristics, winter state salt marshes and young pioneering mangroves. We characterized foreshores by their width and slope. The foreshore profile was the same for simulations with and without vegetation. The foreshore width was determined by calculating the distance between the start and the end of the foreshore. The slope was estimated using a linear regression. This approach has two advantages over detailed modelling of wave attenuation over all transects: it is much quicker, allowing for iterative improvements of the workflow and it does not suggest the precision one would expect from detailed models but cannot be delivered with global data. Average Hs,endforeshore,noveg = 0.6 m (std = 0.5 m) and Hs, endforeshore,veg = 0.3 m (std = 0.4 m).Coastline susceptible to flooding, urban and rural extents and population densityTo assess the need for coastal flood defences, we made a distinction between areas susceptible to coastal flooding and higher, non-susceptible areas. We determined susceptible areas based on possible inundation using coastal flood maps of 1 km resolution for a 1/1000 year surge level. These maps were created with a global geographic information system (GIS) based inundation model that is forced with a spatially varying sea level, accounting for attenuation of the water level due to land surface roughness82. A method that is more sophisticated compared to a simple ‘bathtub’ inundation method. Topographic features, as visible in MERIT, protecting the land from flooding are considered. To classify coastlines as urban or rural a distinction was made based on gridded population from the LandScan database83 using the 2UP model84. A transect is characterized ‘urban’ if it intersects at least one cell with an urban population with a minimum of 1. Populated coasts have been identified by assigning the population density of the population susceptible to flooding in the proximity of the transects. We used WorldPop201785 population data and assigned population to the transects using a buffer of 15 kilometre radius. The population density is the division of the assigned population and the total area of the assigned cells. This procedure is repeated for buffer radius of 5, 10 and 20 km, giving fairly comparable outcomes. Following this approach we found a ratio between rural and urban transects of 73/27.Levee crest heightsThe empirical EuroTop formulations47 gave the required levee heights with respect to water levels and wave heights, assuming the presence of a levee at the end of the vegetated foreshore. We hereby neglected the position and characteristics of levees present in the current situation, as no global dataset of coastal protection structures exists. The assumed levee had a standard 1:3 levee profile without berms and an allowed overtopping discharge of 1 l s−1 m−1. These parameters are representative for simple, low-cost levees in developing countries but conservative for well-constructed and maintained levees. Consequently, savings on levee heights in countries with strict protection standards are overestimated, as reduction in required levee height due to vegetation presence is likely less than predicted here. However, this may be balanced out by the fact that we calculated with an average national construction cost per kilometre and levees applying to stricter protection standards may actually be more expensive (Supplementary Fig. 5).Costs for levee construction and crest height reductionThe calculated levee crest height reductions were monetized using a levee unit price per kilometre length per metre heightening. We used an unit investment costs of levees (metre heightening per kilometre length) of USD 7.0 million42. This estimate represents an average of construction costs in the USA and the Netherlands stated in several studies86,87,88,89. It pertains to all investments costs, including ground work, construction, engineering costs, property or land acquisition, environmental compensation, and project management. Investment costs per metre heightening are well described by a linear function without intercept90. They concluded that for large-scale studies it is sufficient to assume linear costs for each metre of heightening, including the initial costs and the 95% confidence range is between 3x and x/3, where x is the unit cost value. Subsequently we applied three unit levee investment cost prices (low: USD 2.33 million, mid: USD 7.0 million, high: USD 21 million) in line with previous studies42,90. These cost estimates were then adjusted for all other countries by applying construction index multipliers (based on civil engineering construction costs91), to account for differences in construction costs across countries92. Costs were converted to USD2005 power purchasing parity (PPP), to be consistent with the SSPs, using GDP deflators from the World Bank (https://data.worldbank.org/), and annual average market exchange rates between Euros and USD taken from the European Central Bank (unit levee cost per country = unit levee cost x construction index per country / PPP MER rate 2005 index per country). Example: mid unit levee costsUSA = 7.0 ×1 / 1 = 7.0 million USD2005 PPP km m−1. If for a country data was not available in the database, we used the average of all countries in the same World Bank income group. For the reference year 2005, this applies to Western Sahara (ESH), North-Korea (PKR) and Somalia (SOM).ReliabilityA scoring table was used to get insight in the reliability of the results of the global analysis. Results were grouped into four reliability classes ranging from “poor” to “very good”. Transects were placed in these classes based on data accuracy for three characteristics: hydrodynamics, vegetation and profile elevation. In Supplementary Fig. 6 the (sub) results of the analysis are presented. The first category, hydrodynamics, included known inaccuracies in the hydrodynamic data (GTSM and ERA-I). Data from the GTSM model was considered less reliable in areas with a low tidal range and/or with tropical storms, such as cyclones or hurricanes, as those were not included in our analyses. Also wave data from ERA-I are less reliable in these areas, because the effects of tropical storms are flattened due to the relatively coarse grid size. Hence, transects in these areas were pinpointed by linking them to NOAA data of historical hurricane tracks93. In Supplementary Fig. 6B, areas where tropical storms occur can clearly be recognized. In addition, the Mediterranean Sea, the Red Sea, the Black sea and the Caspian sea stand out in inaccuracy, because of limited tidal action.Reliability of vegetation characteristics was determined by data source and vegetation width. For transects with extensive vegetation widths, crest height reduction was less sensitive for possible deviations of the vegetation width, due the non-linear relation between vegetation width and wave reduction. Vegetation cover proved most reliable in areas where data from the salt marsh32—and mangrove map14 were available. Hence, this resulted in a ‘good’ score (Supplementary Fig. 6C). Only in cases of extensive vegetation presence was a ‘very good’ score assigned. Transects were appointed as “very good” if vegetation extended 500 m for mangroves, and 1000 m for salt marshes. These thresholds are chosen based on our model results, which show that after ~500 m (salt marshes) and 1000 m (mangroves) maximum reduced wave transmission by foreshore vegetation is reached. Vegetation cover reliability in Europe was classified as ‘good’, due to reliable vegetation type classification based on CLC30 and the salt marsh map32 in combination accompanied by relatively small vegetation widths. The reliability of the derived vegetation characteristics is especially lacking at the east coast of Canada, at Latin America’s south coast, at Africa’s coasts facing the Mediterranean Sea, coasts along the Red Sea and the Persian Gulf, and along the coasts of China, Japan and Russia. For example, in the Persian Gulf states the vegetation presence map tends to falsely identify foreshores as vegetated.The time-ensemble average (TEA) technique applied for the FAST intertidal elevation map relies on the availability of a reasonable number of images at different tidal stages where the differences in horizontal extent of water coverage can be identified, thus allowing a composite of inundation frequency to be derived. However, the technique is limited by the effective sensor resolution (~30 m, including uncertainty in georeferencing) relative to the horizontal extent of changes in inundation, a function of the tidal range and bed slope. Hence, changes in tidal water extent in microtidal or very high bed-slope regions tend to be too small for reliable discerning differences, leading to poor performance of the technique. However, the merged GEBCO-MERIT dataset was considered less reliable than the FAST intertidal map, based on the resolution and the merging of the two underlying datasets in the intertidal zone. In addition, MERIT tends to overestimate the elevation in mangrove areas, as it measures the canopies as the earth’s surface. Besides the elevation data, the foreshore definition method is used as a profile reliability indicator. The total score per transect is given by the sum of the sub-scores. The sub-scores are normalized to give equal weight to the scoring categories.ValidationFor validation of our method to assess vegetation presence, a comparison of 280 randomly located transects with aerial imagery was carried out. The area accessed in the global assessment was divided in tiles of 90 degrees longitude and 15 degrees latitude. From each tile 6 vegetated and 2 non-vegetated transects were selected. Next, a reference dataset was created by manually identifying vegetation presence using present imagery. Lastly, the vegetation width derived by the model and the manually derived set were compared (Supplementary Fig. 8). For this comparison we made three distinctions, based on (1) vegetation type, (2) foreshore derivation method and (3) vegetation cover source. Comparison showed that the used algorithm on global EO data performs satisfactorily (Supplementary Fig. 8), but in some cases tends to assign a vegetation cover of up to 250 m where there is none. Deviation between observation and the global assessment, is caused by methodological error in the global assessment and inaccuracy in the global datasets, e.g. different timestamps are inevitably compared. This would induce an exaggeration of the effect of vegetation. However, due to the limited dimension of the vegetation extent, the threshold for substantial crest height reduction is falsely exceeded in not more than 2.4% of the cases and the effect is largely balanced out by underestimation of the vegetation cover at larger lengths.To validate wave reduction by vegetation calculated through our lookup table approach, we compared results with local modelling results for the South-Western part of the Netherlands for 38 vegetated transects. The numerical model SWAN94 in stationary mode was used to translate wave conditions from offshore to nearshore. The simulations were performed with a grid size of 0.01 deg and bathymetry from EMODNET95. Extreme water levels were included by a water depth correction, using data from GTSR18. Both wind and wave boundary conditions were derived from the earlier described ERA-I re-analysis. The governing wave direction was based on the average of the fifteenth highest wave events in the available wave data. The wind direction was assumed to be aligned with the wave direction. A parametric JONSWAP spectrum shape was used, using a peak enhancement factor of 3.3 and directional spreading of 20 degrees. Foreshore profiles were constructed using an approach similar to foreshore method 2 in the global study but using local high-resolution bathymetry and topography data. Vegetation width was extracted from the salt marsh map32, which was confirmed locally using aerial imagery. Foreshore wave propagation was determined using XBeach in surfbeat mode79.Our results showed an overestimation of the water depth at the start of the vegetated zone by 0.73 m on average. In addition, the global model derived milder slopes in comparison to the local analysis for narrow vegetated transects. The largest errors were found further away from the mouth of the estuary. Here, the deviation between the wave calculated by SWAN and the depth limited approach is largest. The wave height at the start of the vegetated zone was overestimated on average by 1.12 m, due to the complex geometry and the sheltered configuration of the estuary. The algorithm approximated the wave transmission reduction (RMSE 13%) and the levee crest height reduction relative to the required crest height without vegetation presence (RMSE 19%) with reasonable accuracy (Supplementary Fig. 9).Sensitivity analysisA sensitivity analysis has been performed to provide insight in the uncertainty in the presented potential global levee costs savings. The analysis focused specifically on single key parameters, such as the levee unit cost, the critical overtopping discharge and the wave breaker index. High, mid and low levee unit cost scenarios are taken from previous studies42,90. A high, mid, low for the critical overtopping discharge are respectively 10, 1 and 0.1 l s−1 m−1 to incorporate the quality of the levee cover47. We chose RP10 and RP1000 for, respectively, the low and high storm return period scenario. The uncertainty spread of vegetation width is based on the 75% confidence intervals of the underestimated and overestimated vegetation widths of mangroves (+436 m, −136 m) and salt marshes (+597 m, −104 m) in the vegetation presence validation study. For the breaker index we solely chose a high scenario of 0.78, because the index of the global assessment (0.55) was already quite conservative71. For topography we applied a range corresponding to the typical vertical accuracy of the FAST intertidal elevation dataset (±50 cm). Two representative subsets of 500 transects for respectively mangroves and salt marshes have been derived using the clustering method k-means96, based on hydrodynamic conditions, vegetation cover, profile characteristics and geographical location. With these subsets, we repeated the analysis procedure of the global assessment for the sensitivity scenarios. The results point out that the largest spread is caused by the uncertainty in the unit levee cost with −66% and +200% for, respectively, the low and high scenario with respect to the global reference analysis. The other scenarios: topography (−39%, +47%), critical overtopping discharge (−40%, + 40%), storm return period (−28%, +34%), vegetation width (−28%, +39%), breaker index (+21%) (Supplementary Fig. 10). Larger water depths result in a decrease of depth-induced wave energy dissipation and more dissipation due to wave-vegetation interaction, which explains the outcomes of the topography sensitivity results. Similarly, an increase of the storm return period or the breaker index shifts the ratio of wave energy dissipation by wave-bottom interaction and wave-vegetation interaction. The coastal protection costs by vegetation are sensitive to critical overtopping discharge changes, because of the non-linear relation between the wave height in front of the levee and the overtopping discharge47. More

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