Abstract
This study aims to quantify the dimensions of an oyster reef over two years via low-cost unoccupied aerial vehicle (UAV) monitoring and to examine the seasonal volumetric changes. No current study investigated via UAV monitoring the seasonal changes of the reef-building Pacific oyster (Magallana gigas) in the German Wadden Sea, considering the uncertainty of measurements and processing. Previous studies have concentrated on classifying and mapping smaller oyster reefs using terrestrial laser scanning (TLS) or hyperspectral remote sensing data recorded by UAVs or satellites. This study employed a consumer-grade UAV with a low spectral resolution to semi-annually record the reef dimensions for generating digital elevation models (DEM) and orthomosaics via structure from motion (SfM), enabling identifying oysters. The machine learning algorithm Random Forest (RF) proved to be an accurate classifier to identify oysters in low-spectral UAV data. Based on the classified data, the reef was spatially analysed, and digital elevation models of difference (DoDs) were used to estimate the volumetric changes. The introduction of propagation errors supported determining the uncertainty of the vertical and volumetric changes with a confidence level of 68% and 95%, highlighting the significant change detection. The results indicate a volume increase of 22 m³ and a loss of 2 m³ in the study period, considering a confidence level of 95%. In particular, the reef lost an area between September 2020 and March 2021, when the reef was exposed to air for more than ten hours. The reef top elevation increased from -15.5 ± 3.6 cm NHN in March 2020 to -14.8 ± 3.9 cm NHN in March 2022, but the study could not determine a consistent annual growth rate. As long as the environmental and hydrodynamic conditions are given, the reef is expected to continue growing on higher elevations of tidal flats, only limited by air exposure. The growth rates suggest a further reef expansion, resulting in an increased roughness surface area that contributes to flow damping and altering sedimentation processes. Further studies are proposed to investigate the volumetric changes and limiting stressors, providing robust evidence regarding the influence of air exposure on reef loss.
1 Introduction
The climate crisis entails increasing ocean temperatures, rising sea levels and acidification which affect marine ecosystems, leading to ecological changes (). The ecological shift can partially transform ocean and coastal environments with irreversible degradation and risk of biodiversity losses (). Increasing local ocean temperatures may deteriorate but enhance living conditions () and promote species translocations. Benefiting from the rising water temperatures in the North Sea, the non-native Pacific oyster Magallana gigas (, formerly Crassostrea gigas) has invaded the East Frisian Wadden Sea through larvae drift, mainly from aquaculture plots in the Netherlands in the late 1990s (Wehrmann et al., 2000; ; ).
The Pacific oyster has transformed native blue mussel (Mytilus edulis, von Linné and Salvius (1758)) beds into three-dimensional (3D)-structural reef complexes with highly rough surfaces and replaced the blue mussel in its function as an ecosystem engineering species (; ; ).
Prior to the bioinvasion, the blue mussel dominated the tidal flat sediment surface affecting biological and physical conditions such as the bottom roughness (), the boundary layer and sedimentation processes (; ; ). The blue mussels offer a suitable and often the only accessible hard substrate for the oyster larvae to settle on in the intertidal zone (Wehrmann et al., 2000; ; Wrange et al., 2010). Oysters cement themselves to these hard surfaces and their adjacent conspecifics, building solid oyster-sediment matrices (; ). These structures grow with each generation, and older individuals located deeper can be covered or buried by sedimentation and younger oyster generations. When individuals die, the shells remain and provide further substrate to settle (; ). Depending on the age of a reef, the internal reef structures differ locally in abundance, sediment coverage, shell growth and orientation, forming different patterns and roughness levels within the reef (; ).
The transformation from mussel beds to oyster reefs has caused an irreversible ecological shift in the benthic habitat (; ; ) and may change populations of benthic species. However, the reefs also offer a wide range of ecosystem services, such as filtering water, carbon sequestration, nursery habitats, biological biodiversity and regulation of nutrients (; ; Windle et al., 2022). Besides ecological changes, the complex 3D-reef structures, typically covering nowadays broad spatial areas, affect local hydro- and morphodynamics and vice versa in the German Wadden Sea (; ; ). Oyster reefs can mitigate flow and wave energy as living breakwaters (; ; ) and stabilise surrounding sediment (; ; ). Considering future extreme sea levels (), the flow-attenuating features could be a part of future coastal protection measurements and extend far outside their reefs, affecting tidal flats and protecting the surrounding soft-sediment environment against erosion (Walles et al., 2015). Mitigating erosion and dissipating hydrodynamic energy are anticipated at the lee side that may promote vegetation growth (; ) besides sediment accumulation ().
Oyster reefs comprise densely packed and well-cemented valves; they thus resist remarkably well to forces exerted by waves, tidal currents, and ice drift. This feature is much more pronounced than for blue mussel beds (), resulting in higher survival chances for oysters, once initial reef settlements have been established. However, the habitat must meet specific environmental conditions for the Pacific oyster, including several biological and physical factors. There is only a little understanding of the maximum elevation reached by Pacific oyster reefs at tidal coasts, as their vertical growth depends on the water level. Thus, a possible correlation between reef elevation and the water level should be investigated.
While the Pacific oyster has spread out for more than two decades along the German west coast since its invasion from the Netherlands (Wehrmann et al., 2000), the global oyster population has suffered losses of about 85% worldwide () due to diseases, ocean heating and anthropogenic activities such as overfishing and pollution (; Windle et al., 2022). Across the US Atlantic coastline, remote sensing has been employed to evaluate the American oyster’s (Crassostrea virginica, ) restoration attempts to enhance ecosystem services (; ; ). In contrast, previous studies of the settlement and exponential spread of the Pacific oyster in the North Sea have raised apprehension regarding the possible impact on the native community and surrounding tidal flats (; ; ).
However, recent studies have indicated enhancements such as increased biodiversity (; Zwerschke et al., 2020), coastal protection (; ) and providing a habitat for various species through oyster reefs compared to blue mussel beds ().
In that context, it is paramount to access suitable methods and instrumentation that allow a more extensive spatial coverage and provide accuracies as temporal growth magnitudes across species-related cycles are in the order of <10 cm. In the past, oyster reefs have been surveyed using measurement methods, such as terrestrial laser scanning (TLS) (; ; ), airborne (; ) and satellite surveys (; ; Winter et al., 2016). The time-consuming TLS can yield high-resolution point clouds of reef structures, but more efficient methods cover broader areas. Using aerial photographs, distinguished dead oyster reefs from living oyster reefs by colour as a decisive parameter. As an active optical technique, airborne light detection and ranging (LiDAR) measures geometric structures with three-dimensional information and has become a standard tool for monitoring coastal zones (). Optical airborne and satellite data are limited by daytime, tidal and weather conditions (). To overcome these issues, electro-optical and high-resolution synthetic aperture radar (SAR) sensors of satellites were added to the optical data to measure the electromagnetic reflectance from the surface (Winter et al., 2016), independently from the tide, cloud coverage and poor light conditions (). , and could confirm an improved classification of mussel beds and other intertidal habitats in optical imagery when combined with TerraSAR-X images due to high surface roughness of mussel beds and oyster reefs, which increase the reflecting backscatter of the signal from the rough surfaces. Winter et al. (2016) tested a combination of remote sensing techniques for mapping and assessing intertidal flats in the German Wadden Sea, namely airborne LiDAR, electro-optical (RapidEye) and synthetic radar (SAR) sensors installed on satellites (; , ). Previous studies investigated mussel beds (including oysters), bedforms and other surface types via TerraSAR-X images but were limited in precisely determining the extension of bivalve beds ().
Satellites and airborne surveys from higher altitudes cannot provide structural features such as roughness, slope and elevation on a high-resolution level to identify typical small-scale reef characteristics (). Therefore, adding structural features as training data on cm-scale may increase the classification success of oyster reefs.
In contrast to other monitoring techniques, unoccupied aerial vehicles (UAV) can increase the efficiency of monitoring oyster reefs due to a higher spatial resolution, similar accuracies to TLS and without physically influencing the field of investigation (). In the last two decades, the application of UAVs has increased in mapping coastal and estuarine environments (). The application of UAV has been reported for marine ecosystems like macrophytes (), seagrass (; ; Ventura et al., 2023), mangroves (), corals (; ; ) and for beach morphology studies (; ). While investments in professional UAV systems can be costly but derive data of high accuracy, consumer-grade UAVs have less precision but have so far shown promising results in determining information and data from marine ecology, e.g., in analysing the topography of honeycomb worm reefs () and the annual changes in sediment volume of a fringing reef island ().
Consumer-grade UAVs are often equipped with RGB sensors. In contrast, more sophisticated UAVs are equipped with multi- and hyperspectral sensors providing a broader detection of wavelengths (; ). and stated that a higher spectral resolution has a much larger effect on detecting oysters than increasing the spatial resolution. stated similar appearance of oyster reefs, mudflats and saltmarshes in remote sensing data, which may complicate the mapping of reefs in standard GIS tools. However, investigated different coastal habitats, proving that high-resolution mapping and classification can also be achieved based on simple RGB sensors on a consumer-grade UAV, promoting low-cost monitoring.
The structure from motion (SfM) technique is reliable for generating a digital model for subsequent spatial analysis and mapping of oyster reefs (Windle et al., 2019; ; ) and other coastal environments (). Various approaches have been employed for surveying and mapping oyster reefs, ranging from time-consuming () manual interpretation using satellite imagery (; ) to machine learning algorithms such as Support Vector Machine (SVM) (), with a convolutional neural network (CNN) () and other approaches that learn and detect patterns in data.
Windle et al. (2022) applied an unsupervised classification tool to cluster segmented RGB pixel values of an orthomosaic to classes according to oyster colour. Based on multispectral and RGB imagery, oyster reefs were identified in an estuary in the harbour of Auckland, New Zealand () and Florida () by applying object-based image analysis (OBIA, segmentation technique) of low-spectral and multispectral imagery, respectively. Based on the above literature reviewed, we conclude that multiple studies focus on identifying oyster reefs based on spectral signatures derived from UAV and satellite imagery by supervised and unsupervised classification methods. High spectral resolution sensors of satellites cannot generally offer a spatial resolution, which in turn, the use of UAVs promises. Low-cost consumer-grade UAVs commonly miss a higher spectral resolution, so elevation models are applied to provide additional information for classifying UAV data.
Only very few studies analysed the dynamic of oyster reefs. Still, the seasonal volumetric changes have yet to be examined, while such understanding would facilitate future prognostic capabilities to consider marine environmental conditions and changes. The measurement results of the vertical reef growth and volumetric changes result from various geomorphological processes, including sediment accumulation, biodeposition (), subsidence of the North Sea (; ) and reef consolidation (Wehrmann, 2009; ). However, due to the limitation of separating these processes within this study, positive and negative volumetric reef changes refer to volume increase and loss, respectively, and include all processes in the following.
In our study, we quantified the reef growth of the Pacific oyster in the German Wadden Sea based on high-spatial and low-spectral resolutions derived from consumer-grade UAVs that proved effective for detecting oyster reefs by a machine learning classifier. Despite small-scale changes in oyster reefs, errors through measurement and alignment were hardly recognised as uncertainties in the statement of results by previous studies. In this study, we will pursue the following specific objectives:
• To identify and map oyster reefs using high-resolution digital elevation models (DEMs) and orthomosaics.
• To quantify the reef dimensions over two years and to derive yearly growth parameters.
• To detect and characterise any significant seasonal changes in the volume of the oyster reef.
• To examine the impact of considering the uncertainties of the vertical elevation on the volumetric results due to measurement and alignment errors.
2 Materials and methods
2.1 Study site
This study focuses on the oyster reef (latitude: 53.6470116°N, longitude: 8.2664760°E) at the tidal channel, named “Kaiserbalje”, on the “Hohe Weg” tidal flats in the central Wadden Sea north of the peninsula Butjadingen, Germany (Figure 1). The Kaiserbalje is situated between the Jade and Weser estuaries. The area of interest was approximately 650 m from west to east and 300 m from north to south, whereas the area recorded by the UAV was 270,000 m². A local tidal range of 3.35 m (Pegel Hooksielplate (WSV, 2022a), Figure 1E) gives a mean aerial exposure time of approximately 3.5h at the oyster reef. The climate of the central North Sea is temperate and influenced by westerly winds (). Mainly, the current velocity flows from east to west, and its maximum velocity ranges from 0.2 to 0.4 m/s. The wave direction is primarily from the northeast and east, with a significant wave height below 0.75 m. Waves with significant heights above 0.75 m come from the north and northwest (). At Kaiserbalje, the mean oyster abundance has increased from less than ten individuals per m² in 2003 to around 360 individuals per m² in 2019 (). Even though the Pacific oyster has transformed mussel beds into oyster reefs, the blue mussel has found its spatial niche among oysters, as observed in this study’s field investigations (Figure 2). Observation evidence obtained during field surveys between 2019-2022 suggests that sandy conditions prevail mainly around the Kaiserbalje reef, while muddier flats and soft biodeposits made of faeces and pseudofaeces appeared within and near the reef.
Figure 1
Figure 2

The coexistence of the Pacific oyster (Magallana gigas, blue ring) and the blue mussel (Mytilus edulis, red arrows) shapes the appearance of the Kaiserbalje and the roughness of the reef surface. Barnacles (the brown oval shape) commonly infest the surface of oyster shells, whereas the blue mussel finds its place between oysters.
2.2 Data collection
In six spring/autumn field campaigns, the topography of the Kaiserbalje was measured between October 2019 and March 2022 to obtain high-resolution data sets, employing an unoccupied, drone-style aerial vehicle (UAV) system. Every campaign comprised a two-hour flight mission of the reef, collecting between 1,099 and 2,010 images. Three-hour time windows limited the accessibility to the reef and working time during low tide conditions.
Recording of the study site was accomplished by a UAV (DJI Phantom 4 Pro), equipped with an integrated high-resolution RGB camera 1"-CMOS sensor capturing 20-megapixel images. This gimbal-mounted lightweight quadcopter used the Global Positioning System (GPS) of the US and the Global Navigation Satellite System (GLONASS) of Russia as on-board global navigation satellite system (GNSS) receivers.
The flight app DroneDeploy (https://www.dronedeploy.com) was used for automatic flight missions, specifying a systematic strip-by-strip route with a constant velocity of 3 m/s and a recording frequency of one photo every three seconds. A sufficient front (80%) and side overlap (60%) ensured adequate data processing, according to
In the three-hour time of aerial survey by UAV, up to 21 GCPs were distributed on the ground, evenly across the entire field area, particularly along the edges, corners, and as one line spanning the entire width of the reef. The GCPs used in this study were made of PVC tarpaulins and marked with a measuring cross. No GCPs were located at most 200 m from a neighbouring GCP, as recommended by
The concerned authorities permitted all field investigations: Wadden Sea National Park Authority of Lower Saxony (NLPV) and the Agency for Coastal Defence, National Park and Marine Conservation Schleswig-Holstein (LKN.SH). Under the given regulations of the authorities, any effects on other species were minimised, and no disturbances were noticed during measurements. The drone pilot complied with the competency requirements of the EU regulation 2019/947 and was supported by one trained observer in the field.
2.3 Data processing
2.3.1 Structure from motion
Structure from motion (SfM) was applied to construct DEMs and orthomosaics by identifying characteristic key points in a series of multiview stereo imagery with known camera orientations and geotags (Westoby et al., 2012; Windle et al., 2022), providing robust 3D-structural metrics on a cm-scale (
First, during the study period, the application of the GCPs was improved by increasing the number of GCPs and expanding the distribution across the entire reef. For this reason, the DEM of October 2019 could not be generated without any doming effects and, consequently, was removed from the analysis. These distortions led to non-acceptable data for further processing. Sufficiently good coverage of GCPs was established for the subsequent field campaigns, and the closer vicinity offered an appropriate georeferencing of the models. However, no more GCPs could be placed further north, as the northern part was hardly accessible due to strong currents in tidal creeks.
2.3.2 Classification
All DEM and orthomosaics were exported with a 5 cm/pixel resolution into QGIS (version 3.16 “Hannover”; qgis.org) to arrange the training data set for detecting oysters in the UAV imagery using a machine learning algorithm, as described below. This resolution was chosen to compromise the computing power (processing time and storage capacity) and the accuracy. This study integrated the oyster reef’s representative spectral and structural (roughness, slope and elevation) features into the training data set derived from the RGB bands and DEMs, respectively. Moreover, including a colour band derived from the orthomosaics as a roughness attribute demonstrates an enhanced classification success rate when incorporated into the training data. In this context, colour roughness refers to variances in the colour values between neighbouring pixels.
The supervised method Random Forest (RF) by
The prediction of the unclassified data was performed using the randomForest package (v. 4.7-1.1) (
Applying the OOB error, no further cross-validation nor testing was needed to estimate the classification performance (
The classified output data was achieved with a success rate expressed as OOB error between 2.0% (March 2022) and 6.6% (September 2020), respectively. The OOB errors have reached a final equilibrium after 100-150 trees. The elevation and the roughness of both the DEM and the red channel of the RGB orthomosaic often had the highest impact on the MDA and MDG performance, i.e. the classification accuracy mainly depended on these factors. Figure 3 illustrates the coverage and the successful identification of the oyster reef.
Figure 3

(A) and (B) showcase the oyster distribution (green coverage) determined by the Random Forest algorithm during the March surveys of 2020 and 2022. One can discern the precise classification of oyster populations by examining the insets (red area) of the main maps.
2.4 Error propagation
Determining small-scale changes in topography via remote sensing requires consideration of error propagation and uncertainties. Although the consequences of errors on volumetric results, previous marine and coastal studies have rarely analysed the propagation of errors, particularly for significant elevation changes. A few studies, e.g.
The photogrammetric tool Metashape evaluated the vertical accuracy of the SfM-derived point clouds based on the coordinates of CPs and provided a single global uncertainty σCP as root mean square error (RMSE) covering the entire area of the DEM. RMSE is a standard statistical metric to assess the prediction errors (
where n is the number of CPs. A total vertical RMSE of 2-3 times the GSD is expected to be achieved here, which can be considered well-accepted (
The dGPS device estimated the vertical error using the vertical root mean square error (VRMSE) for each GCP measurement, which appeared to be higher than the manufacturer-specified device precision (s. subsection 2.2). The total vertical uncertainty of the GPS measurements σGPS is given as
and presents a global vertical uncertainty with n number of measurements. From the GPS measurement (Table 1), a σGPS between 1.9 cm (March 2021) and 2.5 cm (March 2020) was derived by the output of the dGPS device.
For each specific DEM, the overall vertical uncertainty σz,mmm−yy was calculated as the square root of the corresponding squares of the vertical uncertainties from both σCP and σGPS using the following formula
where σz ranges between 2.7 (September 2020) and 3.9 cm (October 2021, Table 1).
Table 1
| Survey | σGPS [cm] | σCP [cm] | σz [cm] |
|---|---|---|---|
| 2020 March | 2.5 | 2.5 | 3.6 |
| 2020 September | 2.3 | 1.4 | 2.7 |
| 2021 March | 1.9 | 3.2 | 3.7 |
| 2021 October | 2.2 | 3.2 | 3.9 |
| 2022 March | 2.2 | 1.9 | 2.9 |
The vertical uncertainty of the GPS measurement σGPS, the validation of the alignment σCP and the total vertical uncertainty σz.
When determining the volume growth and loss as well as the growth-related elevation changes of the reef structures over the study period, the total vertical uncertainty must be considered since this enables identifying areas with significant changes (
for the DEMs of difference (DoD) that highlighted the geomorphological changes with a confidence level of 68% (
Concerning the minimum change detection with a confidence level of 95%, the level of detection (LoD), a spatially uniform approach by
where t equals 1.96 under the t-distribution (
The seasonal σDoD ranges between 4.5 and 5.4 cm, whereas the LoD varies between 8.8 and 10.6 cm (Table 2). Due to the smallest propagation errors in September 2020 and March 2022, the smallest uncertainty could be found for the DoD between September 2020 and March 2022, with a value of 4.0 cm and an LoD of 7.7 cm. The largest error occurred for DoD between March and October 2021, with 5.4 cm and 10.6 cm for σDoD and LoD, respectively.
Table 2
| Season | σDoD (68%) [cm] | LoD (95%) [cm] |
|---|---|---|
| March 2020 - September 2020 | 4.5 | 8.8 |
| September 2020 - March 2021 | 4.6 | 9.0 |
| March 2021 - October 2021 | 5.4 | 10.6 |
| October 2021 - March 2022 | 4.9 | 9.5 |
| March 2020 - March 2022 | 4.6 | 9.0 |
The vertical uncertainty of the DEM with a confidence level of 68% (σDoD) and 95% (LoD).
To determine the volumetric uncertainty σV [m³] of a DoD with a confidence level of 68%, the corresponding uncertainty σDoD was multiplied by Apixel and the number n of pixels of the area of either loss or increase.
According to a confidence of 95%, σDoD was replaced by LoD. Since the survey in March 2020 only had one CP, the corresponding RMSEs show low values (<GSD). To improve the reliability of the uncertainty of this survey, the errors were re-calculated and merged as a total vertical RMSE by the Leave-One-Out (LOO) method (Villanueva and Blanco, 2019;
2.5 Calculation of reef top elevation, area and volume change
All reef dimension calculations and the extraction of values were performed within QGIS. For each season survey, the reef area Ammm−yy was determined from the total number of pixels n of the classified raster layer of the DEM multiplied by the pixel area Apixel (0.0025 m²)
where yy and mmm indicate the year and the month. n was determined by subtracting the number of pixels with no values (“NODATA PIXEL COUNT”) from the total pixel count (“TOTAL PIXEL COUNT”). Both counts were yielded by the tool “Raster Layer Unique Values Report” in the “Processing Toolbox” of QGIS.
To determine the reef volume and bottom, we introduced a definition of the reef bottom, applying a lower limit of -0.6 m as a reference elevation zref in the computation. When considering the highest 99% of detected oysters, individuals were still found on this water depth for all maps. As most of the lower-lying oysters did not belong to the reef, the lowest 1% was excluded to eliminate individuals, clusters or other non-reef elements. The reef top (=maximum elevation point of the reef surface) was defined by identifying the 99.9th percentile of oysters within the study field to exclude outliers and individuals outside the central reef structure. So far, no research has established specific upper and lower limits for an oyster reef based on the omission of a specific percentile. However, such an approach is not necessarily required when analysing smaller reefs but for larger reefs (> 100 m²) consisting of complex and interconnected reef components.
To describe the reef volume VDEM,mmm−yy at the respective time of measurement, the sum of the elevation for each pixel that is given by subtracting the pixel elevation zi value from zref was multiplied by Apixel.
If the volumes are computed with an uncertainty, σz (s. Equation 3) is appended to Equation 8. The tool “Raster Surface Volume” of the QGIS Processing toolbox was applied to find the volumes of the DEM raster (Input layer) where zref was used as the “Base level” and “Count Only Above Base Level” as a method. If considering the total vertical uncertainty σz, either σz is subtracted from the “Base level” or added to find the volumetric range. In addition to the volumes at the respective points in time, the volumetric changes were quantified between two survey times, where DoDs (
The DoD raster output contains the elevation change of each pixel where positive values show a growth and negative a decrease in elevation. Subsequently, the volume changes were determined by multiplying the pixel size Apixel with the elevation changes Δz in the DoD pixel-by-pixel (similar to Equation 8)
where σDoD (s. Equation 9) is appended, and t equals either 1 or 1.96 when considering error propagation. To find the volumetric changes, the “Base level” was kept at zero, and either “Count Only Above Base Level” or “Count Only Below Base Level” were chosen in the tool “Raster Surface Volume” tool for volumetric growth and loss, respectively.
Figure 4 illustrates the previously described workflow to highlight the essential processing steps in this study.
Figure 4

The workflow includes field measurements, photogrammetric processing in Metashape and QGIS, classification and statistical analysis in R, and error propagation analysis.
3 Results
3.1 Topographical changes
The elevation of the oyster reef top reaches an average of -0.15 ± 0.02 m NHN between 2020 and 2022, and most of the reef surface rises above the surrounding sediments (Figure 5). This allows the oyster reef to be distinguished from the surrounding tidal flats. The surrounding sediment within the reef boundary (Figure 1A) has an average elevation of -0.49 ± 0.02 m NHN, derived by the DEMs. Depths below -1.0 m NHN are located on the reef’s west and east sides, where tidal creeks were observed from the field measurements that did not fall dry entirely during low tide. The bottom of these tidal creeks pass along the reef edges and run into the bed of larger tidal creeks west and east of the reef. South to the reef, a shallower and narrower bed of a creek can be observed. In all tidal creeks, the water runs from north to south during the ebb and vice versa during the flood. Due to sediment deposition within the tidal creeks, higher surface elevations are observed than the actual bed elevation. When identifying the oyster reef, smaller patches were detected outside the reef, in and beyond the tidal creeks, but neglected in the analysis (<0.05 m²).
Figure 5

Digital elevation model (DEM) of the Kaiserbalje in March 2022 shows the actual elevation of the study site in m NHN. A hillshading effect was added to highlight the rough surfaces, providing a more natural spatial visualisation of the surface. Differences in height from 0.0 to -0.80 m NHN are represented in brown colours, and differences from -1.00 to -1.60 m NHN are illustrated in blue colours. The black contour line presents the rough reef boundary within the tidal creeks and a distance of 10 m to the oyster-identified area. The white points with the black dots present the ground control points (GCPs) distribution, and the black points illustrate the check points (CPs).
Primarily, the oyster reef comprises a lengthy central section surrounded by a transition zone and individual patches, as described in detail in
While the area determination is straightforward, defining the volume needs more attention due to the uncertainties caused by the GPS measurement and the photogrammetric alignment at a magnitude of 1.9 to 2.2 cm and 1.4 to 3.2 cm, respectively. The error propagation reveals elevations with uncertainties that eventually affect each survey’s volumes. Hence, presenting the volumetric results need to include these uncertainties in the following analysis. Maps of change detection with consideration of the confidence level are suitable for visualising the area of elevation changes (Figure 6).
Between March and September 2020 (Figure 6A), the reef grew vertically in a small area of 1,134 m² (3% of the initial reef area), where significant changes of 223 m² (<1%) were included, especially in the southwestern corner. However, erosion predominates in the north-western side over an area of approximately 5,500 m² (15%). In contrast, the reef could gain in growth on an area of >10,000 m² (27%) on the eastern side from September 2020 to March 2021 (Figure 6B). But minor 158 m² (<1%) areas at the edges close to the tidal creek were still eroded. Between March 2021 and March 2022 (Figure 6C), the reef could compromise the losses from the period between March 2020 and September 2020 on the western side (approx. 6,000 m², 18%) and the loss of the reef edge on the eastern side. From October 2021 to March 2022 (Figure 6D), reef expansion on 34 m² and only losses at the edges on the southeastern side (1,846 m², 5%) could be detected, including significant changes of an area of 271 m² (1%).
Figure 6

The subfigures (A–D) illustrate the seasonal and (E) the total vertical changes of the reef between surveys, divided into loss (brown) and increase (blue). (E) shows the topographical changes for the entire period from March 2020 to March 2022. The change detection with a confidence level of 68% is presented with light blue and brown colour and the significant changes with a confidence of 95% with a dark blue and brown colour. Both confidence levels correspond to a different minimum vertical change detection for each DoD (Table 2). Within the range of the uncertainty, the elevation changes are kept transparent, i. e. changes with a confidence of 68% and 95% are included above the uncertainty. The green and magenta arrows present the main direction of the current and waves, and the surveys’ maximum reef top elevations are marked with a black cross. The change detection without considering uncertainties are shown in the maps in Supplementary Material.
In summary, the largest areas of reef elevation increase were observed on the eastern side during the period of September 2020 and March 2021 and on the western side between March and October 2021. The largest area of elevation decrease was observed in the northwest and partially at the edges of the reef between March and September 2020. The DoD of March 2020 and March 2022 (Figure 6E) reveal that the oyster reef principally grew vertically over an area of 11,450 m² (30%) with confidence above 68%, especially at the east and southwest corners close to the tidal creeks. Significant growth in reef elevation was observed over an area of 1,298 m² (3%), whereas small reef parts (254 m², 1%) had decreased elevation along reef edges. The minimum and maximum vertical occurrence of oysters ranges between -1.26 ± 0.23 m and 0.00 ± 0.13 m NHN, including all identified oysters by the Random Forest algorithm.
The maximum surface elevation of the reef top ranged from -18.2 ± 2.7 cm NHN for DEMSep-20 to -12.8 ± 3.7 cm NHN for DEMMar-21, while the average surface elevation of the reef top was -0.15 ± 0.0 cm (47.8 ± 0.5% air exposure). In the final survey in May 2022, the maximum elevation reached -14.8 ± 3.9 cm NHN, resulting in a total vertical growth of 0.7 ± 4.6 cm over two years for the reef top. The largest increase in reef top elevation was observed within the period of September 2020 and March 2021, with an increase of 5.5 ± 4.6 cm (Figure 7). In contrast, the maximum elevation decreased for all other seasons, particularly between March and September 2020 (from -15.5 ± 3.6 cm NHN to -18.2 ± 2.7 cm NHN). The regression line in Figure 7 revealed a modest annual growth trend of 2.1 ± 13.9%, corresponding to 0.3 ± 3.6 cm/y over the two years for the reef top, with a coefficient of determination R2 of 0.21, indicating a limited explanatory power of the model.
Figure 7

Development of the reef top elevation and the oyster reef area at the Kaiserbalje over the entire study period. The blue squares show the elevation at the time, with the associated vertical uncertainties (RMSE), and the red dots present the area size of the reef at the respective times. A regression line was drawn for both developments, with an R² value of 0.21 (elevation) and 0.16 (area).
The reef area development differed from the trend in elevation growth since the surface size had declined constantly from March 2020 (37,765 m²) to March 2021 (34,760 m²), resulting in a loss of 3,006 m² (Figure 7) that was followed by a steady growth until March 2022, the area growth compensates for the loss with 4,784 m² in the second half of the study period. During the study period, the reef area increased by 1,778 m², expanding from 37,765 m² to its maximum recorded size of 39,543 m² in March 2022, within our monitoring. Considering the regression line in Figure 7, the area experienced a growth of 889 m²/y. Since the coefficient of determination R2 indicates a value of 0.16, no reliable trends should be concluded, given the limited explanatory power of the model.
Depending on the development of the reef elevation and area, the total reef volume extended from 8,046 ± 1,350 m³ in March 2020 to 9,370 ± 1,138 m³ in March 2022, with a total growth of 1,324 m³. Using -0.6 m as fixed reference elevation concerning the total vertical uncertainty σz (s. Equation 3), reef volume shows a clear increasing trend over the study period (Figure 8). Like the reef top elevation, the total reef volume decreased to a minimum of 7,561 ± 1,000 m³ in September 2020 but increased until September 2021 with a 9,374 ± 1,483 m³ volume. In contrast to the reef elevation and area, R2 is higher with a value of 0.75, and the growth trend of 662 m³ or 7.9% is more credible for the reef volume.
While Figure 8 focuses on the volumes and net growth of the reef related to the time of the survey, the DoD approach provides insights into reef increase and loss. DoDs divide the volumetric change into loss and increase between two surveys, implying that DoDs do not present the volume at the exact measurement time. The increase or loss did not turn out to be constant, suggesting no seasonal effects on the reef during the study period of two years (Figure 9).
Figure 8

Mean volumetric oyster reef development over two years of observation. Error bars depict root mean square error (RMSE), and the dashed line indicates a linear regression with R² = 0.75.
Figure 9

Volumetric changes illustrated according to their confidence level for each season and the total study period. The increase (blue) and loss (red) include the uncertainty σDoD (squares) and LoD (crosses) of the vertical changes, corresponding to a confidence level of 68% and 95%, respectively.
The results include the loss and increase calculated with a confidence level of 68% (σDoD, s. Equation 4) and a higher confidence level of 95% (LoD, s. Equation 5), highlighting the detection of geomorphological changes (
Between March and September 2020, the reef experienced more loss than increase, followed by the period between September 2020 and October 2021, where the increase exceeded the amount of loss. However, in between October 2021 and March 2022, the loss volume rose relative to the increase. Regarding a confidence level of 95%, the total increase and loss yielded volumetric changes of 22 m³ and 2 m³, respectively, whereas a confidence level of 68% resulted in an increase of 234 m³ and a loss of 6 m³. This means the reef has a net gain of 20 m³ and 228 m³ for 95% and 68% confidence levels, respectively. If no confidence level is considered, the DoDs show broader areas of vertical changes, suggestive of potential growth tendencies on a broader scale. During the study period, the reef experienced a growth of 1,346 m³ covering 34,897 m² in the entire reef and a loss of 88 m³ over 5,348 m³ in the northwestern section when no uncertainties are considered.
3.2 Statistical assessment
The mean reef elevation of the Kaiserbalje ranged between -40.1 ± 2.7 cm (mean ± standard deviation) NHN (September 2020) and -35.7 cm NHN (October 2021), with a standard deviation σ between 9.3 and 10.5 cm, respectively (Figure 10). The mean elevation has shifted throughout the study time from -39.6 to -36.8 cm NHN. The probability distribution of the elevation demonstrates a consistent unimodal distribution with a well-defined peak.
Figure 10

Left: Histograms of the probability distribution of the classified oysters’ elevations with the mean and standard deviation (A, C, E, G and I). Right: The quantile-quantile plots (Q-Q plots) show the probability distribution (B, D, F, H and J).
Regarding an oyster reef elevation distribution, a roughly symmetric distribution was found in March 2020 and for the subsequent surveys (Figures 10B, D, F, H, J), the elevation deviates from a normal distribution. On the one hand, a negative skewness (long left tails) indicates the presence of “extreme” elevation values for September 2020 to March 2022. On the other hand, the light right tails lead to the assumption that a potential upper limit exists. Walles et al. (2015) and
4 Discussion
This study’s results show a dynamic volumetric growth over the observation period of two years on the Kaiserbalje reef, with no evident seasonal effects. However, the magnitude of the results on the volumetric changes strongly depends on the confidence level introduced in the calculation (s. Equation 10 and Equation 11), considering the uncertainty in the elevation changes (
Unlike most previous studies in coastal monitoring, we consider error propagation and incorporate uncertainty in analysing small-scale changes affected by measurement and georeferencing errors. By accounting for these uncertainties, we can identify areas of changes with confidence in maps by applying DoDs, as in Wheaton et al. (2010) and
Studies with large elevation changes relative to vertical uncertainty yield more reliable results in terms of confidence since error propagation has a negligible effect on the minimum level of significant change detection. Most of the elevation changes in this study are within the range of uncertainties between 4.5 and 5.4 cm, especially for a confidence level of 95% (between 8.8 and 10.6 cm). Lower errors in measurement and processing should be pursued to achieve more significant change detection and increase the level of reliable information on topographical changes due to a low error tolerance for small-scale changes. Our results show the effect of uncertainties on volumetric changes and the effect of minor errors during measurements and processing. However, the uncertainties of DEMs and DoDs will unlikely reach values below 2-3 cm and 4-5 cm, respectively, due to the current limitations of the measurement tools. Similar studies (
The Random Forest algorithm classifier has been convincing, featuring simple application and good classification results. The classification accuracy, expressed as OOB error, is below 7%, with the best (2%) results for March 2022. Higher classification results were accomplished because the roughness and texture of oyster reefs stood out on sandbanks, becoming an essential classification feature. The DEM’s high spatial resolution guaranteed the inclusion of the roughness that contributed crucially to the classification success. This aspect could be considered a standard attribute not just for oyster reefs but for other small-scale structures as well. Most studies applied only the spectral reflectance of surfaces as a training feature for RF.
4.1 Reef dynamic
A comparison of the development of reef areas with other studies is limited due to the small number of studies that differ in, e.g. site conditions, initial area sizes, and artificially constructed and not-constructed.
The annual area growth of the reef at the Kaiserbalje was 2.3% (889 m²/y) between 2020 and 2022 and, hence, smaller than in the Oosterschelde, where the Pacific oyster was already introduced in 1964. Notably, direct comparisons of the Kaiserbalje with the Oosterschelde are difficult as the total oyster occurrence in the Oosterschelde consists of multiple reefs, and the environmental conditions vary between these sites. Probably due to the mature state of the Kaiserbalje reef, the constant expansion of the reef has stagnated and achieved a moderate balance of reef area after almost two decades of growth. First distinctive densities occurred in 2002 for the Kaiserbalje (Wehrmann, 2009).
Compared to surface area studies, more detailed investigations exist on the vertical growth of oyster reefs. The literature has shown that the reef’s surface and vertical growth depend on surface elevation or exposure time. When the reef top of the Pacific oyster reaches its growth ceiling, the increasing stress from air exposure will decelerate the vertical reef growth. Both in Walles (2015) and
The volume-to-area ratio can serve as a relative benchmark for comparing other oyster reefs and, possibly, link the reef age. Between March 2020 and March 2022, the ratio of the Kaiserbalje reef varied between 0.21 m³/m² to 0.25 m³/m² (Figure 11) and similar ratios were found for constructed oyster reefs in the Oosterschelde (Walles et al., 2016b) and in North Carolina (
Figure 11

Volume-to-area ratio is presented with the corresponding root mean square error (RMSE, error bar) for all surveys in the study period (black squares). The dashed line presents the mean volume-to-area ratio of four constructed oyster reefs in the Oosterschelde (Walles et al., 2016a), and the triangles present the ratio for the constructed reefs established in 1997 (blue, upward-looking) and 2000 (green, downwardlooking triangle) in
The choice of a 68% or 95% confidence level aligns with statistical principles, ensuring that detected changes are likely to be real changes, not random fluctuations, and providing a robust basis for identifying meaningful geomorphological changes. Confidence levels help to focus on substantial changes while filtering out minor measurement and processing uncertainties. It ensures that only significant changes are considered for the analysis and interpretation. Applying DoDs with σDoD and LoD provides a practical approach to quantify the minimum volumetric changes between seasons or survey periods, as it accounts for the combined error propagation, unlike the previously reported method of individual total DEM volumes.
Considering uncertainties greatly impacts reef elevation and volume results in analysing small-scale changes of the same magnitude as the errors eliminate a large part of the detected changes. Higher confidence levels yield smaller detectable change areas. To increase the areas of change detection, reducing uncertainties in individual DEMs is necessary for geomorphological changes on a cm-scale. Our findings highlight the significant impact of propagation error on volumetric changes, with increase and loss values decreasing substantially when considering uncertainties. Without considering any errors, total increase and loss were 1,346 m³ and 88 m³, respectively, resulting in a total volumetric change of 1,258 m³. However, when applying a confidence level that accounts for uncertainty, the reef growth decreases to 234 m³ at a 68% confidence level and 22 m³ at 95%, a reduction to 17% and 0.3% of the volume without considering uncertainties, respectively. Similarly, loss values decrease to 6 m³ and 2 m³, corresponding to 7% and 2% of the volume without considering uncertainties. We can deduce that the scope for interpreting reef development decreases as the confidence level increases. Other studies have also shown that the degree of confidence and propagation error affects the area and volumes of volumetric changes, resulting in challenges when accurately determining changes (
4.2 Abiotic stressors
The development of oyster reefs depends on numerous abiotic and biotic factors, such as hydrodynamic forces, salinity and temperature, which control growth and mortality (
For further studies, we recommend analysing the reef development and the abiotic stressors, such as the temperature and salinity, since increased acidity endangers the calcification and development of oysters (
Water temperature, however, is a primary driver affecting Pacific oysters’ reproduction, growth and population (
However, extended periods of cold temperatures can increase oyster mortality (
Figure 12

Potential stressors between January and March 2021 (A) and between January and March 2021 (B). The time histories of air (grey) and water (black) temperature and water level (blue) are shown here. Extreme water levels such as extended inundations (> ten hours, yellow) and low water (air exposure > ten hours of half of the reef, deep lilac) are presented as thick lines. Further, several storm events (red dots) occurred in winter. The entire water temperature profile can be found in Supplementary Material.
Global sea surface temperatures (SST) are expected to rise by 0.86° C between 2081-2100 compared to 1995-2014, projected by Coupled Model Intercomparison Project Phase 6 (CMIP6) models under the Shared Socio-economic Pathway 1-2.6 (SSP) (
In addition to ecological stress, oysters encounter constant hydrodynamic stress induced by waves and tidal currents, affecting their settlement and growth. Pacific oysters require suitable areas within the vertical growth zone, but rising sea levels may shift the vertical zone. Therefore, assessing if the reef’s growth rate can keep pace with local sea level rise is crucial. The current results show a yearly increase in the reef top elevation of 3.0 ± 36 mm/y, lagging the current rate of local SLR of 4.0 ± 1.53 mm/y in the North Sea from 1993 to 2009 (Wahl et al., 2013). By 2100, a recent SLR of 0.80 and 0.81 (median of RCP8.5) are projected for Cuxhaven and Hamburg, German North Sea coast cities, by
4.3 Challenges in UAV monitoring and classification of oyster reefs
Low-cost monitoring with high-resolution drone data proved suitable for mapping and analysing reef growth and transferring to other coastal monitoring applications. The RGB bands and spatial resolution of UAVs provided sufficient information to classify the drone data successfully, despite the relatively flat and homogenous surfaces of tidal flats and oyster reefs. In the presence of other surface classes, the influence of seagrass, algae or other marine surfaces on classification success should be further investigated since the overlap of spectral features could result in lower classification success.
In previous studies, the focus was on detecting oyster reefs generally based only on multi- and hyperspectral (
To investigate multiple oyster reefs in the German Wadden Sea on a regular base, the authors recommend investigating the possibilities of combining space-borne satellites and UAV monitoring for future monitoring of oyster reefs or other coastal ecosystems. Satellites can cover the entire German Wadden Sea, independing on personnel, while targeted missions with drones can fly over individual oyster reefs to ensure regular monitoring. Access to the investigation area was challenging and time-consuming, which led to limited spatial coverage and distribution of GCPs 0.8 GCP/ha and CPs (<0.2 CP/ha). However, if environmental and logistical conditions permit, increasing the number of CPs can improve the validity of σCP. In upcoming field campaigns, using an RTK drone may simplify the fieldwork and decrease the need for GCPs and CPs (
Our data provide a basis for small-scale analysis of oyster biomass, abundance and other ecological parameters that vary within the reef depending on the reef’s age and structural class (
So far, no studies have investigated and quantified the wave and current reduction under controlled laboratory conditions, except for
Within this study, we observed sediment accumulation around the reef but did not quantify it since it was outside the scope of this paper. Further research should focus on detecting sediment changes at the Kaiserbalje and comparison to other sites. The ecosystem engineering effects may contribute to coastal protection in the Wadden Sea by reducing the impact of waves and currents on the coast. After all, the flow damping and sediment stabilising need to be further investigated for their extent to evaluate the concrete contribution to coastal protection. It may prove that the Pacific oyster supports coastal protection as a nature-based solution (NbS) in the face of rising sea levels in the North Sea by sediment stabilisation. This can be of great importance when the oyster reefs rise at the same rate as the sea level and stabilise sediment in the longer term. To improve the reliability of the growth trends and limitations of the Kaiserbalje reef, further high-resolution monitoring of the reef via UAV over a longer period is highly recommended. Seasonal monitoring will deliver additional information on which periods the reef further grows and how temperature, air exposure, storm events and other abiotic factors influence the reef growth. So far, an ending of the horizontal expansion cannot be assumed since the surrounding sediment still offers space to settle on, and the reef could increase the coverage.
5 Conclusion
For the first time, seasonal volumetric changes of an oyster reef in the German Wadden Sea were precisely quantified to provide valuable growth parameters for estimating future reef expansion. Using low-cost monitoring through a consumer-grade UAV enabled the acquisition of high-resolution imagery, successfully classified by the Random Forest algorithm, resulting in accurate oyster maps. Despite successful change detection, the uncertainty of vertical growth challenged the quantification of small-scale changes. To address this, we adopted an approach of considering the error propagation through the processing, resulting in a high confidence (95%) of the reported significant changes. However, considering confidence levels reduced the area of detectable changes due to the increasing uncertainties and may eventually not represent the reef evolution sufficiently, resulting in vague interpretations. This study reveals that considering uncertainties ensures confident results, providing a minimum change detection of geomorphological changes. However, the results also show that large reef areas are often excluded from evaluating reef dynamics due to relatively high uncertainties. Even so, results without considering uncertainties can still describe reef dynamics and allow simple interpretations of reef development. As consumer-grade drones only cover small areas and the tide limits the flying time, space-borne systems could support monitoring oyster reefs on a larger scale. Furthermore, while consumer-grade drones offer the high spatial resolution necessary for accurate quantification of reef dynamics and expansion, space-borne systems may be valuable for monitoring oyster reefs on a larger scale, considering drones’ limited coverage and flying time. The approach employed in this study can be adapted for accurate and practicable monitoring of other ecosystems and contributes to marine ecology for sustainable coastal management strategies in the Wadden Sea.
The results indicate a net gain in reef area and volume, suggesting the potential of the reef to continue growing and keeping pace with the current local SLR. Given that the oyster reef or other hard substrate offer appropriate surfaces within the vertical growth zone, larvae can still settle, contributing to further reef expansion in all directions. While we could not define a specific growth ceiling (m NHN or air exposure) regarding reef top elevation for the Pacific oyster, the critical question remains whether the oyster reef will ever reach a growth limitation in light of the ongoing local SLR. While it is assumed that extended air exposure may have contributed to the observed decrease in area, no other stressors and seasonal effects influencing the volumetric growth were identified.
The promising results provided by this study make evident that long-term study periods are required for a better understanding of the growth dynamics and that validation with other oyster reefs to increase the robustness and reliability of results on the volumetric growth, particularly as elevations increase relative to uncertainties, is needed. Understanding the dynamics of oyster reefs is essential for effective coastal management, and this study contributes valuable insights to support such efforts.
Statements
Data availability statement
The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.
Author contributions
TH, MW, TS, MP, AW, and NG contributed to the conceptualisation. TH developed the methodology. TH, JH, KP, LB, AW, and MP carried out the investigation in the field studies. TH conducted the formal analysis. TH wrote the first draft of the manuscript and prepared visualisations. MW, MP, TS, AW and NG provided supervision. NG, AW, TS, and MP organised funding acquisition. NG was responsible for the project administration. All authors contributed to the article and approved the submitted version.
Funding
This project “BIVA-WATT” on which this work is based was funded by the Federal Ministry of Education and Research of Germany (BMBF) under the funding code 03KIS128. The publication of this article was funded by the Open Access Fund of Leibniz Universität Hannover.
Acknowledgments
The authors gratefully acknowledge the captain and crew of the FK Senckenberg for their logistics support and accommodation during the field campaigns and for keeping us safe on the North Sea. A special thank you goes to Torsten Janßen, who managed the field trips and guided us safely on the tidal flats. Regarding photogrammetric evaluation, we would like to thank Alexander Störmer and Dr. Bastian Steinhoff-Knopp (Institute of Physical Geography and Landscape Ecology, Hannover), who gave us hints for the UAV classification, and Prof. Anette Eltner (Institute of Photogrammetry and Remote Sensing, Dresden) for her valuable feedback concerning the analysis of error propagation. Additionally, we extend our appreciation to the students who supported us through their theses (Anne Eilrich, Katharina Michalak, Lennart Stukenborg, Lea Weiß, and Sylvi Junker) and as student assistants (Simon Bruns, Sebastian Köster, David Gläsner, Sharief Füglein, Nina Kohl, and Marten Beiermann). We acknowledge the Lower Saxon Wadden Sea National Park Authority (Wilhelmshaven) for their permits and cooperation.
Conflict of interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Publisher’s note
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.
Supplementary material
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fmars.2023.1245926/full#supplementary-material
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Summary
Keywords
Magallana gigas, remote sensing, monitoring, classification, random forest, error propagation, ecosystem engineer
Citation
Hoffmann TK, Pfennings K, Hitzegrad J, Brohmann L, Welzel M, Paul M, Goseberg N, Wehrmann A and Schlurmann T (2023) Low-cost UAV monitoring: insights into seasonal volumetric changes of an oyster reef in the German Wadden Sea. Front. Mar. Sci. 10:1245926. doi: 10.3389/fmars.2023.1245926
Received
24 June 2023
Accepted
11 September 2023
Published
09 October 2023
Volume
10 - 2023
Edited by
Paolo Ciavola, University of Ferrara, Italy
Reviewed by
Daniele Ventura, Sapienza University of Rome, Italy; Riccardo Brunetta, University of Ferrara, Italy
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Copyright
© 2023 Hoffmann, Pfennings, Hitzegrad, Brohmann, Welzel, Paul, Goseberg, Wehrmann and Schlurmann.
This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Tom K. Hoffmann, hoffmann@lufi.uni-hannover.de
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