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        <gco:CharacterString>This dataset contains predicted seahorse habitat distributions for two species 
(*Hippocampus hippocampus *and *H. guttulatus*) and the genus combined 
(Hippocampus hippocampus MAXENT.asc, Hippocampus guttulatus MAXENT.asc and 
Hippocampus sp. MAXENT.asc, respectively). Rasters of the raw predicted 
habitat suitability outputs from the maximum entropy (MAXENT, Phillips et al. 
2006) species distribution model algorithm are provided.

Additionally, the environmental data layers used for modelling the species 
distributions, including distance to seagrass habitat (degrees) (Distance From 
Seagrass.tif), distance to the coastline (DistCo.tif) bathymetry (m) 
(Bathy.tif), minimum winter and maximum summer SST (oC) (Kriging SST Winter 
Min.tif and Kriging SST Summer Max.tif, respectively) and chlorophyll a 
concentration (mg m-3) (Kriging Chla Winter Smooth.tif and Kriging Chla 
Summer Smooth.tif, respectively).

All raster layers are provided in the WGS84 (EPSG::4326) spatial reference 
system. All raster layers are at a gridded resolution of 0.0042 x 0.0042 
degrees. All layers cover the shelf seas surrounding the UK, including the 
English Channel, Celtic Seas, North Sea, and extend into shallow coastal and 
estuarine habitats. Raster layers of predicted habitat suitability are 
provided in ASCI format, whilst environmental predictor layers are provided in 
TIFF format.

This work was prioritised by Natural England’s Seahorse Working Group and was 
funded by Natural England. This work was conducted in collaboration with The 
Seahorse Trust who provided the seahorse occurrence data. The information 
provided will support marine spatial planning to reduce the broader impact of 
anthropogenic activities and enable better decision-making to protect these 
sensitive species and their habitats.</gco:CharacterString>
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(`https://www.nationalarchives.gov.uk/doc/open-government-licence/version/3/)`_ 
– Please note the applicable attribution statement is as follows: ‘includes 
Crown copyright, Cefas [2023]'**


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            <gco:CharacterString>The outputs from this project advance the development of habitat suitability 
models for two sensitive species of seahorse (*H. hippocampus *and *H. 
guttulatus*) and the combined genus by predicting habitat suitability in 
shallow, nearshore waters (coastal and estuarine sites) that are considered 
important for both species.

Satellite-derived gridded environmental data (sea surface temperature (SST, 
oC) and chlorophyll a concentration (mg m-3), MODIS-Aqua Level-3 
`https://oceancolor.gsfc.nasa.gov/l3/`_[Downloaded May 2018]) were combined 
with Environment Agency field-collected water quality data from the Water 
Quality Archive [Accessed January 2023] using ordinary kriging within the 
"Geostatistical Analyst" tool in ArcGIS (v10.5) to produce environmental 
predictor layers that extend to the coastline. SST and chlorophyll summer 
maximums and winter minimums were calculated. Bathymetry (m) (GEBCO 
Compilation Group (2022) GEBCO 2022 Grid (doi:10.5285/e0f0bb80- 
ab44-2739-e053-6c86abc0289c) [Downloaded January 2023]) was interpolated using 
the natural neighbour method in ArcGIS 10.5. Seagrass (*Zostera marina*, 
*Z. noltii* and *Ruppia *spp.) polygon and point data were collated from 
multiple sources including Natural England’s open-access polygon data on 
seagrass coverage in England. Point data (from the year 2000 onwards) were 
converted to polygons using a buffer and merged with existing polygons to 
ensure all possible seagrass locations were included in the calculation. A 
distance from seagrass meadows layer was calculated in ArcGIS 10.5 using the 
“Euclidean Distance” tool. The final list of six variables was used for 
modelling, all with a Pearson's correlation of less than 0.7, including 
distance to seagrass habitat, bathymetry, minimum winter and maximum summer 
SST and chlorophyll concentrations which are provided.

Seahorse records provided by the Seahorse Trust's National Seahorse Database, 
Natural England and other records from open-source databases described in 
Bluemel et al. 2020 were filtered to remove records that were duplicated or 
fell outside of the study area, or where species identification was dubious. 
Prior to modelling, records were reduced to one point per grid cell to reduce 
sampling biases. All occurrence records from the year 2000 to the present were 
used for model training and testing (*Hippocampus *genus n = 165, 
*Hippocampus hippocampus *= 144, *Hippocampus guttulatus *= 45) (species 
occurrences are not provided). 

Seahorse habitat distributions were modelled using maximum entropy (MAXENT, 
Phillips et al. 2006). The default convergence threshold of 106 and a 
maximum of 5,000 iterations were used. A maximum of 10,000 background samples 
were randomly selected from the study area. Within the model settings, random 
seed was selected, which means that a different random subset was used for 
each model run, and the number of replicates was set to 10 and the replicate 
run type was set to subsample. In addition, the random test percentage was set 
to 25%. All other MAXENT settings were default.

Model performance was examined by computing the Area Under the Curve (AUC) 
score of the receiver operating characteristic (ROC) curve for each species, 
which is calculated by the MAXENT software as part of the modelling process 
and is considered to be an effective measure of model performance (Reiss et 
al. 2011). The AUC represents the relationship between sensitivity and 
specificity and varies between 0 and 1 (Reiss et al., 2011). It is generally 
considered that values &gt;0.9 indicate an excellent model fit, values between 
0.7 and 0.9 represent a good fit, anything &lt;0.7 represents a poor model fit 
and anything &lt;0.5 represents nothing better than random (Reiss et al., 2011). 
The model for *H. guttulatus* had the highest AUC, which had a mean value 
of 0.99 ± 0.0005 across the 10 runs, followed by *H. hippocampus* (0.97 ± 
0.0008) and the genus combined (0.97 ± 0.0006). 

Modelling limitations and sampling biases:

Known sampling biases exist in the seahorse occurrence data used in this study 
and therefore the predictions should be taken with caution. Biases include 
higher sampling effort in the southwest and English Channel, due to high 
recreational SCUBA diving intensity in this area, resulting in uneven sampling 
distributions across the study area. Recorder biases exist due to known 
locations of seahorses being regularly visited and targeted by dive companies, 
and records are often provided by multiple diver trips of the same seahorse 
that has moved to a nearby locality (Garrick-Maidment pers. comm.). MAXENT is 
sensitive to biased data distributions, and there is a strong assumption of 
random sampling distribution for presences when random background data are 
used. Thus the underlying model assumptions are invalidated by the sampling 
bias, which could be restricting predictions to southerly localities where 
sampling biases (increased effort and recorder bias) are known to exist. 

To improve confidence in the habitat predictions for seahorses, further work 
should include testing different modelling algorithms. Variations in model 
predictions were observed by Bluemel et al. (2020), highlighting the need to 
compare outputs and identify consistent robust predictions, especially in 
data-limited, presence-only settings. Additionally, testing different habitat 
suitability thresholds that are tailored to the intended use of the predicted 
distributions (Freeman and Moisen 2008), testing validation approaches more 
suited to presence-only situations, because using AUC in isolation can be 
uninformative when species prevalence is low. Furthermore, sampling bias and 
spatial autocorrelation in the occurrence data require further work. Methods 
to deal with spatial autocorrelation and sampling biases could include 
defining a bias grid to include in the modelling process (i.e., known dive 
sites with higher sampling activity, thus a higher likelihood of observation), 
using other suitable point process models that account for these biases or 
assessing model performance with a block cross-validation approach (Roberts et 
al. 2017)).

Bluemel, J.K., Lynam, C. and Ellis, J. (2020). Fish biodiversity: state and 
pressure indicators. Cefas Project Report for Defra, 67 pp.

Freeman, E.A. and Moisen, G.G. (2008). A comparison of the performance of 
threshold criteria for binary classification in terms of predicted prevalence 
and kappa. Ecological modelling, 217(1-2): 48-58.

Phillips, S.J., Anderson, R.P. and Schapire, R.E., (2006). Maximum entropy 
modelling of species geographic distributions. *Ecological modelling*, 
*190*(3-4): 231-259.

Reiss, H., Cunze, S., König, K., Neumann, H. and Kröncke, I., (2011). Species 
distribution modelling of marine benthos: a North Sea case study. *Marine 
Ecology Progress Series*, *442*: 71-86.

Roberts, D.R., Bahn, V., Ciuti, S., Boyce, M.S., Elith, J., Guillera-Arroita, 
G., Hauenstein, S., Lahoz-Monfort, J.J., Schröder, B., Thuiller, W., others. 
(2017). Cross-validation strategies for data with temporal, spatial, 
hierarchical, or phylogenetic structure. Ecography. 40: 913-929.


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