# Mismatches between the current marine Natura 2000 network and seabird distributions call for enhanced protected areas off metropolitan France

Timothée Poupart, Thibaut de Bettignies, Matthieu Authier, Nicola Baccetti, Luca Börger, Iker Castège, Jacopo G Cecere, Nicolas Courbin, Jamie Darby, Karine Delord, et al.

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Timothée Poupart, Thibaut de Bettignies, Matthieu Authier, Nicola Baccetti, Luca Börger, et al. Mismatches between the current marine Natura 2000 network and seabird distributions call for enhanced protected areas off metropolitan France. Marine Policy, 2025, 180 (106779), 10.1016/j.marpol.2025.106779. hal-05104529

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# Mismatches between the current marine Natura 2000 network and seabird distributions call for enhanced protected areas off metropolitan France

Timothée Poupart<sup>a,1,2</sup>, Thibaut de Bettignies<sup>a,b,*1,3</sup> <sup>●</sup>, Matthieu Authier<sup>c,4</sup>, Nicola Baccetti<sup>d,5</sup>, Luca Börger<sup>e,6</sup>, Iker Castège<sup>f</sup>, Jacopo G. Cecere<sup>d,7</sup>, Nicolas Courbin<sup>g,8</sup>, Jamie Darby<sup>h,9</sup>, Karine Delord<sup>i,10</sup>, Ghislain Doremus<sup>c,11</sup>, Hayley A. Douglas<sup>j,12</sup>, Gilles Faggio<sup>k,13</sup>, Giorgia Gaibani<sup>l</sup>, Fabrice Gallien<sup>m</sup>, Cécile Gicquel<sup>n</sup>, Jacob González-Solís<sup>o,p,14</sup>, David Grémillet<sup>g,q,15</sup>, Simona Imperio<sup>d,16</sup>, Jude V. Lane<sup>r,17</sup>, Amélie Lescroël<sup>s,18</sup>, Maite Louzao<sup>t,19</sup>, Jeanne de Mazières<sup>a</sup>, Noëmie Michez<sup>u,20</sup>, Emilie Milon<sup>f,21</sup>, Ellie Owen<sup>v,22</sup>, Vitor H. Paiva<sup>w,23</sup>, Federico De Pascalis<sup>d,24</sup>, Clara Péron<sup>x,25</sup>, Francesco Pezzo<sup>d,26</sup>, Pascal Provost<sup>y</sup>, Raül Ramos<sup>o,p,27</sup>, Jaime A. Ramos<sup>ab,28</sup>, Solène Robert<sup>a</sup>, Diego Rubolini<sup>z</sup>, Olivier Scher<sup>aa</sup>, Lorenzo Serra<sup>d</sup>

<footnote><sup>a</sup> PatriNat (OFB-MNHN), Muséum national d'Histoire naturelle, Maison Buffon, 36 rue Geoffroy-Saint-Hilaire, Paris 75005, France</footnote>

<footnote><sup>b</sup> Centre de Recherche sur la Biodiversité et l'Environnement (CNRS, Université de Toulouse, INPT) Avenue de l'Agrobiopole, Castanet-Tolosan 31320, France</footnote>

<footnote><sup>c</sup> UAR 3462 CNRS-La Rochelle Université, Observatoire Pelagis, 5 allée de l'Océan, La Rochelle 17000, France</footnote>

<footnote><sup>d</sup> Area Avifauna Migratrice, Istituto Superiore per la Protezione e la Ricerca Ambientale (ISPRA), via Ca' Fornacetta, Ozzano dell'Emilia 9 - 40064, Italy</footnote>

<footnote><sup>e</sup> Department of Biosciences, Swansea University, Swansea, Wales, UK</footnote>

<footnote><sup>f</sup> Centre de la Mer de Biarritz, Etablissement des bains, Boulevard du Prince de Galles, Biarritz 64200, France</footnote>

<footnote><sup>g</sup> CEFE, Université de Montpellier, CNRS, EPHE, IRD, Montpellier, France</footnote>

<footnote><sup>h</sup> School of Biological, Environmental and Earth Sciences, University College Cork, Cork T23 N73K, Ireland</footnote>

<footnote><sup>i</sup> Centre d'Etudes Biologiques de Chizé, UMR 7372 du CNRS, La Rochelle Université, Villiers-en-Bois 79360, France</footnote>

<footnote><sup>j</sup> Clyde Muirshiel Regional Park, Castle Semple Centre Lochlip Road, Lochwinnoch PA12 4EA, UK</footnote>

<footnote><sup>k</sup> Office de l'Environnement de la Corse, 14 avenue Jean Nicoli, Corte 20250, France</footnote>

<footnote><sup>l</sup> Conservation Department, LIPU-BirdLife Italy, Parma, Italy</footnote>

<footnote><sup>m</sup> Groupe Ornithologique Normand (GONm), 181 rue d'Auge, Caen 14000, France</footnote>

<footnote><sup>n</sup> Office Français de la Biodiversité, Parc naturel marin d'Iroise, pointe des renards, Le Conquet 29217, France</footnote>

<footnote><sup>o</sup> Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals (BEECA), Facultat de Biologia, Universitat de Barcelona, Barcelona, Spain</footnote>

<footnote><sup>p</sup> Institut de Recherche de la Biodiversitat (IRBio), Universitat de Barcelona, Barcelona, Spain</footnote>

<footnote><sup>q</sup> FitzPatrick Institute of African Ornithology, Department of Biological Sciences, University of Cape Town, South Africa</footnote>

<footnote><sup>r</sup> RSPB Centre for Conservation Science, Sandy, Bedfordshire, UK</footnote>

<footnote><sup>s</sup> Point Blue Conservation Science, 3820 Cypress Drive, Suite 11, Petaluma, CA 94954, USA</footnote>

<footnote><sup>t</sup> AZTI Marine Research, Basque Research and Technology Alliance (BRTA), Pasaia, Spain</footnote>

<footnote><sup>u</sup> Parc naturel marin du golfe du Lion, Office français de la biodiversité, 2 impasse de Charlemagne, Argelès-sur-Mer 66700, France</footnote>

<footnote><sup>v</sup> The National Trust for Scotland, Balnain House, 40 Huntly Street, Inverness IV3 5HR, UK</footnote>

<footnote><sup>w</sup> University of Coimbra, CFE - Centre for Functional Ecology / TERRA - Science for People & the Planet, Department of Life Sciences, Calçada Martim de Freitas, Coimbra 3000-456, Portugal</footnote>

<footnote><sup>x</sup> Laboratoire de Biologie des Organismes et Ecosystèmes Aquatiques (BOREA), UMR 8067 - MNHN, CNRS, IRD, SU, UCN, UA, Paris 75005, France</footnote>

<footnote><sup>y</sup> Ligue pour la Protection des Oiseaux, réserve naturelle nationale des Sept-Iles, Station LPO Ile-Grande, Pleumeur-Bodou 22560, France</footnote>

<footnote>* Corresponding author at: PatriNat (OFB-MNHN-CNRS-IRD), Muséum national d'Histoire naturelle, Maison Buffon, 36 rue Geoffroy-Saint-Hilaire, Paris 75005, France</footnote>

<footnote>E-mail addresses: timothee.poupart@gmail.com (T. Poupart), thibaut.de-bettignies@mnhn.fr (T. de Bettignies), matthieu.authier@univ-lr.fr (M. Authier), nicola.baccetti@isprambiente.it (N. Baccetti), l.borger@swansea.ac.uk (L. Börger), iker.castege@centredelamer.fr (I. Castège), iacopo.cecere@isprambiente.it (J.G. Cecere), ncourbin@gmail.com (N. Courbin), jamie.darby@ucc.ie (J. Darby), karine.delord@cebc.cnrs.fr (K. Delord), ghislain.doremus@univ-lr.fr (G. Doremus), h.a.douglas@hotmail.co.uk (H.A. Douglas), gilles.faggio@oec.fr (G. Faggio), giorgia.gaibani@lipu.it (G. Gaibani), fabrice.gallien@gonm.org (F. Gallien), cecile.gicquel@ofb.gouv.fr (C. Gicquel), jgsolis@ub.edu (J. González-Solís), david.gremillet@cefe.cnrs.fr (D. Grémillet), simona.imperio@isprambiente.it (S. Imperio), jude.lane@rspb.org.uk (J.V. Lane), alescroel-ra@pointblue.org (A. Lescroël), mlouzao@azti.es (M. Louzao), jeanne.de-mazieres@mnhn.fr (J. de Mazières), noemie.michez@ofb.gouv.fr (N. Michez), emilie.milon@centredelamer.fr (E. Milon), eowen@nts.org.uk (E. Owen), vitorpaiva@uc.pt (V.H. Paiva), federico.depascalis@isprambiente.it (F. De Pascalis), clara.peron@mnhn.fr (C. Péron), francesco.pezzo@isprambiente.it (F. Pezzo), ramos@ub.edu (R. Ramos), jramos@uc.pt (J.A. Ramos), solene.robert@mnhn.fr (S. Robert), diego.rubolini@unimi.it (D. Rubolini), olivier.scher@cen-occitanie.org (O. Scher), lorenzo.serra@isprambiente.it (L. Serra).</footnote>
ARTICLE INFO

Keywords:

Seabirds

Natura 2000

Special Protection Areas

Marine Important Bird Areas

Network assessment

ABSTRACT

Seabirds are among the most threatened vertebrates, under pressure from fisheries bycatch, climate change, overfishing, and human disturbance. In France, demographic studies have highlighted adult survival as a key factor in population trends, which calls for large-scale marine conservation efforts. In this context, the Natura 2000 policy requires the designation of Special Protection Areas (SPAs) to protect seabirds under the Birds Directive. To assess the completeness of the French marine SPA network, data from aerial, boat, and coastal surveys, as well as tracking devices and distribution models, were collected for 57 seabird taxa. This data collection allowed the EU minimum criteria for a coherent SPA network to be spatially implemented, and the most ecologically valuable areas for seabirds around metropolitan France to be identified and prioritised, and overlaid with the current French SPA network and Marine Important Bird Areas (mIBAs) to identify potential inconsistencies. This analysis revealed seabird hotspots outside the existing ecological network, confirming some insufficiencies for coherent seabird conservation. Although data dependent, this analysis highlighted the limitations of using global proportion coverage to assess network coherence when coverage of biodiversity and abundance hotspots was not achieved. Furthermore, these results summarised the main target areas for policy makers to effectively improve seabird conservation around metropolitan France. In a context of increasing demands for marine spatial planning, improvements in this knowledge, the SPA network and conservation actions are required.

1. Introduction

In the context of the continuing human population growth and its demands for natural resources and energy, which reduce the natural areas and their connectivity, conservation biology has emphasized the need to link conservation with land-use planning and economic development [1]. For global effectiveness, species habitat use should be considered not only within protected areas, but also beyond their boundaries to avoid compromising conservation efforts made elsewhere [2–4]. A key operational tool is the development of ecological networks, in which core areas surrounded by buffer zones, and connected by corridors across a heterogeneous landscape, should form a coherent ecosystem that supports more biodiversity than core areas in isolation

[5].

To be functional and achieve its objectives, the spatial configuration of such a network is crucial and requires a systematic approach across landscapes [6]. In Europe, this goal of long-term sustainability for wildlife and natural habitats has been translated into policy by the European Union (EU): the designation of Special Protection Areas (SPA) and Special Areas of Conservation (SAC) under the Birds (Council Directive 79/409/EEC) and Habitats (Council Directive 92/43/EEC) directives respectively, form the Natura 2000 network. This network covers 27 EU Member States and is considered to be the largest integrated ecological network in the world [7].

Initiated on land, the Natura 2000 network has been extended to the marine environment [8], which is also affected by global human threats [9]. At sea, SPA perimeters are dedicated to seabirds, one of the most threatened groups of vertebrates, with declines across taxonomic groups mainly due to the cumulation of threats such as incidental bycatch in fisheries, climate change, overfishing and disturbance [10]. Around France, demographic analysis points to adult survival as a key parameter for populations trajectories, requiring at-sea and international conservation action [11,12]. Such a large-scale effort is not without its challenges, and national approaches vary across Europe [13].

However, scientific criteria for sites designation have been established in order to ensure effectiveness of the network [8,14]. Completeness of the network, also referred to as network sufficiency, is achieved when all habitat types and species listed in the Birds and Habitats Directives in a country, are adequately represented and protected within the national network. Although considered challenging to achieve [15,16], it is a legal obligation for all EU Member States to have a national network in place and managed in accordance with these European criteria. In this context, the achievement of network completeness is being monitored by the European Commission (EC). The Birds Directive requires a scientific procedure, independent of socio-economic considerations, to designate "the most suitable territories, in number and size, as special protection areas" (Art. 4). This non-specific criterion has so far led the EC and the EU Court of Justice to assess the adequacy of SPA designations against the non-legally binding NGO inventory of Important Bird and Biodiversity Areas, and at sea, the marine Important Bird Areas (mIBA) [17].

Although the marine waters covered by SPAs in metropolitan France exceed the 30 % target set by the EU Biodiversity Strategy 2030 [18] and the UN Convention on Biological Diversity [19], the coherence of this

<footnote>z Dipartimento di Scienze e Politiche Ambientali, Università degli Studi di Milano, via Celoria 26, Milano I-20133, Italy</footnote>

<footnote>aa Conservatoire d'espaces naturels d'Occitanie, 26 allée de Mycènes, Montpellier 34000, France</footnote>

<footnote>ab University of Coimbra, MARE - Marine and Environmental Sciences Centre / ARNET - Aquatic Research Network, Department of Life Sciences, Calçada Martim de Freitas, Coimbra 3000-456, Portugal</footnote>

<footnote n="1">¹ These authors contributed equally to this work and share first authorship</footnote>

<footnote n="2">² https://orcid.org/0000-0003-4120-933X</footnote>

<footnote n="3">³ https://orcid.org/0000-0003-0565-1200</footnote>

<footnote n="4">⁴ https://orcid.org/0000-0001-7394-1993</footnote>

<footnote n="5">⁵ https://orcid.org/0000-0001-6579-6060</footnote>

<footnote n="6">⁶ https://orcid.org/0000-0001-8763-5997</footnote>

<footnote n="7">⁷ https://orcid.org/0000-0002-4925-2730</footnote>

<footnote n="8">⁸ https://orcid.org/0000-0002-6428-5012</footnote>

<footnote n="9">⁹ https://orcid.org/0000-0002-9757-7150</footnote>

<footnote n="10">¹⁰ https://orcid.org/0000-0001-6720-951X</footnote>

<footnote n="11">¹¹ https://orcid.org/0000-0001-5873-2851</footnote>

<footnote n="12">¹² https://orcid.org/0009-0004-3998-8789</footnote>

<footnote n="13">¹³ https://orcid.org/0000-0001-6518-9666</footnote>

<footnote n="14">¹⁴ https://orcid.org/0000-0002-8691-9397</footnote>

<footnote n="15">¹⁵ https://orcid.org/0000-0002-7711-9398</footnote>

<footnote n="16">¹⁶ https://orcid.org/0000-0002-9982-205X</footnote>

<footnote n="17">¹⁷ https://orcid.org/0000-0002-5841-6346</footnote>

<footnote n="18">¹⁸ https://orcid.org/0000-0002-9030-6593</footnote>

<footnote n="19">¹⁹ https://orcid.org/0000-0002-9997-5144</footnote>

<footnote n="20">²⁰ https://orcid.org/0000-0003-2298-6723</footnote>

<footnote n="21">²¹ https://orcid.org/0000-0001-7871-6277</footnote>

<footnote n="22">²² https://orcid.org/0000-0003-2073-2420</footnote>

<footnote n="23">²³ https://orcid.org/0000-0001-6368-9579</footnote>

<footnote n="24">²⁴ https://orcid.org/0000-0002-1414-7770</footnote>

<footnote n="25">²⁵ https://orcid.org/0000-0002-6529-4578</footnote>

<footnote n="26">²⁶ https://orcid.org/0000-0003-4437-3805</footnote>

<footnote n="27">²⁷ https://orcid.org/0000-0002-0551-8605</footnote>

<footnote n="28">²⁸ https://orcid.org/0000-0002-9533-987X</footnote>
national network is being questioned [20]. In particular, the EC has raised questions about the completion of the network in coastal waters (compared to mIBAs) where anthropogenic pressure is high [21]. Therefore, in the present study, a spatial implementation of the EU minimum criteria for a coherent SPA network was carried out, to 1) locate and prioritise the most ecologically valuable areas for seabirds around metropolitan France, and 2) overlay them with the current French SPA network and marine Important Bird Areas (mIBA) to identify potential inconsistencies.

## 2. Materials and methods

### 2.1. Biogeographical areas, seabird species and time-periods

The Birds Directive defines the biogeographical regions for its application in Europe. In metropolitan France, this concerns the Atlantic and Mediterranean marine environments, delimited by the boundaries of waters under French jurisdiction [22]. For the analysis, the marine environment polygons were gridded into 0.05 × 0.05° cells, for consistency with the available distribution models [23–25]. The Birds Directive also lists the bird species to be considered in SPAs, which were previously refined with seabirds regularly occurring in metropolitan France [26]. Rare vagrants or species associated with coastal lagoons

were excluded from this list, resulting in 53 taxa for the Atlantic and 27 taxa for the Mediterranean (57 taxa in total including subspecies, Table A.1). The annual life cycle of seabirds can be divided into two main periods, which are associated with drastic changes in their spatial distribution: 1) the breeding period, when breeding individuals become central-place foragers during the pre-nuptial, incubation and chick-rearing stages, and 2) the non-breeding period, when individuals may disperse away from breeding colonies and/or undertake migrations to wintering areas. This temporal distinction was maintained for the distribution data, either by using the information provided by the data producers, or, when at-sea sightings did not provide information on reproductive status, by combining the Julian date with the reported breeding phenology of the species in metropolitan France [27].

### 2.2. Data compilation

Spatial conservation planning requires high quality information on abundance and distribution [8]. In order to include all relevant and available data, we searched for all at-sea seabird data available in 2022 among predicted distribution, tracking, density and sighting data, produced and/or held by various NGOs, universities and national institutions (Table 1, Fig. 1). Tracking studies were searched in the online platforms Seabird Tracking Database [28] and MoveBank [29],

<table><caption>**Table 1**
Types of data and datasets used to map the distribution of seabirds in French waters.</caption><thead><tr><th>Data</th><th>Platform</th><th>Acquisition protocol</th><th>Data format</th><th>Pro(s)</th><th>Con(s)</th><th>Datasets</th></tr></thead><tbody><tr><td rowspan="2">Predicted distribution</td><td>Plane</td><td>Aerial transects</td><td>Density predicted by habitat modelling</td><td>Basin-scale sampling covering all the study areas, enabling large-scale habitat modelling</td><td>Decadal campaigns, species identification often at taxonomic group level, low detectability of elusive species. Modelling requires sufficient sightings for robust results</td><td>SAMM 1, SAMM 2, SCANS III, ASI</td></tr><tr><td>Multi-source</td><td>Boat transects + coastal repeated counts</td><td>Density predicted by habitat modelling</td><td>Regional scale sampling enables habitat modelling</td><td>In the Mediterranean, insufficient sampling leads to limited spatial predictions</td><td>Balearic shearwater National Action Plan 2018</td></tr><tr><td rowspan="2">Interpolated density</td><td>Boat</td><td>Boat transects</td><td>Densities predicted by spatial interpolation</td><td>Annual, seasonal campaigns<br/>Large scale sampling</td><td>Timed for fish studies. Coastal waters under sampled, abyssal waters not sampled.</td><td>MEGASCOPE, PELMED (PELAGIS Observatory)</td></tr><tr><td></td><td></td><td></td><td>Annual, monthly campaigns</td><td>Regional-scale sampling</td><td>ERMMA (Biarritz sea centre)</td></tr><tr><td rowspan="3">Tracking</td><td>Bird</td><td>GPS, Argos</td><td>Bird locations</td><td>Fine-scale recording of trips at sea, from days to months (depending on material and settings). Ability of infer behaviour. Freedom from observer bias</td><td>Biologging impact [83]. Large sample size required for representativeness [84]. Repeated sampling over time and space for robust identification of priority areas [85,86]</td><td>Detail of the numerous studies in Annex II</td></tr><tr><td>Plane</td><td>Transects</td><td></td><td>Annual, seasonal campaigns, sampling of coastal and neritic waters</td><td>Local/Regional scale</td><td>SPEE, survols OFB (Bretagne Nord, PNMI), DepoBio</td></tr><tr><td>Boat</td><td>Transects, repeated counts, unprotocoled sightings</td><td></td><td>Annual, seasonal campaigns, sampling of coastal and neritic waters</td><td>Local/Regional scale</td><td>PelMed (CEFE), ObsEnMer, DepoBio, RN estuaire de Seine</td></tr><tr><td rowspan="3">Sightings</td><td></td><td></td><td>Bird numbers</td><td>Wide coverage of the coast</td><td>Spatio-temporal variation of sampling effort, coverage limited to 3 nautical miles</td><td>OFB grebes and loons survey, Trektellen</td></tr><tr><td></td><td>Repeated point-counts, unprotocoled sightings</td><td></td><td>Knowledge of bird's origin and age</td><td>Spatio-temporal variation of sampling effort, close observation only</td><td>CRBPO</td></tr><tr><td>Coastal</td><td></td><td></td><td>Covers the whole coast, species-level sightings, most reports of elusive species come from here</td><td>Spatio-temporal variation of sampling effort, detection distance limited to 3 nautical miles</td><td>Faune-France, SINP</td></tr><tr><td></td><td></td><td>Sectoral repeated counts</td><td></td><td>Coordinated counts within assigned sectors, annual</td><td>Detection distance limited to 3 nautical miles</td><td>International waterbird census (Wetlands International)</td></tr></tbody></table>
<figure><img src="image_3.png" /><figcaption>**Fig. 1.** Spatio-temporal distribution of seabird data and observation efforts (nautical in blue, aerial in green, tracking in grey and sightings in brown) collected in the waters under French jurisdiction in the Atlantic and Mediterranean during the period 2002–2022.</figcaption></figure>

restricted to GPS or Argos tags for spatial accuracy. This included not only the studies conducted in metropolitan France, but also studies conducted abroad with seabirds using French waters (Table A.2). Only data collected during the last two decades (2002–2022) were considered, given the possible temporal evolution of population sizes and their areas of use (Table 1, Fig. 1). This choice was a compromise between obtaining a large amount of data, maximising the number of repeated observations over time (necessary for a robust selection of priority areas for mobile species, [30,31]), while remaining sufficiently recent to be representative of the current situation.

## 2.3. Data preparation

All data were imported into R version 4.1.3 [32], georeferenced to the WGS 84 geodetic system using the *sf* package [33], and plotted using the *ggplot2* package [34].

### 2.3.1. Filtering

The compilation of the various datasets required an initial filtering process, as the quality of the data may have an impact on the results. In particular, the following data were excluded: observations of stranded individuals following exceptional weather conditions [35] and oiled individuals, whose locations may reflect exhaustion rather than natural occurrence. Data from fisheries bycatch and translocation experiments were excluded as they are likely to reflect attraction to vessels and unnatural trip characteristics, respectively. Counts of breeding colonies or roosting places on land were also excluded because they do not provide accurate information on areas used at sea, as seabirds can travel tens to hundreds of kilometres per day [36–38]. Furthermore, the terrestrial colonies have already guided the first SPA designations in France,

whereas seabird conservation today requires more knowledge at sea [12,39].

### 2.3.2. Species distribution models

Given the species-based approach recommended for Natura 2000, the density at-sea predicted by habitat models available in metropolitan French waters was used (Table 1). When available at a taxonomic group level, they were attributed to the most abundant species in the group (Table A.3). In the Mediterranean, the predicted densities of shearwaters were judged to be biased towards the coast [24], which contradicts the repeated tracking data studies [40–42]. Therefore, these were excluded in favour of the tracking data.

#### 2.3.3. Tracking data

This heterogeneous collection of tracking data included different devices, settings, and sample sizes, so these raw data required preprocessing steps. For each species, tracking data were divided into foraging trips [43]. Anomalous locations with unrealistic speeds above 90 km/h were removed [44] prior to linear interpolation at one-minute intervals (common denominator). From these trips, the time spent within the gridded regions was extracted and transformed as a percentage of the total trip duration, averaged across individuals. This standardised time per area weighted the importance of the cells used, as areas where birds spent more time at sea are indicative of foraging behaviour [45,46]. The result was a distribution layer by species and period, including both French and foreign tracking information.

#### 2.3.4. Interpolated density

Bird density layers from at-sea surveys were either received directly as usable layers (ERMMA dataset, Table A.2), or obtained from raw data
by kriging (SAMM, SCANS dataset, Table A.2) assuming a Poisson distribution and zero nugget model [47] implemented in the Pelakrig package [48]. Other studies that did not provide data on prospecting effort data were treated as sightings.

### 2.3.5. Sightings

Sightings were derived from a variety of counting methods and mostly from citizen reports of opportunistic observations. They are heterogeneous and sensitive to observation effort [49], so they cannot be analyzed using the statistical techniques described above. However, the numbers reported all along the coasts can still provide information on some locations with high concentrations of species, and sometimes represent the majority of the available data (Table A.5, A.6). Therefore, they were integrated as another distribution layer by species and time period. To smooth out their spatio-temporal sampling bias, the sightings were aggregated within the cells of the gridded regions as the mean of the annual maxima. Such a transformation made it possible to map areas of regular and important concentrations for each species by season.

## 2.4. Data processing

### 2.4.1. Collation

The first step was to create a spatial distribution layer for each available data type, species and life cycle period, forming the basis for the network evaluation (Fig. 2). Where a layer was derived from repeated data within cells, they were summarized by keeping the average of the maximum per year. The resulting distribution layers, with different quantitative scales depending on the data type, were transformed as follows:

$$ \text{standardised data (from 0 to 1)} = \frac{(\text{data} - \text{minimum})}{(\text{maximum} - \text{minimum})} $$

This standardization of the layers into a common relative scale allowed the different distribution gradients to be used simultaneously with equal weight for a comprehensive analysis of species distribution (Fig. 2).

### 2.4.2. Process for identifying priority areas

Official European texts require a scientific procedure to designate "the most suitable territories, in number and size, as Special Protection Areas" (Birds Directive, Art. 4). Finding these most important areas, where management efforts would conserve biodiversity in the most cost-effective way, is the goal of systematic conservation planning [6], and is based on distribution data and predefined conservation objectives. In the case of the Natura 2000 network for highly mobile and migratory marine species such as seabirds, these objectives are translated into specific criteria published in European texts for the establishment and evaluation of Natura 2000.

#### 2.4.2.1. European scientific criteria relevant to highly mobile species.
For a given species, the adequacy of a Natura 2000 network is assessed on the basis of by two main criteria: the natural range and the sufficiency. These two criteria are broken down into several sub-criteria. Natural range is assessed by the distribution variability and habitat/genetic variability. Sufficiency is assessed by the proportion, the conservation status and the proportionality (Table 2).

For seabird species in metropolitan French waters, the conservation status, population trend and proportionality sub-criteria have already been integrated into a Responsibility Index (RI) for each species [50], established in the French context of the Marine Strategy Framework Directive (Directive 2008/56/EC) (Table A.5, A.6). This recent index, available by marine sub-region, allowed a geographical refinement of the Atlantic region: Greater North Sea, Celtic Sea, Bay of Biscay and Iberian coast. Therefore, these categorical sub-criteria (natural range) and quantitative sub-criteria (network sufficiency) were applied to the compiled data in order to assess the potential presence of remaining gaps as defined by the EC recommendations.

#### 2.4.2.2. Identification and prioritization of the areas of concentration.
Several decision support tools that use an algorithmic approach to species distribution data have been developed to identify areas of high conservation value [51–53], and have been used in the context of national ecological network assessments [16,54,55]. The performance and results of these decision support tools are not necessarily consistent with

<figure><img src="image_4.png" /><figcaption>Fig. 2. Data processing for the evaluation of the SPA network for seabirds, applying the EU criteria.</figcaption></figure>
<table><caption>**Table 2**
European scientific criteria used to evaluate the Special Protection Areas network for seabirds.</caption><thead><tr><th>Criteria</th><th>Sub-criteria</th><th>Implementation principle</th><th>Reference</th></tr></thead><tbody><tr><td rowspan="3">Natural range</td><td>Distribution variability</td><td>Variability in spatio-temporal distribution</td><td>HD art 3.1<br>BD art 4,1–2<br>ETC/BD<br>2010</td></tr><tr><td>Habitat/Genetic variability</td><td>Natural variability of the habitat itself and species genetic variability (Sub-species and local population with specific ecology/behaviour)</td><td>EC 1997</td></tr><tr><td>Proportion<br>Conservation status</td><td>“Insufficient” when proportion of the species population is ≤ 20 % within the network<br>Higher priority if species evaluated with population in decline or threatened</td><td>ETC/BD<br>2010</td></tr><tr><td>Network<br>sufficiency</td><td>Proportionality</td><td>Proportion of the species population within network must be equivalent to European proportion</td><td>HD art 3.2<br>Evans 2012</td></tr></tbody></table>

one-another, and the choice of software depends on the objective of the analysis [56,57].

Given our objective to localize priority areas for comparison with the existing networks, we used the open-source package *Prioritizr* [53]. Within the 0.05 × 0.05° gridded marine environment, our spatial problem was to extract from the available standardized distribution layers the minimum number of cells necessary (*add_min_set_objective* and *add_binaryDecision* functions) to represent a network of areas meeting the minimum EC proportion criteria (20 %, set with the *add_relative_target* function). For each species, the algorithm identified the cells that fall within the optimal mathematical solution, representing the areas with the highest population concentrations. These selected cells are weighted with the corresponding Responsibility Index (RI) (as a reminder, depending on the species, the marine sub-region and the life cycle period [50]), or cumulative RI when multiple species were encountered in the same cell. By summing the weighted solutions, areas with the highest conservation values are distinguished (Fig. 2). In addition, all available standardized layers for a species were combined into a single averaged layer, to assess the global proportion of the species population covered by the current SPA network.

Within the biogeographical regions, the failure to cover the minimum proportion threshold and/or important(s) area(s) outside the existing SPA perimeters were used to assess the presence of remaining gap(s).

# 3. Results

## 3.1. Data collation

A total of 3 855 346 data observations were collected in the waters under French jurisdiction, 2 857 590 in the Atlantic and 997 756 in the Mediterranean (Fig. 1). In the Atlantic, these raw numbers were mainly derived from direct observations (54.4 %) and tracking data (45.6 %), while in the Mediterranean the opposite was true, with more data coming from tracking studies (71.3 %) and less from direct observations (28.7 %). Detailed regional data contributions are available by dataset (Table A.4) and by species (Table A.5, A.6).

## 3.2. Minimum network and priority areas

In the Atlantic, the algorithm identified 5 034 cells as areas of highest species concentration, covering 38.6 % of the region, in all oceanographic domains (Fig. 3). Within this minimum network, 2 566 cells were located within the existing SPA network, representing a general agreement of 50.9 %. With respect to the mIBAs, 1 313 cells were located within it, representing a general agreement of 26.0 %. Considering the yearly period, the highest cumulative RI values distinguished three main areas: (A1) the plume front of the Picardy estuaries (35.6), (A2) around the head of the submarine canyon off Capbreton (24.8), and (A3) on a tidal coastal front between the islands of Houat and Dumet (19.1) (Fig. 3). These areas corresponded to the non-breeding period hotspots where the cumulative Responsibility Index (RI) was found to be the highest (28.8, 24.8 and 16.1 respectively). Interestingly, areas A1

<figure><img src="image_5.png" /><figcaption>**Fig. 3.** Theoretical minimum network for 53 seabird taxa meeting the representativeness criteria in the Atlantic under French jurisdiction, coloured by the cumulative Responsibility Index (RI). The outer limits of waters under French jurisdiction and territorial waters are shown as dashed and dotted lines respectively. The existing French networks of SPAs and mIBAs are shown in blue and black, respectively. The grey lines represent the -50, -100, -200, -500 and -1000 m isobaths.</figcaption></figure>

and A3 were partially covered by existing SPA perimeters, while A2 was completely outside (Fig. 3). During the breeding season, the highest cumulative RI values were found on other areas: (A4) around the Septîles archipelago (22.9), (A5) Cape Fréhel (14.3) and (A6) the Loire estuary (12.3) (Fig. 3). On the other hand, these areas (A4–6) were located within existing SPA perimeters (Fig. 3) designated for nearby seabird colonies of national importance.

In the Mediterranean, the algorithm identified 1 275 cells as areas of highest species concentration, covering 25.9 % of the region in all oceanographic domains (Fig. 4). Within this minimum network, 199 cells were located within the existing SPA network, representing a general agreement of 15.6 %. For the mIBAs, 367 cells were located
within it, representing a general agreement of 28.7 %. Considering the yearly periods, the highest cumulative RI values distinguished six main areas: (M1) the eastern part of the Languedoc coast, off the Thau lagoon (28), (M2) offshore towards the centre of the Gulf of Lion (22.5), (M3) the Cape Béar (22), (M4) the Gulf of Marseille (20.7), (M5) the Rhône estuary / Gulf of Fos-sur-Mer (16.2) and (M6) the western part of the Languedoc coast (15.2) (Fig. 4). The highest cumulative RI values were found mainly during the breeding season (M1=17.7, M4=15.2, M3=14.7 and M2=13.2) and to a lesser extent during the non-breeding season (M6=11.7, M5=10.5). Of these seabird hotspots, only M3 was within a SPA. The rest were located either partially (M1) or completely (M4, M2, M5) outside the existing SPA network (Fig. 4), as well as outside mIBA boundaries when offshore (M2).

The spatial analysis of the seabird species distributions in regard to the current protection network also revealed that the proportion of the species population with the SPA network was above the critical threshold for all species in the Atlantic, but below that threshold in the Mediterranean for 6 coastal affinity species and 3 pelagic affinity species (Fig. 5).

# 4. Discussion

## 4.1. Contribution of the study

This study integrated available and relevant datasets on seabird at sea data to provide a comprehensive synthesis of the most important areas for seabirds in the Atlantic and Mediterranean waters under metropolitan French jurisdiction. This data-driven work allowed a scientific assessment of the completeness of the French marine Natura 2000 network for seabirds and showed contrasted results between the Atlantic and the Mediterranean regions.

<figure><img src="image_6.png" /><figcaption>**Fig. 4.** Theoretical minimum network for 27 seabird taxa meeting the representativeness criteria in the Mediterranean under French jurisdiction, coloured by the cumulative Responsibility Index (RI). The outer limits of waters under French jurisdiction and territorial waters are shown as dashed and dotted lines respectively. The existing French networks of SPAs and mIBAs are shown in blue and black, respectively. The grey lines represent the -50, -100, -200, -500 and -1000 m isobaths.</figcaption></figure>

### 4.1.1. Priority areas and gaps identified

In the Atlantic, the SPA network met the minimum population proportion and, unlike the mIBA network, covered most of the seabird concentration areas. However, some priority areas remained outside the SPA network, in coastal and territorial waters. In the Mediterranean, the SPA network failed to meet the minimum population proportion for a third of the species, as it covered only a minority of the seabird concentration areas, and even fewer than the mIBA network. Although some priority areas remained outside the SPA network in coastal waters, the largest were found in offshore waters.

The presence of remaining gaps in the French SPA network has already been suggested based on area comparisons [17,20], spatial prioritization using pooled mammal and seabird aerial surveys [58–60] or expert opinion (EU Pilot 8347/16/ENVI). Our novel approach, objectively based on at-sea seabird data only, explained why the present results were not fully consistent with previous criticisms. For example, some low coverage of mIBA by SPA in the absence of concentration areas did not seem problematic, because of their different methodologies [61]. The current French validated mIBA were determined by applying the theoretical foraging radius (2001 data) around breeding colonies [62]. This type of mIBA failed to distinguish fine-scale [63] or colony-specific habitat use [41], but more importantly, failed to cover important areas further offshore [64] and during the non-breeding season. Therefore, their crude use of surface area to assess the sufficiency of an ecological network raises concerns. On the other hand, the present results refined previous offshore areas reported for SPA designation in the Atlantic (GS7, Capbreton) and Mediterranean (GS8, Gulf of Lion) [59,60], which were confirmed for seabirds around the submarine canyon head off Capbreton, and in the central part of the Gulf of Lion, respectively. In addition, the current study identified two new priority areas outside the current SPA network in the Atlantic (Picardy estuaries and between the islands of Houat and Dumet), although they had previously been reported for taxon concentrations by aerial [23,25] or nautical [31] surveys. Overall, our analysis revealed remaining gaps and calls for improvement of the SPA network in both regions, with priority in the Mediterranean.

## 4.2. Limitations and prospects

### 4.2.1. Data collection

The large amount of data collected in the present study (> 3.8 million) must be placed in relation to the vast marine areas considered. In the Atlantic and the Mediterranean, the average number of bird locations per km² was 11 and 9, respectively. This scarcity of at-sea data for seabirds in metropolitan France has already been reported since the seaward extension of the Natura 2000 network [67], prompting the creation of a national marine data acquisition plan. Although this resource-intensive plan has improved the understanding of the general distribution [23,25] and foraging areas of certain seabird species [42], the data available for metropolitan France remain uneven between species and sometimes insufficient (Table A.5, A.6), potentially weakening the results for species with low distributional precision. The reliance on data quantity and quality in the spatial prioritization process is a recognized challenge, but the growing body of knowledge on seabird distribution and at-sea behaviour [65] offers hope for improvement.

In the current national context of offshore wind farm planning, new aerial, nautical transect and/or tracking studies at both basin and regional scales have been initiated in recent years, while long-term studies continue. In addition, local impact studies are now feeding data into the French DepoBio platform. All these projects will significantly increase the amount of data and knowledge in metropolitan French waters in the coming years. Furthermore, statistical development to combine different data sources in distribution modelling [66] (aerial surveys, boat transects, sightings) is underway throughout France. Together with accurate correction factors accounting for the species detection [67], these developments will improve the robustness of
<figure><img src="image_7.png" /><figcaption>**Fig. 5.** Assessment of the proportion of species population within the French network of SPAs. The red dashed line represents the critical threshold below which it is recognised as insufficient (red bars).</figcaption></figure>

distribution models.

The present data collection revealed the need for complementary at-sea data for several species of high conservation value: Storm petrel, Balearic and Manx shearwaters, Black and Roseate terns, Razorbill, Velvet scoter, Puffin, Greater scaup, Fulmar and Little gull. Although the present study provided a readily applicable starting point towards the goal of completing the Natura 2000 network at sea, a future reassessment of the French SPA network should be undertaken once such data

improvements are achieved.

### 4.2.2. Arbitrary quantitative criteria and qualitative criteria requiring expert opinion

The use of European quantitative criteria (i.e. the proportion of the seabird population to be included in the Natura 2000 network) also implies the use of an arbitrary threshold (i.e. the minimum 20 % threshold), which drives the results. Obviously, the use of a higher
threshold would increase the number of concentration areas and the size of the optimal network. The application of this minimum threshold at the biogeographic scale was responsible for the selection of the most important hotspots and the potential to neglect less important ones. For this reason, some areas may appear blank ('white') on the results maps, either inside or outside the SPAs, but this did not mean that birds were not present in these areas. It simply meant that such data did not pass the threshold, despite the potential presence of a breeding colony or a known foraging area. Therefore, our results represented a relative prioritization of the most concentrated areas for seabirds and should only be considered in the present context of network evaluation.

Moreover, this application of quantitative criteria was an important first step in the process of designating concentration and priority areas, but should not be considered in isolation. Indeed, the other European qualitative criteria (i.e. network distribution within the species range, physical and biological essential factors) must also be considered prior to SPA designation, according to the available literature and expert opinion [68]. For example, the inclusion of essential habitat has supported the designation of the large SPA along the Atlantic shelf break (SPA n°FR5212016), where data are very limited.

### 4.2.3. *National vs. pan-European networks*

Establishing coherent and functional ecological networks for highly mobile species remains a scientific challenge [18]. The site-based Natura 2000 approach recommends appropriate siting of protected areas around seasonal aggregations in order to benefit conservation and optimize efforts [73]. This approach has been criticized for limited effectiveness for highly mobile species that require large-scale, coordinated international action [69,70]. National designations are limited to administrative boundaries that seabirds regularly cross [71]. Following the need for transboundary marine spatial planning and management [72,73], SPA and mIBA network assessment could be based on pooled European distribution data and common workflow, to facilitate gap analysis and final network completion at the pan-European level.

### 4.2.4. *A fixed network in a changing world*

To date, the complex, interconnected and spatio-temporally variable marine environment has been protected by fixed SPA boundaries. As environmental variability and climate change impacts on marine ecosystems, among others, have contrasting effects on seabird demography and distribution [74], the fixed boundaries of protected areas have become an issue for sustainable seabird conservation. In the long term, the conservation contribution of current boundaries may be compromised. One proposed solution is to incorporate predicted future distributions into network design and evaluation [75]. Such predictions would allow potential range shifts to be identified and refugia to be pre-emptively secured, thus creating a network that allows for seabird adaptation and resilience [76,77]. However, accurately predicting the future distribution of marine species and ecosystems is another major challenge [78]. Another more practical option, less sensitive to uncertainty by adapting to environmental variability in near real time, has also been developed with dynamic management [79]. In any case, the sustainability of the seabird populations and their adequate coverage by the Natura 2000 network will always depend on strong scientific knowledge integrated into policy [20] and effective management [80].

## 5. Conclusion

The inherently challenging marine environment has slowed down the establishment of protected areas within it, as has the time-consuming and resource-intensive task of collecting data at sea repetitive. Despite the efforts made to monitor seabirds throughout the maritime zone of metropolitan France, it is not surprising that information on their spatio-temporal distribution and ecology still needs to be improved. The valuable data already collected have shown the importance of certain areas that have been repeatedly identified by

various studies. Again, their identification in this study as seabird hotspots outside the French Natura 2000 network of SPAs gave confidence to confirm the existence of remaining gaps in the network. This national example illustrated the challenge of establishing a coherent, complete and functional ecological network for seabirds at the pan-European level. While additional marine data are being collected, the SPA network still needs scientific progress to be completed and to meet its conservation objectives. Continuing pressures on marine ecosystems make this imperative.

## Funding

This work was supported by the French Ministry of Ecological Transition and Territorial Cohesion.

## Credit authorship contribution statement

**Timothée Poupart:** Writing – review & editing, Writing – original draft, Visualization, Software, Project administration, Methodology, Investigation, Formal analysis, Data curation, Conceptualization. **Thi-baut de Bettignies:** Writing – review & editing, Writing – original draft, Visualization, Validation, Supervision, Resources, Project administra-tion, Methodology, Investigation, Funding acquisition, Conceptualiza-tion. **Matthieu Authier:** Writing – review & editing, Data curation. **Nicola Baccetti:** Writing – review & editing, Data curation. **Luca Börger:** Writing – review & editing, Data curation. **Iker Castège:** Writing – review & editing, Data curation. **Jacopo G. Cecere:** Writing – review & editing, Data curation. **Nicolas Courbin:** Writing – review & editing, Data curation. **Jamie Darby:** Writing – review & editing, Data curation. **Karine Delord:** Writing – review & editing, Data curation. **Ghislain Doremus:** Writing – review & editing, Data curation. **Hayley A. Douglas:** Writing – review & editing, Data curation. **Gilles Faggio:** Writing – review & editing, Data curation. **Giorgia Gaibani:** Writing – review & editing, Data curation. **Fabrice Gallien:** Writing – review & editing, Data curation. **Cécile Cicquel:** Writing – review & editing, Data curation. **Jacob González-Solís:** Writing – review & editing, Data curation. **David Grémillet:** Writing – review & editing, Data curation. **Simona Imperio:** Writing – review & editing, Data curation. **Jude Lane:** Writing – review & editing, Data curation. **Amélie Lescroël:** Writing – review & editing, Data curation. **Maite Louzao:** Writing – review & editing, Data curation. **Jeanne de Mazières:** Writing – review & editing, Data curation. **Noëmie Michez:** Writing – review & editing, Data curation. **Emilie Milon:** Writing – review & editing, Data curation. **Ellie Owen:** Writing – review & editing, Data curation. **Vitor H. Paiva:** Writing – review & editing, Data curation. **Federico De Pascalis:** Writing – review & editing, Data curation. **Clara Péron:** Writing – review & editing, Data curation. **Francesco Pezzo:** Writing – review & editing, Data curation. **Pascal Provost:** Writing – review & editing, Data curation. **Raül Ramos:** Writing – review & editing, Data curation. **Jaime A. Ramos:** Writing – review & editing, Data curation. **Solène Robert:** Writing – review & editing, Data curation. **Diego Rubolini:** Writing – review & editing, Data curation. **Olivier Scher:** Writing – review & editing, Data curation. **Lorenzo Serra:** Writing – review & editing, Data curation.

## Declaration of Competing Interest

The authors have no competing interests to declare.

## Acknowledgements

Our thanks go first to the many direct and indirect data contributors, volunteers and professionals, who go out into the field or help with logistics, feeding or maintaining the increasingly complex databases. Their daily actions contribute to improving the knowledge and translating it into policy support. Their contributions are listed in the
summary of the data used (Table A.2). Specifically in the French context, we would like to thank all those who have contributed to the production and accessibility of data, from the field to the shared databases of the

French National Platform of the Natural Heritage Inventory Information System (SINP). Finally, we would like to thank the reviewers for their comments, which significantly improved the manuscript.

Appendices

<caption>Table A.1</caption>

Seabird taxa by biogeographical region (ATL= Atlantic Ocean, MED= Mediterranean Sea)

<table><thead><tr><th rowspan="2">English name</th><th rowspan="2">Latin name</th><th colspan="2">Region</th><th colspan="2">IUCN<br>Conservation status</th></tr><tr><th>ATL</th><th>MED</th><th>Europe</th><th>France</th></tr></thead><tbody><tr><td>Red-throated diver</td><td>Gavia stellata</td><td>☑</td><td>☒</td><td>LC</td><td>na</td></tr><tr><td>Black-throated diver</td><td>Gavia arctica</td><td>☑</td><td>☑</td><td>LC</td><td>na</td></tr><tr><td>Great northern diver</td><td>Gavia immer</td><td>☑</td><td>☒</td><td>LC</td><td>VU</td></tr><tr><td>Great crested grebe</td><td>Podiceps cristatus</td><td>☑</td><td>☑</td><td>LC</td><td>LC</td></tr><tr><td>Red-necked grebe</td><td>Podiceps grisegena</td><td>☑</td><td>☒</td><td>VU</td><td>CR</td></tr><tr><td>Slavonian grebe</td><td>Podiceps auritus</td><td>☑</td><td>☒</td><td>VU</td><td>VU</td></tr><tr><td>Black-necked grebe</td><td>Podiceps nigricollis</td><td>☑</td><td>☒</td><td>VU</td><td>LC</td></tr><tr><td>Fulmar</td><td>Fulmarus glacialis</td><td>☑</td><td>☒</td><td>VU</td><td>NT</td></tr><tr><td>Scopoli's shearwater</td><td>Calonectris diomedea</td><td>☑</td><td>☑</td><td>LC</td><td>VU</td></tr><tr><td>Cory's shearwater</td><td>Calonectris borealis</td><td>☑</td><td>☒</td><td>LC</td><td>-</td></tr><tr><td>Great shearwater</td><td>Puffinus gravis</td><td>☑</td><td>☒</td><td>LC</td><td>na</td></tr><tr><td>Sooty shearwater</td><td>Puffinus griseus</td><td>☑</td><td>☒</td><td>NT</td><td>na</td></tr><tr><td>Manx shearwater</td><td>Puffinus puffinus</td><td>☑</td><td>☒</td><td>LC</td><td>EN</td></tr><tr><td>Balearic shearwater</td><td>Puffinus mauretanicus</td><td>☑</td><td>☑</td><td>CR</td><td>VU</td></tr><tr><td>Yelkouan shearwater</td><td>Puffinus yelkouan</td><td>☒</td><td>☑</td><td>VU</td><td>EN</td></tr><tr><td>Storm petrel (Atlantic ssp.)</td><td>Hydrobates pelagicus pelagicus</td><td>☑</td><td>☒</td><td>LC</td><td>VU</td></tr><tr><td>Storm petrel (Mediterranean ssp.)</td><td>Hydrobates pelagicus melitensis</td><td>☒</td><td>☑</td><td>LC</td><td>VU</td></tr><tr><td>Leach's storm petrel</td><td>Oceanodroma leucorhoa</td><td>☑</td><td>☒</td><td>NT</td><td>na</td></tr><tr><td>Gannet</td><td>Morus bassanus</td><td>☑</td><td>☑</td><td>LC</td><td>NT</td></tr><tr><td>Cormorant</td><td>Phalacrocorax carbo carbo</td><td>☑</td><td>☑</td><td>LC</td><td>LC</td></tr><tr><td>Shag (Atlantic ssp)</td><td>Phalacrocorax aristotelis aristotelis</td><td>☑</td><td>☒</td><td>LC</td><td>LC</td></tr><tr><td>Shag (Mediterranean ss.)</td><td>Phalacrocorax aristotelis desmarestii</td><td>☒</td><td>☑</td><td>LC</td><td>LC</td></tr><tr><td>Dark-bellied brent goose</td><td>Branta bernicla bernicla</td><td>☑</td><td>☒</td><td>LC</td><td>LC</td></tr><tr><td>Light-bellied brent goose</td><td>Branta bernicla hrota</td><td>☑</td><td>☒</td><td>LC</td><td>LC</td></tr><tr><td>Common shelduck</td><td>Tadorna tadorna</td><td>☑</td><td>☒</td><td>LC</td><td>LC</td></tr><tr><td>Greater scaup</td><td>Aythya marila</td><td>☑</td><td>☒</td><td>LC</td><td>NT</td></tr><tr><td>Eider</td><td>Somateria mollissima</td><td>☑</td><td>☑</td><td>EN</td><td>CR</td></tr><tr><td>Long-tailed duck</td><td>Clangula hyemalis</td><td>☑</td><td>☒</td><td>LC</td><td>na</td></tr><tr><td>Common scoter</td><td>Melanitta nigra</td><td>☑</td><td>☑</td><td>LC</td><td>LC</td></tr></tbody></table>

(continued on next page)
<table><caption>Table A.1 (continued)</caption><tr><td>Velvet scoter</td><td>Melanitta fusca</td><td>☑</td><td>☑</td><td>VU</td><td>EN</td></tr><tr><td>Goldeneye</td><td>Bucephala clangula</td><td>☑</td><td>☒</td><td>LC</td><td>na</td></tr><tr><td>Red-breasted merganser</td><td>Mergus serrator</td><td>☑</td><td>☒</td><td>NT</td><td>CR</td></tr><tr><td>Red phalarope</td><td>Phalaropus fulicarius</td><td>☑</td><td>☒</td><td>LC</td><td>na</td></tr><tr><td>Pomarine skua</td><td>Stercorarius pomarinus</td><td>☑</td><td>☒</td><td>LC</td><td>na</td></tr><tr><td>Arctic skua</td><td>Stercorarius parasitus</td><td>☑</td><td>☑</td><td>EN</td><td>na</td></tr><tr><td>Long-tailed skua</td><td>Stercorarius longicaudus</td><td>☑</td><td>☒</td><td>LC</td><td>VU</td></tr><tr><td>Great skua</td><td>Catharacta skua</td><td>☑</td><td>☑</td><td>LC</td><td>na</td></tr><tr><td>Mediterranean gull</td><td>Ichthyaetus melanocephalus</td><td>☑</td><td>☑</td><td>LC</td><td>LC</td></tr><tr><td>Little gull</td><td>Hydrocoloeus minutus</td><td>☑</td><td>☑</td><td>LC</td><td>na</td></tr><tr><td>Sabine's gull</td><td>Xema sabini</td><td>☑</td><td>☒</td><td>LC</td><td>na</td></tr><tr><td>Black-headed gull</td><td>Chroicocephalus ridibundus</td><td>☑</td><td>☑</td><td>LC</td><td>NT</td></tr><tr><td>Audouin's gull</td><td>Ichthyaetus audouinii</td><td>☒</td><td>☑</td><td>VU</td><td>EN</td></tr><tr><td>Common gull</td><td>Larus canus</td><td>☑</td><td>☒</td><td>LC</td><td>EN</td></tr><tr><td>Lesser black-backed gull</td><td>Larus fuscus</td><td>☑</td><td>☑</td><td>LC</td><td>LC</td></tr><tr><td>Herring gull</td><td>Larus argentatus</td><td>☑</td><td>☒</td><td>LC</td><td>NT</td></tr><tr><td>Yellow-legged gull</td><td>Larus michahellis</td><td>☑</td><td>☑</td><td>LC</td><td>LC</td></tr><tr><td>Great black-backed gull</td><td>Larus marinus</td><td>☑</td><td>☒</td><td>LC</td><td>LC</td></tr><tr><td>Kittiwake</td><td>Rissa tridactyla</td><td>☑</td><td>☑</td><td>VU</td><td>VU</td></tr><tr><td>Sandwich tern</td><td>Thalasseus sandvicensis</td><td>☑</td><td>☑</td><td>LC</td><td>NT</td></tr><tr><td>Roseate tern</td><td>Sterna dougallii</td><td>☑</td><td>☒</td><td>LC</td><td>CR</td></tr><tr><td>Common tern</td><td>Sterna hirundo</td><td>☑</td><td>☑</td><td>LC</td><td>LC</td></tr><tr><td>Arctic tern</td><td>Sterna paradisaea</td><td>☑</td><td>☒</td><td>LC</td><td>CR</td></tr><tr><td>Little tern</td><td>Sternula albifrons</td><td>☑</td><td>☑</td><td>LC</td><td>LC</td></tr><tr><td>Black tern</td><td>Chlidonias niger</td><td>☑</td><td>☑</td><td>LC</td><td>EN</td></tr><tr><td>Guillemot</td><td>Uria aalge</td><td>☑</td><td>☒</td><td>LC</td><td>EN</td></tr><tr><td>Razorbill</td><td>Alca torda</td><td>☑</td><td>☑</td><td>LC</td><td>CR</td></tr><tr><td>Puffin</td><td>Fratercula arctica</td><td>☑</td><td>☑</td><td>EN</td><td>CR</td></tr><tr><td>TOTAL : 57</td><td></td><td>53</td><td>27</td><td></td><td></td></tr></table>
<table><caption>**Table A.2**
Datasets compiled for waters under French jurisdiction in the Atlantic Ocean (ATL) and the Mediterranean Sea (MED)</caption><thead><tr><th>Dataset name</th><th>Region</th><th>Producer(s)</th><th>Diffusion platform</th><th>Access</th><th>Linked publication</th><th>URL giving access or contact</th></tr></thead><tbody><tr><td>Argos_Macreuse noire Mont St Michel_2011_2012</td><td>ATL</td><td>P. Provost S. Provost</td><td>-</td><td>request</td><td>-</td><td>pascal.provost@lpo.fr</td></tr><tr><td>Argos_Plongeon catmarin Lithuanie_2013_2017</td><td>ATL</td><td>R. Žydelis</td><td>Movebank</td><td>request</td><td>-</td><td>https://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study5636685 zydelis@ornitela.eu</td></tr><tr><td>Argos_Puffin des Baléares Mor braz_2012_2013</td><td>ATL, MED</td><td>FAME program</td><td>-</td><td>request</td><td>-</td><td>henri.weimerskirch@cebc.cnrs.fr</td></tr><tr><td>Comptages Wetlands</td><td>ATL, MED</td><td>Wetlands International</td><td>LPO, INPN</td><td>request</td><td>-</td><td>https://www.wetlands.org/ gwenael.quaintenne@lpo.fr</td></tr><tr><td>DepoBio transects aérien</td><td>ATL</td><td>Impact assessment studies</td><td>Depobio</td><td>harvest (SINP)</td><td>-</td><td>https://depot-legal-biodiversity.naturefrance.fr/</td></tr><tr><td>Enquête OFB Grèbes-Plongeons GPS Goéland brun 2010_2019</td><td>ATL, MED</td><td>OFB</td><td>-</td><td>request</td><td>-</td><td>sonia.carrier@ofb.gouv.fr</td></tr><tr><td>Faune France 2002_2022</td><td>ATL, MED</td><td>UVA</td><td>Zenodo</td><td>public</td><td>[81]</td><td>https://zenodo.org/records/3565706#.Yz1U-oTP02x</td></tr><tr><td>GPS Cormoran huppé Chausey_2011_2018</td><td>ATL</td><td>D. Grémillet</td><td>-</td><td>request</td><td>[82]</td><td>http://www.faune-france.org/ philippe.jourde@lpo.fr https://doi.org/10.1007/s00227-020-3655-5 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Cormoran huppé Saint Marcouf_2018</td><td>ATL</td><td>D. Grémillet</td><td>-</td><td>request</td><td>[82]</td><td>https://doi.org/10.1007/s00227-020-3655-5 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Fou de Bassan Bass Rock_2018_2019</td><td>ATL</td><td>K. Hamer, J. Lane</td><td>Seabird Tracking Database</td><td>request</td><td>[83]</td><td>https://data.seabirdtracking.org/dataset/1815</td></tr><tr><td>GPS Fou de Bassan Great Saltee_2017_2018</td><td>ATL</td><td>M. Jessopp, A. Bennison</td><td>Seabird Tracking Database</td><td>request</td><td>-</td><td>https://data.seabirdtracking.org/dataset/1543 m.jessop@ucc.ie</td></tr><tr><td>GPS Fou de Bassan Les Etacs_2011_2015</td><td>ATL</td><td>V. Warwick-Evans</td><td>Movebank</td><td>public</td><td>[84]</td><td>https://www.datarepository.movebank.org/handlle/10255/move.697 https://data.seabirdtracking.org/dataset/734 and 1793,1794,1795,1796 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Fou de Bassan Sept Îles_2005_2017</td><td>ATL</td><td>D. Grémillet</td><td>Seabird Tracking Database</td><td>request</td><td>-</td><td>https://doi.org/10.3354/meps13887 jamie.darby@ucc.ie</td></tr><tr><td>GPS Fulmar boréal 2010_2019</td><td>ATL</td><td>UCC</td><td>-</td><td>request</td><td>[85]</td><td>https://data.seabirdtracking.org/dataset/1237eowen@nts.org.uk</td></tr><tr><td>GPS Fulmar boreal St Martins_2011</td><td>ATL</td><td>RSPB, FameStar-NOFU-STM</td><td>Seabird Tracking Database</td><td>request</td><td>[85]</td><td>https://zenodo.org/record/3541812#.Yz1niITP02x</td></tr><tr><td>GPS Goéland argenté Oostende_2013_2018</td><td>ATL</td><td>UVA</td><td>Zenodo</td><td>public</td><td>[86]</td><td>https://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study261292168 h.a.douglas@hotmail.co.uk http://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study277815715 gilles.faggio@oec.fr https://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study297847376 nicola.baccetti@isprambiente.it https://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study603876591 frederic.robin@lpo.fr https://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study1131915461 lorenzo.serra@isprambiente.it https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Goéland brun Clyde Muirshiel_2017_2018</td><td>ATL</td><td>H. Douglas</td><td>Movebank</td><td>request</td><td>-</td><td>https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Goéland d'Audouin Aspretto_2014_2016</td><td>MED</td><td>B. Recorbet, G. Faggio</td><td>Movebank</td><td>public</td><td>-</td><td>https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Goéland d'Audouin Pianosa_2015_2020</td><td>MED</td><td>N. Bacetti, M. Zenatello</td><td>Movebank</td><td>request</td><td>-</td><td>https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS LARUS LPO Ré_2017_2022</td><td>ATL</td><td>LPO</td><td>Movebank</td><td>request</td><td>-</td><td>https://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study603876591 frederic.robin@lpo.fr https://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study1131915461 lorenzo.serra@isprambiente.it https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Mouette mélanocéphale Cervia_2016_2017</td><td>ATL, MED</td><td>J.G.Cecere, L.Serra, F.De Pascalis</td><td>Movebank</td><td>request</td><td>-</td><td>https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Mouette tridactyle Boulogne_2014</td><td>ATL</td><td>D. Grémillet</td><td>-</td><td>request</td><td>[87]</td><td>https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Mouette tridactyle Fécamp_2014</td><td>ATL</td><td>D. Grémillet</td><td>-</td><td>request</td><td>[87]</td><td>https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Mouette tridactyle St Pierre du Mont_2014</td><td>ATL</td><td>D. Grémillet</td><td>-</td><td>request</td><td>[87]</td><td>https://doi.org/10.1007/s00227-017-3151-8 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Océanite tempête Banneg_2020_2021</td><td>ATL</td><td>B. Cadiou</td><td>Movebank</td><td>request</td><td>-</td><td>https://www.movebank.org/cms/webapp? gwt_fragment=page=studies path=study1401942981 bernard.cadiou@bretagne-vivante.org https://data.seabirdtracking.org/dataset/1556 and 1617, 1853 federico.depascalis@unimi.it</td></tr><tr><td>GPS Océanite tempête Capo Caccia_2019_2021</td><td>MED</td><td>J.G. Cecere, D. Rubolini, F. De Pascalis</td><td>Seabird Tracking Database</td><td>request</td><td>[88]</td><td>https://data.seabirdtracking.org/dataset/1033</td></tr><tr><td>GPS Puffin cendré Cima islet_2011_2015</td><td>ATL</td><td>V. Paiva, J. Ramos</td><td>Seabird Tracking Database</td><td>request</td><td>-</td><td>https://data.seabirdtracking.org/dataset/785 iacopo.cecere@isprambiente.it</td></tr><tr><td>GPS Puffin de Scopoli LaMaddalena_(inc)_2011_2019</td><td>MED</td><td>J.G. Cecere, G. Gaibani, S. Imperio</td><td>Seabird Tracking Database</td><td>request</td><td>[89]</td><td>https://data.seabirdtracking.org/dataset/1618 and 1619, 1623, 1624, 1643,1644 jgsolis@ub.edu https://data.seabirdtracking.org/dataset/1803 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Puffin de Scopoli Cala Morell_2014_2020</td><td>MED</td><td>J. Gonzalez-Solis</td><td>Seabird Tracking Database</td><td>request</td><td>-</td><td>https://data.seabirdtracking.org/dataset/1618 and 1619, 1623, 1624, 1643,1644 jgsolis@ub.edu https://data.seabirdtracking.org/dataset/1803 david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS Puffin de Scopoli Frioul_2011</td><td>MED</td><td>D. Grémillet C. Péron</td><td>Seabird Tracking Database</td><td>request</td><td>[40]</td><td>https://data.seabirdtracking.org/dataset/1618 and 1619, 1623, 1624, 1643,1644 jgsolis@ub.edu https://data.seabirdtracking.org/dataset/1803 david.gremillet@cefe.cnrs.fr</td></tr></tbody></table>

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<table><caption>Table A.2 (continued)</caption><tbody><tr><td>GPS_Puffin de Scopoli<br>Giraglia_2011_2012</td><td>MED</td><td>D. Grémillet<br>C. Péron<br>J. G. Cecere,<br>S. Imperio,<br>D. Rubolini,<br>F. De Pascalis<br>J. G. Cecere,</td><td>Seabird Tracking<br>Database</td><td>request</td><td>[41]</td><td>http://seabirdtracking.org/mapper/index.php<br>david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS_Puffin de Scopoli<br>LaMaddalena_(chick-rear)<br>_2013</td><td>MED</td><td>D. Grémillet<br>C. Péron<br>J. G. Cecere,<br>S. Imperio<br>N. Bacetti<br>D. Grémillet<br>C. Péron</td><td>Seabird Tracking<br>Database</td><td>request</td><td>[90]</td><td>https://data.seabirdtracking.org/dataset/1554 and 1555<br>iacopo.cecere@isprambiente.it</td></tr><tr><td>GPS_Puffin de Scopoli<br>LaScola_2010_2011</td><td>MED</td><td>D. Grémillet<br>C. Péron<br>J. G. Cecere,<br>S. Imperio<br>N. Bacetti<br>D. Grémillet<br>C. Péron</td><td>Seabird Tracking<br>Database</td><td>request</td><td>[91]</td><td>https://data.seabirdtracking.org/dataset/786<br>iacopo.cecere@isprambiente.it</td></tr><tr><td>GPS_Puffin de Scopoli<br>Lavezzi_2011_2012</td><td>MED</td><td>D. Grémillet<br>C. Péron<br>J. G. Cecere,<br>S. Imperio<br>N. Bacetti<br>D. Grémillet<br>C. Péron</td><td>Seabird Tracking<br>Database</td><td>request</td><td>[41]</td><td>http://seabirdtracking.org/mapper/index.php<br>david.gremillet@cefe.cnrs.fr<br>https://data.seabirdtracking.org/dataset/1804 and 1805,<br>1806<br>david.gremillet@cefe.cnrs.fr</td></tr><tr><td>S_Puffin de Scopoli<br>Riou_2011_2020</td><td>MED</td><td>D. Grémillet<br>C. Péron<br>J. G. Cecere,<br>S. Imperio<br>N. Bacetti<br>D. Grémillet<br>C. Péron</td><td>Seabird Tracking<br>Database</td><td>request</td><td>[40,41]</td><td>https://data.seabirdtracking.org/dataset/784<br>iacopo.cecere@isprambiente.it</td></tr><tr><td>GPS_Puffin de Scopoli<br>Spargiotto_2013_2019</td><td>MED</td><td>D. Grémillet<br>C. Péron<br>J. G. Cecere,<br>S. Imperio<br>N. Bacetti<br>D. Grémillet<br>C. Péron</td><td>Seabird Tracking<br>Database</td><td>request</td><td>[91]</td><td>https://data.seabirdtracking.org/dataset/784<br>iacopo.cecere@isprambiente.it</td></tr><tr><td>GPS_Puffin des Anglais<br>2008_2016</td><td>ATL</td><td>T. Guilford</td><td>Movebank</td><td>public</td><td>[92]</td><td>https://www.movebank.org/cms/webapp?<br>gwt_fragment=page=studies.path=study961039869<br>https://www.movebank.org/cms/webapp?<br>gwt_fragment=page=studies.path=study1958889088<br>nicola.baccetti@isprambiente.it</td></tr><tr><td>GPS_Puffin yelkouan<br>Montecristo_2021_2022</td><td>MED</td><td>M. Zenatello,<br>N. Baccetti</td><td>Movebank</td><td>request</td><td>-</td><td>http://seabirdtracking.org/mapper/index.php<br>david.gremillet@cefe.cnrs.fr</td></tr><tr><td>GPS_Puffin yelkouan<br>Port-Cros_2011_2012</td><td>MED</td><td>D. Grémillet,<br>C. Péron</td><td>Seabird Tracking<br>Database</td><td>request</td><td>[42]</td><td>https://www.datarepository.movebank.org/handle/<br>10255/move.1326<br>http://www.movebank.org/cms/webapp?<br>gwt_fragment=page=studies.path=study1546386720<br>olivier.scher@cen-occitanie.org</td></tr><tr><td>GPS_Puffin yelkouan<br>Tavolara_2011_2015</td><td>MED</td><td>M. Zenatello,<br>N. Baccetti</td><td>Movebank</td><td>public</td><td>[93]</td><td>https://www.observatoire-pelagis.cnrs.fr/<br>solene.robert@mnhn.fr</td></tr><tr><td>GPS_Sterne caugek<br>EFGL/EolMed_2021</td><td>MED</td><td>O. Scher,<br>CEN Occitanie</td><td>Movebank</td><td>request</td><td>-</td><td>elodie.giacomini@ofb.gouv.fr<br>cecile.gicquel@ofb.gouv.fr</td></tr><tr><td>ObsEnMer</td><td>ATL,<br>MED</td><td>GECC</td><td>ObsEnMer</td><td>request</td><td>-</td><td>https://www.maisondelestuaire.org/<br>simon.guilbaud@maisondelestuaire.org</td></tr><tr><td>PNM Golfe du Lion</td><td>MED</td><td>OFB</td><td>-</td><td>request</td><td>-</td><td>https://www.maisondelestuaire.org/<br>simon.guilbaud@maisondelestuaire.org</td></tr><tr><td>RN estuaire de la Seine</td><td>ATL</td><td>Natural reserve of the<br>Seine estuary</td><td>-</td><td>request</td><td>-</td><td>https://inpn.mnhn.fr/informations/sinp/presentation<br>solene.robert@mnhn.fr</td></tr><tr><td>SINP</td><td>ATL,<br>MED</td><td>PatriNat</td><td>SINP</td><td>request</td><td>-</td><td>elodie.giacomini@ofb.gouv.fr<br>cecile.gicquel@ofb.gouv.fr</td></tr><tr><td>Survols OFB Bretagne Nord</td><td>ATL</td><td>OFB</td><td>-</td><td>request</td><td>-</td><td>https://www.observatoire-pelagis.cnrs.fr/<br>ghislain.doremus@univ-lr.fr</td></tr><tr><td>Survols OFB PNMI</td><td>ATL</td><td>OFB</td><td>-</td><td>request</td><td>-</td><td>http://www.observatoire-pelagis.cnrs.fr/<br>ghislain.doremus@univ-lr.fr</td></tr><tr><td>Transects aériens SAMM,<br>SCANS</td><td>ATL,<br>MED</td><td>PELAGIS Observatory</td><td>PelaObs</td><td>request</td><td>[23,24]</td><td>https://campagnes.flotteoceanographique.fr/series/19/<br>fr/<br>david.gremillet@cefe.cnrs.fr</td></tr><tr><td>Transects aériens SPEE</td><td>ATL</td><td>PELAGIS Observatory</td><td>PelaObs</td><td>request</td><td>-</td><td>https://www.observatoire-pelagis.cnrs.fr/<br>ghislain.doremus@univ-lr.fr</td></tr><tr><td>Transects nautiques DepoBio</td><td>ATL</td><td>Impact assessment<br>studies</td><td>Depobio</td><td>harvest<br>(SINP)</td><td>-</td><td>https://depot-legal-biodiversite.naturefrance.fr/<br>iker.castege@centredelamer.fr</td></tr><tr><td>Transects nautiques<br>MEGASCOPE<br>2003_2020<br>(CGFS, EVHOE, IBTS,<br>PELGAS)</td><td>ATL</td><td>PELAGIS Observatory</td><td>PelaObs</td><td>request</td><td>-</td><td>https://www.observatoire-pelagis.cnrs.fr/<br>ghislain.doremus@univ-lr.fr</td></tr><tr><td>Transects nautiques PELMED<br>2002_2016</td><td>MED</td><td>Ifremer, CEFE</td><td>-</td><td>request</td><td>-</td><td>https://campagnes.flotteoceanographique.fr/series/19/<br>fr/<br>david.gremillet@cefe.cnrs.fr</td></tr><tr><td>Transects nautiques<br>PELMED, MOOSE</td><td>MED</td><td>PELAGIS Observatory</td><td>PelaObs</td><td>request</td><td>-</td><td>https://www.observatoire-pelagis.cnrs.fr/<br>ghislain.doremus@univ-lr.fr</td></tr><tr><td>Trektellen</td><td>ATL,<br>MED</td><td>Trektellen</td><td>Trektellen</td><td>request</td><td>-</td><td>https://www.trektellen.nl/?language=french<br>gerard.troost@sovon.nl</td></tr></tbody></table>

<caption>Table A.3</caption>

Assignment of species to predicted distribution layers

<table><thead><tr><th rowspan="2">Predicted distribution<br>layers on taxonomic groups<br>from PELAGIS</th><th colspan="2">Attributed species</th></tr><tr><th>Atlantic Ocean</th><th>Mediterranean Sea</th></tr></thead><tbody><tr><td>Alcidae</td><td>Guillemot</td><td>NA</td></tr><tr><td>CATSKU</td><td>Great skua</td><td>NA</td></tr><tr><td>Grey gulls</td><td>Herring gull</td><td>Yellow-legged gull</td></tr><tr><td>Black gulls</td><td>Lesser black-backed gull</td><td>NA</td></tr><tr><td>LARMIN</td><td>Little gull</td><td>Little gull</td></tr><tr><td>LARMEL</td><td>NA</td><td>Mediterranean gull</td></tr><tr><td>LARSPP</td><td>Black-headed gull</td><td>NA</td></tr></tbody></table>

(continued on next page)
<table><caption>Table A.3 (continued)</caption><thead><tr><th rowspan="2">Predicted distribution<br>layers on taxonomic groups<br>from PELAGIS</th><th colspan="2">Attributed species</th></tr><tr><th>Atlantic Ocean</th><th>Mediterranean Sea</th></tr></thead><tbody><tr><td>Hydrobatidae</td><td>Atlantic Storm petrel</td><td>Mediterranean Storm petrel</td></tr><tr><td>Procellaridae</td><td>Manx shearwater</td><td>NA</td></tr><tr><td>Small procellariforms</td><td>NA</td><td>Not used</td></tr><tr><td>Large procellariforms</td><td>NA</td><td>Not used</td></tr><tr><td>Sternidae</td><td>Sandwich tern</td><td>Sandwich tern</td></tr><tr><td>SULBAS</td><td>Gannet</td><td>Gannet</td></tr><tr><td>RISTRI</td><td>Kittiwake</td><td>NA</td></tr></tbody></table>

<caption>Table A.4</caption>

Datasets compiled in this study

<table><caption>Table A.5</caption><thead><tr><th rowspan="2">Dataset</th><th rowspan="2">Producer(s)</th><th colspan="2">Number of data</th><th rowspan="2">Remarks</th></tr><tr><th>ATL</th><th>MED</th></tr></thead><tbody><tr><td>National citizen science database Faune-France</td><td>LPO and contributors</td><td>1 274<br>954</td><td>250<br>750</td><td>Marine environment + 500 m coastal strip to include all land-based observations</td></tr><tr><td>Tracking in France</td><td>see Annex II</td><td>872 908</td><td>635<br>786</td><td>GPS, Argos</td></tr><tr><td>Tracking abroad</td><td>see Annex II</td><td>432 607</td><td>75 909</td><td>GPS</td></tr><tr><td>Trektellen</td><td>Trektellen</td><td>7427</td><td>8 437</td><td>Stationnary birds only</td></tr><tr><td>SINP</td><td>SINP</td><td>54 466</td><td>3 405</td><td>Duplicates removal</td></tr><tr><td>MEGASCOPE<br>(CGFS, EVHOE, IBTS, PELGAS)</td><td>PELAGIS Observatory</td><td>52 785</td><td>-</td><td>-</td></tr><tr><td>Natural reserve of the Seine estuary</td><td>Natural reserve of the Seine estuary</td><td>58 632</td><td>-</td><td>-</td></tr><tr><td>Aerial transects (SAMM, SCANS, SPEE)</td><td>PELAGIS Observatory</td><td>20 427</td><td>3 222</td><td>-</td></tr><tr><td>National citizen science database<br>ObsEnMer</td><td>GECC</td><td>16 277</td><td>4 360</td><td>-</td></tr><tr><td>ERMMA</td><td>Biarritz Sea Centre</td><td>23 771</td><td>-</td><td>-</td></tr><tr><td>Nautical transects (DepoBio)</td><td>DepoBio</td><td>29 636</td><td>-</td><td>-</td></tr><tr><td>Wetlands census</td><td>LPO and contributors</td><td>14 553</td><td>1 733</td><td>Mediterranean Sea:<br>Atlantic Ocean: sub-sectors accuracy, 2019-2022</td></tr><tr><td>Aerial transects (DepoBio)</td><td>DepoBio</td><td>7 993</td><td>-</td><td>-</td></tr><tr><td>Nautical transects PELMED</td><td>Ifremer - CEFE</td><td>-</td><td>9 867</td><td>2002-2016</td></tr><tr><td>Aerial transects OFB Bretagne Nord</td><td>OFB</td><td>2 202</td><td>-</td><td>-</td></tr><tr><td>Aerial transects OFB PNMI</td><td>OFB</td><td>1 923</td><td>-</td><td>-</td></tr><tr><td>Ring controls</td><td>CRBPO</td><td>2 911</td><td>88</td><td>-</td></tr><tr><td>Nautical transects (PELMED, MOOSE)</td><td>PELAGIS Observatory</td><td>-</td><td>2 357</td><td>2017-2020</td></tr><tr><td>National marine parc of the Gulf of Lion</td><td>National marine parc of the Gulf of Lion</td><td>-</td><td>1 842</td><td>-</td></tr><tr><td>Census OFB Grebes and Loons</td><td>OFB</td><td>108</td><td>-</td><td>-</td></tr><tr><td>TOTAL</td><td></td><td>2 857<br>590</td><td>997<br>756</td><td>-</td></tr></tbody></table>

At-sea data by species for the French Atlantic

<table><thead><tr><th>Species</th><th>Maximum Responsibility Index [50]</th><th>Sighting</th><th>Tracking (France)</th><th>Tracking (abroad)</th><th>Interpolated density (nautical transects)</th><th>Predicted density (aerial transects)</th><th>TOTAL</th></tr></thead><tbody><tr><td>Herring gull</td><td>3.3</td><td>131 749</td><td>192 629</td><td>176 158</td><td>4 181</td><td>1 377</td><td>506 094</td></tr><tr><td>Gannet</td><td>4.3</td><td>71 997</td><td>256 452</td><td>103 421</td><td>32 931</td><td>16 995</td><td>481 796</td></tr><tr><td>Lesser black-backed gull</td><td>2.8</td><td>67 314</td><td>95 062</td><td>143 097</td><td>6 474</td><td>1 230</td><td>313 177</td></tr><tr><td>Great black-backed gull</td><td>2.5</td><td>95 721</td><td>181 730</td><td>0</td><td>3 668</td><td>1 238</td><td>282 357</td></tr><tr><td>Black-headed gull</td><td>3</td><td>155 772</td><td>0</td><td>0</td><td>222</td><td>290</td><td>156 284</td></tr><tr><td>Shag</td><td>3</td><td>28 549</td><td>125 792</td><td>0</td><td>1 293</td><td>494</td><td>156 128</td></tr><tr><td>Common shelduck</td><td>3.3</td><td>123 659</td><td>0</td><td>0</td><td>29</td><td>47</td><td>123 735</td></tr><tr><td>Dark-bellied brent goose</td><td>5</td><td>92 605</td><td>0</td><td>0</td><td>111</td><td>162</td><td>92 878</td></tr><tr><td>Cormorant</td><td>2</td><td>87 060</td><td>0</td><td>0</td><td>739</td><td>178</td><td>87 977</td></tr><tr><td>Sandwich tern</td><td>3.3</td><td>74 387</td><td>0</td><td>0</td><td>1 290</td><td>503</td><td>76 180</td></tr><tr><td>Mediterranean gull</td><td>2.3</td><td>66 501</td><td>0</td><td>5 160</td><td>674</td><td>528</td><td>72 863</td></tr><tr><td>Great crested grebe</td><td>2.3</td><td>55 278</td><td>0</td><td>0</td><td>42</td><td>12</td><td>55 332</td></tr><tr><td>Kittiwake</td><td>5</td><td>16 197</td><td>15 698</td><td>0</td><td>4 399</td><td>5 737</td><td>42 031</td></tr><tr><td>Yellow-legged gull</td><td>1.3</td><td>40 294</td><td>0</td><td>0</td><td>155</td><td>5</td><td>40 454</td></tr><tr><td>Common gull</td><td>4.3</td><td>39 040</td><td>0</td><td>0</td><td>234</td><td>155</td><td>39 429</td></tr></tbody></table>

(continued on next page)
<table><caption>**Table A.5 (continued)**</caption><tbody><tr><td>Common tern</td><td>1.5</td><td>35 427</td><td>0</td><td>0</td><td>425</td><td>86</td><td>35938</td></tr><tr><td>Black scoter</td><td>2</td><td>26 704</td><td>4 350</td><td>0</td><td>245</td><td>123</td><td>31 422</td></tr><tr><td>Guillemot</td><td>6.5</td><td>21 073</td><td>0</td><td>0</td><td>10 096</td><td>36</td><td>31 205</td></tr><tr><td>Black-necked grebe</td><td>1.3</td><td>28 876</td><td>0</td><td>0</td><td>19</td><td>0</td><td>28 895</td></tr><tr><td>Balearic shearwater</td><td>8</td><td>19867</td><td>411</td><td>0</td><td>1 813</td><td>86</td><td>22 177</td></tr><tr><td>Great northern diver</td><td>5</td><td>16944</td><td>0</td><td>0</td><td>288</td><td>0</td><td>17 232</td></tr><tr><td>Great skua</td><td>3.3</td><td>11585</td><td>0</td><td>0</td><td>2 380</td><td>658</td><td>14 623</td></tr><tr><td>Razorbill</td><td>7.8</td><td>10 467</td><td>0</td><td>0</td><td>3 254</td><td>4</td><td>13 725</td></tr><tr><td>Fulmar</td><td>5.8</td><td>6 966</td><td>0</td><td>2 645</td><td>2 863</td><td>707</td><td>13 181</td></tr><tr><td>Little gull</td><td>3.8</td><td>10 612</td><td>0</td><td>0</td><td>542</td><td>1 616</td><td>12 770</td></tr><tr><td>Red-throated diver</td><td>5</td><td>11 403</td><td>0</td><td>106</td><td>149</td><td>1</td><td>11 659</td></tr><tr><td>Arctic skua</td><td>5.3</td><td>9 290</td><td>0</td><td>0</td><td>127</td><td>0</td><td>9 417</td></tr><tr><td>Manx shearwater</td><td>6.5</td><td>7 859</td><td>0</td><td>309</td><td>638</td><td>18</td><td>8 824</td></tr><tr><td>Slavonian grebe</td><td>4.3</td><td>8 625</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8 625</td></tr><tr><td>Little tern</td><td>1</td><td>7 039</td><td>0</td><td>0</td><td>4</td><td>5</td><td>7 048</td></tr><tr><td>Storm petrel</td><td>5.3</td><td>4 444</td><td>784</td><td>0</td><td>1 722</td><td>6</td><td>6 956</td></tr><tr><td>Eider</td><td>5</td><td>6 416</td><td>0</td><td>0</td><td>16</td><td>5</td><td>6 437</td></tr><tr><td>Black-throated diver</td><td>5</td><td>5 487</td><td>0</td><td>0</td><td>126</td><td>0</td><td>5 613</td></tr><tr><td>Black tern</td><td>6.5</td><td>5 370</td><td>0</td><td>0</td><td>49</td><td>0</td><td>5 419</td></tr><tr><td>Sooty shearwater</td><td>1</td><td>4 563</td><td>0</td><td>0</td><td>314</td><td>0</td><td>4 877</td></tr><tr><td>Long-tailed duck</td><td>1</td><td>2 969</td><td>0</td><td>0</td><td>1</td><td>0</td><td>2 970</td></tr><tr><td>Velvet scoter</td><td>6.3</td><td>2 962</td><td>0</td><td>0</td><td>0</td><td>6</td><td>2 968</td></tr><tr><td>Light-bellied brent goose</td><td>5</td><td>2 965</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2 965</td></tr><tr><td>Red phalarope</td><td>1</td><td>2 571</td><td>0</td><td>0</td><td>28</td><td>0</td><td>2 599</td></tr><tr><td>Roseate tern</td><td>7</td><td>2 518</td><td>0</td><td>0</td><td>3</td><td>0</td><td>2 521</td></tr><tr><td>Pomarine skua</td><td>5.3</td><td>2 387</td><td>0</td><td>0</td><td>94</td><td>0</td><td>2 481</td></tr><tr><td>Sabine's gull</td><td>1</td><td>2 002</td><td>0</td><td>0</td><td>86</td><td>3</td><td>2 091</td></tr><tr><td>Cory's/Scopoli's shearwater</td><td>3.5</td><td>1 741</td><td>0</td><td>0</td><td>18</td><td>221</td><td>1 980</td></tr><tr><td>Arctic tern</td><td>1</td><td>1 815</td><td>0</td><td>0</td><td>12</td><td>0</td><td>1 827</td></tr><tr><td>Goldeneye</td><td>1.8</td><td>1 714</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1 714</td></tr><tr><td>Cory's shearwater</td><td>3.5</td><td>0</td><td>0</td><td>1 711</td><td>0</td><td>0</td><td>1 711</td></tr><tr><td>Great shearwater</td><td>1</td><td>1 297</td><td>0</td><td>0</td><td>386</td><td>11</td><td>1 694</td></tr><tr><td>Puffin</td><td>7.8</td><td>1 327</td><td>0</td><td>0</td><td>255</td><td>0</td><td>1 582</td></tr><tr><td>Greater scaup</td><td>6</td><td>1 525</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1 525</td></tr><tr><td>Red-breaster merganser</td><td>2.5</td><td>1 467</td><td>0</td><td>0</td><td>4</td><td>2</td><td>1 473</td></tr><tr><td>Leah's storm petrel</td><td>1</td><td>1 372</td><td>0</td><td>0</td><td>16</td><td>0</td><td>1 388</td></tr><tr><td>Red-necked grebe</td><td>1</td><td>1 062</td><td>0</td><td>0</td><td>1</td><td>0</td><td>1 063</td></tr><tr><td>Long-tailed skua</td><td>5.3</td><td>275</td><td>0</td><td>0</td><td>5</td><td>0</td><td>280</td></tr><tr><td></td><td></td><td>1 437</td><td></td><td></td><td></td><td></td><td>2 857</td></tr><tr><td>TOTAL</td><td></td><td>109</td><td>872 908</td><td>432 607</td><td>82 421</td><td>32 545</td><td>590</td></tr></tbody></table>

<caption>**Table A.6**</caption>

At-sea data by species for the French Mediterranean

<table><thead><tr><th>Species</th><th>Maximum Responsibility Index [50]</th><th>Sighting</th><th>Tracking (France)</th><th>Tracking (abroad)</th><th>Interpolated density (nautical transects)</th><th>Predicted density (aerial transects)</th><th>TOTAL</th></tr></thead><tbody><tr><td>Scopoli shearwater</td><td>5.3</td><td>5 874</td><td>521 261</td><td>45 501</td><td>2 226</td><td>202</td><td>575<br>064</td></tr><tr><td>Sandwich tern</td><td>2.5</td><td>23 263</td><td>59 731</td><td>0</td><td>683</td><td>184</td><td>83 861</td></tr><tr><td>Yellow-legged gull</td><td>3.5</td><td>76 117</td><td>0</td><td>0</td><td>6 163</td><td>600</td><td>82 880</td></tr><tr><td>Black-headed gull</td><td>3.3</td><td>57 137</td><td>0</td><td>0</td><td>36</td><td>52</td><td>57 225</td></tr><tr><td>Yelkouan shearwater</td><td>6.5</td><td>2 661</td><td>17 748</td><td>24 226</td><td>896</td><td>0</td><td>45 531</td></tr><tr><td>Audouin's gull</td><td>6.5</td><td>1 808</td><td>37 046</td><td>3 153</td><td>127</td><td>0</td><td>42 134</td></tr><tr><td>Cormorant</td><td>2</td><td>34 176</td><td>0</td><td>0</td><td>3</td><td>9</td><td>34 188</td></tr><tr><td>Mediterranean gull</td><td>2.8</td><td>10 547</td><td>0</td><td>2 569</td><td>287</td><td>462</td><td>13 865</td></tr><tr><td>Great crested grebe</td><td>1.5</td><td>12 464</td><td>0</td><td>0</td><td>0</td><td>0</td><td>12 464</td></tr><tr><td>Common tern</td><td>1.5</td><td>10 032</td><td>0</td><td>0</td><td>823</td><td>0</td><td>10 855</td></tr><tr><td>Little tern</td><td>2</td><td>9 003</td><td>0</td><td>0</td><td>12</td><td>16</td><td>9 031</td></tr><tr><td>Gannet</td><td>1.8</td><td>7 058</td><td>0</td><td>0</td><td>399</td><td>156</td><td>7 613</td></tr><tr><td>Shag</td><td>3.5</td><td>6 168</td><td>0</td><td>0</td><td>6</td><td>18</td><td>6 192</td></tr><tr><td>Little gull</td><td>5.5</td><td>1 753</td><td>0</td><td>0</td><td>0</td><td>1 328</td><td>3 081</td></tr><tr><td>Lesser black-backed gull</td><td>1</td><td>2 646</td><td>0</td><td>70</td><td>4</td><td>0</td><td>2 720</td></tr><tr><td>Black-throated diver</td><td>3</td><td>1 704</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1 704</td></tr><tr><td>Balearic shearwater</td><td>7</td><td>1 008</td><td>0</td><td>0</td><td>376</td><td>0</td><td>1 384</td></tr><tr><td>Black tern</td><td>4.3</td><td>1 257</td><td>0</td><td>0</td><td>19</td><td>0</td><td>1 276</td></tr><tr><td>Storm petrel</td><td>7.8</td><td>593</td><td>0</td><td>390</td><td>90</td><td>97</td><td>1170</td></tr><tr><td>Arctic skua</td><td>3</td><td>1 114</td><td>0</td><td>0</td><td>14</td><td>0</td><td>1 128</td></tr><tr><td>Razorbill</td><td>4.3</td><td>985</td><td>0</td><td>0</td><td>1</td><td>89</td><td>1 075</td></tr><tr><td>Eider</td><td>3</td><td>906</td><td>0</td><td>0</td><td>0</td><td>0</td><td>906</td></tr><tr><td>Velvet scoter</td><td>4.3</td><td>802</td><td>0</td><td>0</td><td>0</td><td>0</td><td>802</td></tr></tbody></table>

(continued on next page)
<table><caption>Table A.6 (continued)</caption><tr><td>Kittiwake</td><td>3</td><td>602</td><td>0</td><td>0</td><td>1</td><td>1</td><td>604</td></tr><tr><td>Black scoter</td><td>1.3</td><td>392</td><td>0</td><td>0</td><td>0</td><td>0</td><td>392</td></tr><tr><td>Great skua</td><td>1</td><td>311</td><td>0</td><td>0</td><td>36</td><td>8</td><td>355</td></tr><tr><td>Puffin</td><td>4.3</td><td>234</td><td>0</td><td>0</td><td>22</td><td>0</td><td>256</td></tr><tr><td>TOTAL</td><td></td><td>270 615</td><td>635 786</td><td>75 909</td><td>12 224</td><td>3 222</td><td>997</td></tr></table>

## Data availability

The authors do not have permission to share all the data, but available data are shown in Table A.2.

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