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Satellite tagging of Mediterranean fin whales: working towards the identification of critical habitats and the focussing of mitigation measures

  • Scientific Reports 7, Article number: 3365 (2017)
  • doi:10.1038/s41598-017-03560-9
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Mediterranean fin whales comprise a genetically distinct population, listed as Vulnerable (VU) in the IUCN Red List. Collisions with vessels are believed to represent the main cause of human-induced mortality. The identification of critical habitats (including migration routes) incorporating satellite telemetry data is therefore crucial to develop focussed conservation efforts. Between 2012 and 2015 thirteen fin whales were equipped with satellite transmitters, 8 in the Pelagos Sanctuary (although two ceased within two days) and 5 in the Strait of Sicily, to evaluate movements and habitat use. A hierarchical switching state-space model was used to identify transiting and area-restricted search (ARS) behaviours, believed to indicate foraging activities. All whales undertook mid- to long-distance migrations, crossing some of the world’s busiest maritime routes. Areas where the animals predominantly engaged in ARS behaviour were identified in both study areas. The telemetry data were compared with results from ecosystem niche modelling, and showed that 80% of tagged whale positions was near (<7 km) the closest suitable habitat. The results contribute to the view that precautionary management should include establishment of a coordinated and dynamic basin-wide management scheme; if appropriate, this may include the establishment of protected areas by specific regional Conventions.


Considerable effort has been devoted to understanding the ecology and conservation status of fin whales (Balaenoptera physalus) in the Mediterranean Sea over the past two decades1,2,3,4,5,6. The species is particularly abundant in the Corso-Liguro-Provençal Basin and Gulf of Lions and congregates in the offshore waters of the Pelagos Sanctuary for Mediterranean Marine Mammals7, a well-documented summer feeding ground8, as well as in the adjacent waters9. Winter feeding aggregations have been reported sporadically around the Island of Lampedusa, Strait of Sicily10, but no additional information exists on other wintering destinations or on seasonal movements.

Stable isotope and acoustic studies11,12,13,14,15,16,17 show the presence of two distinct, seasonally overlapping, sub-populations in the Western Mediterranean. ‘True’ Mediterranean fin whales, which are genetically distinct from North Atlantic conspecifics18, 19, remain year-round in the central and eastern part of the Basin, while North East North Atlantic (NENA) fin whales, seasonally travel between the North Atlantic Ocean and the south of Spain20. This paper focuses on ‘true’ Mediterranean fin whales. Nevertheless, conclusive evidence on the potential input of NENA fin whales to the region - either in the past or at present - remains uncertain.

‘True’ Mediterranean fin whales are listed as Vulnerable (VU) in the IUCN Red List21. This population is exposed to several actual and potential threats, with ship strikes being the main cause of human-induced mortality in the Region22. Further actual or potential threats include chemical pollution3, 23, 24, acoustic disturbance from seismic surveys25 and climate change26, which may negatively influence the population.

Animal-borne telemetry has been increasingly used during the last decade in a variety of environments and for diverse taxa, contributing important information towards the management of species and their environment, and more in general targeting and informing effective conservation27.

ACCOBAMS (the Agreement on the Conservation of Cetaceans in the Black Sea Mediterranean Sea and Contiguous Atlantic Area, under the Convention for Migratory Species) has identified the need to develop a ‘conservation management plan’ for fin whales in the Mediterranean, similar to the approach for such plans developed by the International Whaling Commission (IWC; Any such plan requires inter alia good knowledge of the distribution, movements and important habitat28 of the population in question. Reliable data are essential for the development and success of conservation plans; at the same time it has been recognised that precautionary management, especially for long-lived whales with slow dynamics, may require action before conclusive proof of cause-effect relationships with potential threats29, 30. Satellite telemetry can and has provided valuable information on movements and important habitats of cetaceans that can inform management actions31,32,33,34. This paper provides preliminary information on movements and on the presence of potentially important feeding areas for fin whales in different times of the year, and it also discusses the use of satellite telemetry data to assist in the development of focussed ship strikes mitigation measures, providing indications on the degree of overlap between tagged fin whales and ships (see also supplementary material). While additional data are required to fully address the above mentioned issues, these data contribute to the development and implementation of a proper management and conservation plan for this species in the Mediterranean Sea.



Tag deployment occurred in the Pelagos Sanctuary in September 2012, and around the Island of Lampedusa in March 2013 and 2015 (Fig. 1). Tagging occurred as late in the summer as possible in the Pelagos Sanctuary to gather information outside known summer feeding areas and to observe movements towards winter destinations. In the Strait of Sicily transmitters were deployed in March, when small numbers of whales are known to concentrate for feeding purposes10. When possible, individuals equipped with satellite transmitters were visually monitored after tag deployment to qualitatively assess behavioural changes and verify the correct position of the transmitter on the body of the whales. Only adult individuals, visually assessed by the principal investigator, were tagged.

Figure 1
Figure 1

The two study areas selected for the research project: the Pelagos Sanctuary and the Strait of Sicily. The map was produced with ArcGIS Desktop 9.3 (

Transdermal (Wildlife Computers molds 177 and 193) and Low Impact Minimally Percutaneous External Electronic (LIMPET, molds 260B and 260C) location-only Argos satellite tags35,36,37 were placed on the dorsum or the dorsal fin of whales. The transdermal tags were deployed with a custom-modified pneumatic line thrower (the Air Rocket Transmitter System – ARTS), while the LIMPET ones with a 150 lb crossbow.


Animals feeding on patchy resources are expected to engage in area-restricted search (ARS) behaviour38, 39, characterised by increased turning angles and decreased autocorrelation in direction and speed to maximise searching effort in the most profitable areas. Therefore, Bayesian hierarchical switching state-space models (hSSSM)40,41,42 were fit to Argos location data to differentiate transiting43 from ARS behaviours (details in Supplementary Information) and potentially infer preferred habitats. ARS is believed largely to correspond to foraging behaviour, and its occurrence has been used in several studies to identify putative foraging areas44.

Although recognising the small sample size, the ARS and tracking data were compared to the results of previous ecological niche modelling (ENM) used to predict the potential feeding habitat of fin whales in the Mediterranean Sea45 (see Supplementary Information). This ENM was previously built to integrate knowledge on ecological traits of fin whales (e.g. mobility, feeding strategy) with patterns of selected environmental variables (chlorophyll-a fronts and concentration, water depth) that are thought to be explanatory (or proxy) variables for fin whale distribution. Fin whales were mostly sighted off the continental shelf in the vicinity of chlorophyll-a fronts with low chlorophyll levels (<0.5 mg.m−3). Chlorophyll-a fronts are highly dynamic features that have been shown to be hotspots of marine productivity as they remain long enough (weeks to months) to efficiently transfer the energy in marine food webs through zooplankton growth and to attract higher trophic levels (e.g., the Atlantic bluefin tuna46 loggerhead sea turtles, albacore tuna47, hake recruits48).

Using the derived positions of all the whales equipped with satellite transmitter, a Minimum Bounding Geometry (MBG) enclosing each whale location was calculated. The Utilisation Distribution49, 50 (UD) was calculated across the MBG to gather information on the spatial extent of animal’s home range, as well as a measure of the spatial intensity of use. UD was calculated through Kernel Density Analysis (KDE) to obtain the likelihood of an animal being present at a given point within the home range51. Following the methodology presented by Sprogis et al.52 KDE accounting for physical barriers to movement was calculated using different toolboxes from the ArcMap 10.1 Software53. From KDE values, isopleths (i.e. contours of the UD) were calculated using the Geospatial Modelling Environment54 (GME; Version and the statistical software R55 to determine Core Home Ranges (50% isopleths; CHR) and Total Home Ranges (90% isopleths; THR)56, 57.

As one approach to identifying possible areas of high ship-strike risk, CHRs and THRs were overlaid to density maps of commercial shipping activities58 (merchant ships >1000 gross tonnage at sea). Vessel density maps59,60,61 (1 km2 raster cells; raster values ranging from 0 to 409; available from were georeferenced and vectorised in ArcMap 10.1 and high density areas across the MBG were extracted. The percentage of overlap between traffic areas and THR and CHR areas was then calculated.


The research was conducted in accordance with the guidelines and authorizations of the competent authorities of France, Italy, and the Principality of Monaco recognising both legal and ethical animal welfare criteria.

Research permits and experimental protocols were issued and approved by:

  • Italy: Ministero dell’Ambiente e della Tutela del Territorio e del Mare, Direzione Generale per la Protezione della Natura e del Mare, Divisione II, tutela della biodiversità.

  • France: Direction régionale de l’Environnement, de l′Aménagement et du Logement des Provence-Alpes-Côte d’Azur, upon request from the Ministère de l’Écologie, du Développement Durable, des Transports et du Logement.

  • Monaco: Département de l′Équipement, de l′Environnement et de l′Urbanisme, Direction des Affaires Maritimes.


Between 2012 and 2015, thirteen fin whales were tagged with transdermal (average duration = 45.6 days, SD = 51.0) or LIMPET tags (mean = 29.6 days, SD = 10.3). Eight transmitters (5 transdermal and three LIMPET) were deployed in 2012 in the Pelagos Sanctuary, while 3 transdermal and 2 LIMPET tags were deployed in the Strait of Sicily, in 2013 and 2015, respectively (Table 1).

Table 1: Summary of tag performances.

Corso-Liguro-Provençal Basin

Two out of eight tags deployed in 2012 transmitted for less than 2 days and are not further considered here. The remaining tags revealed consistent movements within the Corso-Liguro-Provençal Basin (Fig. 2), with whales mainly remaining in the waters of the Pelagos Sanctuary and adjacent waters until transmissions stopped. Two individuals (PTT: 112716, 112708, Fig. 2) moved towards the Gulf of Lions and the Balearic Islands at the end of October 2012, remaining in this area for approximately one month before transmissions stopped.

Figure 2
Figure 2

Telemetry tracks of fin whales tagged in the Ligurian Sea in 2012 in the Strait of Sicily in 2013 and 2015, reconstructed from the posterior estimates of the parameters of the hSSSM, and inferred transiting, ARS and uncertain behaviours (based on Jonsen et al.’s62 conservative cut-off values). The light grey points represent model uncertainty around each location, as indicated by the latitude and longitude values at each MCMC iteration. The map was generated using R software, version 3.2.3 (R Core Team, 2015) with packages PBSmapping (Schnute et al. 2015) and rgdal (Bivand et al. 2016). R Foundation for Statistical Computing, Vienna, Austria (

The hSSSM showed fin whales predominantly engaged in ARS behaviour (66%), with potential feeding areas being revealed within the Corso-Liguro-Provençal Basin and towards the Balearic Islands (Fig. 2). Behavioural state estimates indicated a 27% of locations classified as uncertain under the conservative cut-off values proposed by Jonsen et al.62 (see Supplementary material). Inspection of the diagnostic plots and posterior distributions suggested this may reflect the small spatial scale of the animals’ displacement, compared to the accuracy and frequency of the Argos satellite fixed locations63. These locations could also correspond to transition periods between the two behaviours or to short periods of directed movement within longer ARS intervals62, but these sources of uncertainty are confounded63.

The overlap between the whales’ satellite-derived positions and potential foraging habitats obtained from the ENM in the Corso-Liguro-Provençal Basin is shown in Fig. 3 (see discussion).

Figure 3
Figure 3

Seven Argos satellite derived positions overlaid on monthly mean potential feeding habitat (frequency of occurrence from September 2012 to February 2013). Note that only one position per day is shown. The potential habitat is derived from the daily detection of chlorophyll-a front, a range of surface chlorophyll-a content and a minimum water depth (see Supplementary materials for details). The 200 m depth contour is shown. The maps were prepared off-line using a commercial software package: MATLAB and Statistics Toolbox Release 2015b, The MathWorks, Inc., Natick, Massachusetts, United States (

Strait of Sicily

Animals tagged in the Strait of Sicily remained around the Island of Lampedusa for a significant portion of the time they were tracked (March) (Fig. 2). While tags deployed in 2013 did not last long enough for tracking mid- or long-range movements, the longevity of tags improved in 2015 and movements towards the southern coast of Sicily and northern Tunisia were observed. Shorter tracks could be included in the analysis because of the hierarchical nature of the state-space model, which integrated information from longer tracks to classify behavioural states. Most of the whales sighted off Lampedusa in 2013–2015 were observed actively feeding at the surface on large swarms of krill, most likely of the species Nyctiphanes couchii10 (personal observations).

The hSSSM analysis revealed a high rate of ARS behaviour (65%), suggesting several potential feeding areas in the Strait of Sicily (Fig. 2), including the one previously reported around the Island of Lampedusa10. Two other areas, one south-east of Lampedusa and another closer to the coast of Tunisia, may also represent important habitat for fin whales. In fact, after spending 19 days in the Strait of Sicily, one fin whale (PTT 87776) moved north towards the Southern Tyrrhenian Sea and the east coast of Sardinia Island. Then it headed back towards the south of Sardinia, between Tunisia and the Island of Lampedusa where it remained for a few days engaging in intense feeding before transmissions were interrupted.

After remaining in the Sicily Strait area for a month, the second whale (PTT 87780) moved northwards to the area of the Pelagos Sanctuary, remaining for eight days north of Corsica Island before moving south-westward. The hSSSM analysis highlighted the presence of potential feeding areas in both the Strait of Sicily and the Ligurian Sea, corroborating previous knowledge and further supporting the importance of these areas for the ecology of the species in the Region (Fig. 3).

The overlap between the whales’ satellite-derived positions and potential foraging habitats obtained from the ENM in the Strait of Sicily and in the South-western Tyrrhenian Basin, between Sicily and Sardinia, is shown in Fig. 4 (see discussion).

Figure 4
Figure 4

Two Argos satellite derived positions overlaid on fortnights mean potential feeding habitat (March-April 2015). Note that only one position per day is shown. The potential habitat of fin whale is derived from the daily detection of chlorophyll-a fronts, a range of surface chlorophyll-a content and a minimum water depth (see Supplementary materials for details). The 200 m depth contour is shown. The maps were prepared off-line using a commercial software package: MATLAB and Statistics Toolbox Release 2015b, The MathWorks, Inc., Natick, Massachusetts, United States (

Traffic and whale movements

When considering both the 90% total home range and 50% core home range areas, the 17.4% of the former and 14.0% of the latter overlap with areas of high traffic volumes within the minimum bounding geometry (MBG) (Fig. 5).

Figure 5
Figure 5

Overlapping of the 90% total home range areas and 50% core home range with regions of high traffic usage. Green dashed line represents the minimum bounding geometry enclosing all whale locations. Orange dashed area corresponds to areas of high traffic densities. Kernel derived total home ranges (90% isopleths) in blue and core home ranges (50% isopleths) in red. The map was produced with ArcGIS Desktop 9.3 (


Satellite telemetry data such as that from Mediterranean fin whales in the Pelagos Sanctuary and in the Strait of Sicily between 2012 and 2015 – further complemented with spatial modelling, for example with results from ecological niche modelling and from aerial surveys5 – can provide an important contribution with other data (e.g. from cetacean abundance surveys and information on human activities) to begin to prioritise and help develop conservation actions27, 64,65,66,67,68 (e.g. suggesting areas where conservation actions such as reduced ship speed58, 69, 70 or regulation of whale watching activities) within the species’ range, both in summer and winter seasons. Understanding why whales are where they are when they are is a vital component of developing effective mitigation measures and focussing where they are needed. This is an important task but it must be recognised that for long-lived, wide-ranging and difficult-to-study species such as whales, this will be inevitably a cumulative process over a number of years. We fully recognise that this will require the collection of more satellite derived data from fin whales, as well as building up of additional data on explanatory variables and improved modelling (e.g. ref. 71) from a number of sources, but for illustrative purposes of this paper, comparison of the tracks of the whales with the results of predictive feeding areas from one modelling approach40 for the same time periods has been presented. This does not imply that that this particular ENM approach and the use of chlorophyll-a data is the best proxy for fin whale feeding habitat, but it rather provides a useful start to an ongoing process, as does work on spatial modelling from ship and aerial surveys (e.g. refs 5, 72 and 73).

Corso-Liguro-Provençal Basin

The overlap between the whales’ satellite-derived positions and potential foraging habitats obtained from the ENM in the Corso-Liguro-Provençal Basin, is shown in Fig. 3.

The ENM used chlorophyll-a fronts as a proxy for food availability45. Based upon this, the potential feeding habitat for fin whales in the north-western Mediterranean Sea during the 2012 study period expanded from a limited extent during late summer in the Corso-Liguro-Provençal Basin (Fig. 3a,b) to most of that area from November 2012 (Fig. 3c) to the entire western Mediterranean by January 2013 (Fig. 3d,e), prior to tracing the general circulation of the Western Mediterranean in February 2013 in the northern part of the basin (Fig. 3f). The surface area of suitable habitat for the period 2003–2014 ranged from 7 to 8% of the western Mediterranean Sea in July-September (4–6% for 2012) to 27% in January 2003–2014 (37% for 2013). The analysis showed that 80% of tagged whale positions (n = 415) were closer than 7 km to the closest suitable habitat, which is consistent with the overall habitat model performance of 8 km (n = 1287)45. The spatial dispersion of the six whales from the Ligurian Sea in September 2012 to most of the northern basin in the following months matches the habitat expansion, with clear phases of transit when successive positions are distant, and ARS behaviour otherwise (Fig. 2). Feeding habitat contraction within Pelagos during summer, and successive expansion during winter, could provide insights into the mechanisms driving fin whales’ concentration there in summer and dispersion later20.

Krill in the Ligurian Sea has been reported in spring, summer and autumn with strandings of M. norvegica swarms along the coast of western Liguria, southern France and Monaco during the winter months74, 75. Nonetheless, fin whales’ presence in the area drastically drops during winter5, with animals primarily occurring during the early summer and engaging in feeding8. This seasonal occurrence of fin whales in the Ligurian Sea has been related to the fact that, despite the overall year-round presence of krill as possible prey, potential feeding habitats require high density swarms and not simply high average concentrations; these are substantially more spread over the basin in winter45, leading whales to migrate outside the well-known summer feeding grounds to take advantage of higher concentration of food in other areas76. The tagged whales remained in the Pelagos Sanctuary and adjacent waters longer in the season than described in the available literature, engaging in ARS behaviour for a significant portion of time (66%). Due to the errors associated with ARGOS fixes and the small scale of the displacement in this region, behaviour was classified as uncertain (under the conservative criteria proposed by Jonsen et al.62) in almost a third of the locations63. The poor quality of ARGOS locations contributed to these moderate levels of uncertainty in the behavioural classification by the state-space model. While disentangling ARGOS error (intrinsic in our data) from the uncertainty associated with the behavioural process would only be possible with an independent index of behavioural state (which is not currently available), our conclusions regarding the whales’ ecology are not affected by the locations classified as uncertain.

Notwithstanding these limitations, the whales clearly showed a complex movement pattern in the area. It is interesting to note that the longer transiting movements corresponded with areas that were not identified as potential feeding habitat by the ENM (Fig. 3). This also supports the view that whales persisted in using the Pelagos Sanctuary at the end of summer due to favourable conditions allowing for prolonged feeding.

The results presented here, even if based on a small sample size, emphasize that potentially important fin whale habitat extends westward of the Pelagos Sanctuary9, 77, well within the north-western Mediterranean Pelagic Ecosystems Ecologically or Biologically Significant Marine Area (EBSA,, identified by the Convention on Biological Diversity (CBD). Moreover,  previous research along the eastern borders of the Sanctuary78, as well as spatial modelling of aerial survey data, showed a longitudinal gradient in the occurrence of fin whales within the northern Pelagos Sanctuary, predicting higher densities west of Sardinia, primarily in deep offshore waters79.

Strait of Sicily

The overlap between the whales’ satellite-derived positions and potential foraging habitats obtained from the ENM in the Strait of Sicily and in the South-western Tyrrhenian Basin, between Sicily and Sardinia, is shown in Fig. 4.

The ENM and tracking results from the Strait of Sicily in 2015 (Fig. 4) showed that potential feeding habitat extended well beyond the near-shore waters of the Island of Lampedusa10. Whales remained in this area for almost a month after tagging, engaging in ARS behaviour 65% of the time, prior to moving northwards. In addition to being a hotspot for Mediterranean biodiversity (, the Strait of Sicily also represents the main deep-water vessel traffic channel connecting the Eastern and Western basins of the Mediterranean, with severe traffic volumes between the Suez Canal and the Strait of Gibraltar80 (see Fig. S1 in the Supplementary Information). This may have potentially serious impacts to fin whales, both with respect to generated underwater noise81,82,83 and ship strike risk22.

General observations on movements/migration

The observed longitudinal movements of fin whales tagged in the Ligurian Sea in the late summer and the latitudinal migration recorded in early spring, support the hypothesis that the whales summering in the north-western Mediterranean Sea travel southward towards the winter feeding grounds in the Strait of Sicily, and possibly towards non identified breeding areas in the Southern Mediterranean Sea8, 12. One additional hypothesis arising out of our limited information that requires additional research effort is that whales would later move northbound towards the Pelagos Sanctuary and adjacent waters during the early- mid-spring, following the marked feeding habitat concentration in the area described by Notarbartolo di Sciara et al.20.

The results of the hSSSM analysis for one whale tagged in the Strait of Sicily (PPT 87780), and, to a lesser extent, of the whales tracked in the Ligurian Sea, suggest that, despite the presence of potential feeding areas along their tracks, whales may travel consistently between different locations without suspending their movements to engage in ARS behaviour. Although clearly a sample size of one is insufficient, this pattern has been observed for fin whales in other areas44 and is consistent with the idea that for energetic reasons, whales typically exploit areas of high concentration of food resources, without capitalising on small patches of prey38. The ENM takes into account the size of chlorophyll-a fronts - the hypothesis is that the larger the productivity of the frontal feature and the longer its stability over time, the higher the probability that dense aggregations of prey will form, attracting feeding fin whales.

The movements between the waters of the Strait of Sicily and the Ligurian Sea indicate that at least some of the whales that visit the Strait of Sicily in winter will congregate later in spring and summer in the North-Western Mediterranean. This supports the hypothesis of seasonal movements of Mediterranean fin whales between the two areas, probably related at least in part to feeding8, 12. The seasonal presence of fin whales further to the east, including in the Ionian84 and Southern Adriatic Seas85, can also be explained by more complex searching for prey concentrations across the central Mediterranean.

The connection between whales seen off Lampedusa and those that spend the summer in Pelagos Sanctuary – also witnessed by photo-identification data and suggested by aerial survey data5, 86 - illustrates large and defined seasonal movements of fin whales through the Mediterranean. It is therefore evident that a substantial portion of the Central and North-Western Mediterranean plays an important role in the ecology of the species. By extension, especially in light of the high level of anthropogenic stressors affecting the area, it is also an important area for conservation efforts. The northbound migration routes shown here reveal that the animals move through areas with high human activity. Overlap of whale presence with such activity gives rise to increased exposure to threats, such as ship strikes, where at present no protection or mitigation schemes are in place. This new information, whilst preliminary, highlights the need for early consideration of a comprehensive basin-wide mitigation scheme to complement any national measures, in conjunction with additional research to provide a fuller understanding of whale distribution and behaviour at appropriate geographical and temporal scales.

The movements described here, assuming that chlorophyll-a is a reasonable proxy for potential feeding areas, appears to match the dynamics of the habitat. It suggests that whales have the ability to track food resources over time in order to exploit seasonally and spatially restricted habitats during the peaks of prey abundance and perhaps ensure prolonged access to higher-quality foraging opportunities. Thus, the movements observed within the north-Western Mediterranean Sea, between the Gulf of Genoa and the Balearic Islands, as well as between the Strait of Sicily and the Pelagos Sanctuary, is an effective response to resource fluctuations, both within and across seasons and areas.

Although of limited sample size, the data show a synchrony between the start of the northbound migration and the shifting in the occurrence of potential feeding habitats. For example, two whales remained in the area off Lampedusa in March, when the probability of feeding habitat is high in the Strait of Sicily and very low in the Tyrrhenian and Ligurian Seas. As feeding habitat decreased in the Strait of Sicily and increased in the South Tyrrhenian Sea during the first half of April, both whales started to move northbound towards richer areas. The whale with the remaining transmitter in the second half of April pursued its northward migration to the area of the Pelagos Sanctuary as the habitat became more productive, before transmissions stopped. Movements of both whales demonstrate the switch of favourable habitat occurrence in the first half of April from the area south of Sicily to the western Mediterranean Sea. In particular, the second whale went directly to the summer feeding grounds (900 km apart in a straight line distance) in about five days.

Conservation implications

Ship strikes represent the main cause of concern for the conservation of fin whales across the Mediterranean basin87. This issue is being addressed at the regional and international levels through a joint effort by ACCOBAMS, the Pelagos Sanctuary Agreement and the International Whaling Commission (IWC). These organizations work towards the identification of high-risk areas where to apply and test mitigation actions. Among the measures currently in place in other areas of the world, mainly implemented through the International Maritime Organization (IMO), there are Areas To Be Avoided (ATBA), Particularly Sensitive Sea Areas (PSSA) and the establishment or the shifting of Traffic Separation Schemes (TSS). A proposal by France is currently envisaging the establishment of a PSSA within the Pelagos Sanctuary borders, while the presented results would argue in favour of a wider area, possibly encompassing more critical habitats for fin whales, such as CBD’s ‘north-western Mediterranean Pelagic Ecosystems’ EBSA. The data collected here, once supplemented by additional deployments and integrated with data from other whale studies and information on human activities such as vessel traffic, are informative for the implementation and assessment of these initiatives. The approach illustrated here (Fig. 5), identifying areas used by fin whales and areas of high traffic volumes, is one that can be improved and developed with additional data.

We recognise that the present findings alone are insufficient to identify robustly the most important areas for fin whales. However, they do provide an important start to this task and support the importance of beginning political efforts to develop an effective multi-national conservation management plan for this species with a designated action plan to address actual and potential threats. Approaches to consider include the establishment of a seasonal/dynamic focussed protection regime (e.g. a Marine Protected Area or a Specially Protected Area of Mediterranean Importance (SPAMI) under the auspices of the Barcelona Convention). Static MPAs often fail to encompass the dynamic features that many pelagic and mobile species exploit88. Hence, despite substantial practical and legal challenges, spatially and temporally dynamic protection schemes, intended to change their location and size based on habitat characteristics, species occurrence and movements, as well as water column features, could be effective in protecting highly mobile species, provided that they are accompanied by the necessary research to develop robust models of whale occurrence at the appropriate spatial and temporal scales89,90,91.


The Pelagos Sanctuary and its surrounding bodies of water represent the major summer feeding grounds for the species8 and host critical feeding habitats also for a large portion of the autumn and early winter months. It is therefore crucial that mitigation of detrimental activities, such as maritime traffic (see Fig. S1 in the Supplementary Information), and overall actions to warrant the protection and conservation of fin whales and their critical habitats extend beyond the present boundaries of the Sanctuary.

Satellite telemetry combined with habitat modelling is one important tool to assess critical and highly used habitats of fin whales in the Mediterranean Sea and important migration routes. Such information is essential to prioritise mitigation of human-induced threats, including, for example, ship strikes.

As models become more robust - and more data are collected - the results can feed into coordinated and dynamic management schemes92 – as foreseen by the Marine Spatial Planning (MSP) - to try to effectively protect fin whales in the Mediterranean. Information on the temporal and geographical distribution and abundance of fin whales (and thus fin whales’ critical habitats) are also required for identification of Important Marine Mammal Areas (IMMAs), which could lead to the proposal for the establishment of Specially Protected Areas of Mediterranean Importance (SPAMIs) under the framework of the Barcelona Convention. It is therefore recommended that further studies, such as the one proposed and described here, be conducted also in other targeted areas of the Mediterranean Basin such as the Ionian Sea and the North African coasts, to provide a more complete picture of the whales’ distribution and movements within the Mediterranean and the presence of their critical habitats.

Additional Information

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  1. 1.

    Azzellino, A., Gaspari, S., Airoldi, S. & Nani, B. Habitat use and preferences of cetaceans along the continental slope and the adjacent pelagic waters in the western Ligurian Sea. Deep Sea Res. Part Oceanogr. Res. Pap. 55, 296–323 (2008).

  2. 2.

    Forcada, J., Aguilar, A., Hammond, P., Pastor, X. & Aguilar, R. Distribution and abundance of fin whales (Balaenoptera physalus) in the western Mediterranean sea during the summer. J. Zool. 238, 23–34 (1996).

  3. 3.

    Fossi, M. C. et al. A multi-trial diagnostic tool in fin whale (Balaenoptera physalus) skin biopsies of the Pelagos Sanctuary (Mediterranean Sea) and the Gulf of California (Mexico). Mar. Environ. Res. 69, S17–S20 (2010).

  4. 4.

    Laran, S. & Gannier, A. Spatial and temporal prediction of fin whale distribution in the northwestern Mediterranean Sea. ICES J. Mar. Sci. 65, 1260–1269 (2008).

  5. 5.

    Panigada, S., Lauriano, G., Burt, L., Pierantonio, N. & Donovan, G. Monitoring winter and summer abundance of cetaceans in the Pelagos Sanctuary (northwestern Mediterranean Sea) through aerial surveys. PloS One 6, e22878 (2011).

  6. 6.

    Panigada, S., Notarbartolo di Sciara, G. & Zanardelli Panigada, M. Fin whales summering in the Pelagos Sanctuary (Mediterranean Sea): Overview of studies on habitat use and diving behaviour. Chem. Ecol. 22, S255–S263 (2006).

  7. 7.

    Notarbartolo di Sciara, G., Agardy, T., Hyrenbach, D., Scovazzi, T. & Van Klaveren, P. The Pelagos Sanctuary for Mediterranean marine mammals. Aquat. Conserv. Mar. Freshw. Ecosyst. 18, 367–391 (2008).

  8. 8.

    Notarbartolo di Sciara, G., Zanardelli, M., Jahoda, M., Panigada, S. & Airoldi, S. The fin whale Balaenoptera physalus (L. 1758) in the Mediterranean Sea. Mammal Rev. 33, 105–150 (2003).

  9. 9.

    Notarbartolo di Sciara, G., Panigada, S. & Agardy, T. Is the Pelagos Sanctuary sufficiently large for the cetacean populations it is intended to protect? Rapp Comm Int Mer Médit 623 (2013).

  10. 10.

    Canese, S. et al. The first identified winter feeding ground of fin whales (Balaenoptera physalus) in the Mediterranean Sea. J. Mar. Biol. Assoc. UK 86, 903 (2006).

  11. 11.

    Bentaleb, I. et al. Foraging ecology of Mediterranean fin whales in a changing environment elucidated by satellite tracking and baleen plate stable isotopes. Mar Ecol Prog Ser 438, 285–302 (2011).

  12. 12.

    Castellote, M., Clark, C. W. & Lammers, M. O. Fin whale (Balaenoptera physalus) population identity in the western Mediterranean Sea. Mar. Mammal Sci 28, 325–344 (2012).

  13. 13.

    Castellote, M., Clark, C. W. & Lammers, M. O. Dispersal of North Atlantic fin whales (Balaenoptera physalus) into the Mediterranean Sea and exchange between populations: response to Giménez et al., Rapid Commun. Mass Spectrom. 2013, 27, 1801–1806. Rapid Commun. Mass Spectrom. RCM 28, 665–667 (2014).

  14. 14.

    Cotté, C. Scale-dependent interactions of Mediterranean whales with marine dynamics. Limnol. Oceanogr. 56, 219–232 (2011).

  15. 15.

    Giménez, J., Gómez-Campos, E., Borrell, A., Cardona, L. & Aguilar, A. The uncertain status of the Mediterranean and northeastern North Atlantic fin whale subpopulations: reply to Castellote et al., Rapid Commun. Mass Spectrom. 2014, 28, 665–667. Rapid Commun. Mass Spectrom. RCM 28, 668–670 (2014).

  16. 16.

    Giménez, J., Gómez-Campos, E., Borrell, A., Cardona, L. & Aguilar, A. Isotopic evidence of limited exchange between Mediterranean and eastern North Atlantic fin whales. Rapid Commun. Mass Spectrom. 27, 1801–1806 (2013).

  17. 17.

    Ryan, C., McHugh, B., O’Connor, I. & Berrow, S. Lipid content of blubber biopsies is not representative of blubber in situ for fin whales (Balaenoptera physalus). Mar. Mammal Sci. 29, 542–547 (2013).

  18. 18.

    Palsbøll, P. J., Bérubé, M., Aguilar, A., Notarbartolo di Sciara, G. & Nielsen, R. Discerning Between Recurrent Gene Flow and Recent Divergence Under a Finite-Site Mutation Model Applied to North Atlantic and Mediterranean Sea Fin Whale (Balaenoptera physalus) Populations. Evolution 58, 670–675 (2004).

  19. 19.

    Bérubé, M. et al. Population genetic structure of North Atlantic, Mediterranean Sea and Sea of Cortez fin whales, Balaenoptera physalus (Linnaeus 1758): analysis of mitochondrial and nuclear loci. Mol. Ecol. 7, 585–599 (1998).

  20. 20.

    Notarbartolo di Sciara, G., Castellote, M., Druon, J.-N. & Panigada, S. In Advances in Marine Biology (ed. Notarbartolo di Sciara, G., Podestà, M. and Curry, B. E.) 75, 75–101 (Academic Press, 2016).

  21. 21.

    Panigada, S. & Notarbartolo di Sciara, G. Balaenoptera physalus (Mediterranean subpopulation). The IUCN Red List of Threatened Species. Version 2014.3 Available at: (Accessed: 12th April 2015) (2012).

  22. 22.

    Panigada, S. et al. Mediterranean fin whales at risk from fatal ship strikes. Mar Pollut Bull 52, 1287–1298 (2006).

  23. 23.

    Fossi, M. C. et al. The use of a non-lethal tool for evaluating toxicological hazard of organochlorine contaminants in Mediterranean cetaceans: new data 10 years after the first paper published in MPB. Mar Pollut Bull 46, 972–982 (2003).

  24. 24.

    Fossi, M. C., Casini, S. & Marsili, L. Potential toxicological hazard due to endocrine-disrupting chemicals on Mediterranean top predators: state of art, gender differences and methodological tools. Environ. Res. 104, 174–182 (2007).

  25. 25.

    Castellote, M., Clark, C. W., Colmenares, F. & Esteban, J. A. Mediterranean fin whale migration movements altered by seismic exploration noise. J. Acoust. Soc. Am. 125, 2519 (2009).

  26. 26.

    Simmonds, M. P., Gambaiani, D. & Notarbartolo di Sciara, G. In Life in the Mediterranean Sea: A Look at Habitat Changes 685–701 (2012).

  27. 27.

    McGowan, J. et al. Integrating research using animal-borne telemetry with the needs of conservation management. J. Appl. Ecol. doi:10.1111/1365-2664.12755 (2016).

  28. 28.

    Notarbartolo di Sciara, G. et al. Place-based approaches to marine mammal conservation. Aquat. Conserv. Mar. Freshw. Ecosyst. 26, 85–100 (2016).

  29. 29.

    Grand, J., Cummings, M. P., Rebelo, T. G., Ricketts, T. H. & Neel, M. C. Biased data reduce efficiency and effectiveness of conservation reserve networks. Ecol. Lett. 10, 364–374 (2007).

  30. 30.

    Possingham, H., Ball, I. & Andelman, S. In Quantitative Methods for Conservation Biology 291–306 (Springer New York, 2000).

  31. 31.

    Gendron, D., Serrano, I. M., Cruz, A. U., de la Calambokidis, J. & Mate, B. Long-term individual sighting history database: an effective tool to monitor satellite tag effects on cetaceans. Endanger. Species Res. 26, 235–241 (2015).

  32. 32.

    Mate, B. R. et al. Critically endangered western gray whales migrate to the eastern North Pacific. Biol. Lett. 11, 20150071 (2015).

  33. 33.

    Double, M. C. et al. Migratory Movements of Pygmy Blue Whales (Balaenoptera musculus brevicauda) between Australia and Indonesia as Revealed by Satellite Telemetry. PLoS ONE 9, e93578 (2014).

  34. 34.

    Sveegaard, S. et al. Defining management units for cetaceans by combining genetics, morphology, acoustics and satellite tracking. Glob. Ecol. Conserv. 3, 839–850 (2015).

  35. 35.

    Zerbini, A. N. et al. Satellite-monitored movements of humpback whales Megaptera novaeangliae in the Southwest Atlantic Ocean. Mar Ecol Prog Ser 313, 295–304 (2006).

  36. 36.

    Andrews, R. D., Pitman, R. L. & Ballance, L. T. Satellite tracking reveals distinct movement patterns for Type B and Type C killer whales in the southern Ross Sea, Antarctica. Polar Biol. 31, 1461–1468 (2008).

  37. 37.

    Kennedy, A. S. et al. Local and migratory movements of humpback whales (Megaptera novaeangliae) satellite-tracked in the North Atlantic Ocean. Can J Zool 92, 9–18 (2013).

  38. 38.

    Kareiva, P. & Odell, G. Swarms of Predators Exhibit ‘Preytaxis’ if Individual Predators Use Area-Restricted Search. Am. Nat. 130, 233–270 (1987).

  39. 39.

    Tinbergen, N., Impekoven, M. & Franck, D. An Experiment on Spacing-Out as a Defence against Predation. Behaviour 28, 307–321 (1967).

  40. 40.

    Jonsen, I. D. et al. State-space models for bio-loggers: A methodological road map. Deep Sea Res. Part II Top. Stud. Oceanogr 88–89, 34–46 (2013).

  41. 41.

    Jonsen, I. D., Myers, R. A. & Flemming, J. M. Meta-analysis of animal movement using state-space models. Ecology 84, 3055–3063 (2003).

  42. 42.

    Jonsen, I., Flemming, J. & Myers, R. A. Robust state-space modeling of animal movement data. Ecology 86, 2874–2880 (2005).

  43. 43.

    Zollner, P. A. & Lima, S. L. Search Strategies for Landscape-Level Interpatch Movements. Ecology 80, 1019–1030 (1999).

  44. 44.

    Silva, M. A., Prieto, R., Jonsen, I., Baumgartner, M. F. & Santos, R. S. North Atlantic Blue and Fin Whales Suspend Their Spring Migration to Forage in Middle Latitudes: Building up Energy Reserves for the Journey? PLoS ONE 8, e76507 (2013).

  45. 45.

    Druon, J. et al. Potential feeding habitat of fin whales in the western Mediterranean Sea: an environmental niche model. Mar. Ecol. Prog. Ser. 464, 289–306 (2012).

  46. 46.

    Druon, J.-N. et al. Habitat suitability of the Atlantic bluefin tuna by size class: An ecological niche approach. Prog. Oceanogr. 142, 30–46 (2016).

  47. 47.

    Polovina, J. J., Howell, E. A., Kobayashi, D. R. & Seki, M. P. The Transition Zone Chlorophyll Front updated: Advances from a decade of research. Prog. Oceanogr, doi:10.1016/j.pocean.2015.01.006.

  48. 48.

    Druon, J.-N. et al. Modelling of European hake nurseries in the Mediterranean Sea: An ecological niche approach. Prog. Oceanogr. 130, 188–204 (2015).

  49. 49.

    Van Winkle, W. Comparison of Several Probabilistic Home-Range Models. J. Wildl. Manag. 39, 118–123 (1975).

  50. 50.

    Worton, B. J. Kernel Methods for Estimating the Utilization Distribution in Home-Range Studies. Ecology 70, 164–168 (1989).

  51. 51.

    Kernohan, B. J., Gitzen, R. A. & Millspaugh, J. J. In Radio Tracking and Animal Populations 125–166 (Elsevier, 2001).

  52. 52.

    Sprogis, K. R., Raudino, H. C., Rankin, R., MacLeod, C. D. & Bejder, L. Home range size of adult Indo-Pacific bottlenose dolphins (Tursiops aduncus) in a coastal and estuarine system is habitat and sex-specific. Mar. Mammal Sci. 32, 287–308 (2016).

  53. 53.

    ESRI. ArcGIS Desktop: Release 10 (Environmental Systems Research Institute, 2011).

  54. 54.

    Beyer, H. L. Geospatial Modelling Environment (Version (software) (2012).

  55. 55.

    R Development Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0, (2013).

  56. 56.

    Börger, L. et al. An integrated approach to identify spatiotemporal and individual-level determinants of animal home range size. Am. Nat. 168, 471–485 (2006).

  57. 57.

    Clapp, J. G. & Beck, J. L. Evaluating distributional shifts in home range estimates. Ecol. Evol 5, 3869–3878 (2015).

  58. 58.

    Irvine, L. M. et al. Spatial and temporal occurrence of blue whales off the US west coast, with implications for management. PloS One 9, e102959 (2014).

  59. 59.

    Halpern, B. S. et al. Spatial and temporal changes in cumulative human impacts on the world’s ocean. Nat. Commun. 6, 7615 (2015).

  60. 60.

    Halpern, B. S. et al. A Global Map of Human Impact on Marine Ecosystems. Science 319, 948–952 (2008).

  61. 61.

    Micheli, F. et al. Cumulative Human Impacts on Mediterranean and Black Sea Marine Ecosystems: Assessing Current Pressures and Opportunities. PLOS ONE 8, e79889 (2013).

  62. 62.

    Jonsen, I. D., Myers, R. A. & James, M. C. Identifying leatherback turtle foraging behaviour from satellite telemetry using a switching state-space model. Mar. Ecol. Prog. Ser. 337, 255–264 (2007).

  63. 63.

    Breed, G. A., Costa, D. P., Goebel, M. E. & Robinson, P. W. Electronic tracking tag programming is critical to data collection for behavioral time-series analysis. Ecosphere 2, art10 (2011).

  64. 64.

    Nowacek, D. P., Christiansen, F., Bejder, L., Goldbogen, J. A. & Friedlaender, A. S. Studying cetacean behaviour: new technological approaches and conservation applications. Anim. Behav. 120, 235–244 (2016).

  65. 65.

    Carter, M. I. D., Bennett, K. A., Embling, C. B., Hosegood, P. J. & Russell, D. J. F. Navigating uncertain waters: a critical review of inferring foraging behaviour from location and dive data in pinnipeds. Mov. Ecol. 4, 25 (2016).

  66. 66.

    Hays, G. Key Questions in Marine Megafauna Movement Ecology. Trends Ecol. Amp Evol. 31, 463–475 (2016).

  67. 67.

    Allen, A. M. & Singh, N. J. Linking Movement Ecology with Wildlife Management and Conservation. Front. Ecol. Evol. 3 (2016).

  68. 68.

    McGowan, J. & Possingham, H. P. Commentary: Linking Movement Ecology with Wildlife Management and Conservation. Front. Ecol. Evol. 4 (2016).

  69. 69.

    Guzman, H. M., Gomez, C. G., Guevara, C. A. & Kleivane, L. Potential vessel collisions with Southern Hemisphere humpback whales wintering off Pacific Panama. Mar. Mammal Sci. 29, 629–642 (2013).

  70. 70.

    Kennedy, A. S., Zerbini, A. N., Rone, B. K. & Clapham, P. J. Individual variation in movements of satellite-tracked humpback whales Megaptera novaeangliae in the eastern Aleutian Islands and Bering Sea. Endanger. Species Res. 23, 187–195 (2014).

  71. 71.

    International Whaling Commission. Report of the Joint NMFS-IWC preparatory workshop ‘Towards Ensemble Averaging of Cetacean Distribution Models’. J. Cetacean Res. Manag. 17(Suppl), 601–609 (2016).

  72. 72.

    Lauriano, G., Pierantonio, N., Donovan, G. & Panigada, S. Abundance and distribution of Tursiops truncatus in the Western Mediterranean Sea: An assessment towards the Marine Strategy Framework Directive requirements. Mar. Environ. Res. 100, 86–93 (2014).

  73. 73.

    Lauriano, G., Panigada, S., Casale, P., Pierantonio, N. & Donovan, G. P. Aerial survey abundance estimates of the loggerhead sea turtle Caretta caretta in the Pelagos Sanctuary, northwestern Mediterranean Sea. Mar Ecol Prog Ser 437, 291–302 (2011).

  74. 74.

    Franqueville, C. Macroplancton profond (invértebrés) de la Méditerranée Nord-Occidentale. Tethys 11–56 (1971).

  75. 75.

    Orsi Relini, L. et al. Meganyctiphanes norvegica and fin whales in the Ligurian Sea: new seasonal patterns. Eur. Res. Cetaceans 179–182 (1994).

  76. 76.

    Geijer, C. K. A., Notarbartolo di Sciara, G. & Panigada, S. Mysticete migration revisited: are Mediterranean fin whales an anomaly? Mammal Rev. 46, 284–296 (2016).

  77. 77.

    Bauer, R. K., Fromentin, J.-M., Demarcq, H., Brisset, B. & Bonhommeau, S. Co-Occurrence and Habitat Use of Fin Whales, Striped Dolphins and Atlantic Bluefin Tuna in the Northwestern Mediterranean Sea. PLoS ONE 10, e0139218 (2015).

  78. 78.

    Arcangeli, A., Marini, L. & Crosti, R. Changes in cetacean presence, relative abundance and distribution over 20 years along a trans-regional fixed line transect in the Central Tyrrhenian Sea. Mar. Ecol. 34, 112–121 (2013).

  79. 79.

    Panigada, S. et al. Estimating cetacean density and abundance in the Central and Western Mediterranean Sea through aerial surveys: implications for management. Deep Sea Res. Part II Top. Stud. Oceanogr (2017).

  80. 80.

    Vaes, T. & Druon, J.-N. Mapping of potential risk of ship strike with fin whales in the Western Mediterranean Sea. A scientific and technical review using the potential habitat of fin whales and the effective vessel density (European Commission Joint Research Centre Institute for the Protection and Security of the Citizen) (2013).

  81. 81.

    Castellote, M., Clark, C. W. & Lammers, M. O. Acoustic compensation to shipping and airgun noise by Mediterranean fin whales (Balaenoptera physalus). Acoust. Soc. Am. J. 127, 1726 (2010).

  82. 82.

    Castellote, M., Clark, C. W. & Lammers, M. O. In The Effects of Noise on Aquatic Life (eds. Popper, A. N. & Hawkins, A.) 321–321 (Springer New York, 2012).

  83. 83.

    Castellote, M., Clark, C. W. & Lammers, M. O. Acoustic and behavioural changes by fin whales (Balaenoptera physalus) in response to shipping and airgun noise. Biol. Conserv. 147, 115–122 (2012).

  84. 84.

    Tringali, L. M., Caltavuturo, G., Gurrieri, G. & Di Martino, V. Osservazioni preliminari sulla presenza estiva di balenottera comune (Balaenoptera physalus) e tursiope (Tursiops truncatus) nelle acque dello Stretto di Messina. Biol. Mar. Mediterr. 6, 619–622 (1999).

  85. 85.

    Pierantonio, N. & Bearzi, G. Review of fin whale mortality events in the Adriatic Sea (1728–2012), with a description of a previously unreported killing. Mar. Biodivers. Rec. 5, e109 (2012).

  86. 86.

    Aïssi, M. et al. Large-scale seasonal distribution of fin whales (Balaenoptera physalus) in the central Mediterranean Sea. J. Mar. Biol. Assoc. U. K. 88, 1253–1261 (2008).

  87. 87.

    International Whaling Commission. JOINT IWC/ACCOBAMS WORKSHOP ON reducing risk of collisions between vessels and cetaceans (2010).

  88. 88.

    Hartel, E. F., Constantine, R. & Torres, L. G. Changes in habitat use patterns by bottlenose dolphins over a 10-year period render static management boundaries ineffective. Aquat. Conserv. Mar. Freshw. Ecosyst. 25, 701–711 (2015).

  89. 89.

    Hyrenbach, K. D., Forney, K. A. & Dayton, P. K. Marine protected areas and ocean basin management. Aquat. Conserv. Mar. Freshw. Ecosyst. 10, 437–458 (2000).

  90. 90.

    Lascelles, B. et al. Migratory marine species: their status, threats and conservation management needs. Aquat. Conserv. Mar. Freshw. Ecosyst. 24, 111–127 (2014).

  91. 91.

    Briscoe, D. K., Maxwell, S. M., Kudela, R., Crowder, L. B. & Croll, D. Are we missing important areas in pelagic marine conservation? Redefining conservation hotspots in the ocean. Endanger. Species Res. 29, 229–237 (2016).

  92. 92.

    Hazen, E. L. et al. WhaleWatch: a dynamic management tool for predicting blue whale density in the California Current. J. Appl. Ecol. doi:10.1111/1365-2664.12820 (2016).

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We are grateful to Ygor Geyer for tag deployment during 2012 and 2013. Many thanks to Giulia Bergamaschi for the support during fieldwork in the Pelagos Sanctuary and to Joan Gonzalvo Villegas and Valentina Corrias for the support in the Strait of Sicily. Thanks to Paolo Pinto and Roberto Raineri (Flash Vela d’Altura) for providing the sailing boat for fieldwork in the Pelagos Sanctuary. We are in debt with the personnel of Area Marina Protetta Isole Pelagie, namely to Fabio Giardina, Salvatore Taranto, Giovanni Raptis and Pietro de Rubeis, for their invaluable support in Lampedusa and for providing the inflatable for the research. This research was funded thanks to a series of grants from the Italian Ministry of the Environment. We are thankful to the Regional Activity Centre for Specially Protected Areas (RAC/SPA) in Tunis, for additional support to fieldwork in the Strait of Sicily.

Author information


  1. Tethys Research Institute, c/o Acquario Civico, Viale G.B. Gadio 2, 20121, Milan, Italy

    • Simone Panigada
    • , Nino Pierantonio
    • , Margherita Zanardelli
    •  & Giuseppe Notarbartolo di Sciara
  2. International Whaling Commission, The Red House, 135 Station Road, Impington, CB24 9NP, Cambridge, United Kingdom

    • Gregory P. Donovan
  3. European Commission, DG Joint Research Centre, Directorate D – Sustainable Resources, Unit D.02 Water and Marine Resources, Via Fermi, TP 051, 21027, Ispra, (VA), Italy

    • Jean-Noël Druon
  4. Institute for Environmental Protection and Research - ISPRA, Via V. Brancati 60, 00144, Rome, Italy

    • Giancarlo Lauriano
  5. School of Mathematics, Washington State University, 14204 NE Salmon Creek Ave, Vancouver, WA, 98686, USA

    • Enrico Pirotta
  6. National Marine Mammal Laboratory, Alaska Fisheries Science Center – NOAA, 7600 Sand Point Way N.E., Seattle, WA, 98115-6349, USA

    • Alexandre N. Zerbini
  7. Cascadia Research Collective, Olympia, WA, USA

    • Alexandre N. Zerbini
  8. Instituto Aqualie, Juiz de Fora, Minas Gerais, Brazil

    • Alexandre N. Zerbini


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S.P., N.P., M.Z., G.D., A.N.Z. and G.N.S. conceived the study. S.P., G.L., N.P., M.Z., and G.D. collected the cetacean telemetry data. S.P., E.P., A.N.Z., and J.N.D. conducted the habitat and movement analysis. S.P. and G.D. wrote the manuscript and N.P., J.N.D., E.P. and G.L. prepared the figures. All authors reviewed and edited the manuscript.

Competing Interests

The authors declare that they have no competing interests.

Corresponding author

Correspondence to Simone Panigada.

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