Basking Shark

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Marine Protected Areas and Basking sharks

Position paper Summary

1. 2. 3. 4. 5. 6. 7. Basking sharks are included on the MPA search feature list for Scottish territorial waters only (Marine Scotland, 2011a). Currently there are no existing protected area measures that help to safeguard basking sharks and they have been identified as a search feature priority in the South-west and West MPA regions (Carruthers et al., 2011). On the basis of the available evidence, one search location is proposed for basking sharks to take forward for further assessment against the MPA Selection Guidelines - Skye to Mull (see Figure 1 overleaf). Proposed solely for basking shark and minke whale, the role of the Skye to Mull search location is to contribute to the protection of the following (as cited on page 15 of the MPA Selection Guidelines):  significant aggregations or communities of important marine species in Scottish waters;  essential areas for key life cycle stages of important mobile species that persist in time, including habitats known to be important for reproduction and nursery stages; and,  areas contributing to the maintenance of ecosystem functioning in Scottish waters (currently of more direct relevance to observed use of parts of the Skye to Mull search location by minke whales). The evidence used to support the Skye to Mull search location for basking sharks is:  presence of consistent (seasonal) large numbers of sharks based on effort corrected data;  presence of large groups or aggregations of basking sharks (i.e. only observed in the proposed search location despite surveys covering large areas around the west of Scotland); and,  presence of social and courtship-like behaviour (i.e. only observed in this area despite surveys covering large areas around the west of Scotland). Further work is required to refine our understanding of critical habitats for basking sharks. This will help determine whether current evidence is sufficient to underpin further assessment against the MPA Selection Guidelines. Basking sharks also depend on areas in deeper water that support high productivity; however, there are currently insufficient data to enable the identification of search locations for such areas within Scottish waters (the principle could also apply to the offshore area). Ensuring replication and coherence of the MPA network for basking sharks is difficult to ascertain on the current evidence available, perhaps placing heightened importance to the Skye to Mull area. It may be possible to add value to relevant deep water MPAs in the future (by including basking sharks as a protected feature), therefore providing some degree of replication. This would be subject to the conclusions of any new survey / tagging work and ongoing work to define the large-scale fronts MPA search feature. 1

Figure 1 The Skye to Mull MPA search location. Hatched areas represent the Small Isles and Coll and Tiree MPA search locations. The Skye to Mull search location superseded the Coll and Tiree location (dropped) following discussions at the 3 rd MPA stakeholder workshop (October 2011)

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Purpose of document

8. This paper outlines the rationale for, and the progress made to date in the identification of MPA search locations for basking sharks within Scottish waters. The paper has been prepared to support discussions at the 4 th national MPA stakeholder workshop on March 14 th and 15 th , 2012.

Background

9. Two initial MPA search locations identified for basking sharks were discussed at the 3 rd national MPA stakeholder workshop in October 2011; the Small Isles and, Coll and Tiree (Figure 1). The Small Isles were also identified as a search location for minke whale, shelf deeps, burrowed mud, and fan mussel and northern feather star aggregations. Additional MPA search features within the Coll and Tiree search location included minke whale, black guillemot, shelf banks and mounds, and shelf deeps. 10. During discussions at the 3 rd workshop, participants identified wider areas around the initial search locations that were appropriate for further investigation for both basking shark and minke whale. The new Skye to Mull search location was developed from these discussions, for the two mobile species interests only and based on available effort-corrected data. The Skye to Mull search location has subsumed the basking shark and minke whale features from the original Small Isles proposal. The Small Isles MPA search location is progressing to a more detailed assessment against the Scottish MPA Selection Guidelines for a range of other seabed habitats and species.

Distribution in Scottish waters

11. Basking sharks are found globally in both the Pacific and Atlantic oceans (see Figure 2), predominantly in coastal waters and over the continental shelf of temperate waters (Compagno, 2001).

Figure 2 Global basking shark distribution (from FAO website)

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12. Around the UK, basking sharks are sighted all around the coast but predominantly around the south-west peninsula of England and around the west coast of Scotland (MCS website; Bloomfield and Soldant, 2007). Sightings are seasonal with increased numbers of basking sharks observed from May through to September (Speedie et al., 2009). 13. There are currently no population estimates for basking sharks, however, there are well documented declines in historic fisheries landings data with no evidence of recovery to the high levels once seen (OSPAR, 2009).

The three-pillar approach to marine nature conservation

14. Marine Scotland ’s Marine Nature Conservation Strategy (Marine Scotland, 2011b) advocates a 3-pillar approach to conservation in the marine environment; with the 3 pillars representing species-specific measures, site protection and a wider seas policy approach. For basking sharks there are currently species measures in place which afford some protection. 15. Basking sharks are an OSPAR Threatened and Declining (T&D) species, listed as vulnerable worldwide and endangered in the north-east Atlantic (International Union for Conservation of Nature 2004 Red List). 16. Basking sharks are listed on Schedule 5 of the Wildlife and Countryside Act (1981) (as amended) and as such are protected against killing, taking or injuring as well as the intentional or reckless disturbance or harassment. EC Regulation 2555/2001 also provides protection with a zero total allowable catch in European waters. In summary, the current species protection measures focus on targeted fishing pressure and avoiding disturbance. 17. In terms of wider seas policies and measures, there is potential for marine planning to recognise the sensitivities of particular habitats and species and identify locations where they might be considered vulnerable. The extent to which regional marine planning in Scotland might be able to deliver protection of basking shark habitat is currently unknown. 18. Under the protected sites pillar, Nature Conservation MPAs are a tool specifically designed to provide habitat protection and could therefore be used to complement existing species protection for basking sharks. Further consideration of basking shark habitat requirements is required to illustrate the role that MPAs might play in their conservation.

Role of MPAs for basking shark

19. In relation to mobile species, the MPA Selection Guidelines consider that Nature Conservation MPAs are appropriate for contributing to the protection of the following:  significant aggregations of communities of important marine species in Scottish waters;  essential areas for key life cycle stages of important mobile species that persist in time, including habitats known to be important for reproduction and nursery stages; and,  areas contributing to the maintenance of ecosystem functioning in Scottish waters. 4

20. This approach is intrinsically linked to the concept of ‘critical habitats’, areas upon which species are strongly dependent, or areas where species show high fidelity (essential for day-to-day well-being and survival and to maintain healthy populations). 21. For basking sharks these three broad categories can be more clearly articulated according to a number of different types of areas as follows (reflecting our understanding of the ecology of the species): i Significant aggregations of communities of important marine species in Scottish waters; a. Places used regularly for feeding, breeding, calving / spawning and or raising young and socialising. b. Locations where associated and supporting activities (e.g. courtship, resting, playing, communication) take place. c. Locations with regular seasonal concentrations. ii Essential areas for key life cycle stages of important mobile species that persist in time, including habitats known to be important for reproduction and nursery stages; a. Places used regularly for feeding, breeding, calving / spawning and or raising young and socialising. b. Locations where associated and supporting activities (e.g. courtship, resting, playing, communication) take place. c. Locations with regular seasonal concentrations. iii Areas contributing to the maintenance of ecosystem functioning in Scottish waters; a. Areas where important ecosystem processes occur that support high (primary) productivity of (pelagic) prey species (e.g. thermal fronts). 22. Within territorial waters, the perceived MPA role in relation to basking sharks is to provide protection to significant aggregations of sharks and to essential areas for key life cycle stages. The MPA designation will ensure specific conservation objectives are met and that an assessment is carried out for activities likely to impact basking sharks. This will offer heightened and appropriate protection in those areas where basking sharks are potentially most sensitive.

Identifying MPA search locations for basking sharks

23. In order to identify MPA search locations for basking sharks (i.e. where basking sharks are drivers / the primary reason for designation), consistent observations of all three of the following parameters are required:  presence of large numbers of basking sharks (based on effort-corrected data);  presence of large groups or aggregations of basking sharks; and,  presence of social and / or courtship-like behaviour. 24. These three parameters have been used to identify important areas previously (Speedie et al., 2009). 25. Basking sharks also utilise areas at depth within territorial waters e.g. feeding in areas of high productivity such as the Islay front or within the Sea of Hebrides. However, there are currently insufficient data to draw any conclusions about the relative importance of any specific locations. Areas of high productivity associated with the continental slope edge (where sharks are presumed to be feeding), are also used by 5

basking sharks and may represent essential areas for key life cycle stages. It is therefore recommended that basking sharks could add value (recognition as a protected feature) to search locations identified primarily for other search features, particularly where evidence indicates they are associated with large-scale MPA search features such as fronts (areas contributing to the maintenance of ecosystem functioning in Scottish waters).

Evidence base

26. A summary of all the information available which provides an evidence base for the existence of areas which may be appropriate for safeguarding basking sharks is discussed in this section.

Historical fishing effort

27. Historic fishing areas for basking sharks in Scottish waters are presented in Sims et al., (2005). The three key areas were the Firth of Clyde, around the Hebrides / throughout the Minch and in waters to the west of Shetland (see Figure 3).

Figure 3

Historic fishing areas (from Sims et al., 2005) 6

Surface sightings and feeding behaviour

28. The Marine Conservation Society collates sightings data for basking sharks in the UK (Figure 5). Whilst sightings data provide some information on the distribution of basking sharks, there is a considerable degree of bias towards those areas that are easy to get to and / or are visible from land, increasing the chances of sighting sharks. 29. Effort-corrected data reduces the bias that exists in sightings data. Observations are made along a predetermined transect line and expressed as a unit of effort (time or distance), i.e. sharks observed per unit effort (SPUE). Some bias is also shown in effort-corrected data as observations are of sharks at the surface of the water, and do not take account of shark distribution at all other times (e.g. when feeding at depth). 30. However, effort-corrected data are considered the best representation of distribution while basking sharks are at the surface of the water. It is for these reasons that effort corrected data have been used to provide the evidence base for identifying MPA search locations. 31. Effort-corrected sightings from boat transect surveys were carried out between 2002 and 2004 within the Clyde Sea and 2002 - 2006 in the Sea of the Hebrides, as part of The Wildlife Trusts Basking Shark Project (Speedie et al., 2009). The Hebridean Whale and Dolphin Trust (HWDT) has also collected effort-corrected data on the west coast of Scotland since 2003. 32. Results indicate that high numbers of surface feeding sharks were found consistently and seasonally in a number of areas on the west coast of Scotland. Both surveys recorded high SPUE in the areas around Coll and Tiree, and Hyskeir and Canna (Figure 4). Further detail on the annual SPUE from both surveys can be found in Annex 1. 33. The area to the east of the Bishop’s Isles (south of Barra) showed the next highest levels of SPUE (Speedie et al., 2009, and Figure 4). The HWDT data also highlights the area to the north-west of Lewis for medium levels of SPUE (Figure 4). 34. Historically, the Clyde Sea has been an area with high numbers of basking shark observations. However, this is not reflected in the effort-corrected data available for the period 2002 - 2004 (Speedie et al., 2009). There maybe some more recent data available from this area (Rupert Ormond pers. comm. 2011), and this will be followed up as detailed in the Issues and Opportunities section.

Feeding behaviour

35. Basking sharks feed on zooplankton, selectively foraging for Calanus helgolandicus, and shark presence is strongly linked with fronts where nutrients and plankton are concentrated (Sims and Quayle, 1998). The behaviour of basking sharks occupying waters that are well-stratified is different from that of sharks occupying tidal front regions and mimics the behaviour of their prey (Sims et al., 2003a). This is attributed to Calanus sp. which can show reverse diel migration in response to presence of their predators (chaetognaths). There is therefore, a clear link between basking sharks and underlying environmental features. Further work will help to clarify these links for specific search locations (see the Issues and Opportunities section). 7

Figure 4 Effort-corrected basking shark observation data on the west coast of Scotland. 4a. Number of sharks observed per km (from The

Wildlife Trusts Basking Shark Project - Speedie et al., (2009)), & b. Number of sharks per km (HWDT data). 8

Figure 5 a. MCS sightings data for the UK. Figure 6 Average SPUE (sharks per unit effort) for areas around Scotland, Area 15 is Coll

and Tiree and Area 17 is Canna and Hyskeir (adapted from Speedie et al., 2009) Area 11 12 13 14 15 16 17 18 19 20 Sharks observed 11 0 19 22 251 9 268 2 0 11 Hours on survey 225.20 0.88 82.17 41.68 144.52 185.22 95.03 82.12 80.77 18.27 SPUE Sharks / hr 0.05 0.00 0.23 0.53 1.74 0.05 2.82 0.02 0.00 0.60 9

Relative abundance

36. Average SPUE values of 1.74 and 2.82 were recorded in Canna and Hyskeir, and Coll and Tiree (areas highlighted in Figure 6). These are SPUE values calculated within the area over the whole survey period (2002 to 2006). The highest average SPUE value recorded in other parts of the UK using comparable survey methodology and area is 0.48 (The Wildllife Trusts, 2008). The high average values demonstrate that high SPUE have been found consistently throughout the survey period and that the relative abundance of basking sharks is particularly high in Scotland.

Large groups of sharks

37. In addition to the consistently high average SPUE values recorded throughout the study period, Hyskeir and Canna along with Coll and Tiree were two areas where there were consistent observations of large shoals of basking sharks, with counts reaching over 50 individuals (Speedie et al., 2009). Such observations have only been made from these two areas in Scottish waters. Further detail on the annual aggregations of basking sharks observed from both surveys can be found in Annex 1.

Social behaviour

38. Observations of basking shark behaviour have been limited to sharks at the sea surface. Courtship-like behaviour (e.g. nose to tail following, parallel synchronised swimming, close approach involving rostral contact) was observed only around Hyskeir and Canna, and Coll and Tiree (Speedie et al., 2009). Breaching was performed by individuals that were part of a group of sharks and breaching sharks were also involved in courtship-like behaviour. The majority of breaching behaviour also occurred around Hyskeir and Canna, and Coll and Tiree (Speedie et al., 2009). Behavioural data was not routinely collected on the HWDT surveys, however, observations of breaching and courtship-like behaviour that were recorded were only in the areas around Hyskeir and Canna, and Coll and Tiree in the years 2005 to 2011. 39. Areas that are important for social behaviour such as courtship could also be important for breeding (Sims et al ., 2000). Similar ‘nose to tail following’ behaviour in other sharks are known to be followed by copulation (e.g. Carrier and Pratt, 1998). The importance of such behaviour to basking shark populations should be a key consideration in their protection.

Breeding

40. There is an unconfirmed report of a basking shark foetus measuring 1.7 m long, suggesting that this is the size of individuals near birth, considerably larger than other sharks (FAO species fact sheet). Sightings records indicate that basking sharks below 2 m in length have been recorded all around the UK coast (Bloomfield and Solandt, 2007) and particularly in SW England and the Sea of the Hebrides (Speedie pers. comm.); this could indicate that the first year class of basking shark populations utilise our coastal waters - if pups are indeed around 1.7 m in length. 10

Sharks feeding at depth

41. Evidence suggests that sharks migrate south during autumn and winter, spending more time in deeper water tracking the continental shelf edge to various hotspots of productivity (e.g. Sims et al., 2003b; Sims et al., 2005). Only one shark from these satellite tagging studies spent a relatively long period in Scottish waters; this shark moved from Plymouth to Scotland via the west coast of Ireland tracking the continental shelf edge, where it spent 1.5 months (August to October) in the Sea of the Hebrides area, suggesting it is important for feeding (Figure 7).

Figure 7

a. Location of tidal and shelf break fronts in UK waters (Sims et al., 2005). b.

Tagged shark track showing position individual spent 1.5 months in the Sea of

Hebrides (Sims et al., 2005). c. Tagged shark track from Stephen et al., 2011. d.

Front locations from GeMS (Geodatabase of Marine features around Scotland)

a. b. c. d. 11

42. The most recent published work on satellite tagging of basking sharks showed one shark that moved northwards from the Iroise Sea in France to the Hebrides via the west of Ireland. Other tagged sharks in this study spent significant periods of time in deeper waters on the continental shelf edge of the Irish and Celtic seas during winter months (Stephan et al., 2011). Tagging work in Scotland has shown an individual basking shark spent time in deeper water around the Islay front (Ormond pers comm. 2011). 43. Whilst the link between deeper water high primary productivity and shark presence is clear, the identification of particular fronts in deeper water that basking sharks use in Scotland is difficult to establish based on the current tagging results. It is recommended that the evidence showing basking sharks utilise deeper water for feeding, and therefore may be essential to their life cycle, should be used to add value to MPA proposals (rather than serving as a driver) where other features underpin the identification of search locations.

Basking shark population structure

44. Basking sharks in Scotland may not be discrete populations as tagging research indicates that sharks can travel the length of the UK in time periods of a few weeks (Sims et al., 2005). There is one recorded incident of basking shark transatlantic migration (Gore et al., 2008) and genetic studies have shown no significant difference between the mitochondrial DNA of Atlantic and Pacific basking sharks (Hoelzel et al., 2006).

Summary of evidence base available to inform MPA identification

45. On the basis of the evidence available, the following conclusions can be drawn regarding basking shark presence and distribution in Scottish waters:  The relative abundance of basking sharks is particularly high in Scotland, with the highest average SPUE values recorded being much greater than in other parts of the UK;  Basking sharks feed on zooplankton, selectively foraging for Calanus helgolandicus, and shark presence is strongly linked with fronts where nutrients and plankton are concentrated;  Basking sharks also spend time away from the sea surface, and highly productive deep water frontal areas are also considered to serve as essential areas for key life cycle stages. However, there is currently insufficient direct evidence to identify particularly important deep water locations around Scotland;  Two areas show high average SPUE values (number of sharks encountered on transects) - the areas around Coll and Tiree, and Hyskeir and Canna;  Consistently high SPUE data is available for 4 and 9 consecutive years (Speedie et al., 2009 and HWDT data respectively) for the Coll and Tiree, and Hyskeir and Canna areas;  Large shoals of basking sharks have been recorded consistently from these two locations; as have,  High levels of social behaviour including breaching and courtship-like behaviour. 46. Whilst populations may span the globe, it is considered that MPAs for basking sharks in Scotland could make a valuable contribution to the protection of this OSPAR T&D species. 12

47. Table 1 summarises the evidence base for each of the types of areas that could be

used to help contribute to the protection of basking sharks (listed in paragraph 16) and recommends which should be considered further in terms of supporting identification of MPA search locations for basking sharks in Scottish territorial waters.

Table 1 Summary of the evidence base for the types of areas that could be used to help contribute to the protection of basking sharks in territorial waters

Type of area

Places used regularly for feeding

and

Locations with regular seasonal presence Locations where associated and supporting activities take place

Evidence base for area type

Consistent high levels of SPUE of surface feeding of sharks for 4 and 9 years (Speedie et al., 2009 and HWDT data respectively) Functional link to zooplankton at locations (Sims and Quayle, 1998; Sims et al., 2003a) Tagging data showing sharks spend time in deeper water corresponding to high productivity (Sims et al, 2005 and Ormond pers comm.) High SPUE relative to rest of UK (Speedie et al., 2009) Presence of aggregations or large groups (50+ individuals) Courtship-like behaviour (Speedie et al., 2009 and HWDT data) Areas where important ecosystem processes occur that support high productivity of prey species Pupping in UK waters (FAO website) Sightings of first year class around UK (Bloomfield and Soldant, 2007) Fronts of mixed water supporting high productivity of prey zooplankton Calanus sp. linked to basking shark feeding behaviour (Sims and Quayle, 1998)

Recommendation

Consider locations where high levels of SPUE occur Confirms link between feeding behaviour and prey presence Limited evidence in Scottish waters therefore not considered further Consider locations where high levels of SPUE occur Consider locations where large aggregations occur Consider locations where social and or courtship-like behaviour occur Lack of evidence so not considered further Linked to assumption of pup size so not considered further Consider areas supporting high productivity of prey (zooplankton) which are linked to basking shark feeding behaviour

MPA search locations

48. One search location is proposed for basking sharks to take forward for further assessment - Skye to Mull (Table 2 and Figure 8). The boundary of this location has been altered slightly since that proposed in the post-workshop 3 interim report 1 , to reflect new data from the HWDT. 49. Table 2 also presents summary conclusions regarding 3 other areas considered as part of this review (on the basis of historical or more recent effort-corrected data indicating the presence of large numbers of basking sharks). There is currently insufficient evidence to identify an MPA search location for basking sharks within any of these areas. 1 Available online at - http://www.scotland.gov.uk/Resource/Doc/295194/0123094.doc

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Table 2

Area

Skye to Mull Clyde Sea

Summary of areas where basking sharks were assessed as a potential driver in the identification of MPA search locations; the unshaded row is the proposed Skye to Mull search location, shaded rows are areas not recommended for progression on the basis basking sharks

Bishop Isles (east of Barra) North-west Lewis

OSPAR region

III III III III

Other mobile species MPA search features

Minke whale n/a n/a n/a

Comments / Evidence

MPA search location developed following 3 rd MPA workshop Two key areas show high SPUE, large groups of animals and social behaviour Historic data but poor recent SPUE - discount Medium SPUE with limited data (Speedie et al., 2009) - discount Medium SPUE with limited data (HDWT) - discount

Developing approach to management

50. Work on MPA management requirements is being considered in parallel to the identification of the search locations. Work is underway to develop a tool which will help identify key pressures that search features may be exposed to in the marine environment. Key pressures identified for basking sharks include physical pressures (e.g. loss of, or displacement from, essential areas for key life stages; death or injury by collision with vessels); biological pressures (e.g. entanglement in fishing gear, acoustic impacts); and hydrological changes (e.g. changes in temperature, salinity and water flow that may alter prey availability in essential areas for key life cycle stages). 51. Our current knowledge of the interactions between basking sharks and the key pressures highlighted varies. The identification of these pressures is the first step in the process and further investigation of the interactions will be necessary, particularly when considering potential management measures (requiring input from stakeholders and basking shark ecologists). 52. Land based observations from the Isle of Man indicate that surface feeding and courtship-like behaviour, in both individuals and large groups of basking sharks, may be disrupted by the presence of boats (Galpin, 2009). Such displacement may go unnoticed by boat users due to their lower vantage point and the limited wider context they have during their observations. However, observations during many boat surveys in Scotland indicate that sharks engaged in courtship are less troubled by boats than Galpin suggests, although difficult to confirm (Speedie pers comm.). 53. Voluntary management measures are in place for boat operators in the two areas where high concentrations of basking sharks have been identified previously (Speedie et al., 2009). Work through the WiSe scheme (a marine wildlife training scheme for tour operators) and the Royal Yachting Association has ensured publicity of these areas and voluntary measures to commercial ecotourism operators and recreational boat users. However, the effectiveness of these measures is currently uncertain. 14

Figure 8 The Skye to Mull MPA search location with underpinning effort-corrected basking shark observation data

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54. Recent analyses from results of effort-corrected data (Bradbury, 2011) have linked SPUE with environmental parameters such as distance from shoreline, time of day and tidal cycle, indicating that it may be possible to further define essential areas for key life stages. Zooplankton availability and water temperature were also important parameters, but additional work is required to clarify their significance.

Issues and opportunities

55. The main issue is the gap in effort-corrected data coverage. There could be other areas essential to key life stages for basking sharks elsewhere in Scotland, which have not been highlighted for consideration in the MPA process so far. 56. Replication in the MPA network for basking sharks is not currently considered possible, due to the lack of data and evidence. It may be possible to achieve replication within the network in the future by adding value to other MPA search locations (by recognising basking sharks as a protected feature). 57. There are new data being gathered from renewables work around the south-west of Tiree which could aid in the subsequent more detailed assessment of the Skye to Mull search location against the MPA Selection Guidelines. However, these data are not yet available to the Scottish MPA Project (commercially sensitive information). 58. Further data may also be available from land based observations in Orkney (still to be investigated) and additional effort-corrected data from the Clyde Sea. Similarly, further work is underway on defining and mapping the distribution of the large scale MPA features including fronts. The results of these studies will clarify overlaps with other MPA search locations. 59. A scoping study is planned in 2012 to determine how we can improve our understanding of why and how basking sharks are using the areas highlighted as essential for key life cycle stages to see if such areas can be defined on the basis of environmental parameters (habitat modelling).

Next steps

60. Following discussion at the 4 th MPA workshop, there may be revisions to the MPA search location boundary. Once agreed, the search location will be subject to a more detailed assessment against the Scottish MPA Selection Guidelines. This will take account of the results of any habitat modelling work, new data and further investigation of basking shark interactions with key pressures.

References

Bloomfield, A and Soldant, J.L. (2007). The Marine Conservation Socierty Basking Shark Watch. 20 years report (1987-2006). Marine Conservation Society report. Bradbury, C. (2011). Bio-physical characteristics of basking shark, Cetorhinus maximus, habitats and sightings in the Sea of the Hebrides, Scotland. MSc Thesis, University of Aberdeen.

Carrier, J.C. and Pratt, HL. (1998) Habitat management and closure of a nurse shark breeding and nursery ground. Fisheries Research. 39. (2) 209-213.

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Carruthers, M., Chaniotis, P.D., Clark, L., Crawford-Avis, O., Gillham, K., Linwood, M., Oates, J., Steel, L., and Wilson, E. (2011). Contribution of existing protected areas to the MPA network and identification of remaining MPA search feature priorities. Internal report produced by SNH, JNCC and MS for the Scottish MPA Project. Compagno, L.V.J. (2001). Sharks of the world. An annotated and illustrated catalogue of

shark species known to date. Vol 2. Bullhead, mackerel and carpet sharks

(Heterodontiformes, Lamniformes and Orectolobiformes). Rome: FAO Species Catalogue for fisheries Purposes, FAO. FAO species fact sheet - http://www.fao.org/fishery/species/2005/en Galpin, J. (2009). Land based observation of annual basking shark behaviour and responses to disturbance, Dalby Point, Isle of Man. 1995 to 2009. Unpublished Basking shark conference paper, Isle of Man, 2009 Gore, M., Rowet, D., Hall, J., Gell, F.R. and Ormond, R. (2008). Transatlantic migration and deep mid-ocean diving by basking shark. Biology Letters, 4. 394-398. Hoelzel, A.R., Shivji, M.S., Magnussen, J. and Francis, M.P. (2006). Low worldwide genetic diversity in the basking shark (Cetorhinus maximus). Biology Letters, 2, 639-642. Marine Scotland (2011a). Marine Protected Areas in the Seas around Scotland. Guidelines on the selection of MPAs and development of the MPA network. Available from Marine Scotland (2011b). A Strategy for Marine Nature Conservation in Scotland. Available from MCS web http://www.mcsuk.org/what_we_do/Wildlife+protection/Reports+and+downloads/ Reports+and+downloads OSPAR Commission (2008). Case Reports for the OSPAR List of Threatened and/or Declining Species and Habitats. OSPAR Commission. Biodiversity Series. Available from < http://qsr2010.ospar.org/media/assessments/p00358_case_reports_species_and_ habitats_2008.pdf

> OSPAR Commission (2009). Background document for Basking shark Cetorhinus maximus. Biodiversity series, 2009. Available from: < http://qsr2010.ospar.org/media/assessments/Species/P00419_basking_shark.pdf

> Sims, D.W. and Quayle, V.A. (1998). Selective foraging behaviour of basking sharks in a small-scale front. Nature, 393, 460-464. Sims, D.W., Southall, E.J., Quayle, V.A. and Fox, A.M. (2000). Annual social behaviour of basking sharks associated with coastal front areas. Proceedings of the Royal Society of London. 267. 1897-1904. Sims, D.W., Southall, E.J., Merrett, D. and Saunders, J. (2003a). Effects of zooplankton density and diel period on surface-swimming duration of basking sharks. Journal of the Marine Biological Association of the UK, 83, 643-646. Sims, D.W., Southall, E.J., Richardson, A.J., Reid, P.C. and Metcalfe, J.D., (2003b). Seasonal movements and behaviour of basking sharks from archival tagging: no evidence of winter hibernation. Marine Ecological Progress Series, 248, 187-196. Sims, D.W., Southall, E.J., Metcalfe, J.D., and Pawson, M.G. (2005). Basking shark 17

population assessment. Final report for Global Wildlife Division of Defra. Speedie, C.D., Johnson, L.A., and Will, M.J. (2009). Basing shark hotspots on the West Coast of Scotland: Key sites, threats and implications for conservation of the species. SNH Commissioned Report No.339. Stephan, E., Gadenne, H., and Jung, A. (2011). Satellite tracking of basking sharks in the North-east Atlantic ocean. APECS Final report. The Wildlife Trusts. (2008). Basking shark hotspots in the UK. Results from The Wildlife Trusts’ basking shark survey. 18

Annex 1.

Figure 9: Annual SPUE (basking sharks per km on transect) from The Wildlife Trusts Basking

Shark Project (Speedie, et al., 2009) and the HWDT surveys. 19

Figure 9 continued: Annual SPUE (basking sharks per km on transect) from the HWDT surveys.

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Figure 10: Annual aggregations of basking sharks (groups of sharks) from The Wildlife Trusts

Basking Shark Project (Speedie, et al., 2009) and the HWDT surveys. 21

Figure 10 continued: Annual aggregations of basking sharks (groups of sharks) from the HWDT survey.

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