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Biological Conservation

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Missing native oyster (Ostrea edulis L.) beds in a European Marine ProtectedArea: Should there be widespread restorative management?

Jose M. Fariñas-Francoa,f, Bryony Pearcea,b, James M. Maira, Dan B. Harriesa,Rebecca C. MacPhersona, Joanne S. Porterd, Paula J. Reimerc, William G. Sandersona,e,⁎

a Centre for Marine Biodiversity and Biotechnology, ILES, School of Energy, Geoscience, Infrastructure and Society, Heriot-Watt University, Edinburgh EH14 4AS, UKb Pelagica Limited, 2 School House Cottages, Market Street, Hoylake, Wirral CH47 3EP, UKc 14CHRONO, Centre for Climate, the Environment and Chronology, Queen's University Belfast, Belfast BT7 1NN, UKd International Centre for Island Technology, School of Energy, Geoscience, Infrastructure and Society, Heriot-Watt University, The Old Academy, Stromness, OrkneyKW16 3AW, UKe St. Abbs Marine Station, St Abbs, Scottish Borders TD14 5QF, UKf Plant Systems Biology Laboratory, School of Natural Sciences and Ryan Institute, National University of Ireland, Galway, University Road, Galway, Ireland

A R T I C L E I N F O

Keywords:OysterOstrea edulisConservation managementMPA, baseline, restoration

A B S T R A C T

Anthropogenic pressures on the marine environment have escalated and shellfish habitats have declined sub-stantially around the world. Recently, Marine Protected Areas (MPAs) have rapidly increased in number, butmanagement baselines rarely account for historical conditions. Marine examples of habitat restoration aretherefore unusual.

An interdisciplinary review of management baselines was undertaken for the Dornoch Firth protected area(NE Scotland) as well as three adjacent inlets and 50 km of open coastline. The protected area has low levels ofindustrial development, is sparsely populated, and previously achieved management objectives.

Here we systematically searched for historical evidence of native oyster (Ostrea edulis) beds, a habitat nowrare and of conservation importance throughout Atlantic Europe. Archaeological records, navigational charts,historical maps, museum collections, land-use records, fisheries records, public online databases and naturalists'records were searched. We conducted intertidal and subtidal surveys and sample oyster shells were radiocarbondated.

The combined interdisciplinary sources showed that O. edulis occurred in the inlets and open coast areas of NEScotland, and specifically in the protected area: Probably since the end of the last glaciation to the late 1800swhen they were likely over-fished. Present environmental conditions are also suitable for oyster restoration.

Habitat restoration in protected areas is an emerging global theme. However, European oyster restorationeffort is currently confined to remnant populations with a clear history of exploitation or dwindling associatedfisheries. An interdisciplinary review of baselines will probably show scope for the restoration of O. edulis, fornature conservation, in many other European MPAs.

1. Introduction

Human populations have long depended on coastal zones and theecosystem services they provide; particularly in terms of food provision,transport, recreation and cultural activities (Mehvar et al., 2018;Neumann et al., 2015). As human populations have grown, so too haveglobal pressures on coastal areas such as overfishing, physical damageto benthic habitats, pollution and eutrophication and these pressuresare regionally intense in places such as Atlantic Europe (Halpern et al.,2015). Among marine habitats, shellfish reefs are especially sensitive to

anthropogenic disturbance (see Cook et al., 2013; Fariñas-Franco et al.,2018) and are one of the most “imperilled” marine ecosystems on Earth(Beck et al., 2009).

In the last 25 years there has been an increasing recognition of theimportance of protecting the marine environment through the desig-nation of Marine Protected Areas (MPAs); now a mainstream globalmanagement tool applied to 7% of the world's marine environment(UNEP-WCMC and IUCN, 2018). In Atlantic European countries such asthe UK, around half of these MPAs are Special Areas of Conservation(SACs) designated under the EU Habitats Directive (Frost et al., 2016).

https://doi.org/10.1016/j.biocon.2018.03.010Received 28 June 2017; Received in revised form 1 March 2018; Accepted 8 March 2018

⁎ Corresponding author at: Centre for Marine Biodiversity and Biotechnology, ILES, School of Energy, Geoscience, Infrastructure and Society, Heriot-Watt University, Edinburgh EH144AS, UK.

E-mail address: w.g.sanderson@hw.ac.uk (W.G. Sanderson).

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0006-3207/ © 2018 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/BY/4.0/).

T

The condition and management plans of SACs, as with almost all MPAs,are typically judged against the ecological condition of the site whenthe legislation was introduced (1992 in the case of the EU HabitatsDirective). These ‘baselines’ thus provide the benchmark against whichchanges are measured (Piazzi et al., 2016).

Since baselines have usually been defined at the time a protectedarea was designated, it is plausible that they might not reflect pastconditions when certain species and habitats were more common,biodiversity was higher or when locally extinct species were present.Indeed, it has previously been observed that conservation and MarineSpatial Planning usually make no reference to historical conditions,specifically those prior to the influence of large scale anthropogenicchanges since the Industrial Revolution (Plumeridge and Roberts,2017). However, assessing historical conditions can be difficult becauseimpacts often pre-date readily available documentation or any kind ofscientific study.

The ‘shifting baselines’ paradigm was first referred to in the scien-tific literature to suggest that the concept of ‘pristine’ in the marineenvironment evolves across different generations of sea users (Pauly,1995; Sáenz-Arroyo et al., 2005). As a result, the past-history of the seacan inform management policies (Roberts, 2007) which are otherwisefounded on baseline conditions derived from limited historical data orcurrent conditions (Gatti et al., 2015). Although the shifting baselinesconcept has been commonly applied within a fisheries context (Ferrettiet al., 2015), it is also applicable to habitats and whole ecosystemswhere losses in biodiversity are typically measured against very recentconditions with little regard to the ‘millennia of habitat loss’ (Airoldiand Beck, 2007; Howarth et al., 2014). Recent initiatives to improve themanagement of the marine environment call for careful considerationof environmental baseline and reference conditions (see OSPARCommission, 2011) because, in the context of ‘sustainable manage-ment’, it is logical to consider what could be achieved as well as what isbeing achieved in terms of benefits to human society.

Shellfish reefs, and oyster reefs in particular, have been severelyimpacted by overexploitation, disease and other human induced pres-sures throughout their global distribution; mostly in the late 1800's andearly 1900's (Beck et al., 2011; Gatti et al., 2015; Howarth et al., 2014).In the USA declines of up to 88% of the American oyster (Crassostreavirginica) biomass have been documented in some waterbodies (e.g.,Chesapeake Bay) with consequent losses in associated ecosystem ser-vices such as water filtration (Zu Ermgassen et al., 2012b). In Europe,the native oyster Ostrea edulis has been a documented source of foodsince at least Roman times and in the North Sea over-exploitation hasled to the loss of extensive beds (Anon., 1885–1886, 1882).

O. edulis is found in Atlantic Europe and North Africa from Moroccoto Norway as well as the Black Sea and in the Mediterranean (Canoet al., 1997; Zaitsev and Alenxandrov, 1998). It usually inhabits rela-tively shallow sheltered estuaries but has been historically recorded indeeper waters of up to 80m off the Channel Islands and Grimsby (UK)and the wider North Sea (Laing et al., 2005). Native O. edulis beds werealso present in Scotland but the advent of industrialised trawling(Roberts, 2007) and the development of the railway system at the endof the 19th century, facilitated a sharp increase in demand by urbanisedmarkets and a corresponding extirpation in the Firth of Forth (Scotland,UK; Fig. 1A) where an oyster (O. edulis) bed was reported to cover anarea of about 10×30 km and to produce up to 30 million oysters insome years (Olsen, 1883; Fulton, 1896; Thurstan et al., 2013). Oysterpopulations in the UK still exist in remote sheltered sea lochs on theScottish west coast, most notably Loch Ryan (University MarineBiological Station Millport, 2007), and in a few inlets and coastal wa-ters of southern England and Wales (Woolmer et al., 2011). Overall, O.edulis habitats are endangered in Europe, losses and threats have beenoverlooked and there is scope for restoration (cf Airoldi and Beck,2007). As a result, in areas with well documented historical O. edulisfisheries, as well as remnant fisheries and populations, the concept oflocalised restoration is gaining traction (Gercken and Schmidt, 2014;

Sawusdee et al., 2015; Smaal et al., 2015; Smyth et al., 2018) and O.edulis beds are now identified as a priority marine habitat for protectionin European MPAs (OSPAR Commission, 2011). However, the EuropeanAtlantic MPA network covers a substantial area and O. edulis popula-tions may well have been ubiquitous in what is now the MPA network:over and above those places where the recent history of fisheries isapparent in the modern context.

Although a commercial fishery for oysters existed in the past inScotland (notably the Firth of Forth; Thurstan et al., 2013), there is nomention of O. edulis in conservation objectives or management planningdocuments for any of the marine protected areas of North-East Scot-land, including the Dornoch Firth (SNH, 2016).

Through an inter-disciplinary review and novel first-hand field in-vestigation, we aimed to determine the likely historical presence ofnative O. edulis beds in the Dornoch Firth protected area and the threeneighbouring inlets and open coast in northeast Scotland. The ultimategoal was to determine if oyster restoration could be relevant to theconservation objectives of the wider European Atlantic MPA network,with its long history of environmental degradation (Gilbert et al.,2014); especially where there are now no remnant populations orcontemporary knowledge of historical fisheries.

2. Materials and methods

2.1. Site details

The Dornoch Firth is a little-studied marine protected area locatedin the remote northeast coast of Scotland with low fishing pressure anda sparse human population. Unlike many areas in the North Sea, theDornoch Firth (Fig. 1A–C) has been virtually unaffected by industrialdevelopment and is considered of high environmental quality (Mackayet al., 2004). The Dornoch Firth is a Special Area of Conservation (SAC)and a Special Protected Area (SPA) under European Directive and, assuch, it is also part of the OSPAR network of MPAs. In addition, theintertidal zone bounding the Dornoch Firth is a Site of Special ScientificInterest (SSSI). Two Nature Conservation Orders banning collection ofshellfish and other invertebrates (except for mussels) provide statutoryprotection.

The Dornoch Firth is an estuarine inlet and, along with theInverness, Beauly and Cromarty Firths, is part of the much larger MorayFirth system (Fig. 1A, B; Hunter and Rendall, 1986). The Firth is asheltered, semi-enclosed embayment of glacial origin, ca. 23 km longand occupying 12,273 ha (Stapleton and Pethick, 1996). The mainfreshwater inputs are from relatively small rivers (Balls, 1994; SEPA,2011). The subtidal zone is shallow, not exceeding 10m, with the ex-ception of the entrance channel between Gizzen Briggs and WhitenessSands (Fig. 1C) where 17m depth has been recorded (UK HydrographicOffice, 1978). Minimum water temperatures are 5 to 6 °C in Februaryand in July to September temperatures range from 12 °C to 14 °C (Balls,1994; unpublished SEPA Shellfish Growing Waters data 2006 to 2008).

The outer Moray Firth is a fully saline (> 34 psu), open watersystem with a decreasing salinity gradient towards the inner firths(Balls, 1994). Most of the inner Dornoch Firth can be regarded asbrackish, with salinities ranging from<9 psu in the uppermost sectionto 25–31 psu in the main channel. The outer sections east of the Dor-noch Bridge are less influenced by freshwater inputs and conditionshere are regarded as fully marine.

The waters in the Dornoch Firth are well mixed from tidal currentsand occasional wave action from easterly winds. According to Balls(1994) oxygen saturation at or exceeding atmospheric levels was foundthroughout the Dornoch Firth whilst dissolved oxygen levels wereconsistently above 5mg l−1 from 2006 to 2008 (7–12mg l−1 in mostsampling dates), particularly in the winter months (SEPA, 2009).

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2.2. Searches

A variety of information sources were searched, including naviga-tional charts, historical maps, museum collections (biological and ar-chaeological), land-use records, fisheries records, statistical records,public online databases and naturalists' descriptions (see Appendix ATable A.1). Searches for records of Ostrea edulis in Scotland (live

specimens and oyster bed habitats) were conducted using the NationalBiodiversity Network online atlas (NBN, www.nbn.org.uk), the Marlinwebsite (www.marlin.ac.uk), the Marine Environmental Data Network(MEDIN; www.oceannet.org) and UK Marine Recorder (July 2014snapshot; http://jncc.defra.gov.uk/page-1599).

The archaeological online tool “Canmore” (http://canmore.org.uk),compiled and managed by Historic Environment Scotland, and the

Fig. 1. Study area. (A) Location of North East Scotland; (B) and (C) Map of the Dornoch, Moray and adjacent Firths indicating the main places relevant to the presentstudy. Sites with O. edulis shell or records and all field sites investigated are shown. Letter codes for survey sites (diamonds) indicate SACFOR abundance estimates.Map composed using qGIS v.2.10.1.

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Highland Council Historical Environment Record database (HER;http://her.highland.gov.uk/) were interrogated using “midden”, “shellmidden”, “shells”, “Ostrea” and “oyster” as search terms. UK grid re-ferences (OSGB1936) for possible midden sites were plotted onto ter-rain maps using the open source GIS software qGISv2.10.1 (QGISDevelopment Team, 2014) and three sites in the Dornoch Firth werevisited to verify records (below).

Government reports, statistical records and grey literature in localand university libraries and the National Library of Scotland weresystematically searched for references to oysters (see search termsabove) in the Moray and Dornoch Firths (e.g., Anon., 1885–1886;Groome, 1901, 1884–1985). Similar online searches were performedusing resources such as The Internet Archive (www.archive.org),Google Scholar and Google Books, where Public Domain 19th century(and earlier) references to “oysters” and “oyster fisheries” were avail-able (e.g., Blaeu, 1654; Dunbar et al., 1837; Eyton, 1858; Groome,1884–1985 p76; Lizars, 1850). The electronic versions of the StatisticalAccounts of Scotland for the Dornoch and Tain parishes (available fromEDINA http://edina.ac.uk/stat-acc-scot/ and Google Books) were con-sulted (Sinclair, 1791; Anon., 1845). Historical and/or current in-formation on O. edulis landings were obtained from the UK's MarineManagement Organisation (MMO), the Annual Reports of the FisheriesBoard of Scotland (Anon., 1882) and the Scottish Sea Fisheries Statistics(Appendix A, Table A.1).

Published accounts were complemented by consultation with ex-perts from the universities of Aberdeen and York, inshore fisheriesgroups, local fishers and conservation and environment agencies(Scottish Environment Protection Agency, Scottish Natural Heritage).Museum curators and independent researchers on palaeontology, bi-valve ecology, archaeomalacology, palaeoanthropology, Iron Age andmedieval archaeology and natural history, were also asked for accountsof oysters and oyster fisheries in the study area (see acknowledgementsfor list of consultees). Specimens were sought in collections of bivalves,archaeological finds and fossils held at The National Museum ofScotland Edinburgh (NMS), the Hunterian Museum in Glasgow, theGlasgow Museum and the Natural History Museum of London. Localmuseums and field societies in NE Scotland were also consulted in-cluding Elgin Museum, Inverness Museum and Art Gallery, DornochHistorylinks Museum, Tain Museum and Tarbat Discovery Museum(Portmahomack). Overall our searches were focused on the DornochFirth and the wider Moray Firth, but contextual accounts from NorthEast Scotland were also captured during this process.

2.3. Fieldwork

Between 2014 and 2017 shell middens recorded in the DornochFirth area were visited (e.g., Ness of Portnaculter, Cuthill Links;Fig. 1C). In the same period, beach transects, subtidal SCUBA surveys,and drop-down video surveys were conducted throughout the DornochFirth and opportunistically in adjacent areas (see Fig. 1). The in-vestigators walked from the upper littoral to the lower infralittoralzones, swam and searched for 45min on SCUBA surveys or towed acamera for 10min on drop-down video surveys to record the presenceand abundance of live O. edulis or their shells. Subtidal O. edulis shelldensities were estimated using the standard semi quantitative SACFORabundance scale (S= Superabundant; A=Abundant; C=Common;F= Frequent; O=Occasional; R=Rare; Joint Nature ConservationCommittee, 1997). All O. edulis shell collected during these surveyswere catalogued and deposited in the National Museum of Scotlandcollections in Edinburgh.

2.4. Radiocarbon dating

A subsample of five oyster (O. edulis) shells collected during theintertidal and subtidal fieldwork was aged using radiocarbon datingtechniques. Three oyster shells were used from the subtidal and two

from the intertidal (see Fig. 1C). The shells were selected to be re-presentative of the geographical distribution of the collection sites. Anyencrusting fauna, such as barnacles and bryozoans found to be growingon the dead oyster shells were scraped off and a sub-sample of shell(~1 cm2) next to the ligament was cut out using a Dremel saw. Thisincorporated multiple growth increments, which has been found toreduce the errors associated with intra-shell variability (Culleton et al.,2006; Rick et al., 2012) and is more consistently preserved in oystershells than the comparatively fragile outer edges. The O. edulis shellsub-samples were rinsed in fresh water, air dried and sent to the14CHRONO Centre at Queen's University Belfast (QUB) for radiocarbondating. Prior to analysis, the outer layer of shell was removed byetching with 1% hydrochloric acid (HCl) to ensure that any potentialcontaminant carbon from post-depositional exchange and/or adsorbedmodern CO2 was removed. The samples were then hydrolysed to CO2

with phosphoric acid. The CO2 was converted to graphite on an ironcatalyst using the hydrogen reduction method (Vogel et al., 1987). The14C/12C ratio and the 13C/12C ratio were then measured by acceleratormass spectrometry (AMS). The sample 14C/12C ratio was backgroundcorrected and normalised to the HOXII standard (SRM 4990C; NationalInstitute of Standards and Technology). The radiocarbon ages werecorrected for isotope fractionation using the AMS measured 13C/12C,which accounts for both natural and machine fractionation.

Radiocarbon dates of the oyster shells were calibrated using CALIB7.0.4 with the Marine13 calibration curve (Reimer et al., 2013; Stuivertand Reimer, 1993), which includes a correction for the global surfaceocean reservoir effect (ca. 400 years), and a local reservoir effect cor-rection (ΔR) (−29 ± 53 years; Russell et al., 2015).

No permits were required for the described study, which compliedwith all relevant regulations. Field studies did not involve endangeredor protected species.

3. Results

Archaeological records indicate that the first known human in-habitants of the Dornoch Firth area can be traced to the MesolithicPeriod after the last glaciations, when the shoreline had retreated toapproximately its present position (Doody et al., 1996; Fletcher, 1998).Twenty-two native oyster shells from this period collected at the CuthillLinks midden (Canmore Site No. 85364; Figs. 1C and 2A; Appendix ATable A.2) were located at the Dornoch Historylinks Museum (seeBatey, 1993). The Glasgow Museum also has a collection of Mesolithicand Neolithic remains from the Meikle Ferry midden (Canmore site No.14654; Fig. 1C; Appendix A Table A.2) that are dated 3000 to 6000 BCE(ca. 5–8000 BP) and consisted of man-made artefacts (e.g., arrowheads,flint and ash), animal remains, cockle, periwinkle and oyster shell(Batey, 1993).

Oyster, cockle and mussel shell were ubiquitous in middens asso-ciated with Mesolithic, Neolithic and Bronze Age settlements, datingfrom ca. 8000 BP, e.g., Meikle Ferry, Rosskeen (near Invergordon) andCuthill Links (Batey, 1993) through to the 1800s (Appendix A, TableA.2; Figs. 2A and 3), suggesting that O. edulis was common in coastalareas close to where they were found. In general, these middens con-sisted primarily of periwinkle and cockle shell with less mussel andoyster shell (Historic Environment Scotland, 2014; Fig. 2B). Neolithicchambered cairns and Bronze and Iron Age ‘brochs’ and hillforts havealso been recorded in archaeological surveys on both sides of theDornoch Firth at Tarlogie and Castle Corbert, some of them with as-sociated shell middens (C. Hatherley pers. com. 2014).

O. edulis shell has also been recorded (as ‘oysters’) from more recenthistorical periods at several archaeological sites excavated in the sur-rounding area. For example, excavations at an early medieval settle-ment at Portmahomack (Fig. 3) yielded oyster shell associated withmiddens from the early Christian monastic period (6–9th centuries)through to civilian settlements in the 13–15th centuries (Carver et al.,2016; Holmes, 2017). Two hundred and twenty oyster shells (along

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with other marine shells and animal bones) from late medieval andearly modern times (16–18th centuries) were found at the Brora salt-works (Fig. 3) in waste heaps from the construction and post-con-struction levels (Hambly, 2012). These foods are thought to have beenopportunistically gathered from the shore by workers involved in theconstruction and operation of the salt houses.

The first written account of oysters near the Dornoch Firth wasobtained from Joan Blaeu's Atlas of Scotland (Blaeu, 1654) from the1600s which states that “oysters […] and other shellfish are abundantat the river-mouths and sea-cliffs [of the northern Moray Firth]”. Blaeufurther described the Moray Firth area as being “fished for turbot, ray,dogfish, flounder and also lobsters, oysters and mussels, among otherspecies” (p110). In the 19th century, Olsen's 1883 Piscatorial Atlas(Fig. 4; Olsen, 1883) showed oyster beds along the southern shores ofthe wider Moray Firth, to the east of Lossiemouth (Figs. 1 and 4) wherethere are also records of extensive oyster shell deposits (Duff, 1842).There are remains of ancient oyster shell beds in Munlochy, Nigg Bayand near Tain (Miller, 1872). In 1837 in Nigg Bay and the Cromarty

Firth (Fig. 1B), William Macgillivray, a geologist and naturalist, re-called “an oyster scalp above Balintraad and Invergordon” and “…found oysters and about thirty or forty other species of shellfish, allindicating salt water” (Dunbar et al., 1837, pp. 411–413). There areother published accounts of oyster shell in the area, suggesting thatdense beds of oysters were present in the Cromarty and Moray Firths(University Marine Biological Station Millport, 2007).

The Statistical Accounts of Scotland (SAS) have one relevant entry forthe period 1835–1845 (Vol. 2, p 24) under Nigg Parish, where therewere: “…few oysters of large size at the Cromarty Firth but they areseldom dredged and do not promise to multiply.” There were no recordsfor oyster fishing in the Dornoch or Tain Parishes in either SAS volumes(for 1791 and 1834–1845) or in the annual reports of the FisheriesBoard of Scotland (FBS) from 1882 to 1938 but the 1883 FBS reportsuggested that the Dornoch Firth would be well suited to oyster culti-vation. However, the Cromarty District which encompasses thesouthern shores of the Dornoch Firth, is listed as one of three districtscontributing to Scotland's oyster fishery in 1883 and 1884 (Anon.,

Fig. 2. (A) Neolithic O. edulis shells, Cuthill Links midden (Dornoch Historylinks Museum). (B) Ness of Portnaculter shell midden (Fig. 1C). (C) O. edulis shell,Tarlogie; (D) O. edulis shell, Tarlogie (Fig. 1C). Left and right valves were present and attached. (E) O. edulis shell with intact ligament. (F) O. edulis shell collectedfrom Portmahomack beach.

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1884) the other two districts being Leith (encompassing the Firth ofForth, see Fig. 1) and Ballantrae (encompassing Loch Ryan, on the westcoast of Scotland). Thirty thousand oysters were recorded in fisherieslandings in the Cromarty District in 1883 and 26,000 in 1884. No oysterlandings were recorded in the Cromarty, or adjacent districts, after thistime and there is no evidence of industrial processing in this region(that would account for any large shell deposits). Oyster landings havesince been largely restricted to the west coast of Scotland with LochRyan accounting for most recorded Scottish oyster landings for theperiod of 1886–1938 and since.

A review of the state of Scottish oyster fisheries published in the1888 FBS report also notes that between 1876 and 1878, the wholeMoray Firth bed, inside Fort George (i.e., Beauly Firth; Fig. 1B), wasfound to have good quality oysters (Anon., 1882). This oyster bed wasidentified and subsequently fished to extinction by fishermen fromColchester, England (based ~800 km away), such that by 1885 the bedswere too sparse for dredging to be financially viable (Anon., 1882;Young, 1888).

Live O. edulis from the Moray Firth area were collected in 1879 and1881 during a dredging expedition in the Cromarty and Inverness Firths(Inverness Scientific Society, 1881, 1879); these specimens were de-posited in the Inverness Museum collections. In the early 1900s somelive specimens were reported from Findhorn (Moray Firth) (Bell, 1920)but the most recent verified scientific record of living oysters within theMoray Firth (Beauly Firth) is from 1975 (McKay and Smith, 1979).However, there is also a non-specific reference to a hobby fishery in theMoray Firth Partnership Management Plan in 2011 (MFIFG, 2011). Inaddition to historical accounts, interviews with local naturalists, mu-seum curators and fisheries experts indicated that (to their knowledge)

no local oyster fishery has been based in or immediately around theDornoch Firth in recent times (i.e., the last 50–100 years).

From 45 intertidal survey sites (Fig. 1B, C), O. edulis shells werefound at five locations in the west of the mid Dornoch Firth: emergentfrom sediment in an area scoured by freshwater run-off from the Tar-logie Spring, and scattered among mussel shell, cobble and pebble inthe upper infralittoral zone and along the high tide mark (Figs. 1C and2C). The shells were all intact and in good condition, and the majorityhad full sculpture. Out of a sample of 39 specimens that were randomlycollected, 16 had the hinge ligament intact, with the remains of theperiostracum still present in some shells and two with articulated valves(Fig. 2D–E). Occasional (between 1 and 9 in 100m−2) oyster shellswere also found on the beach west of Portmahomack (Fig. 1B); thesehad substantial wear, almost polished in appearance with little remnantsculpture (Fig. 2F). Opportunistic shore surveys (2017) of eleven fur-ther sites found “common” oyster shells along the Cromarty Firth(Cromarty Harbour Bay and Alness Point) and the Beauly Firth (Ar-dersier) but not Loch Fleet (Fig. 1B). O. edulis shell was also found atnine out of 18 random dive surveys and none of the 34 drop-down videolocations undertaken throughout the Dornoch Firth from the GizzenBriggs west to Edderton Sands (Fig. 1C). Rapid flow, shallow water andpoor visibility precluded further SCUBA survey up-stream of this point.Shell abundances ranged from ‘Common’ (1–9m−2) to ‘Rare’ (between1 and 9 in 1,000m−2) as per SACFOR scales (Joint Nature ConservationCommittee, 1997). The five shells that were sub-sampled for radio-carbon dating were found to have calibrated ages of between 4203 and8241 years before present (BP; Table 1).

Fig. 3. Timeline summarising key O. edulis records from the Dornoch Firth and surrounding areas. Note that archaeological records have not been compiled forMoray Firth and East Scotland regions. ⁎Radiocarbon dates obtained from the present study.

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Fig. 4. Entry for O. edulis in Olsen's (1883) Piscatorial Atlas of the North Sea. Fishermen accounts at the time indicated abundant oyster beds (in orange) in eastScotland along the south shore of the Moray Firth (1), the Firth of Tay (2) and the Firth of Forth (3). (For interpretation of the references to color in this figure legend,the reader is referred to the web version of this article.)

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4. Discussion

Our study shows that Ostrea edulis have been reported from shellmiddens in the Dornoch Firth, adjacent Firths, and the wider MorayFirth, from the Mesolithic Period to the Modern Ages (16th–19th cen-turies; Figs. 1–4). Furthermore, shell remains are still presentthroughout the system: some carbon-dated between 4000 and8000 years BP from the sites where they were most common in thepresent study. Literary sources and fishing records provided historicalevidence of live O. edulis present in the same areas as recently as the19th century. Therefore, it can be concluded that a long-standing O.edulis population existed in the Dornoch Firth and the encompassingMoray Firth system (Cromarty and Beauly Firths, the coastline fromInverness to Lossiemouth; Fig. 1A, B). However we only found a smallnumber of records of wild O. edulis in north-east Scotland after the late1800s: Limited fisheries statistics showed that oysters have not beenlanded on a commercial basis since 1884 and the accompanying reportsfurther indicated that, as elsewhere in Europe and the world, oysterpopulations appear to have been extirpated (Beck et al., 2011; Cranfieldet al., 2004; Zu Ermgassen et al., 2012a).

Oysters were absent in the Scottish fossil record at the end of thePliocene, appearing at the beginning of the Pleistocene only to becomescarcer again as a result of sea temperature fluctuation and ice abrasionduring the glaciations (Duff, 1842; Jamieson, 1866; Tooley and Smith,2005) and freshwater influxes in the post-glacial period (Bell, 1920).The presence of oyster shell in cultural remains from subsequent Me-solithic and Neolithic human deposits in the Dornoch Firth andneighbouring areas (Figs. 1–3; Appendix A Tables A.1 and A.2) showsthat oysters repopulated the marine habitats following the retreat of theglaciers as early as 10,000 years BP. This view is in keeping with ar-chaeological opinion that, because O. edulis shells have been recordedin discard heaps for thousands of years, oysters were relatively commonand likely to have been easily gathered from the foreshore nearby(Carver et al., 2016; Holmes, 2017).

None of the oyster shells found in the Tarlogie stream were fossi-lised, and 16 out of 39 specimens collected had remains of the hingeligament, which we initially interpreted as an indication of these oy-sters not being very old (i.e. recent historical times). This assumptionwas challenged by the radiocarbon dating which placed these shells inthe Mesolithic Period, at over 7000 years BP (Figs. 1 and 2C–E;Table 1). Freshwater can have a large radiocarbon offset from the at-mosphere due to the input of ancient carbon from weathering sedi-mentary rocks in the watershed or remineralisation of terrestrial or-ganic material (Ingram and Southon, 1996). Thus, since the Tarlogieoysters (Table 1) have been exposed to marine and fresh water influ-ences during their lifetimes it is possible that the age, which is based ona fully marine calibration, is exaggerated (see Ingram and Southon,1996). However, our finding is not unusual, as hinge ligaments havebeen found well preserved in oyster shells collected from archaeologicaldigs under certain conditions, i.e. waterlogged environments (Winder,

2017). On balance, despite initial assumptions, the age of these shellscorroborates the age of shell middens around the Dornoch Firth (Fig. 3)and is likely to be relatively accurate.

Blaeu's (Blaeu, 1654) literary accounts of oysters might be con-sidered problematic because the translation of the Atlas (originallywritten in Latin) refers, on several occasions, to freshwater mussels as“oysters”. However, because oysters are also included in a descriptionof commercial species in the coastal areas of Sutherland, Blaeu's recordsshould not be disregarded. Despite evidence to the contrary in thepresent study, T.J. Olsen's, 1883 maps (Olsen, 1883; Fig. 4) indicateoyster beds around Lossiemouth and not in the wider Moray Firth (i.e.,Dornoch, Beauly, Inverness and Cromarty Firths). Nonetheless, Olsen'swork was based on consultation with fishermen in SE England, ca.600 km to the south of the Moray Firth. Consequently, it might be ex-pected that these relatively small northern water bodies would not havebeen as well documented as those closer to home ports or to majorhuman conurbations, cf the Firth of Forth (Thurstan et al., 2013). In-deed, from the timing, it seems likely that the Colchester oystermenmoved up the coast to the Moray Firth area once stocks in the Firth ofForth became depleted (University Marine Biological Station Millport,2007). Uncertainties over bed ownership contributed to the failure toregulate fishing in the Moray Firth in the 1880s and these roamingfishermen also appear to have targeted oysters laid by locals (Young,1888). Exhaustion of the native oyster beds in both the Cromarty andMoray Firths occurred soon after (University Marine Biological StationMillport, 2007) and it seems possible that other small, undocumented(to fisheries managers) O. edulis beds, such as those in the DornochFirth, would have suffered the same fate as a result of harvesting, ascenario similar to that reported for C. virginica in the USA by Lenihanand Peterson (1998).

The conservation status of European Special Areas of Conservation(SAC) have commonly been determined, as with other types of marineprotected areas, against baselines at the time when conservation leg-islation (e.g., the 1992 EU ‘Habitats Directive’: 92/43/EEC) was im-plemented (Johnson, 2008; OSPAR Commission, 2012). However, thesebaselines do not necessarily represent ‘reference conditions’, i.e., “astate at which anthropogenic pressures are absent or negligible”(OSPAR Commission, 2011). In Scotland, as in most industrialisedcountries in Europe, it is highly improbable that examples of pristinemarine ecosystems can be found and used as an environmental re-ference because most areas have experienced some form of decline inecological condition (Halpern et al., 2015; Thurstan and Roberts, 2010;Thurstan et al., 2013). Even areas currently meeting management ob-jectives, such as the Dornoch Firth have been modified by anthro-pogenic influences. Therefore, the current conservation status of thesemarine protected areas probably does not reflect their full ecologicalpotential (Costello, 2014; van Leeuwen et al., 2012).

Across Europe, centuries of heavy fishing have affected large ex-panses of the seabed (Airoldi and Beck, 2007; Worm et al., 2006;Halpern et al., 2015) and the baseline (see Pauly, 1995) for the

Table 1AMS 14C dates from five oyster (Ostrea edulis) shells sampled from the Dornoch Firth between 2014 and 2015.

Sample ID (UBA-) Date collected Location Shell condition 14C age BP Cal age ranges

(14C year BP) ± 1σ (cal BP) 2σ

DFR24 (30646) 24.06.2015 Subtidal, East of DornochBridge

Substantial wear, polished appearance 4241 ± 28 4203–4555

DFR27 (30647) 24.06.2015 Subtidal, North of Tain Little wear, full sculpture present 4386 ± 28 4420–4777DFR32 (30648) 24.06.2015 Subtidal, West of Dornoch

BridgeModerate wear, shell sculpture degraded 6520 ± 44 6892–7228

Tarlogie I (30649) 20.03.2014 Intertidal, Tarlogie Spring Moderate wear, some shell sculpture still present 7604 ± 35 7950–8241Tarlogie II (32868) 20.03.2014 Intertidal Tarlogie Spring Light-moderate wear, some shell sculpture. Both valves and hinge

ligament present.7536 ± 33 8301–8289

Radiocarbon dates of the oyster (O. edulis) shells (14C age BP) were calibrated using CALIB 7.0.4, with the Marine13 calibration curve, which corrects for the globalocean reservoir effect (ca. 400 years), and a local reservoir effect to give calibrated age ranges expressed as cal BP at 2σ (Russell et al., 2015).

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management of marine protected areas (including SACs) has un-doubtedly shifted in many cases (Plumeridge and Roberts, 2017). Ashifted baseline might be expected for a more developed system such asthe Clyde or the Firth of Forth, where industrial and fishing impactshave been well documented (Thurstan and Roberts, 2010; Thurstanet al., 2013). In the Dornoch Firth, however, the history of humanpressures and historical conditions have hitherto been opaque at bestand might have mistakenly been considered negligible (cf OSPARCommission, 2011). Examination of historical data and novel survey forO. edulis (this study) has shown that present biological communities inthe Dornoch Firth are unlikely to correspond with unimpacted re-ference communities.

Despite the EU Habitats Directive requiring Member States to “…maintain or restore the natural habitats and the populations of wildfauna and flora at a favourable status”, there are few, if any of theensuing marine protected areas (SACs) where pre 1992 [restorative]baselines or targets have been considered. In the wider Europeanmaritime context, the Marine Strategy Framework Directive (MSFD;2008/56/EC) now seeks to promote the sustainable use of seas and theconservation of marine habitats. Because O. edulis beds are “specialhabitats” identified elsewhere under International agreements (EUHabitats Directive; OSPAR), they should attract widespread manage-ment and monitoring attention under MSFD (Cochrane et al., 2010).The historical presence and extent of key habitats such as O. edulis bedstherefore needs to be considered widely in reference conditions andbaselines for sustainable management if the aims of the MSFD are to beachieved.

Whilst it is known from the recovery of non-extractive marine re-serves throughout the world that there is scope for biodiversity increasefrom the point of designation (Costello, 2014), extinct or extirpatedpopulations and habitats have received very little, if any, considerationin Marine Protected Areas. Globally, the literature on habitat restora-tion has increased two orders of magnitude since 2015 (Nilsson et al.,2016) with many authors considering the practicalities and cost-bene-fits in forest, riparian and, to a lesser extent in marine (mostly tropical)habitats (e.g., Aradóttir et al., 2013; Crouzeilles et al., 2016; Fariñas-Franco and Roberts, 2014; Mayfield, 2016; Nunes et al., 2016). Therestoration of the European oyster has started to attract attention (asdid the American oyster, C. virginica, over the last 20 years; Brumbaughet al., 2006) particularly where remnant populations, well documentedhistorical fisheries and remnant small-scale fisheries have persisted(Gercken and Schmidt, 2014; Sawusdee et al., 2015; Smaal et al., 2015).What our study illustrates, and what has not yet been considered, is thatthere is probably a widespread case for restoration throughout theEuropean Atlantic MPA network where populations are long since ex-tirpated and records are not immediately available without a detailedinterdisciplinary review. The strength of the inter-disciplinary review isthe complementarity of evidence that might otherwise be consideredweak in isolation: the location of shell deposits, for example, might beconsidered entirely the product of long-term physical processes, exceptthat the archaeology also places oysters in the same location, as doesthe ancient literature and modern-day oceanographic information (seebelow). Likewise, any doubt that environmental conditions and presentclimate might be appropriate is mitigated by persistent archaeologicalremains, carbon dated shell, ancient literature references and fisheriesrecords in the region: collectively showing evidence of long-term suit-ability.

The systematic reintroduction of O. edulis throughout its formerdistributional range in the European Atlantic Marine Protected Areanetwork is a concept that would require a shift in conservation base-lines and could re-instate beneficial ecosystem services (see Coen et al.,2007; Kent et al., 2016). Existing data (Appendix A Table A.3) suggestthat present day environmental conditions in the Dornoch Firth aregenerally suitable for restoration: high water quality, adequate sub-stratum and hydrodynamic conditions and the potential for the site tobe a sink for larvae. In addition, there is a relatively low risk of physical

anthropogenic impacts from fishing, partly because of the aforemen-tioned protected status and legislation relating to the area but alsobecause access from the outer Dornoch Firth is restricted by a militaryfiring range five days per week. Despite the ancient right to fish bluemussels (Mytilus edulis) in the Dornoch Firth, the centre of this fisheryhas historically been in the eastern mid-section of the site, whereas thecentre of oyster shell deposits, and likely area for restoration, aregeographically separated to the west of this part of the Dornoch Firth(Fig. 1), in keeping with ‘channel beds’ formerly described from thesubtidal flanks of tidal channels in the Wadden Sea (Hagmeier andKändler, 1927; Möbius, 1877).

There are other practical challenges in the restoration of oysterpopulations. Source brood stock for restoration is one potential problembut O. edulis has persisted in aquaculture sites, albeit at low rates ofproduction. Genotype selection and the availability of locally fit stock isprobably not one of the challenges, however, because remnant O. edulispopulations show high levels of genetic similarity at the regional seascale (Vera et al., 2016): not limiting supply options. The introductionof disease agents such as Marteilia refringens and Bonamia ostreae withrestorative oyster brood-stock and/or oyster spat movements, however,is a significant consideration (Culloty and Novoa, 1999) and wouldrequire stringent biosecurity measures, especially because restorativeactivities can involve the movement of millions of oysters (Brumbaughet al., 2008; Schulte et al., 2009). In Scotland, shellfish farms aremanaged to help prevent disease outbreaks and allow for the control ofdamaging species, including those considered to be invasive (cf TheAquatic Animal Health (Scotland) Regulations 2009 and The Aqua-culture and Fisheries (Scotland) Act 2013). Aquaculture movementshave long been identified as vectors for invasive non-native species anddisease (Eno et al., 1997) therefore whatever the present regulatorysystems are, the scale of restoration programmes would represent apotential step-change to the movement of oysters from nursery and on-growing operations and therefore demand levels of biosecurity that arecompatible with the conservation management of protected areas ob-jectives in the Dornoch Firth.

Reintroduction of oyster habitat is entirely compatible with con-temporary conservation management policy and legislation. TheMarine (Scotland) Act 2010, for example, has provision for oyster bedhabitats as desired features in new marine protected areas (Tyler-Walters et al., 2012). However, restoration operations could sig-nificantly affect marine protected areas and the habitats and species forwhich they are presently being managed (in this case against 1992baselines). Designated protected sites, and with a ‘sliding baseline’(above), might therefore require their objectives to be revised if, ir-onically, restoration is to avoid being perceived as a conflict with thepresent-day protected features.

5. Conclusions

Overall, this study has shown that the native European oyster(Ostrea edulis) was historically present in the Dornoch Firth SAC andthroughout the region and the conditions in the protected area remainwithin its current potential distributional range. The analysis of severaltypes of evidence in an interdisciplinary review has enabled confidencein historical presence, distribution and persistence of oysters. As an areawith low anthropogenic pressures and suitable environmental condi-tions for O. edulis, reintroduction could be successful. European oysterpopulations have been lost throughout their range (Airoldi and Beck,2007; Zu Ermgassen et al., 2012a) but there has recently been a rapidinternational expansion of MPAs (Costello, 2014) and legislative driversfor sustainable marine management. Interdisciplinary review of pro-tected area baselines will probably show scope for O. edulis restorationin many more protected areas, throughout the European MPA networkin addition to areas that support the few remnant populations andfisheries.

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Acknowledgements

This work is part of the Dornoch Environmental EnhancementProject (DEEP), a partnership between Heriot-Watt University, theMarine Conservation Society and the Glenmorangie Whisky Company.The enthusiasm and support of Hamish Torrie, Peter Nelson, ChasMcEwan, Peter Murphy, Calum Duncan, Iain Russell (who also pro-vided historical data), Nicola Jamieson, John McMullen, Mark Bruntonand Andrew Macdonald are especially acknowledged. The authors alsowish to thank the following experts consulted during the project: AlanMackenzie (Cromarty oyster and mussel grower), Alex Adrian (CrownEstates); Sankurie Pye, Alison Sheridan, Sarah Stewart (NationalMuseums of Scotland); Andy Woolmer (Mumbles Oyster Company);Michelle Cadger, Anthony Watson (Tarbat Discovery Museum); JohnBaxter, David Donnan, Ben James (SNH); Janet Khan, MoragMacLachlan (SEPA); Candy Hatherley and Gordon Noble (AberdeenUniversity); David McKay (Malacologist); Dai Roberts (Queen'sUniversity Belfast); James Bromham (Highland Council); Janet Trythall(Elgin Museum); Jessica Winder; Kate Gormley and Flora Tulbure(Heriot Watt University); Mary Spencer Jones, Jon Todd and KathieWay (Natural History Museum of London); Kathryn Logan (Moray FirthPartnership), Maggie Reilly (Hunterian Museum Glasgow); Mark James(St. Andrew's University); Martin Carver (University of York); Nick Lake

(Moray Inshore Fisheries Group), Shelagh Smith; Sarah Heaton(Inverness Museum and Art Gallery); Simon Taylor (ConchologicalSociety); Stuart Anderson (Anderson Marine); Sue Higgings(Historylinks Museum at Dornoch); Richard Sutcliffe (GlasgowMuseum). Diving and field support provided by Matt Barnes, GeorgeBrown, Rob Cook, Ash Cordingley, Bill Ruck, George Stoyle, RichardShucksmith, Simon and David Exley were much appreciated. Editorialscrutiny and review of the manuscript by three anonymous referees wasmuch appreciated.

Role of the funding source

The Glenmorangie Whisky Company funded the present research.The company commissioned an environmental review but played norole in the present study design, data collection, analysis, interpreta-tion, writing or decision to publish.

Funding

The work was supported by the Glenmorangie Whisky Company(A15R10520) and the MASTS pooling initiative (the Marine Alliance forScience and Technology for Scotland) funded by the Scottish FundingCouncil (grant reference HR09011).

Table A.1Sources consulted, and information used to investigate the historical presence of O. edulis in the Dornoch Firth and Northeast Scotland.

Data resource Source/Location web link/Expertconsulted

Data type Data used (ifany)

Period Example of query/Information found

NBN Atlas NationalBiodiversityNetwork (NBN)

www.nbn.org.uk Biological records Records of O.edulis inScotland

1870–2014 https://records.nbnatlas.org/occurrences/search?q=lsid:NHMSY-S0020975316

MARLINspeciesrecordsdatabase

MARLIN www.marlin.ac.uk Biological records Records of O.edulis inScotland

UK MarineDatadiscoveryportal

MarineEnvironmental DataNetwork (MEDIN)

www.oceannet.org Biological records/Environmentalparameters

Records of O.edulis inScotland/Abioticparameter datain the DornochFirth

1950–2009 http://www.oceannet.org/finding_data/search/full/catalogue?q=%22ostrea+edulis%22&t=co&Graticule=Graticule

MarineRecorder

Joint NatureConservationCommittee (JNCC)

http://jncc.defra.gov.uk/page-1599

Biological records/Environmentalparameters

Records of O.edulis inScotland/Abioticparameter datain the DornochFirth

1990–2014 National records forO. edulis presenceand O. edulis bedhabitats (e.g.http://www.jncc.gov.uk/marine/biotopes/biotope.aspx?biotope=JNCCMNC-R00000788)

ScotMap -InshoreFisheriesMappingProject inScotland

ScottishGovernment/Marine Scotland

http://www.gov.scot Fisheries landingsdata/maps

No relevantdata found/used

http://www.gov.scot/Topics/marine/science/MSInteractive/Themes/ScotMap

www.scotland.gov.uk 1922–2016

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Table A.1 (continued)

Data resource Source/Location web link/Expertconsulted

Data type Data used (ifany)

Period Example of query/Information found

AnnualScottish SeaFisheriesStatistics/FisheriesBoard ofScotlandReports

ScottishGovernment

Fisheries landingsdata

Oyster landingsonly on westcoast ofScotland. Norecords in studyarea.

http://www.scotland.gov.uk/Topics/Statistics/Browse/Agriculture-Fisheries/PubFisheries

Canmore-NationalRecord ofthe HistoricEnvironme-nt

HistoricEnvironmentScotland

http://canmore.org.uk Archaeologicaldatasets

Shell middens,oyster shell

Mesolithic-Modern Age

http://canmore.rcahms.gov.uk/en/search/?keyword=midden&submit=search

HighlandCouncilEnvironme-nt Record(HER)

Highland Council http://her.highland.gov.uk/

Archaeologicaldatasets

Shell middens,oyster shell

Mesolithic-Modern Age

Searches for“midden”, “oyster”,“shell”

Archaeologydata service

Archaeological DataService (UK)

http://archaeologydataservice.ac.uk

Archaeologicaldataset/Publishedliterature.

Shell middens,oyster shell

Mesolithic-Modern Age

http://archaeologydataser-vice.ac.uk/archives/view/des/index.cfm?decade=1990&

The SCAPETrustExcavationsat the Brorasaltpans

Shorewatch www.shorewatch.co.uk Published literature. Brora Saltpansarchaeologicaldig reports

1400s–1700s http://www.shorewatch.co.uk/brora/html/reports.asp

Digital Libraryof theNationalLibrary ofScotland

National Library ofScotland

http://digital.nls.uk/gazetteers-of-scotland-1803-1901

Published historicalaccounts

OrdnanceGazeteers ofScotland

1800s–1900s http://digital.nls.uk/gazetteers-of-scotland-1803-1901/pageturner.cfm?id=97392450&mode=transcription

NationalLibrary ofScotlandMapsService

National Library ofScotland

http://maps.nls.uk Admiralty and otherhistorical charts

Blaeu Atlas/Olsen'sPiscatorial Map

1600–1900 http://maps.nls.uk/atlas/blaeu/page.cfm?seq=109;http://maps.nls.uk/view/74412462

The InternetArchive

The InternetArchive

www.archive.org Historical Accounts/Fisheries records

Annual Reportsfor theFisheries Boardof Scotland

1800s–1922 https://archive.org/search.php?query=creator%3A%22Fishery+Board+for+Scotland%22

Hathi TrustDigitalLibrary

Hathi Trust https://www.hathitrust.org/

Historical Accounts/Fisheries records

Report to theCommissioners

1885–1886 https://catalog.hathitrust.org/Record/101710112

Google Scholar Google Inc. https://scholar.google.co.uk/schhp?hl=en

Current andhistorical records

Relevantpublished peerreviewedliterature

1900s–2000s

Google Books Google Inc. https://books.google.com/

Grey Literature/Public DomainBooks

Mostly 1800sand early 1900s

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Table A.1 (continued)

Data resource Source/Location web link/Expertconsulted

Data type Data used (ifany)

Period Example of query/Information found

Referencehistoricalaccounts foroyster presenceand fisheries inthe UK andIreland

https://play.google.com/books/reader?printsec=frontcover&output=reader&id=WGI9AAAAYAAJ&pg=GBS.PA9;https://play.google.com/books/reader?printsec=frontcover&output=reader&id=KDoDAAAAQAAJ&pg=GBS.PR1

OriginesParochialesScotiae/AnnualReports fortheFisheriesBoard ofScotland.

Forgotten Books www.forgottenbooks.org

Grey Literature/Public DomainBooks

OriginesParochialesScotiae/AnnualReports for theFisheries Boardof Scotland.Other greyliterature

1800s–1900s http://www.forgottenbooks.org/readbook_text/Origines_Parochiales_Scotiae_1000684228/449

The UniversityofEdinburgh/EDINA

EDINA http://stat-acc-scot.edina.ac.uk/

StatisticalAccounts ofScotland

1800s-Early1900s

http://stat-acc-scot.edina.ac.uk/sas/sas.asp/?account=1&accountrec=007404&action=publicdisplay&county=Ross%20and%20Cromarty&monospace=&naecache=66&navbar=&nohighlight=&pagesize=&parish=Nigg&session-id=0c743c5692f7012c-d0208100ac21147f&transcript=&twoup=#pageimage

MMO AnnualStatisticsArchive

MarineManagementOrganisation(MMO)

www.marinemanagement.org.uk

Fisheries Statistics/Historical records(Report to theCommissionersappointed to enquireinto the sea fisheriesof the UnitedKingdom)

AnnualStatisticsArchive/O.edulis landingsand historicalrecords

1800–1950 http://webarchive.nationalarchives.gov.uk/20140109182227/http://www.marinemanage-ment.org.uk/fisheries/statistics/documents/ukseafish/archive/1866.pdf

Essex Naturalist http://www.essexfieldclub.org.uk

Historical Records 1900–1950

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Table A.1 (continued)

Data resource Source/Location web link/Expertconsulted

Data type Data used (ifany)

Period Example of query/Information found

EssexNaturalist,Vol. 19(1920):BritishOysters:Past andpresent.

O. edulisdistributionand fisheries inScotland in the1900s

http://www.essexfieldclub.org.uk/portal/p/Archive/s/020/o/0211

The NationalMuseum ofScotland

Edinburgh Sankurie Pye/SarahStewart

Museum Collections O. edulisspecimensincludingfossils

No specimensheld from NEScotland

GlasgowMuseumCollections

Glasgow http://collections.glasgowmuseums.com

Museum Collections O. edulisspecimens frommiddens(Meikle Ferry)

8000–6000 YBP(Mesolithic)

http://collections.glasgowmuseums.com/starobject.html?oid=235248

The HunterianMuseumGlasgow

University ofGlasgow

Maggie Reilly Museum Collections O. edulisspecimensincludingfossils

No specimensheld from theEast coast ofScotland.

Natural HistoryMuseumLondon

London, England John Todd/Kathie Way Museum Collections O. edulisspecimensincludingfossils

Contacted/Nodata used

Elgin Museum Elgin, Scotland Jane Trythall Museum Collections O. edulis shells Possiblyspecimens heldfrom theLadyhill site, inElgin and theRev. GeorgeGordonCollection (Notvisited)

InvernessMuseumand ArtGallery

Inverness, Scotland Sarah Heaton Museum Collections O. edulis shells(Sutherlandexpeditions,Cromarty andInvernessFirths)

1879–1881

DornochHistorylinksMuseum

Dornoch, Scotland Sue Higgings Museum Collections O. edulis shells Oyster shellsfrom Neolithicand Pictishperiod middens

http://www.historylinksarchive.org.uk/search.asp?id=&page=754&start=9443&q=and in situ visits

Tain Museum Tain, Scotland http://www.tainmuseum.org.uk

Museum Collections O. edulis shells No informationobtained

TarbatDiscoveryMuseum

Tarbat, Scotland http://www.tarbat-discovery.co.uk/Michelle Cadger/Anthony Watson

Museum Collections O. edulis shells/information

No oysterspecimens keptin the collection.Information onPortmahomackdig by MartinCarver.

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Table A.1 (continued)

Data resource Source/Location web link/Expertconsulted

Data type Data used (ifany)

Period Example of query/Information found

Tarlogie Dunand otherIron Ageforts in thearea

University ofAberdeen

Dr. Candy Heatherley/Dr. Gordon Noble

Expert Account Iron Age Most shells frommiddens seenwere periwinkleand cockle. Nooysters.

Dr. MartinCarver

University of York http://martincarver.com/publications/portmahomack-tarbat-ness-changing-ideologies-north-east-scotland-sixth-sixteenth

Published records/Expert Account

Medieval (8thto 13thcenturies)

O. edulis shellsreported frommonasticsettlement atPortmahomack

(Carver et al., 2016)

Dr. JessicaWinder

Independentarchaeomalacologist

http://oystersetcetera.wordpress.com/

No recollectionof oysters in theDornoch Firth.Usefulinformation onoyster dating

Winder (2017)

Table A.2Archaeological sites with shell middens in the Dornoch Firth and the surrounding area (see Fig. 2). Source: http://canmore.gov.uk/; HighlandCouncil HER http://her.highland.gov.uk/.

Site name OS grid CanmoreRef

Date estimation (bysource)

Site description Ostrea edulis shellpresence

ArdjachiePoint

NH74988478 HERreferenceMHG29337.

Not available Shells at the NW part of the point. Discovered byDavid Fiddimore, Customs Officer, Glenmorangie.

Oysters notconfirmed, onlyscallops, winklesand cockles.

ArdjachieFarm

NH746845 14736 Early Neolithic toPictish

Notable for a Pictish cup-marked stone. A DGPSsurvey revealed midden layers (NH78SW 25),probably Neolithic.

Unknown

Ness ofPortnacult-er

NH744848 85379 Not known. Disturbed midden, fire-cracked stone andcompacted shell layers. Periwinkle and cockle shellrecorded during field visits.

Unknown.

Cuthill LinksWest

NH743871 85364 Mesolithic toNeolithic

Eroding shell midden. Artefacts including stonehammer and leaf arrowheads.

Yes. Some depositedin the DornochMuseum collections

Castlehaven NH931873 15630 Probably Neolithic Shell midden. Littoral shellfish remains, cod bonesand spines of skate.

Not known.

Castle Corbet NH902833 15633 Mesolithic/Neolithic

Earliest remains of littoral shell-fish in the area.Knife-like forms of quartz.

Not known.

Cuthill LinksEast

NH749873 14647 Neolithic to BronzeAge

Large shell midden, Arrowhead(s) (flint), Brooch(bronze)

Sources describe avariety of shells.

Meikle Ferry NH72808700 14654 6000 BCE–3000 BCE Midden with shells, split bones and teeth showingthe action of fire. Remains of deer, ox, horse, pigand goat or sheep as well as flakes and a knife orscraper of sandstone.

Yes. Deposited inGlasgow Museumcollections.

Cyderhall NH754887 14633 Unknown, possiblyNeolithic.

Possible mound of midden shell. Unknown

Portmahomack NH9150484170 15656 800–1300 CE Icehouse and shell midden. YesEarl's Cross NH80469026 15631 Possibly Neolithic. Alleged ancient shell midden. Mussel shells and

worked stones tools.Not recorded.

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Table A.2 (continued)

Site name OS grid CanmoreRef

Date estimation (bysource)

Site description Ostrea edulis shellpresence

LittleferryLinks

NH814966 15339 Mesolithic to EarlyMedieval (Pictish)

Numerous stone tools, carved stones and shellmounds.

Yes

Brora Salt Pans NC905034 294450 1500–1800 CE Remains of former saltworks. Extensive middensincluding oyster shell.

Yes

EddertonSands

NH720847 85378 Not known Lithic scatter and eroded shell midden. Unknown

Tarlogie Dun NH7616 8389 14684 Iron Age to Pictish(800 BCE to 600 CE)

Dun located in a small natural hillock. Cockles,mussels and periwinkles.

No oysters (GordonNoble & CandyHatherley pers. com.2014)

Rosskeen NH6893 6944 85350 Bronze Age A large ditch with a secondary fill of oyster shelland animal occupation.

Yes (see Batey,1993)

Rosskeen NH69116945 85350 Bronze Age A fragmentary shell midden on W bank of theRosskeen Bum. Oyster and winkle shells.

Yes (see Batey,1993)

Table A.3Recommendations for site selection in shellfish restoration projects (adapted from Brumbaugh et al., 2006) and conditions encountered in theDornoch Firth during the present study in relation to Ostrea edulis reintroduction.

Site selectioncriteria

O. edulis requirements Conditions encountered in the Dornoch Firth

Historical presenceof the targetspecies

Identify areas where reefs or target shellfish populations historicallyexisted.

Records show O. edulis presence and exploitationsince Mesolithic times. O. edulis shells are foundintertidally and subtidally.

Environmentalconditions:Hydrodynamics

Locate restoration projects within small, replicable sub-estuaries.Determine whether this area is a “sink” for larvae.Assess the current velocity. Shellfish growth is generally higher wherecurrents are greater, delivering food and oxygenated water andcarrying away waste by-products. Moderate currents are preferredover strong currents (> 2 knots) because they help prevent excesssiltation and promote the supply of food. Strong currents are notrecommended because they increase the risk of dislodgment, thuspreventing sustainability and local recruitment.

The Dornoch Forth can be considered a “sub-estuary” within the larger Moray Firth system.Existing data indicates high levels water retentionand recirculation and prolonged flushing times.Sublittoral sediments of the Dornoch Firth aredominated by scoured coarse gravel and cobbleand relatively well sorted sands, an indication of atidally dynamic environment.Stapleton and Pethick (1996) described aneast–west, ebb-flood reversing current pattern,with peak spring velocities of 1m s−1 (~2 knots)and 0.3ms−1 during neaps which are usuallyhigher on the ebb side of the tide. Flows thereforewithin recommended range.

Environmentalconditions:Substratum

Evaluate bottom conditions to determine if the bottom will supportaddition of shell or other materials used for habitat enhancement.Oysters occur on a variety of bottoms from mud to rock, muddy sand,muddy gravel with shell and hard silt but all of them need to be firm(Laing et al., 2005).

Substratum is mostly medium to fine compactsands with gravels and a coarser bottom in thedeep, tide swept channels at its mouth. Silt isdominant in the innermost part of the firth only.Acoustic surveys undertaken on behalf of SNH in1996 concluded that the most common substratein the Dornoch Firth was medium sand whilst mudwas the rarest, a reflection of the dynamism of thesystem and low input from the rivers in itscatchment.

Environmentalconditions:Oxygen

The overlying waters must be well oxygenated. Optimal levels ofdissolved oxygen in the water ensure growth, feeding activity andsurvival. No values found for O. edulis. As an indication, the minimumdissolved oxygen for C. virginica is set 2.3mg l−1 (Newell et al., 1999).

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Table A.3 (continued)

Site selectioncriteria

O. edulis requirements Conditions encountered in the Dornoch Firth

Waters are well mixed as a result of tidal currentsand occasional wave action from easterly winds:hypoxia therefore unlikely an issue for O. edulisrestoration. Dissolved oxygen data in the DornochFirth ranges from 7 to 12mg l−1 in most months(Balls, 1994; SEPA, 2011).

Environmentalconditions:Temperature

One of the most important environmental parameters controlling thedistribution and physiological adaptations of marine bivalves is seawater temperature (Laing and Spencer, 2006; Lucas, 2003). Growth inO. edulis usually starts when temperatures reach 8–9 °C in the springand picks up in the summer at temperatures of 16 °C to 18 °C (Lainget al., 2005). Gametogenesis in oysters usually starts at temperaturesof 7 °C to 10 °C with a spawning triggered by temperatures of at least15 °C (see references in University Marine Biological Station Millport,2007).

Historical data shows water temperatures in theDornoch Firth range from ~5 to 15 °C, within theoptimal range for survival and growth of O. edulis.The 15 °C spawning temperature threshold for O.edulis might not be reached every year. Availabletemperature data was roughly collected every fourmonths and it is entirely possible that peaktemperatures throughout the Firth may have beenmissed. In one year (2006) higher peaktemperatures were recorded upstream and it ispossible that this also occurs more often than thedata suggests.

Environmentalconditions:Salinity

According to Laing et al. (2005), native oysters can be found fromestuaries with salinities as low as 16 psu to fully saline open waters,i.e.> 30 psu. The filtration mechanism is impaired at salinities lowerthan 14 psu (Zu Ermgassen et al., 2012b).

The outer Moray Firth is a fully saline (> 34 psu),well mixed open water system with a decreasingsalinity gradient towards the inner firths,including the Dornoch Firth (Barne et al., 1996).Most of the inner Dornoch Firth can be regarded asbrackish, with salinities ranging from<9 psu inthe upper sections towards Bronar Bridge to25–31 psu in the main channel.

Environmentalconditions:Phytoplankton

Food is one of the most important factors limiting growth,reproduction and survival of bivalves (Gosling, 2003).

Targeted primary production data is scarce andfurther site specific monitoring needs to beundertaken to determine seasonal and spatialvariability to inform the site selection process. Thechlorophyll concentrations reported for the regionare within the range for conditioning bivalves1.68mg l−1 and up to 6 μg l−1 and similar to thoserecorded in areas that support bivalve populations(Da Costa et al., 2008; Fariñas-Franco and Roberts,2014).

Environmentalconditions:Turbidity

Jackson and Wilding (2009) consider O. edulis to have high tolerance,very low sensitivity and high recoverability from elevated turbiditylevels.

Seabed substrata in the Dornoch Firth aredominated by mobile medium sand, particularlyeast of Meikle Ferry, and turbidity levels are likelyto be elevated in the central channels during thepeak tidal flows. Highest turbidity levels up to40mg l−1 found in more brackish conditions(e.g.< 10 ppu) and low sediment loads(0–20mg l−1) in more saline areas (Eyre and Balls,1999). These values are similar to those recordedin sheltered sea lochs, which usuallyhave<50mg l−1 (Laing and Spencer, 2006).

Legislative context Consider placement of restoration projects in areas where illegalimpacts can be deterred.

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Table A.3 (continued)

Site selectioncriteria

O. edulis requirements Conditions encountered in the Dornoch Firth

The Dornoch Firth is protected by several pieces ofshellfish and conservation legislation. It is aSpecial Area of Conservation (SAC), a SpecialProtected Area (SPA), a Site of Special ScientificInterest (SSSI) and part of the OSPAR network ofMPAs. The Firth is also listed as Shellfish GrowingWaters.An oyster restoration project falls withinconservation and management strategies ofconservation agencies and other competentauthorities consulted during the study (SNH,SEPA, and Highland Council).With the exception of dredging for mussels, fishingusing mobile gears is banned in the Dornoch Firth.Protection is provided by two Nature ConservationOrders banning collection of shellfish and otherinvertebrates from the exposed and submergedforeshore. Further protection can be obtainedthrough a Regulating or Several Order toeffectively prevent any unauthorised extraction ofshellfish.

Pollution/Anthropogenicpressures(threats)

Determine what threats exist in areas formerly populated by shellfish. Mussel dredging occurs in the Dornoch Firth.There is evidence of unregulated shellfishgathering in the intertidal.The main sources of pollution are the regulateddischarges from two whiskey distilleries, onesalmon fishfarm located on land and two wastewater treatment works. The discharges from thefishfarm and distilleries have consistentlycomplied with legislation with a score of Good orExcellent. However, the WWTWs have a poorrecord of compliance which only affects thequality of shellfish for human consumption.

Disease andpredation

The most pernicious disease in UK native oysters is Bonamiosis, causedby a protozoan Bonamia ostrea. Bonamiosis has caused massmortalities in the past (Culloty and Novoa, 1999). Because thisparasite prefers brackish conditions siting of translocation sites foroysters in fully saline areas is thought to afford some degree ofprotection (Laing et al., 2005). The parasite Marteillia perfringens alsofavours low salinity environments.Predation loses of reintroduced juvenile oysters can be as high as 91%and starfish and seabirds can decimate mussel beds (Gaymer andHimmelman, 2002; Gosselin and Qian, 1997; Laing et al., 2005).

Ostrea edulis probably disappeared from theDornoch Firth in the 1800s. Potential broodstockfrom Loch Ryan in the west coast of Scotland isdisease free.In the Dornoch Firth predation from birds in theintertidal zone and starfish in the subtidal arelikely to be the most important natural threats to atranslocated shellfish population. The DornochFirth is host to an important population of wadersand sea ducks that prey on the mussel scalps whilststarfish have been recorded in the subtidal beds bySNH and the Highland Council in the past(unpublished data, 2004). According to the MorayFirth Fisheries Management Plan (2011) resourceshave already been deployed to manage starfishpredation. On-growing oyster seed prior todeployment (to reduce mortality) is probably thebest strategy for restoration.

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