# Lessons from Lyme Bay (UK) to inform policy, management, and monitoring of Marine Protected Areas

Chloe Renn<fnref n="1" />, Sian Rees<fnref n="1" />, Adam Rees<fnref n="1" />, Bede F.R. Davies<fnref n="2" />, Amy Y. Cartwright<fnref n="1" />, Sam Fanshawe<fnref n="3" />, Martin J. Attrill<fnref n="1" />, Luke A. Holmes<fnref n="1" />, Emma V. Sheehan<fnref n="1" />,*

## Abstract

This decade represents a critical period to profoundly rethink human-nature interactions in order to address the interwoven climate and biodiversity crises. Marine Protected Areas (MPAs) demonstrate promise for increasing ecosystem resilience and reversing habitat and population declines, but outcomes vary considerably from context to context. Partially protected areas offer a compromise between ecological recovery and the social needs of local communities, but their success is contingent on an array of factors. This in-depth review summarizes 15 years of marine conservation research and impact in Lyme Bay (southwest UK), to serve as a model for the future adoption of partially protected MPAs. The findings from the UK's longest integrated socioecological monitoring MPA study are presented and supplemented by an evaluation of the whole-site management approach as a core element of Lyme Bay's achievements. The journey from research to improved monitoring and ambitious policy is illustrated within and interspersed with stories of novel discoveries, ongoing challenges, and method developments. What started as a dedicated group of community members has grown into an immense collaboration between fishers, scientists, NGOs, and regulators, and their combined efforts have sent ripple effects of positive change across the globe.

**Keywords:** Marine Protected Areas; marine policy; fishery management; fisheries; socioeconomic; ecosystem-based management; marine conservation; partnerships

## Introduction

Marine ecosystems face a range of increasingly stark threats (Halpern et al. 2019, He and Silliman 2019) each cumulatively contributing to global biodiversity loss that rivals the extent of past mass extinction events (Ceballos et al. 2015, Cowie et al. 2022). The beginning of this decade marks a critical time point to fundamentally rethink the nature of human-ocean interactions, as global temperatures are predicted to approach 1.5°C of warming as early as 2030 (IPCC 2018, UNEP 2022a). Since the effects of climate change-induced perturbations (e.g. ocean acidification, warming, deoxygenation, and extreme storm events) will inevitably intensify in the marine environment (IPCC 2019), there is a need to limit additional anthropogenic stressors to ensure the greatest chance of species survival (Sumaila and Tai 2020).

Marine Protected Areas (MPAs) are known to have interconnected social and economic dynamics (Rees et al. 2018), and thus, together with their respective resource users, MPAs form a complex social-ecological system (SES; Ostrom 2009). In an ecological sense, MPAs can be used to restore and potentially future-proof marine ecosystems (Roberts et al. 2017, Worm 2017), but their success varies dramatically depending on several biophysical and socioeconomic factors, from enforcement to species mobility (Edgar et al. 2014, MacKeracher et al. 2019).

Management approach also differs vastly between MPAs (Grorud-Colvert et al. 2021). While some MPAs exclude all

extractive activities from an entire area [e.g. UK No Take Zones and Highly Protected Marine Areas (HPMAS)], others selectively permit activities if they are deemed nonthreatening to designated features (Sciberras et al. 2013, Zupan et al. 2018). When employed in tandem with effective threat management legislation, MPAs can facilitate, sometimes rapid, population recovery (Lester et al. 2009, Caselle et al. 2015) even for large, mobile species (García-Rubies et al. 2013). Further, when destructive fishing practices are excluded from the entire area via whole-site protection, benefits extend beyond individual species populations and create ecosystem-wide improvements (Sheehan et al. 2013a, Solandt et al. 2020, Davies et al. 2022a). Such improvements can also expand the availability of essential fish habitats through increased habitat complexity and quality, leading to additional fishery benefits, including spillover effects (Gell and Roberts 2003, Kerwath et al. 2013, Da Silva et al. 2015).

MPA establishment has accelerated across the globe, expanding at a rate of ~8% per year since the 1960s (Worm 2017, Duarte et al. 2020), although estimates are lower when areas with ineffectual levels of protection are excluded (Sala et al. 2018). However, when adopted in the absence of stakeholder engagement, enforcement, and sufficient planning, MPAs signify nothing but lines on a map with no meaningful socio-ecological impacts (Giakoumi et al. 2018, Turnbull et al. 2021). Dramatic disparities in MPA performance relative to objectives emphasize the importance of learning

<footnote n="1">¹School of Biological and Marine Sciences, University of Plymouth, Drake Circus, Plymouth PL4 8AA, United Kingdom</footnote>

<footnote n="2">²Nantes Université, Institut des Substances et Organismes de la Mer, ISOMer, UR 2160, F-44000 Nantes, France</footnote>

<footnote n="3">³Blue Marine Foundation, Somerset House, London WC2R 1LA, United Kingdom</footnote>

<footnote>*Corresponding author. School of Biological and Marine Sciences, University of Plymouth, Drake Circus, Plymouth PL4 8AA, United Kingdom. E-mail: emma.sheehan@plymouth.ac.uk</footnote>
from past successes and failures, to maximize future conservation outcomes (e.g. Coleman et al. 2004, Hays et al. 2020, Stewart et al. 2020). While no-take zones are generally perceived as most effective (Edgar et al. 2014, Sala and Giakoumi 2018), partially protected areas (Day et al. 2019) offer advantages as a feasible middle ground (Rees et al. 2010a), balancing ecological recovery with the socioeconomic needs of local communities by limiting only certain activities (e.g. destructive demersal-towed fishing methods). Permitting some level of extraction within MPAs, however, leaves considerable room for error, and without stringent monitoring and enforcement, they can create the illusion of conservation success while failing to meet MPA management objectives (Turnbull et al. 2021).

The challenges associated with managing partially protected areas highlight the importance of identifying well-implemented examples to both showcase their potential and act as a blueprint for their successful implementation elsewhere. The Lyme Bay MPA (southwest UK) represents a good example of a partially protected area that has benefitted from collaborative community effort and produced ecological and social gains through its whole-site approach to management and monitoring (Sheehan et al. 2013a, Davies et al. 2021, Rees et al. 2021a). However, concerns remain over high levels of static fishing (potting and netting) inside the MPA, particularly from nomadic vessels, and establishing legislative control over these fishing methods represents an ongoing challenge (Mangi et al. 2011, Rees et al. 2021a). With increasing international appetite for MPAs (evidenced by the ‘30 by 30’ initiative; Belote et al. 2021) and the recent designation of HPMAs in the UK (Defra, 2019), this study aims to provide a timely synthesis of the main outcomes and lessons learned from 15 years of marine conservation research and collaborative stakeholder engagement in Lyme Bay. This in-depth case study will outline the socioeconomic and ecological effects of the Lyme Bay MPA, to serve as an example for the future adoption of partially protected MPAs. This will be supplemented by an evaluation of the whole-site approach as a core component of Lyme Bay’s achievements, while highlighting the development of innovative research techniques, novel scientific discoveries, the collaborative management approach, and the national and international reach of the Lyme Bay model.

## Lyme Bay MPA

Lyme Bay is a coastal inlet (~2460 km²) in the southwest UK, spanning the Devon-Dorset border. The seafloor hosts rich biodiversity, supported by a mosaic of habitats, from muddy, soft sediments to rocky and biogenic reef features. The associated marine fauna is nationally and internationally significant (Gubbay 1988) and colloquially referred to as the UK’s ‘coral garden’. Species of interest include the following sessile bioconstructors: ross coral, *Pentapora foliacea*, pink sea fans, *Eunicella verrucosa*, and nationally rare species such as sunset coral, *Leptopsammia pruvoti*. For many years, these reefs were subject to intensive demersal-towed fishing (trawling and scallop dredging) activity (Mason 1983, Andrews 2008). These gears inflicted adverse environmental and social impacts through removal of epibenthic biodiversity, structural damage to underlying mudstone habitats (Hiddink et al. 2008), and damage and loss of local static gear pots (Clover 2022). The environmental destruction caught the attention of several local groups, including divers who witnessed the impacts (e.g. piles of uprooted sea fans) first-hand, and was later systematized

ically documented through the Devon Wildlife Trust (DWT) led ‘Lyme Bay Reefs’ project (DWT 2007).

Concerns about demersal-towed fishing damage were first raised in the early 1990s, which led to a voluntary closure in 2001 (Hiddink et al. 2008, Mangi et al. 2011). The South-ern Sea Fisheries Committee considered a byelaw to protect reef features in 2001, but following opposition from large vessel representatives, a voluntary agreement was developed between DWT, the South West Fish Producers’ Organisation, and local fishermen (Rees et al. 2010a) to protect two key areas (Saw Tooth Ledges and Lanes Ground) of rocky reef from demersal-towed fishing gear (Fig. 1). In 2006, Natural England applied for a Ministerial Stop Order to exclude dredging from 60 square nautical miles of the seabed, which resulted in an expansion of existing voluntary closures (DWT 2007, Hiddink et al. 2008) by the then Fisheries Minister to include two further important areas (East Tennants and Beer Home Ground). On 11 July 2008, voluntary closures were extended to create a single continuous 206 km² protected area, which was ratified through a Statutory Instrument (SI) [The Lyme Bay Designated Area (Fishing Restrictions) Order 2008] by the UK government (Defra 2008), under guidance from Natural England. This created a legal framework through which to enforce the demersal-towed fishing ban. The primary aim was to conserve Lyme Bay’s reef substrate and associated biodiversity from the impacts of trawling and dredging (for shellfish) with heavy demersal-towed fishing gear (Defra 2008), through whole-site protection. This created, at that time, the UK’s largest and most strongly protected MPA for reef features. Within the SI boundary, static gear fishing (pots and nets) and SCUBA diving to collect King scallops (*Pecten maximus*, Pectinidae) are permitted.

In 2011, Lyme Bay was granted further and extended protection by the creation of a Site of Community Importance, which later became an SAC to meet the obligations of the European Union Habitats Directive [(92/43/EEC) European Commission 2000] for the Annex I reef habitat through a feature-based management approach (Natural England 2015). The SAC overlapped the area encompassed by the SI and correspondingly excluded demersal-towed fishing from the conservation feature [Annex I reef: bedrock reef, stony reef, and biogenic reef (Fig. 1)] outside the SI while allowing the same activity to take place between the reef features (Jackson and Mcleod 2000). Similar to the SI, static gear fishing and fishing with SCUBA gear are permitted across the SAC. The management objectives for the SAC are to ‘maintain’ the reef features. The resultant overall spatial management regime in Lyme Bay presently consists of a 206 km² SI and an overlapping ~270 km² SAC (Fig. 1). The combination of the SI and the SAC forms the boundary of the Lyme Bay MPA (Rees et al. 2021a). The governance of the Lyme Bay MPA has been described as ‘government-led’ (Singer and Jones 2021) as the conservation objectives for the MPA are enforced by the two local Inshore Fisheries and Conservation Authorities (IFCAs) using the legal tools of byelaws and permitting byelaws.

## Methods and method development

### Ecological monitoring regime

Despite MPA establishment accelerating around the globe, accurate performance assessments and subsequent adaptive management have been impeded by a lack of robust monitor-
<figure><img src="image_1.png" /><figcaption>**Figure 1** Map of Lyme Bay spatial management regimes, including (1) the SI (dashed line; 2008–present), (2) the Special Area of Conservation (SAC) (dotted line; 2011–present), and (3) the Previous Voluntary Closures (shaded polygons; 2001–08 for Lanes Ground and Saw Tooth Ledges; 2006–08 for East Tennants and Beer Home Ground).</figcaption></figure>

ing, due to limited management resources (Gill et al. 2017). In Lyme Bay, an ecological monitoring scheme, designed to evaluate epibenthic and nektonic community restoration, was embedded into MPA management (Stevens et al. 2014). Cost-effective video methods were selected to minimize damage to the recovering ecosystem. Static baited remote underwater video (BRUV) surveys documented mobile species recovery, and a novel high-definition towed flying video array (Fig. 2) was purposefully designed to nondestructively quantify sedentary and sessile species by obtaining standardized 200 m video transects of the seabed, with minimal benthic contact (Sheehan et al. 2010). This method underwent comparison with traditional towed video methods, which maintain seabed contact (Sheehan et al. 2016). The towed flying video array outperformed in terms of adaptability across habitat types, ease of deployment, and minimizing seabed disturbance, while providing high-quality ecological measurements. Joint Nature Conservation Committee also assessed the method as the current best practice at a European level to evaluate the marine habitat condition in response to management interventions (Hitchin et al. 2015). Since then, the towed flying video array has been adopted widely by regional fishery regulators such as the Devon & Severn IFCA to ground truth and map various benthic habitats (Ocean Ecology Limited 2015, 2016).

Baseline MPA monitoring is desirable to fully capture species and habitat recovery trajectories from a pre-exploited state (Claudet and Guidetti 2010). However, resource limitations often create time-lags between MPA establishment and monitoring initiation (Ahmadia et al. 2015, Gill et al. 2017). In Lyme Bay, baseline surveys, funded by Defra and Natural England, were conducted as soon as feasible, 6 weeks after MPA implementation in Summer 2008. The ecological monitoring regime in Lyme Bay (Stevens et al. 2014) was designed to follow a Before–After Control–Impact (BACI) experimen-

tal survey design (Underwood 1994), with the Summer 2008 treatment acting as the Before treatment due to the low recoverability of the surveyed species (Glasby 1997).

The annual surveys include the following samples sites: (1) inside the whole-site SI (SI); (2) inside the whole-site SI and within previous voluntary closures (PVC); (3) within the feature-based SAC but outside the whole-site MPA in preidentified Annex I reef habitats (SAC); and (4) in Open Controls where demersal-towed fishing is still permitted (Table 1). Historical fishing effort and substrate distributions were incorporated into site selection to strive for ecological analogy between MPA and Open Control sites (Stevens et al. 2014). Meanwhile, PVC sites represented relatively ‘unimpacted’ areas, as benchmarks of recovery potential. Video surveys have been conducted annually, in the summer months since 2008 (Table 1).

## Developing a novel socioeconomic monitoring framework

MPAs are recognized as having linked social and economic dynamics (Rees et al. 2018) and meeting socioeconomic outcomes is increasingly acknowledged as a key component of MPA success (MacKeracher et al. 2019). Despite the many descriptive arguments for conservation-driven economic benefits, an overarching obstacle to improving marine environmental management is the limited empirical evidence of MPA impacts on socioeconomic outcomes (Wells et al. 2016, Rees et al. 2021a).

Lyme Bay presented an opportunity to ensure socioeconomic outcomes retained adequate attention by developing a world-first integrated socioeconomic and ecological monitoring regime. A standardized protocol was developed, using questionnaires to evaluate the monetary and nonmonetary
<figure><img src="image_2.png" /><figcaption>**Figure 2** (a, b) Towed flying video array and (c) frame grab example with quadrat overlay and scaling lasers. Grab taken from a 50 m transect of video from the 2022 survey.</figcaption></figure>

value of Lyme Bay MPA to various marine stakeholder groups (Rees et al. 2010b). These were initially piloted at small scale to determine the MPAs worth to: dive businesses, dive clubs, sea anglers, and charter boat operators, which provided a baseline against which socioeconomic outcomes were repeatedly assessed over time [2008–11 (Rees et al. 2015)]. Questionnaires were supplemented by the measurement of additional indicators, including visit frequency to key recreational sites, subjective fisher wellbeing, and fishery landings quantity and value (Rees et al. 2015, 2021a). This socioeconomic evaluation framework has since been applied to various MPAs across the British Isles, including Berwickshire, Jersey, and Sussex (Blue Marine Foundation 2023a).

## Expansion of existing monitoring regime

Marine monitoring in the Lyme Bay MPA has resulted in the longest ecological dataset integrated with socioeconomic research on MPAs (Hitchin et al. 2015). Ecological surveys are ongoing and have expanded to include tagging and tracking of various fish species with acoustic telemetry (through

the FISH INTEL project) and analysis of the benthic soundscape (Davies et al. 2020) using the Acoustic Complexity Index (ACI), which demonstrated potential to cost-effectively signal complex changes to the benthos. The monitoring regime now also includes surveillance of the UK's largest offshore rope-cultured mussel farm, which is co-located within Lyme Bay adjacent to the MPA. Multimethod analysis with the same nondestructive technologies has demonstrated the mussel farm's value as a potential OECM (other effective area-based conservation measure), which provides *de-facto* protection from fishing and has facilitated infaunal community recovery (Mascorda-Cabre et al. 2023), and the establishment of a mussel bed reef on the previously degraded seabed beneath the farm (Bridger et al. 2022). The unique mussel farm structure mandated the creation of the 'PelagiCam', a novel video method suitable for studying pelagic communities aggregated by various marine structures (Sheehan et al. 2020). Understanding the unique interactions between the coexisting MPA and mussel farm OECM is a key avenue for future research in Lyme Bay.
<table><caption>**Table 1.** Details of the towed flying video array and BRUV surveys conducted annually in the Summer, for the continuous ecological monitoring regime in Lyme Bay MPA.</caption><thead><tr><th rowspan="2">Monitoring method</th><th rowspan="2">Video analysis approach</th><th rowspan="2">Deployment/video length</th><th colspan="4">Number of survey sites (sampled annually)</th></tr><tr><th>SI</th><th>SAC</th><th>PVC</th><th>OC</th></tr></thead><tbody><tr><td>Towed flying video array</td><td>Two-tiered analysis approach:<br>(1) The video transect is analysed in full to calculate the density (per m<sup>2</sup>) of all rare and conspicuous taxa present across the entire transect.<br>(2) A subsample of 30 screen grabs per transect is exhaustively analysed to calculate the density (per m<sup>2</sup>) of all individual and discrete colonial organisms.</td><td>20-min deployment (~200 m transect)<br>20 min analysed one replicate per site</td><td>18</td><td>15</td><td>15</td><td>18</td></tr><tr><td>BRUVs (baited remote underwater video)</td><td>All individuals of all mobile taxa are documented per minute. The MaxN per taxon is then calculated as the maximum number of individuals observed in any 1-min segment over the 30-min video [see Davies et al. (2021, 2022a) for further details]</td><td>45-min deployment 30 min analysed (after 5-min settling period)<br>Three replicates per site</td><td>6</td><td>6</td><td>6</td><td>6</td></tr></tbody></table>

Surveys are conducted in the following treatments: Statutory Instrument (SI), Special Area of Conservation (SAC), Previous Voluntary Closures (PVC), and Open Controls (OC).

## Ecological findings

### Ecological recovery and alternative biodiversity indicators

Benthic recovery in Lyme Bay following the demersal-towed fishing ban has been regularly assessed throughout the past decade (Sheehan et al. 2013a, Davies et al. 2021, 2022a). This has revealed novel insights into species recovery rates and contributed to the previously scarce body of evidence around temperate MPA efficacy. The first study (Sheehan et al. 2013a) assessed the ‘passive recovery’ (Elliott et al. 2007) of macro epi-benthic species, from 2008 to 2011. Towed video data for four treatments (MPA, Previous Voluntary Closure, Near Open Control, and Far Open Control) were compared to statistically assess whether the reef ecosystem in the MPA had become more alike the Previous Voluntary Closure treatment and more distinct from the Open Controls over time. Definitive evidence of recovery (signified by a significant Year × Treatment interaction) was apparent for species richness, two of the nine sessile indicator taxa (*P. foliacea* and *Phallusia mammillata*), and one of the four mobile indicator taxa [*Pecten maximus* (Sheehan et al. 2013a)]. Meanwhile, five indicator species (*Alcyonium digitatum*, *E. verrucosa*, grouped hydroids, *Asterias rubens*, and *Necora puber*) exhibited significant temporal population increases inside the MPA [but displayed no significant Year × Treatment interaction (Sheehan et al. 2013a)]. These results were among the earliest empirical evidence for ecological recovery timescales in temperate MPAs following demersal-towed fishing bans. Conclusions broadly aligned with the wider literature by detecting early stages of recovery within the first few years of MPA establishment (Idjadi et al. 2006, Mumby and Harborne 2010, Caselle et al. 2015). However, full recovery is thought to occur over decadal

timescales (Babcock et al. 2010), and measuring full recovery rates *in-situ* remains a priority for future research in Lyme Bay.

Continuous annual video monitoring and yearly reports have revealed a more complete picture of ecological recovery. Analysis of data from both the towed and baited surveys found increases in trait diversity and functional redundancy within the MPA (Davies et al. 2022b). In doing so, the study linked ecosystem-based management to increased ecosystem functions and services and suggested this may increase the areas ecological resilience when confronted by additional threats. Another study examined the MPA effects on species beyond designated priority sessile reef features (Davies et al. 2021). It was hypothesized that mobile species would indirectly benefit from the whole-site MPA approach due to increased availability of resources and essential fish habitats created by benthic recovery (Rabaut et al. 2010).

Trends in exploited and nonexploited mobile species populations were compared [using BRUV abundance data (2009–19)], to determine whether ongoing exploitation of targeted species with static fishing gears was limiting their recovery (Davies et al. 2021). Relative to Open Controls, overall taxonomic richness of all mobile species significantly increased inside the MPA over time, while total abundance increased in both treatments (Fig. 3). When exclusively examining exploited fish, 430% increases in taxon richness and 370% increases in total abundance were evident inside the MPA over 11 years (Fig. 4). However, number of exploited fish taxa also significantly increased over time in the Open Controls, potentially signifying spillover effects from the MPA (Fig. 4; Davies et al. 2021). Conversely, exploited invertebrates (crab, lobster, cuttlefish, and whelk) exhibited no signs of recovery in the MPA (Davies et al. 2021), potentially explained by increased potting within the MPA over this period (Rees et al. 2021a).
<figure><img src="image_3.png" /><figcaption>**Figure 3** Number of taxa (a) and total abundance (fourth root transformed) (b) by Year across treatments (Marine Protected Area [MPA] = circles; and Open Controls [OC] = triangles). Lines and equations show linear regression equation coefficients. Points with errors bars show mean values and SEs. Image (c) displays the baited remote underwater camera rigs and (d, e) show sample stills from baited video analysis (in the MPA treatment) (from Davies et al. 2021).</figcaption></figure>

Nevertheless, these findings demonstrated that when implemented effectively, the benefits of a whole-site approach can extend beyond priority features and facilitate a win-win scenario for conservation and fisheries alike (Davies et al. 2021).

## Understanding the value of the 'whole-site approach'

Feature-based protection signifies an MPA management approach, which protects specific features (habitats or species) of interest, based on visible boundaries. It poses challenges that limit the recovery potential of associated species, but it remains the default approach for UK MPAs. For instance, despite Brexit, the European Union Habitats Directive (92/43/EEC; European Commission 2000) remains a highly influential legislative framework for establishing and enforcing ecologically meaningful MPAs, both in the UK [through the Habitat Regulations, 2017 (Defra 2021)] and Europe (European Commission 2000).

However, it incentivizes feature-based protection by outlining an exclusive list of priority habitats and species, for which it necessitates the creation of SACs and Special Protected Areas (SPAs). Therefore, characterizing the risks of feature-based protection has been critical in optimizing MPA management.

MPAs are often established in exploited ecosystems, whose current conditions rarely reflect the potential extent of priority features in pristine condition. This is limiting when feature-based management is interpreted to mean delineating the boundaries of designated features and conserving them in their current state. In Lyme Bay, observations of recovering sessile Reef Associated Species (RAS) on pebbly sand between conspicuous rocky reef patches during towed video analysis prompted an investigation into the true spatial extent of these species. A study tested the hypothesis that, significantly more sessile RAS would be supported by inter-reef pebbly sand habitats protected from demersal-towed fishing, than similar,
<figure><img src="image_4.png" /><figcaption>**Figure 4** Number of taxa (a) and total abundance (c) of exploited fish by year and treatments, and number of taxa (b), and total abundance (fourth root transformed: (d) of non-exploited fish by year and treatments (Marine Protected Area [MPA]: circles and Open Controls [OC]: triangles). Lines and equations show linear regression equation coefficients. Points with error bars show mean values and SEs (from Davies et al. 2021).</figcaption></figure>

unprotected habitats (Sheehan et al. 2013b). All sessile RAS found on pebbly sand habitats were enumerated from towed flying video array surveys. A BACI survey design (Underwood 1994) compared RAS in the MPA with those in Open Control sites, Before (2008) and After (2011) the closure (Sheehan et al. 2013b). Mean abundance of overall RAS was significantly greater (+158%) in the After MPA treatment than both the Before and Open Control (*P* < .05) treatments (Sheehan et al. 2013b). Similarly, four out of six indicator species (*P. foliacea*, *P. mammillata*, *A. digitatum*, and branching sponges) abundances demonstrated significant temporal increases inside the MPA relative to Open Controls (*P* < .05). Although the remaining two species (*E. verrucosa* and hydroids) exhibited no significant Time × Treatment interaction, they increased sig-

nificantly over time inside the MPA [*P* < .05 (Sheehan et al. 2013b)]. These seminal results had major marine conservation policy implications by demonstrating how the true extent of a depleted priority feature can be significantly greater when allowed to re-colonize surrounding space in the absence of disturbance (Sheehan et al. 2013b). This revealed fundamental pitfalls in feature-based protection and demonstrated the benefits of the whole-site approach as an alternative to counteract the effects of shifting baselines (Pauly 1995) by building in additional space for recovery and allowing protected species to define their own limits.

Further comparison of the whole-site approach with feature-based management was facilitated by Lyme Bay's unique combination of two co-located MPAs with contrast-
ing management styles (SAC = feature-based and SI = whole-site protection). Since the SAC was established after the SI, the recovery of taxa was compared for the first 7 years following respective MPA implementation (SI = 2008–14 and SAC = 2012–18) relative to Open Controls (Davies et al. 2022a). Changes in assemblage composition, number of taxa, and total abundance were quantified from both towed and baited annual video surveys and Functional Richness and Redundancy were measured to explore differences in ecosystem function and resilience (Davies et al. 2022a). Both management approaches resulted in benthic recovery over time when compared to Open Control sites. However, total abundance of taxa increased by just 15% in the SAC compared to 95% in the whole-site SI. Furthermore, functional redundancy of mobile taxa was 7% greater inside the whole-site MPA compared to the feature-based SAC (Davies et al. 2022a), potentially offering greater protection against additional threats [e.g. biological invasions and extreme climatic events (McLean et al. 2019, Sheehan et al. 2021)]. Overall, evidence from this study supported the advantages of a whole-site approach for achieving generally faster ecological recovery and greater environmental resilience, and for protecting biodiversity beyond preselected target features (Davies et al. 2022a).

## Climate resilience

As human-driven climate change continues to impact the marine environment, threats such as rising water temperatures, ocean acidification, and deoxygenation are intensifying (Garcia-Soto et al. 2021, Gong et al. 2021). MPAs have the potential to protect against environmental threats by limiting additional anthropogenic pressures on marine systems (Graham et al. 2013). However, few studies have quantified this buffering potential in situ, in the context of climate change (Freedman et al. 2020, Jacquemont et al. 2022). Storm events represent one potential climate threat to benthic assemblages in coastal subtidal habitats, which are often affected by sea surface wave action (Connell et al. 1997). In the northeast Atlantic, the most-severe wave heights are projected to increase under climate change and tropical cyclones have been increasing in frequency (Wolf et al. 2020, Lima et al. 2021); however, few studies have examined whether the impacts of extreme, high energy weather events on subtidal temperate reefs can be reduced by MPA protection (Choat et al. 1988, Holbrook et al. 1994). An opportunity to explore MPA-driven climate resilience arose when severe storms, of unusual intensity inflicted substantial damage to the benthos in Lyme Bay in 2013–14. The storms took place over an 8-week period from mid-December to mid-February and constituted the most energetic storm period since 1953 (Masselink et al. 2016). The events effectively reduced the benthic diversity inside the MPA to resemble a preprotected state.

Long-term monitoring in Lyme Bay provided a novel opportunity to quantify the magnitude of natural disturbance caused by the extreme storm events, and understand subsequent recovery (Sheehan et al. 2021). The study explored whether ecological resilience to extreme climatic disturbance was greater inside Lyme Bay MPA relative to Open Control sites subject to continued demersal-towed fishing. Results indicated that the MPA broadly supported a greater number of taxa and overall abundance of individuals than the Open Control sites over the monitoring period (2008–18). When patterns of recovery over postdemersal-towed fishing and post-

<figure><img src="image_5.png" /><figcaption>**Figure 5** Number of taxa (a) and abundance (b) over Years Since Impact in the Lyme Bay Marine Protected Area (MPA; blue) and Open Control treatments (Open; purple) and compared for post-Demersal-Towed Fishing (Post-DTF) dashed lines and triangles) and post-Storm (solid lines and diamonds) periods. Lines show GAMMs estimates with shading showing 95% CIs. Points with errors bars show mean values and 95% CI (from Sheehan et al. 2021).</figcaption></figure>

storm periods were compared, both demonstrated an increase in abundance of taxa inside the MPA with years since impact (Fig. 5). However, the overall abundance and number of taxa documented inside the MPA were relatively stable during the first 3 years after MPA implementation. Conversely, following the storm events, both response metrics exhibited an immediate increase between 2014 and 2017. This demonstrated that benthic ecosystem recovery was faster inside the MPA after the storm damage, than in the period following the demersal-towed fishing ban, indicating the potential of MPAs to improve the resilience of benthic fauna to storm events (Sheehan et al. 2021). A supplementary analysis of wave exposure observations found that 29% of UK coastal reef MPAs may experience similar wave climate extremes and may therefore be comparably impacted in the future (Sheehan et al. 2021). Overall, these findings demonstrate the increasing need for further protection of coastal habitats against climate change-related threats, which may be aided by MPA protection. How-
ever, by focusing on a single climate stressor, and a single stochastic event, this study provides a limited glance into the interactions between MPAs and the broad range of chronic and acute environmental threats caused by climate change. Thus, a key avenue for future research is to examine how the full range of climate stressors and their interactions may influence the future efficacy of MPAs and the resilience of protected features.

## Investigating fish territory size

The nonspecies-specific nature of video analysis provides flexibility to adapt to emerging management needs, as archived footage can be reanalysed in response to new questions. For instance, in 2015, a live wrasse (Labridae) fishery developed in Devon to supply cleaner fish to Scottish salmon farms (Davies 2016). To understand more about the behaviour of these fishes and the potential impacts of the live fishery, a novel method of territory size was developed (Davies and Sheehan 2019). The opportunity arose after 'laser-chasing' behaviour in wrasse was observed during the Lyme Bay annual towed video analysis (Davies and Sheehan 2019). Several wrasse species could be witnessed clearly pursuing the lasers projected onto the seabed (used to standardize the video transect width). Inter- and intraspecific differences in behaviour were explored by comparing chase distance (the estimated length of laser pursuit) and chase likelihood in cuckoo wrasse (*Labrus mixtus*) and goldsinny wrasse (*Ctenolabrus rupestris*) of different sizes and colour presentations (representing either 'dominant males' or 'females/subordinate males'). Various behaviours were identified, and each wrasse-laser interaction was classified as: flee, neutral, interest, or chase, each signalling differing levels of aggression and territoriality (Benelli et al. 2015). Results found significant interspecific differences in response type and chase distance whereby cuckoo wrasse were significantly more likely to chase the laser and for longer distances than goldsinny wrasse [$P < .001$ (Davies and Sheehan 2019)]. Intraspecific differences in behaviour were only observed in goldsinny wrasse, with significantly longer chase distances associated with larger fish. Inferring territory size from laser chase behaviour relies on various assumptions (e.g. chase response is exclusively motivated by territorial defence), which require further testing. However, overall, these results demonstrate promise for applying this method as a proxy for territory size in fish spatial ecology studies.

## Socioeconomic outcomes

### Quantifying socioeconomic benefits

A series of studies were conducted to quantify the socioeconomic effects of the demersal-towed fishing closure in Lyme Bay in 2008. Profound impacts, both positive and negative, within the social and economic system were detected, including the proliferation of commercial static fishing gears inside the MPA and increased conflicts between static and mobile fishermen outside MPA boundaries (Mangi et al. 2011, Rees et al. 2021a). The static fishery consists of pots for whelk (*Buccinum undatum*), brown crab (*Cancer pagurus*), cuttlefish (*Sepia officinalis*), and lobster (*Homarus gammarus*), and nets for skates (Rajidae) and flatfish, particularly lemon sole (*Microstomus katτ*). Reef-based recreational activities (diving and angling) and associated businesses (charter boats and dive businesses) were valued and found to be highly important to

the local economy, and because of their dependence on the reef ecosystem (Rees et al. 2010b), value estimates contributed to the rationale for the demersal-towed fishing closure. In the 3 years following the closure, recreation participants increased their use of the MPA resulting in a £2.2 million increase in expenditure (divers and anglers) and associated turnover [dive businesses and charter boat operators (Rees et al. 2015)].

A follow-up study undertaken 10 years post-MPA designation, compared fishing activity in the years before and after the demersal-towed fishing ban using 3-year averages between 2005 and 2017. A demonstrable increase in vessels using static gear inside and outside the MPA was found, with a significant increase in static-gear vessels inside the MPA over time (Rees et al. 2021a). Despite an increase in static fishing activity, mean weight of static gear landings per vessel per month decreased across the same time periods by 110 kg, while landings value per vessel per month significantly increased by £1452 (Rees et al. 2021a). The increased value was thought to be due to greater catches of high value species such as lobster, which are associated with the reef ecosystem.

Demersal-towed fishing activity within the MPA, which previously provided high volume and value landings, ceased in July 2008. Comparing the time periods before and after MPA establishment (2005–08 and 2014–17), demersal-towed fishing effort increased outside the MPA in terms of weight and value (Rees et al. 2021a). While the spatial distribution of mobile fishing changed in response to the MPA, the overall value of mobile demersal gear landings in 2017 was comparable to fishing activity prior to the MPA designation. However, in terms of social effects, fishers predominantly using mobile demersal gear reported material losses and lower subjective well-being, meanwhile static gear fishers reported comparatively higher levels of subjective wellbeing over time (Rees et al. 2021a). Crucially, the wellbeing benefits experienced by static fishers were attributed to their involvement with the Lyme Bay Fisheries and Conservation Reserve Consultative Committee demonstrating the importance of community collaboration and stakeholder engagement in maximizing social benefits for local communities and increasing engagement with management strategies (Rees et al. 2021a).

## Optimal potting effort

Restrictions on demersal-towed fishing inside MPAs can reduce spatial overlap between different commercial fishing sectors while providing new opportunities for alternative and novel fishing methods, particularly static gear fishing (Mangi et al. 2011, Enever et al. 2022). In Lyme Bay, this has led to increases in commercial pot fishing, targeting nonquota high-value species such as brown crab (*C. pagurus*) and European lobster (*H. gammarus*). Pot fishing can cause discernible physical disturbance to long-lived, slow growing benthic taxa, during the hauling and retrieval process (Gall et al. 2020) and fishermen feared that increases in fishing effort would jeopardize the sustainability of future stocks. To address these concerns, a manipulative field experiment was designed to characterize potential environmental impacts of potting (Rees et al. 2021b). In collaboration with local fishermen, areas of reef were experimentally exposed to a pot fishing density gradient [unpotted, low (5–10 pots per 0.25 km²), medium (15–25 pots per 0.25 km²), and high (≥30 pots per 0.25 km²)] over 4 years. Impacts on the seabed were assessed using the towed flying video array and BRUV surveys, while the effect on catch
<figure><img src="image_6.png" /><figcaption>**Figure 6** Mean abundance (PERMANOVA main and pairwise results from Table 3 in Rees et al. 2021b) + SE of (a) brown crab (*Cancer pagurus*) [Control (No): *n* = 609, low: *n* = 583, medium: *n* = 602, and high: *n* = 481] and (b) European lobster (*Homarus gammarus*) [Control (No): *n* = 106, low: *n* = 109, medium: *n* = 104, and high: *n* = 69] by pot density treatment. Main test significance (*P* ≤ .05) is denoted with an asterisk, and lines above bars indicate the location of significant differences (from Rees et al. 2021b).</figcaption></figure>

was measured seasonally using an experimental, standardized potting protocol (outlined in Rees et al. 2021b), which quantified differences in catch per unit effort between potting densities.

Negative effects of medium and high pot fishing densities, on both the benthic epibiota and the fishery were evident. Mean abundance of *P. foliacea* was 83% greater in the unpotted treatment compared to the potted treatments. Abundance of *P. mammillata* was significantly lower in the medium and high treatments relative to the low and unpotted treatments (Rees et al. 2021b). High potting effort also reduced the catch per unit effort of commercially targeted species, which was 19% and 35% lower for *C. pagurus* and *H. gammarus*, respectively, when compared to medium potting densities (Fig. 6). Importantly, this study demonstrated seminal evidence of a pot fishing density 'threshold' (medium density, 15–25 pots per 0.25 km²), above which, adverse benthic impacts became significantly greater and the abundance of commercially important crustaceans was significantly reduced (Fig. 6; Rees et al. 2021b). This suggested that commercial pot fisheries can be compatible with conservation objectives inside MPAs when maintained at low, sustainable levels. Estimating optimal fishing thresholds represents a key avenue for future research, which could be applied across various fisheries to improve management. In Lyme Bay specifically, further exploration is warranted to determine the effects of other static gears such as netting and whelk potting, which have also increased in the area (Rees et al. 2021a).

## Collaborative, multisector, management partnerships

Research demonstrates the importance of active local communities and multisector partnerships in achieving conservation outcomes (Dodson 2014, Stewart et al. 2020). Research in Lyme Bay has been conducted in partnership between the University of Plymouth and local fishermen, with funding from Defra, Natural England, NERC, Blue Marine Foundation,

the European Maritime and Fisheries Fund (EMFF), the South West Inshore Fishermen's Association, South Western Fish Producers' Organisation, the Pig Shed Trust, FLAG, Devon Wildlife Trust, and INTERREG France (Channel) England. Additional expertise was sought from the Marine Biological Association and Plymouth Marine Laboratory to identify appropriate indicator species (Jackson et al. 2008) and support socioeconomic benchmark development (Mangi et al. 2011). Further, local knowledge from fishermen has been integral in informing and delivering successful long-term monitoring and all field research has been conducted on local fishing boats (Morgan et al. 2020). Forming these partnerships was initially challenging due to the broad range of marine interests involved and the delicate nature of existing relationships between the fishing industry and conservationists (Fleming and Jones 2012). However, critical relationships were developed over time through frequent stakeholder engagement and aided further by the co-design approach to MPA monitoring. Future replication of these partnerships in other locations should dedicate time to build trust through transparent and early communication of results and project changes.

To facilitate collaborative conservation and fishery management within Lyme Bay MPA, Blue Marine Foundation established the Lyme Bay Working Group in 2011, uniting a core group of fishermen (from Beer, Axmouth, West Bay, and Lyme Regis) with nongovernmental organizations and regulators. The Working Group developed a 'Memorandum of Understanding' to balance the long-term needs and cultural importance of the fishing community with habitat protection goals. In 2013, the Working Group joined with additional stakeholders to form the Lyme Bay Fisheries and Conservation Reserve Consultative Committee and determined that further intervention was required to manage static fishing increases (Mangi et al. 2011, Rees et al. 2021a) and unquantified recreational angling impacts. Two voluntary codes of conduct were agreed to supplement statutory regulations set by the regional IFCAs in Lyme Bay. These voluntary codes set pot and net limits for static gear fishermen operating within the MPA and
promoted best practice for recreational angling (Blue Marine Foundation 2023b).

Providing incentive to comply with conservation measures and gaining support from local fishermen are considered critical aspects of MPA success (MacKeracher et al. 2019). Collaborative management in Lyme Bay has maintained open communication channels between fishermen, scientists, regulators, and environmental groups, which has increased fisher support and involvement (Rees et al. 2013). Many of the benefits achieved by the MPA can be attributed to the sense of place and resultant stewardship exhibited by local fishermen who participated in collaborative management (Morgan et al. 2020). However, positive local perceptions towards the MPA and support for voluntary conservation measures have been encouraged further by additional interventions from Blue Marine Foundation (with funding from the European Maritime Fisheries Fund). Static gear fishermen abiding by the voluntary codes of conduct, were supplied with facilities to optimize catch quality. Insulated fish boxes, chiller units, ice machines, and a dedicated delivery van allowed fish to be kept in prime condition and transported to market quickly, commanding competitive market prices (Blue Marine Foundation 2023b, Appendix 1). However, voluntary codes exhibit limitations and unregulated static fishing remains a major challenge in Lyme Bay. Future statutory legislation will likely be required to manage the activities of nonlocal boats, who do not adhere to voluntary measures.

## Wider implications of the Lyme Bay model

### Informing national and international conservation frameworks and policy

The work in Lyme Bay represents the UK's longest ecological dataset on MPAs, with integrated socioeconomic research and is considered internationally as a leading example of a long-term MPA monitoring study (Hitchin et al. 2015). Due to seminal findings on ecological recovery rates and the benefits of whole-site MPAs, the impacts of work in Lyme Bay can be detected at a national and international scale. For instance, UK parliamentary documents have referred to Lyme Bay, to inform the UK government on fisheries management and climate adaption (POST 2018, 2022). Meanwhile, research also featured as a case study in the MadeAtUni Climate Action campaign (MadeAtUni 2022), which showcases the contributions of UK universities to meaningful climate action, and Lyme Bay research was the topic of a dedicated session at the International MPA Congress in 2023.

The Lyme Bay model demonstrates the value of collaboration between local fishermen, NGOs, regulators, and scientists through a co-management approach to balance the needs of fisheries with marine conservation objectives (Appendix 2) and has been applied as a blueprint around the globe [e.g. in the Aeolian Islands and the Maldives (M. Robertson, personal communication)]. Fisher collaborations were also integral in identifying the sustainable potting threshold (Rees et al. 2021b), which is now available to fisheries regulators to evaluate sustainability of potting levels in other UK regions. Furthermore, involvement with regional fisheries regulators helped ensure research findings were fed into local byelaws and management plans. For instance, evidence of ecological recovery in Lyme Bay (Attrill et al. 2012, Mangi et al. 2012, Sheehan et al. 2013a, b) supported the implementation of the

Nearshore Trawling Byelaw, developed by the Sussex IFCA (Sussex IFCA 2019).

## Informing a paradigm shift towards the 'Whole-Site' approach

Research from Lyme Bay provided pivotal empirical evidence for the advantages of the whole-site approach in driving linked socioecological gains and helped embed the approach into UK marine policy. Regionally, the science from Lyme has been used by NGOs (J-L. Solandt, personal communication) to defend arguments for advanced protection measures for >20 MPAs around England and Scotland since 2013. Meanwhile, the Lyme Bay model (Fig. 7) has inspired a new tranche of effectively managed whole-site MPAs including those in: Sussex, the Isles of Scilly, and Jersey (Sussex IFCA, 2019, Blampied et al. 2022). The benefits provided by whole-site protection in three Jersey MPAs have already become evident (Blampied et al. 2022). A greater diversity of taxa and a higher proportion of important (Labridae spp. and juvenile bream *Spondyliosoma cantharus*) and vulnerable taxa are present inside whole-site MPAs relative to trawled reference sites (Blampied et al. 2022). Meanwhile, Lyme Bay research has also influenced whole-site-based regional fisheries byelaws, such as the ban on demersal-towed fishing in South Dorset Marine Conservation Zone (MMO 2022a). The call for evidence (MMO 2022b) explicitly cited Rees et al. (2020) and Solandt et al. (2020) as evidence for multisectoral support of the whole-site approach.

At a national scale, Natural England (the government advisory board on managing the environment) has used evidence from the Lyme Bay MPA research to inform its Conservation Advice Packages (Natural England 2017). These documents will be used to guide future management of MPAs around the UK and help meet the obligations of The Conservation of Habitats and Species Regulations (2017). Meanwhile, Lyme Bay research was integral to the inclusion of the whole-site approach (and additional progressive policies) in the UK Governments' 25-year Environment Plan (Defra 2018). This document represents a blueprint for the UK's approach to sustainable fisheries management and meeting international conservation goals.

Furthermore, a major cornerstone in UK marine conservation has been designation of HPMAs, within which extractive, destructive, and depositional activities will be prohibited. An integral step towards designating HPMAs was the Benyon review (Defra, 2019) within which Lyme Bay featured heavily as a key case study and one of few empirical examples of the importance of strong protection within MPAs. The review advised that HPMAs follow a whole-site approach to encourage multispecies benefits and resulted in the designation (in July 2023) of three pilot sites (North East of Farnes Deep, Allonby Bay, and Dolphin Head) as HPMAs, encompassing a total area of 986 km². The announcement of these HPMAs directly referred to evidence from Lyme Bay MPA to illustrate the importance of strong protection for enhancing marine biodiversity (Defra 2023).

## Spreading the word

Translating research findings into policy and disseminating information to wider audiences has relied on science-industry partnerships and the maintenance of diverse communication channels. Online and multimedia engagement has been facilitated by the Lyme Bay MPA website (www.lymebayreserve.
<figure><img src="image_7.png" /><figcaption>**Figure 7** Summary of Lyme Bay research and impact.</figcaption></figure>

co.uk, live since 2013), and active social media channels on Twitter and Facebook, facilitated by Blue Marine Foundation. The MPA achievements have also garnered attention from local and national news outlets (e.g. BBC and ITV news), for instance, Lyme Bay was hailed as a 'world first' by Fishing News and twice been a finalist for the Natura 2000 award in 2014 and 2016. Furthermore, YouTube videos were created to disseminate findings about sustainable potting and storm recovery, and to celebrate the Lyme Bay MPA 10-year anniversary, a short documentary-film—'Lyme Bay: The Road to Recovery' was launched (Murray 2020). The film, produced by Blue Marine Foundation, showcased the importance of building reciprocal relationships of knowledge sharing between fishing communities and conservationists to create a win-win model for sustainable marine management. This 10-year milestone was further commemorated by an in-person community event (MPArty), which was held at the University of Plymouth to

celebrate ongoing accomplishments and express gratitude to the members of the fishery-science-regulator partnerships responsible for Lyme Bay's achievements.

## Multigenerational learning

Findings from Lyme Bay have been translated into alternative formats to aid multigenerational learning. For instance, the story of Lyme Bay was presented as a book chapter to exemplify the roles of 'social learning' and 'place attachment' in fostering stewardship and driving sustainable fisheries development (Morgan et al. 2020). This contributed to the book's wider objective of encouraging a deeper examination of human-nature reciprocity to inspire sustainable interactions. Furthermore, a chapter of the book 'Rewilding the Sea' is dedicated to Lyme Bay's journey to protect 'England's coral garden' and the people involved (Clover 2022). Lyme
Bay is featured among other positive examples to illustrate how letting nature lead benefits both people and industry.

Meanwhile, research findings on storm resilience (Sheehan et al. 2021) and the ecological benefits of the mussel farm (Bridger et al. 2022) have been published in the *Environmental Science Journal for Teens*, to make study results accessible to younger demographics. Further engagement with future generations was facilitated by a Schools Outreach Programme (established by Blue Marine Foundation with funding from the Rita Lila Howard Foundation), delivered by fishermen 'ambassadors' who engaged local school children with the story of marine conservation in Lyme Bay.

## Looking forward

This decade signifies a crucial window of opportunity for marine conservation progress. As we navigate through the UN Decade of Ocean Science for Sustainable Development (2021–30) and move towards the targets of the Sustainable Development Goals, marine conservation is becoming increasingly prominent in public consciousness. This opportunity for intervention is strengthened by coinciding ambitious targets to protect 30% of global biodiversity by 2030, through the ‘30 by 30’ commitment, included in the Kunming–Montreal Global Biodiversity Framework, agreed at the Convention on Biological Diversity (CBD) at COP15 in December 2022 (Belote et al. 2021, UNEP 2022b), and in the UK more specifically through the 25-Year Environment Plan (YEP) (Defra 2018). To help seize this opportunity, funding for continued monitoring in Lyme Bay must be secured to facilitate a greater understanding of the timescales of full ecological recovery. Comparison with HPMAs represents another key area for future research, which could elucidate differences in recovery rates at varying degrees of protection. Further, several new Defra-funded Fisheries–Industry–Science Partnership (FISP) projects applying acoustic telemetry to discern patterns of connectivity between the MPA and adjacent mussel farm have been launched in 2023. This will feed into the wider network of acoustic telemetry studies across the English Channel and situate Lyme Bay MPA within the system of essential fish habitats along the South Coast. Connectivity between and relative importance of habitats for different key species [e.g. thornback ray (*Raja clavata*) and pollack (*Pollachius pollachius*)] will be explored. These projects will support advances in ecosystem-based fishery management.

Collaborative, cross-sector management in Lyme Bay continues through the Lyme Bay Fisheries and Conservation Reserve Consultative Committee, which remains a regular forum for fishermen, regulators, scientists, and NGOs to address emerging challenges and maintain ongoing ecosystem recovery in the Lyme Bay MPA. In 2022, the Lyme Bay Fisherman’s Community Interest Company (CIC) was established, with a collective membership of >45 fishermen. The fishermen have adopted the principles of the original Lyme Bay Working Group to implement best practices in biodiversity protection and fishery management, to create long-term benefits for local communities. The CIC is a pioneering approach that exemplifies how the Lyme Bay fishing community has proactively embraced conservation and sustainable fishing practices within the MPA to safeguard fish stocks and the marine environment for future generations (Blue Marine Foundation 2023b).

Finally, the last major roadblock preventing effective marine conservation and sustainable fisheries management in

Lyme Bay is unregulated static fishing (Rees et al. 2021a). Nonlocal vessels acting outside the voluntary code of conduct are permitted to operate beyond the sustainable limits agreed by local fishermen. Further research is required to better understand the extent of benthic impacts, define sustainable limits, and inform the implementation of additional statutory interventions to ensure that nonlocal boats adhere to sustainable static fishing practices within the MPA.

## Conclusions

Reflecting on 15 years of marine conservation research in Lyme Bay has provided an avenue to explore the sequential progression of events and a platform to highlight the opportunities and challenges associated with managing partially protected MPAs. There are no other MPA studies that expand on this time frame and variety of interdisciplinary research. On one hand, Lyme Bay showcases the potential of spatial protection to create intertwined social and ecological benefits. On the other hand, this review demonstrates how MPA management must continually respond to ever-evolving pressures on the marine environment and exhibits the care, rigour, and expansive collaboration required to deliver successful conservation outcomes.

Research in Lyme Bay highlights the importance of protecting whole sites from threats, to build in space for rewilding and recovery, beyond existing exploited bounds and exclusive priority features. Further, Lyme Bay’s robust monitoring, complete with ecologically analogous control sites, an early baseline, and enumeration of all visible taxa, has allowed a detailed picture of benthic recovery to be drawn and statistically interrogated. While this comprehensive monitoring is a luxury only possible through tirelessly securing ongoing funding commitments, it has resulted in the UK’s leading long-term MPA monitoring study, aided novel method developments (e.g. towed flying video array, pelagicam), and revealed scientific discoveries (e.g. recovery rates, sustainable potting, MPA storm resilience).

The inclusion of previously overlooked socioeconomic monitoring and benchmarks in Lyme Bay monitoring has raised the profile of the human dimension of MPA effects, including well-being and income changes, which ultimately affect ecological outcomes through local support and buy-in. Integral to this approach has been the development of multisector partnerships and involvement of local fishermen, whose local knowledge and practical experience have underpinned ongoing annual monitoring. Finally, diverse outreach and communication methods have been crucial in aiding transgenerational engagement and translating findings into ambitious national and international MPA frameworks and policy changes.

Social–ecological theory suggests that transformations to sustainability occur in ‘niches’ at the local level (Lotz-Sisitka et al. 2015). Lyme Bay is an example of such a ‘niche’, where the original desired social and ecological outcomes were born from a dedicated and active local community, who used their sense of place to defend their local area. However, ongoing successes are now owed to the collective efforts of fishermen, researchers, NGOs, and regulators, whose shared knowledge is continually fed into the adaptive co-management cycle through continuing partnerships. It is hoped that the telling of this story, and the lessons within, will help enhance the protection of existing MPAs and guide the effective design of new
MPAs as we progress through the Ocean Decade towards the targets of the ‘30 by 30’ commitment.

## Acknowledgements

We thank all of the Lyme Bay fishermen who have supported and engaged with this research. We especially thank John Walker, Robert King, and Kieran Perree, who also helped us to develop research methods, and have patiently trained count- less UoP staff and students over the years to become safe, seaf- worthy marine scientists. We thank all our funders, stakehold- ers, technical staff, and students who have contributed to this research since 2008, with special thanks to Natural England’s Gavin Black who has tirelessly ensured funding for Lyme Bay’s continued monitoring.

*Conflict of interest:* None declared.

## Data availability

There are no new data associated with this article.

## Funding

This work was funded by the University of Plymouth, Blue Marine Foundation, Defra, and Natural England. These fun- ders have also contributed towards the Lyme Bay monitoring research as well as the Natural Environment Research Coun- cil (NERC NE/M005208/1), Pig Shed Trust, South West In- shore Fishermen’s Association, South Western Fish Producers’ Organisation, the European Maritime Fisheries Fund, Dorset & East Devon Fisheries Local Action Group (FLAG), Devon Wildlife Trust, and Interreg France (Channel) England.

## Author contributions

E.S., M.A., and S.R. conceived this research with Tim Glover from the Blue Marine Foundation. C.R. and E.S. led the writ- ing (visualisation, reviewing, editing) of this article with sup- port from all authors who also approved the submitted ver- sion. E.S., M.A. and S.R. were responsible for funding acqui- sition and methodology development. Formal analyses dis- cussed in this paper were conducted by A.R., B.D., L.H. and E.S.

## References

* Ahmadia GN, Glew L, Provost M *et al.* Integrating impact evaluation in the design and implementation of monitoring marine protected areas. *Philos Trans R Soc Lond B Biol Sci* 2015;370:20140275. https://doi.org/10.1098/rstb.2014.0275.

* Andrews T. *Impact assessment of measures to protect marine biodi- versity in Lyme Bay*. Report to Defra. Department for Environment Food & Rural Affairs, London, 2008, 57.

* Attrill MJ, Austen MC, Cousens SL *et al.* Lyme Bay—a case study: mea- suring recovery of benthic species; assessing potential “spillover” effects and socio-economic changes, three years after the closure. Report 1: response of the benthos to the zoned exclusion of bot- tom towed fishing gear in Lyme Bay, March 2012. University of Plymouth-led consortium, Plymouth, 2012, 82.

* Babcock RC, Shears NT, Alcala AC *et al.* Decadal trends in ma- rine reserves reveal differential rates of change in direct and indi- rect effects. *Proc Natl Acad Sci USA* 2010;107:18256–61. https://doi.org/10.1073/pnas.0908012107.

* Belote RT, Barnett K, Dietz MS *et al.* Options for prioritizing sites for biodiversity conservation with implications for “30 by 30”. *Biol Conserv* 2021;264:1–13.

* Benelli G, Romano D, Desneux N *et al.* Sex differences in fighting- induced hyperaggression in a fly. *Anim Behav* 2015;104:165–74. https://doi.org/10.1016/j.anbehav.2015.02.026.

* Blampied SR, Rees SE, Attrill MJ *et al.* Removal of bottom-towed fishing from whole-site Marine Protected Areas promotes mobile species biodiversity. *Estuar Coast Shelf Sci* 2022;276:1–14. https://doi.org/10.1016/j.ecss.2022.108033.

* Blue Marine Foundation. *BLUEprint for MPAs: A Guide to Establish- ing and Managing Marine Protected Areas*. London: Blue Marine Foundation, 2023a, 60.

* Blue Marine Foundation. *Lyme Bay Reserve Case Studies: BLUEPrint for MPAs*. London: Blue Marine Foundation, 2023b, 25.

* Bridger D, Attrill MJ, Davies BFR *et al.* The restoration potential of offshore mussel farming on degraded seabed habitat. *Aquac Fish Fish* 2022;2:437–49. https://doi.org/10.1002/aff.277.

* Caselle JE, Rassweiler A, Hamilton SL *et al.* Recovery trajectories of kelp forest animals are rapid yet spatially variable across a network of temperate marine protected areas. *Sci Rep* 2015;5:1–14. https://doi.org/10.1038/srep14102.

* Ceballos G, Ehrlich PR, Barnosky AD *et al.* Accelerated modern human-induced species losses: entering the sixth mass extinction. *Sci Adv* 2015;1:9–13. https://doi.org/10.1126/sciadv.1400253.

* Choaot JH, Ayling AM, Schiel DR. Temporal and spatial variation in an island fish fauna. *J Exp Mar Biol Ecol* 1988;121:91–111. https://doi.org/10.1016/0022-0981(88)90249-3.

* Claudet J, Guidetti P. Improving assessments of marine protected areas. *Aquatic Conserv Mar Freshw Ecosyst* 2010;20:239–42. https://doi.org/10.1002/aqc.1087.

* Clover C. 2022 *Rewilding the Sea: How to Save Our Oceans*. London: Witness Books, 261.

* Coleman FC, Baker PB, Koenig CC. A review of Gulf of Mexico marine protected areas. *Fisheries* 2004;29:10–21. https://doi.org/10.1577/1548-8446(2004)29%5b10:AROGOM%5d2.0.CO;2.

* Connell JH, Hughes TP, Wallace CC. A 30-year study of coral abun- dance, recruitment, and disturbance at several scales in space and time. *Ecol Monogr* 1997;67:461–88. https://doi.org/10.1890/0012-9615(1997)067%5b0461:AYSOCA%5d2.0.CO;2.

* Cowie RH, Bouchet P, Fontaine B. The Sixth Mass Extinction: fact, fiction or speculation? *Biol Rev* 2022;97:640–63. https://doi.org/10.1111 brv.12816.

* Da Silva IM, Hill N, Shimadzu H *et al.* Spillover effects of a community- managed marine reserve. *PLoS One* 2015;10:1–18.

* Davies BFR, Attrill MJ, Holmes L *et al.* Acoustic Complexity Index to assess benthic biodiversity of a partially protected area in the south- west of the UK. *Ecol Indic* 2020;111:106019. https://doi.org/10.1016/j.ecolind.2019.106019.

* Davies BFR, Holmes L, Attrill MJ *et al.* Ecosystem benefits of adopt- ing a whole-site approach to MPA management. *Fish Manag Ecol* 2022a;29:790–805. https://doi.org/10.1111/fme.12581.

* Davies BFR, Holmes L, Bicknell A *et al.* A decade implementing ecosystem approach to fisheries management improves diversity of taxa and traits within a marine protected area in the UK. *Divers Distrib* 2022b;28:173–88. https://doi.org/10.1111/ddi.13451.

* Davies BFR, Holmes L, Rees A *et al.* Ecosystem approach to fisheries management works—how switching from mobile to static fishing gear improves populations of fished and non-fished species inside a marine-protected area. *J Appl Ecol* 2021;58:2463–78. https://doi.org/10.1111/1365-2664.13986.

* Davies P, Sheehan EV. Laser chasing behaviour of wild fishes exploited as a tool to compare space use between size, sex and species. *J Appl Ichthyol* 2019;35:1225–33. https://doi.org/10.1111/jai.13982.

* Davies S. *A Review of Wrasse Ecology and Fisheries Interactions*. Brix- ham: Devon & Severn Inshore Fisheries and Conservation Author- ity, 2016.

* Day J, Dudley N, Hockings M *et al.* Guidelines for Applying the IUCN Protected Area Management Categories to Marine Protected Ar-
eas. Best Practice Protected Area Guidelines Series No.19, 2nd edn, IUCN, Gland, 2019, 36. https://portals.iucn.org/library/node/48887

Defra. *A Green Future: Our 25 Year Plan to Improve the Environment.* London: Department for Environment Food & Rural Affairs, 2018, 151.

Defra. *Changes to the Habitats Regulations 2017. Policy Paper.* London: Department for Environment Food & Rural Affairs, 2021.

Defra. Benyon review into Highly Protected Marine Areas. Final Report. London: Department for Environment Food & Rural Affairs 2019

Defra. *Explanatory Memorandum to the Lyme Bay Designated Area Fishing Restrictions Order 2008* 1584. London: Department for Environment Food & Rural Affairs, 2008, 60.

Defra. *First Three Marine Areas Receive Highest Levels of Protection.* Press Release. 5 July 2023, London: Department for Environment Food & Rural Affairs, 2023.

Dodson G. Co-governance and local empowerment? Conservation partnership frameworks and marine protection at Mimiwhangata, New Zealand. *Soc Nat Res* 2014;27:521–39. https://doi.org/10.1080/08941920.2013.861560.

Duarte CM, Agusti S, Barbier E et al. Rebuilding marine life. *Nature* 2020;580:39–51. https://doi.org/10.1038/s41586-020-2146-7.

DWT. Lyme Bay reefs: a 16 year search for sustainability. *Devon Wildlife Trust Report*. UK: Exeter, 2007.

Edgar GJ, Stuart-Smith RD, Willis TJ et al. Global conservation outcomes depend on marine protected areas with five key features. *Nature* 2014;506:216–20. https://doi.org/10.1038/nature13022.

Elliott M, Burdon D, Hemingway KL et al. Estuarine, coastal and marine ecosystem restoration: confusing management and science—a revision of concepts. *Estuarine Coastal Shelf Sci* 2007;74:349–66. https://doi.org/10.1016/j.ecss.2007.05.034.

Enever R, Doherty PD, Ashworth J et al. Scallop potting with lights: a novel, low impact method for catching European king scallop (*Pecten maximus*). *Fish Res* 2022;252:106334. https://doi.org/10.1016/j.fishres.2022.106334.

European Commission. *Managing Natura 2000 sites: The provisions of Article 6 of the Habitats Directive*, In 92/43/EEC. Ed. by European Commission, 2000.

Fleming DM, Jones PJS. Challenges to achieving greater and fairer stakeholder involvement in marine spatial planning as illustrated by the Lyme Bay scallop dredging closure. *Mar Policy* 2012;36:370–7. https://doi.org/10.1016/j.marpol.2011.07.006.

Freedman RM, Brown JA, Caldow C et al. Marine protected areas do not prevent marine heatwave-induced fish community structure changes in a temperate transition zone. *Sci Rep* 2020;10:21081. https://doi.org/10.1038/s41598-020-77885-3.

Gall SC, Rodwell LD, Clark S et al. The impact of potting for crustaceans on temperate rocky reef habitats: implications for management. *Mar Environ Res* 2020;162:105134. https://doi.org/10.1016/j.marenvres.2020.105134.

García-Rubies A, Hereu B, Zabala M. Long-term recovery patterns and limited spillover of large predatory fish in a Mediterranean MPA. *PLoS One* 2013;8:1–10. https://doi.org/10.1371/journal.pone.0073922.

Garcia-Soto C, Cheng L, Caesar L et al. An overview of ocean climate change indicators: sea Surface Temperature, ocean heat content, ocean pH, dissolved oxygen concentration, Arctic Sea Ice extent, thickness and volume, sea level and strength of the AMOC (Atlantic Meridional Overturning Circulation). *Front Mar Sci* 2021;8:642372.

Gell FR, Roberts CM. *The Fishery Effects of Marine Reserves and Fishery Closures*. Washington, DC: World Wildlife Fund, 2003.

Giakoumi S, McGowan J, Mills M et al. Revisiting “Success” and “Failure” of marine protected areas: a conservation scientist perspective. *Front Mar Sci* 2018;5:1–5. https://doi.org/10.3389/fmars.2018.00223.

Gill DA, Mascia MB, Ahmadia GN et al. Capacity shortfalls hinder the performance of marine protected areas globally. *Nature* 2017;543:665–9. https://doi.org/10.1038/nature21708.

Glasby TM. Analysing data from post-impact studies using asymmetrical analyses of variance: a case study of epibiota on marinas. *Aust J Ecol* 1997;22:448–59. https://doi.org/10.1111/j.1442-9993.1997.tb00696.x.

Gong H, Li C, Zhou Y. Emerging global ocean deoxygenation across the 21st century. *Geophys Res Lett* 2021;48:1–9. https://doi.org/10.1029/2021GL095370.

Graham NA, Bellwood DR, Cinner JE et al. Managing resilience to reverse phase shifts in coral reefs. *Front Ecol Environ* 2013;11:541–8. https://doi.org/10.1890/120305.

Grorud-Colvert K, Sullivan-Stack J, Roberts C et al. The MPA Guide: a framework to achieve global goals for the ocean. *Science* 2021;373:eabf0861. https://doi.org/10.1126/science.abf0861.

Gubbay S. *A Coastal Directory for Marine Nature Conservation*. Ross-on-Wye: Marine Conservation Society, 1988, 319.

Halpern BS, Frazier M, Afflerbach J et al. Recent pace of change in human impact on the world's ocean. *Sci Rep* 2019;9:1–8. https://doi.org/10.1038/s41598-019-47201-9.

Hays GC, Koldewey HJ, Andrzejaczek S et al. A review of a decade of lessons from one of the world's largest MPAs: conservation gains and key challenges. *Mar Biol* 2020;167:1–22. https://doi.org/10.1007/s00227-020-03776-w.

He Q, Silliman BR. Climate change, human impacts, and coastal ecosystems in the anthropocene. *Curr Biol* 2019;29:R1021–35. https://doi.org/10.1016/j.cub.2019.08.042.

Hiddink JG, Kaiser PMJ, Hinz H et al. Quantification of epibenthic fauna in areas subjected to different regimes of scallop dredging activity in Lyme Bay, Devon. *NERC Funded Report*, Conducted by the School of Ocean Sciences, College of Natural Sciences, Bangor University, 2008, 1–69.

Hitchin R, Turner JA, Verling E. Epibiota remote monitoring from digital imagery: operational guidelines. *JNCC Report*. Peterborough: JNCC, 2015, 29.

Holbrook SJ, Kingsford MJ, Schmitt RJ et al. Spatial and temporal patterns in assemblages of temperate reef fish. *Am Zool* 1994;34:463–75. https://doi.org/10.1093/icb/34.3.463.

Idjadi JA, Lee SC, Bruno JF et al. Rapid phase-shift reversal on a Jamaican coral reef. *Coral Reefs* 2006;25:209–11. https://doi.org/10.1007/s00338-006-0088-7.

IPCC. Summary for policymakers. In: V Masson-Delmotte, P Zhai, HO Pörtner, D Roberts, J Skea, PR Shukla, A Piraniet al. (eds.), *Global Warming of 1.5°C: An IPCC Special Report on Impacts of Global Warming of 1.5°C above Pre-industrial Levels in Context of Strengthening Response to Climate Change, Sustainable Development, and Efforts to Eradicate Poverty*. Cambridge and New York, NY: Cambridge University Press, 2018.

IPCC. Summary for policymakers. In: H-O Pörtner, DC Roberts, V Masson-Delmotte, P Zhai, M Tignor, E Poloczanska, K Mintenbeck, et al. (eds.), *IPCC Special Report on the Ocean and Cryosphere in a Changing Climate*. Cambridge and New York, NY: Cambridge University Press, 2019, 18–9.

Jackson DL, Mcleod CR. Handbook on UK status of EC habitats directive interest features. *JNCC Report No. 312*. Peterborough: JNCC, 2000, 206

Jackson EL, Langmead O, Barnes M et al. Identification of indicator species to represent the full range of benthic life history strategies for *Lyme Bay and the consideration of the wider application for monitoring of Marine Protected Areas*. Report to the Department of Environment, Food and Rural Affairs from the Marine Life Information Network (MarLIN). Plymouth: Marine Biological Association of the UK, 2008, 60.

Jacquemont J, Blasiak R, Le Cam C et al. Ocean conservation boosts climate change mitigation and adaptation. *One Earth* 2022;5:1126–38. https://doi.org/10.1016/j.oneear.2022.09.002.

Kerwath SE, Winker H, Götz A et al. Marine protected area improves yield without disadvantaging fishers. *Nat Commun* 2013;4:2347. https://doi.org/10.1038/ncomms3347.
Lester SE, Halpern BS, Grorud-Colvert K et al. Biological effects within no-take marine reserves: a global synthesis. *Marine Ecol Prog Ser* 2009;384:33–46. https://doi.org/10.3354/meps08029.

Lima MM, Hurduc A, Ramos AM et al. The increasing frequency of tropical cyclones in the Northeastern Atlantic sector. *Front Earth Sci* 2021;9:745115. https://doi.org/10.3389/feart.2021.745115.

Lotz-Sisitka H, Wals AE, Kronlid D et al. Transformative, transgressive social learning: rethinking higher education pedagogy in times of systemic global dysfunction. *Curr Opin Environ Sustain* 2015;16:73–80. https://doi.org/10.1016/j.cosust.2015.07.018.

MacKeracher T, Diedrich A, Simpfendorfer CA. Sharks, rays and marine protected areas: a critical evaluation of current perspectives. *Fish Fisheries* 2019;20:255–67. https://doi.org/10.1111/faf.12337.

MadeAtUni. *How UK Universities are Tackling the Climate Emergency: Case Studies*. London: Universities UK, 2022.

Mangi SC, Gall SC, Hattam C et al. Lyme Bay—a case-study: measuring recovery of benthic species; assessing potential “spillover” effects and socio-economic changes; 3 years after the closure. Report 2: assessing the socio-economic impacts resulting from the closure restrictions in Lyme Bay. Plymouth: University of Plymouth-led consortium, 2012, 96.

Mangi SC, Rodwell LD, Hattam C. Assessing the impacts of establishing MPAs on fishermen and fish merchants: the case of Lyme Bay, UK. *Ambio* 2011;40:457–68. https://doi.org/10.1007/s13280-011-0154-4.

Mascorda-Cabre L, Hosegood P, Attrill MJ et al. Detecting sediment recovery below an offshore longline mussel farm: a macrobenthic Biological Trait Analysis (BTA). *Mar Pollut Bull* 2023;195:1–11. https://doi.org/10.1016/j.marpolbul.2023.115556.

Mason J. *Scallop and Queen Fisheries in the British Isles*. Farnham: Fishing News Books Ltd, 1983, 144.

Masselink G, Scott T, Poate T et al. The extreme 2013/2014 winter storms: hydrodynamic forcing and coastal response along the south-west coast of England. *Earth Surf Process Landforms* 2016;41:378–91.

McLean M, Auber A, Graham NAJ et al. Trait structure and redundancy determine sensitivity to disturbance in marine fish communities. *Global Change Biol* 2019;25:3424–37. https://doi.org/10.1111/gcb.14662.

MMO. *Decision Document: The South Dorset Marine Conservation Zone*. Dorset: Marine Management Organisation, 2022b, 26.

MMO. *The South Dorset Marine Conservation Zone (Specified Area) Bottom Towed Fishing Gear Byelaw*. Dorset: Marine Management Organisation, 2022a, 6.

Morgan A, Sheehan E, Rees A et al. Towards a marine socio-ecology of learning in the South West of England, Greenwood, D. (eds) Place-based Learning for the Plate., vol 6. Springer, Cham.. In: J Pontius, M Mueller, D Greenwood (eds.), *Place-based Learning for the Plate. Environmental Discourses in Science Education*, Vol. 6. Cham: Springer, 2020, 141–67.

Mumby PJ, Harborne AR. Marine reserves enhance the recovery of Corals on Caribbean Reefs. *PLoS One*, 2010;5:e8657. https://doi.org/10.1371/journal.pone.0008657.

Murray. *Lyme Bay: The Road to Recovery, Short Film*, 2020. (11 August 2023, date last accessed). https://www.lymebayreserve.co.uk/a bout/road-to-recovery.php.

Natural England. *Marine Conservation Advice: Project Summary*. Corporate Report. London: Natural England, 2017.

Natural England. *Site Improvement Plan: Lyme Bay and Torbay*. Worcester: Natural England, 2015, 10.

Ocean Ecology Limited. Pinnacles & Bigbury Bay Seabed Imagery Analysis– Summary Report. Report No. DSISPE0116 Prepared for Devon and Severn IFCA. Gloucester: Ocean Ecology Limited, 2016, 18.

Ocean Ecology Limited. *Start Point to Plymouth Sound & Eddystone SAC Seabed Imagery Analysis & Eunicella verrucosa Condition Assessment—Summary Report*. Report No. DSISPE0215 prepared for Devon and Severn IFCA & Natural England. Gloucester: Ocean Ecology Limited, 2015, 21.

Ostrom E. A general framework for analyzing sustainability of social-ecological systems. *Science* 2009;325:419–22. https://doi.org/10.126/science.1172133.

Pauly D. Anecdotes and the shifting baseline syndrome of fisheries. *Trends Ecol Evol* 1995;10:430.

POST. *Climate Adaptation for Nature*. POSTnote to the House of Parliament, No. 679. London: Parliamentary Office of Science & Technology, 2022, 2–4.

POST. *UK Fisheries Management*. POSTnote to the House of Parliament, No. 572. London: Parliamentary Office of Science & Technology, 2018, 3–4.

Rabaut M, Van de Moortel L, Vincx M et al. Biogenic reefs as structuring factor in *Pleuronectes platessa* (Plaice) nursery. *J Sea Res* 2010;64:102–6. https://doi.org/10.1016/j.seares.2009.10.009.

Rees A, Sheehan EV, Attrill MJ. Optimal fishing effort benefits fisheries and conservation. *Sci Rep* 2021b; 11:1–15. https://doi.org/10.1038/s41598-021-82847-4.

Rees SE, Ashley M, Evans L et al. An evaluation of the social and economic impact of a Marine Protected Area on commercial fisheries. *Fish Res* 2021;235:1–10. https://doi.org/10.1016/j.fishres.2020.10.05819.

Rees SE, Attrill MJ, Austen MC et al. A thematic cost-benefit analysis of a marine protected area. *J Environ Manage* 2013;114:476–85. https://doi.org/10.1016/j.jenvman.2012.10.048.

Rees SE, Attrill MJ, Austen MC et al. Is there a win-win scenario for marine nature conservation? *Ocean Coast Manag* 2010;53:135–45. https://doi.org/10.1016/j.ocecoaman.2010.01.011.

Rees SE, Mangi SC, Hattam C et al. The socio-economic effects of a Marine Protected Area on the ecosystem service of leisure and recreation. *Mar Policy* 2015;62:144–52. https://doi.org/10.1016/j.marpol.2015.09.011.

Rees SE, Pittman SJ, Foster N et al. Bridging the divide: social-ecological coherence in Marine Protected Area network design. *Aquatic Conserv Mar Freshw Ecosyst* 2018;28:754–63. https://doi.org/10.1002/aqc.2885.

Rees SE, Rodwell LD, Attrill MJ et al. The value of marine biodiversity to the leisure and recreation industry and its application to marine spatial planning. *Mar Policy* 2010;34:868–75. https://doi.org/10.1016/j.marpol.2010.01.009.

Rees SE, Sheehan EV, Stewart BD et al. Emerging themes to support ambitious UK marine biodiversity conservation. *Mar Policy* 2020;117:2–10. https://doi.org/10.1016/j.marpol.2020.103864.

Roberts CM, Leary BCO, Mcc>auley DJ et al. Marine reserves can mitigate and promote adaptation to climate change. *Proc Natl Acad Sci USA* 2017;114:6167–75.

Sala E, Giakoumi S. No-take marine reserves are the most effective protected areas in the ocean. *ICES J Mar Sci* 2018;75:1166–8. https://doi.org/10.1093/icesjms/fsx059.

Sala E, Lubchenco J, Grorud-colvert K et al. Assessing real progress towards effective ocean protection. *Mar Policy* 2018;91:11–3. https://doi.org/10.1016/j.marpol.2018.02.004.

Sciberras M, Jenkins SR, Kaiser MJ et al. Evaluating the biological effectiveness of fully and partially protected marine areas. *Environ Evid* 2013;2:1–31. https://doi.org/10.1186/2047-2382-2-4.

Sheehan EV, Bridger D, Nancollas SJ et al. PelagiCam: a novel underwater imaging system with computer vision for semi-automated monitoring of mobile marine fauna at offshore structures. *Environ Monit Assess* 2020;192:1–13. https://doi.org/10.1007/s10661-019-7980-4.

Sheehan EV, Cousens SL, Nancollas SJ et al. Drawing lines at the sand: evidence for functional vs. visual reef boundaries in temperate Marine Protected Areas. *Mar Pollut Bull* 2013;76:194–202. https://doi.org/10.1016/j.marpolbul.2013.09.004.

Sheehan EV, Holmes LA, Davies BFR et al. Rewilding of protected areas enhances resilience of marine ecosystems to extreme climatic events. *Front Marine Sci* 2021;8:1–16. https://doi.org/10.3389/fmar s.2021.671427.

Sheehan EV, Stevens TF, Attrill M, J. A quantitative, non-destructive methodology for habitat characterization and benthic monitoring
at offshore renewable energy developments. *PLoS One* 2010;5:1–8.
https://doi.org/10.1371/journal.pone.0014461.

Sheehan EV, Stevens TF, Gall SC *et al*. Recovery of a temperate reef assemblage in a marine protected area following the exclusion of towed demersal fishing. *PLoS One* 2013a;8:1–12. https://doi.org/10.1371/journal.pone.0083883.

Sheehan EV, Vaz S, Pettifer E *et al*. An experimental comparison of three towed underwater video systems using species metrics, benthic impact and performance. *Methods Ecol Evol* 2016;7:843–52. https://doi.org/10.1111/2041-210X.12540.

Singer R, Jones PJS. Lyme Bay marine protected area: a governance analysis. *Mar Policy* 2021;127:103201. https://doi.org/10.1016/j.marpol.2018.07.004.

Solandt J, Mullier T, Elliott S *et al*. Managing marine protected areas in Europe: moving from ‘feature-based’ to ‘whole-site’ management of sites. In: J Humphreys, RWE Clark, S Fletcher. (eds.), *Marine Protected Areas: Science, Policy and Management*. Amsterdam: Elsevier Ltd, 2020, 157–82.

Stevens TF, Sheehan EV, Gall SC *et al*. Monitoring benthic biodiversity restoration in Lyme Bay marine protected area: design, sampling and analysis. *Mar Policy* 2014;45:310–7. https://doi.org/10.1016/j.marpol.2013.09.006.

Stewart BD, Howarth LM, Wood H *et al*. Marine conservation begins at home: how a local community and protection of a small Bay sent waves of change around the UK and beyond. *Front Mar Sci* 2020;7:1–14. https://doi.org/10.3389/fmars.2020.00076.

Sumaila UR, Tai TC. End overfishing and increase the resilience of the ocean to climate change. *Front Mar Sci* 2020;7:1–8. https://doi.org/10.3389/fmars.2020.00523.

Sussex IFCA. Nearshore Trawling Byelaw 2019. Sussex: Sussex Inshore Fisheries and Conservation Authority, 2019, 10.

Turnbull JW, Johnston EL, Clark GF. Evaluating the social and ecological effectiveness of partially protected marine areas. *Conserv Biol* 2021;35:921–32. https://doi.org/10.1111/cobi.13677.

Underwood A. On beyond BACI: sampling designs that might reliably detect environmental disturbances. *Ecol Appl* 1994;4:3–15. https://doi.org/10.2307/1942110.

UNEP. Emissions Gap Report 2022: The Closing Window—Climate Crisis Calls for Rapid Transformation of Societies. Nairobi: UN Environment Programme, 2022a, 16–7.

UNEP. Kunming-Montreal global biodiversity framework. In: *Conference of the Parties to the Convention on Biological Diversity*, CBD/COP/15/L.25. Canada: UN Environment Programme, 2022b, 14.

Wells S, Ray GC, Gjerde KM *et al*. Building the future of MPAs—lessons from history. *Aquat Conserv Mar Freshw Ecosyst* 2016;26:101–25. https://doi.org/10.1002/aqc.2680.

Wolf J, Woolf D, Bricheno L. Impacts of climate change on storms and waves relevant to the coastal and marine environment around the UK. *MCCIP Sci Rev* 2020;2020:132–57.

Worm B. How to heal an ocean. *Nature* 2017;543:630–1. https://doi.org/10.1038/nature21895.

Zupan M, Fragkopoulou E, Claudet J *et al*. Marine partially protected areas: drivers of ecological effectiveness. *Front Ecol Environ* 2018;16:381–7. https://doi.org/10.1002/fee.1934.

## Appendix 1: Quote from Lyme Bay static MPA fisherman (S. Fanshawe, personal communication)

‘Before the scheme, I would cut up many white fish I caught for bait or would have to drive two hours to market as they would not keep. Being able to get fish on ice the minute it’s caught and stored in the chiller unit until it can be picked up has increased the amount I can earn and taken time off my working day.’ Reserve Fisherman, October 2016

## Appendix 2: Book quote from *Rewilding the Sea: How to save our Oceans*, authored by Charles Clover (Blue Marine Foundation Executive Director)

‘The project, at its best, brought significantly greater agreement between conservationists and fishermen than exists elsewhere and put the fishermen in the driving seat in deciding conservation methods. They are the experts on what works with the grain of fishing. The goodwill involved, unusual between fishermen and conservationists, brought about a significant recovery that would probably not have happened otherwise. There are many things still to tackle but Lyme Bay is an example of getting things right, most of the time, provided that a protected area which encompasses fishing is fished in a small-scale and sustainable way.’ Charles Clover, in *Rewilding the Sea: How to save our Oceans*. 2022

Handling editor: Linwood Pendleton