Introduction
Marine ecosystems are at risk from the effects of climate change on marine species physiology, population and species diversity, and ecological interactions (
Hoegh-Guldberg and Bruno 2010;
Doney et al. 2012). As ectotherms, fish and invertebrate species are especially vulnerable to ocean warming, as their body temperature is largely determined by the surrounding environment (
Sunday et al. 2011;
Pinsky et al. 2019). Species ranges, an outcome of a species’ potential and realized habitat niche, are driven by environmental conditions and moderated by biological interactions such as competition, predation, and long-term interactions among species (such as mutualism, commensalism, parasitism, and others) (
Ackerly et al. 2010). The distribution of many species and populations has already changed as species move in space, such as poleward or to deeper waters (marine species), higher altitudes (terrestrial species), and/or in time as the seasonality of species lifecycles shifts to earlier or later times of the year (
Parmesan and Yohe 2003;
Tingley and Beissinger 2009;
Brown et al. 2015). Marine species range shifts are expected to continue under projected warming and other changes in ocean conditions, with consequences for ecosystems, economies, societies, and management (
Cheung et al. 2015;
Patrizzi and Dobrovolski 2018).
Climate change impacts on species ranges can change where marine protected areas (MPAs) should be situated (
McLeod et al. 2009;
Gerber et al. 2014) and can also disrupt connectivity between protected areas by changing dispersal pathways and species physiology (
Álvarez‐Romero et al. 2017) and affecting adult movement (
Friesen et al. 2021). Well-established conservation planning tools have been applied in response to climate change predictions, such as emphasizing MPA networks, increasing spatial connectivity, habitat heterogeneity, and improving management of the core and edges of reserves (
Hannah et al. 2002). Designating new MPAs could augment the existing global network of MPAs and provide potential benefits of connectivity and redundancy for existing species ranges (
Hannah 2008;
Araújo 2009). However, adding more MPAs today might not provide future benefits to the specific species or habitats they were intended to protect because of climate change.
To date, others have proposed a range of methods to incorporate climate change into conservation planning (see reviews by
Magris et al. 2014;
Jones et al. 2016). Previous research has aimed to identify thermal refugia, or areas that may warm less rapidly and thus offer some protection from increasing temperatures (
Ban et al. 2016;
Lima et al. 2016). However, data limitations—especially in terms of understanding how protecting future habitat might increase species adaptive capacity to climate change—make these methods challenging and uncertain (
Groves et al. 2012;
Magris et al. 2014). Others make a case for “conserving the geophysical stage”, whereby conversation plans are defined by geophysical indicators such as topography as surrogates for biodiversity features (
Groves et al. 2012), an appropriate approach for some but not all species. Others still promote incorporating ecological processes into systematic conservation planning, such as river flows, flood patterns, or animal migration patterns (
McCook et al. 2009;
Groves et al. 2012;
D'Aloia et al. 2017). Typically, conservation planning methods and data have remained temporally static based on the current state of biodiversity; with climate change, much more adaptive and proactive adaptation strategies are necessary (
Groves et al. 2012). Another technique is to ensure the protection of habitat distributions over time (temporal connectivity), which would allow species to track their climatic niche as habitats change with climate change (
Hodgson et al. 2009). Ecological niche theory—the environmental conditions that an organism is dependent upon to survive and reproduce (
Wiens et al. 2009)—can be applied to models to describe how species may respond to future environmental change by identifying habitats that are likely to be used in the future. These forecasts are called species distribution models (SDMs) or bioclimatic niche models (e.g.,
Cheung et al. 2015). These SDMs can then be applied to a spatial decision support tool such as Marxan or zonation to prioritize actions to protect those future habitat needs and species of interest (
Magris et al. 2014; e.g.,
Alagador et al. 2014).
In this paper, we explored two ways that marine conservation planning could incorporate projected changes in species distributions using global climate projections available globally. We used the outputs of an existing dynamic bioclimate envelope model of shifting species distributions (
Weatherdon et al. 2016b) to (1) determine where species ranges overlap within MPAs in the Northern Shelf Bioregion (see below) in the present and future (2060) and (2) use them as inputs into spatial prioritization software (Marxan) to identify priorities for MPAs to represent biodiversity now and into the future across the entire coast of BC (within the Canadian Pacific Exclusive Economic Zone (EEZ) and including the transboundary region of southeastern Alaska and Washington States). We also tracked challenges encountered and reflected on the usefulness of the results for MPA network planning.
We focused on two scales: (1) the Northern Shelf Bioregion, and (2) all of Canada's Pacific EEZ and northern neighbouring regions in Alaska and northern Washington State. The smaller focal region, the Northern Shelf Bioregion, is relevant because this is the part of Canada's Pacific EEZ where a network of MPAs is currently actively being pursued jointly by Federal, Provincial, and First Nations government representatives (
Gale et al. 2019) (
Fig. 1). The bioregion, approximately 100,000 square kilometres in size, is one of 13 ecologically defined bioregions in Canada's EEZ (
Government of Canada 2011). It is the only bioregion (out of 4) in Pacific Canada that has an active MPA network planning process underway and one that involves multiple governments (
McGee et al. 2022;
Reid et al. 2022). There are 118 conservancies, ecological reserves, and parks with a marine component that are under the jurisdiction of the Provincial BC government through BC Parks (hereafter referred to as BC MPAs) and 6 Federal MPAs within the bioregion included in this analysis. We focused on Provincial MPAs as part of this project, which was funded by BC Parks. We worked closely with members of the technical team planning the Northern Shelf Bioregion MPA network, including sharing our approaches, results, and associated data. Understanding how well existing MPAs in the region might fare under climate change is important for the development of the planning process, which includes a climate change sub-committee. The broader focus, on Canada's Pacific EEZ and beyond, is important for understanding how future range shifts might affect Canada's ability to protect marine species in its Pacific EEZ.
Discussion
Considering climate change impacts is important for marine conservation planning, as climate changes will affect species’ ranges, larval dispersal, and population connectivity, and thus compromise the performance of MPAs and MPA networks (
Álvarez‐Romero et al. 2017). MPAs and networks have largely lacked consideration for climate change (
Bruno et al. 2018;
Sala and Giakoumi 2018). Rapidly accelerating climate impacts make the Pacific region especially vulnerable to current and future impacts of global change (
Okey et al. 2014;
Asch et al. 2018). We used two approaches to incorporate existing climate change projections for species of cultural and economic importance within a coastal region. In the first of such analyses in this region, we showed that BC MPAs are likely to protect fewer species by 2060 than they do currently, as species ranges are projected to shift poleward and “leave” current MPAs. Similarly, spatial priority areas as selected by Marxan shifted north as species ranges shifted north, and more spatial priority areas were required for species to reach conservation targets. Knowing this, strategically adding new MPAs farther north as species ranges shift north would help to support conservation goals and objectives, although our analysis did not consider new species moving into the region or species interactions. It is important to note that species abundance is also projected to decline during this time period (
Weatherdon et al. 2016b), and as such, it could become harder for the same number of MPAs to protect the same abundance of species. Weighting MPA zones that are projected to protect the highest number of species from analyses such as these could help to meet conservation objectives. More broadly, strategically working with fisheries management and other adaptation policies, and across jurisdictional borders, could help to improve the conservation effectiveness of MPAs considering climate-change related changes in species-specific thermal niches.
Our efforts to use global data to analyze and inform MPA planning and management in BC highlighted two key challenges. First, there are many uncertainties embedded in our analyses and associated data. By necessity, bioclimatic models incorporate a great deal of complexity and species data, but this adds uncertainty to the predictions of future species distributions (
Ackerly et al. 2010). The projected species ranges model outputs that we used as inputs into both analyses have high uncertainties around emissions, projected changes in ocean conditions, and also around species responses to climate change. The projections also did not consider other socioeconomic or human drivers such as fishing and habitat changes that may affect species’ abundance (
Heikkinen et al. 2006;
Wiens et al. 2009;
Cheung et al. 2016;
IPCC 2018). More specifically, in the Marxan analyses, we used modelled relative abundance data available through SDMs (
Weatherdon et al. 2016b). Applying SDM abundance data to systematic conservation planning has several assumptions and potential associated errors (
Tulloch et al. 2016). These include the assumption that occurrence data that are used as inputs into SDMs reflect the preferred habitat conditions of the species that we explicitly included in our SDMs; some abiotic (e.g., fishing) and/or biotic (e.g., food availability and predation) factors not represented in the models may be driving the past and projected future distributions. As well, these projections assume that species detectability is constant across species, whereas there are known irregularities in species-specific data availability for model inputs (
Weatherdon et al. 2016b).
Future research should take species-specific responses to changing habitat conditions beyond the thermal niche into account, including species interactions and motility (
Montoya and Raffaelli 2010;
Parks et al. 2023). As well, there is uncertainty in real-world future emissions that will affect biological outcomes and adaptation strategies that could be developed at local, regional, or national scales to mitigate these impacts (
Gattuso et al. 2015;
IPCC 2018). Using climate velocity, or the localized speed and direction of climate contours as they are predicted to shift, as a proxy for how species’ distributions will shift to track those thermal niches, could be a simpler method of incorporating climate effects on species movement than modelling species ranges (
Loarie et al. 2009;
García Molinos et al. 2015), although these data are not yet available for this region. Climate velocities can also be combined with species traits where those data are available to build more robust predictions of species range shifts (
Sunday et al. 2015). In addition, future research could incorporate the effects of MPAs on species population dynamics and how conservation targets could be met through spatial planning (e.g., increasing species’ abundance). For example, MPAs could buffer or offset the impacts of climate change on declining species abundance (
Fox et al. 2012).
The second challenge was the coarse resolution of the outputs of the bioclimate modelling used, and hence also our Marxan analysis. The global data we used for the analysis were at a relatively coarse resolution relative to the area of BC MPAs and current MPA network planning efforts in the region. Such a mismatch in spatial resolution increases the projection uncertainties at the grid cell level that are used by the existing management and planning efforts. While our results could not directly inform the design of BC MPA networks, they might be suitable for other broader planning efforts in the region. Downscaled projections are being developed and refined for Canada's Pacific waters, but at present these are unable to resolve the complex nearshore waters and thus exclude most waters where BC MPAs are located (
Masson and Fine 2012;
Holdsworth et al. 2021). In the future, downscaled climate projections will be available to model species range shifts and/or climate velocities relevant for the scale of MPA planning, but this will come too late for the development of the current MPA network. Once available, these should be considered in the adaptive management of the MPA network. Species-specific analyses will also be helpful to understand whether they are being protected by MPAs in a changing climate.
Despite these challenges, the conclusions from our results are still useful for planners and managers within BC to understand the conservation potential of coastal MPA networks now and in the future and to advocate for increased protection measures that can support the resilience of marine ecosystems in an increasingly uncertain climate. In particular, our results highlight that climate change confronts the assumptions of conventional spatial approaches to conservation (
Lawler et al. 2015). To maintain 30% of the baseline (2016) relative abundance of species within BC waters, large portions of the NE Pacific coast may require some type of spatial protection. Given the uncertainty inherent in projecting species range shifts and the effects of ocean warming on the dispersal capacity of many marine species, larger MPAs or a well-connected network of MPAs reduce the reliance on accurate predictions of biodiversity now and in the future and will also be necessary to maintain population recruitment and recolonization after discrete disturbances (
Álvarez‐Romero et al. 2017). However, given the rapid pace of change and the numerous biological outcomes that are not accounted for in these range shift projections (e.g., changing species interactions), static area-based conservation efforts are unlikely to be sufficient on their own to support species persistence in the long term (
Tittensor et al. 2019), and managers will also have social implications to consider. For example, while large MPAs may have high ecological value (
Edgar et al. 2014), smaller MPAs may support human wellbeing, which is important in the coastal context (
Ban et al. 2019). Unconventional approaches, such as temporally dynamic MPAs (
Alagador et al. 2014) and assisted migration (
Swan et al. 2015), might be worth considering in addition to static MPAs (
D'Aloia et al. 2019). Overall, adaptation strategies and unconventional conservation approaches will be necessary as part of a portfolio response to support resilience to climate change (
Millar et al. 2007;
Galatowitsch et al. 2009).
Maintaining biological diversity for resilience to environmental change is a key policy issue for local and national governance in this era of accelerating climate change. Given our finding that it will be difficult to maintain species abundances within MPAs in the future, a key challenge is to understand and develop management approaches across jurisdictions (Canadian and American; Federal, Provincial, and First Nations in Canada's Pacific waters) that can support adaptation and response to global environmental change. While conservation planning cannot prevent the impacts of climate change, the general effects can be predicted and integrated into conservation planning decisions. Depending on the adaptive capacity of species and populations, based on thermal tolerance estimates, some species will be adaptable to changing climates over time, while others will disperse rapidly, if possible, in response to changing habitats and environmental conditions (
Sunday et al. 2012). Managing for resilience through precautionary conservation planning would suggest that incorporating what we do know about climate change impacts, rather than focusing on what we do not, is a more appropriate management choice.