Research

INTRODUCTION

Species that have significant importance to the structure and function of an ecological community are classed as foundation species (Dayton 1972). Ecological foundation species often create complex habitat for many associated species (Stachowicz 2001, Angelini et al. 2011, Beheshti et al. 2022, Smith et al. 2023) and can have direct and indirect effects on biodiversity and ecosystem processes (Ellison 2019). As such, the loss of these species can have cascading effects throughout the system (Ellison et al. 2005, Valiente-Banuet et al. 2015, Sorte et al. 2017, Detmer et al. 2021). Similarly, human communities can be heavily connected to a single species; as a result, population collapse of that species can have direct and indirect social, cultural, ecological, and/or economic effects that ripple throughout the community (Marshall et al. 2007, McCay et al. 2011, Steneck et al. 2011, Ritzman et al. 2018). Species that are important to the cultural stability and identity of people have been identified as “cultural keystone species” or “cultural foundation species” (Cristancho and Vining 2004, Garibaldi and Turner 2004). Cristancho and Vining (2004) highlighted that species can have ecological, economic, and cultural centrality to communities. In this paper we build on these literatures, exploring the impacts of oyster loss within a coastal Florida community and highlighting the extent of economic, social, and cultural interactions with the species. We propose that its individual and collective importance to the community demonstrates its role as a “multidimensional foundation species” with ecological, economic, social, and cultural relevance.

Connection of human communities to multidimensional foundation species can occur through harvest of the foundation species and other taxa supported by the ecosystem structuring, and more indirectly through dependence on the non-provisioning ecosystem services the species provides (Ellison et al. 2005, Springer et al. 2010, McClanahan et al. 2015, Ritzman et al. 2018, Scyphers et al. 2019, Detmer et al. 2021). Because benefits from such foundation species are multi-faceted and can be difficult to quantify, the full extent of the connections to these species can often be under-estimated. Exploring the individual and collective dependence of community members on a foundation species, and the multi-dimensional impacts of species loss, can reveal the full importance of the species and contextualize community needs for sustainable harvest and resource management.

Oysters are known for their importance in both ecological and social realms; they are important ecological foundation species in estuaries around the world, providing a wide range of ecosystem services (Coen et al. 2007, Grabowski et al. 2012), including water filtration and shoreline stabilization (Thomas et al. 2022) and providing feeding and nursery habitat for many juvenile and adult estuarine species (Grabowski et al. 2022). The diverse ecosystems supported by the complex oyster reef structures draw fishers targeting a wide range of commercially and recreationally valuable species and generate opportunities for ecotourism (Thomas et al. 2022).

Oysters are a significant fishery species in many locations, including the eastern and southern coasts of the United States (Kirby 2004). Within these regions people have been harvesting oysters for thousands of years, evidenced by the shell middens that are still present throughout landscapes today (Reeder-Myers et al. 2022). Post-colonization oyster harvest began near population centers as early as the 1600s and spread along the eastern coast of the United States (Kirby 2004). Increased demand led to increasing harvest, which, coupled with other stressors such as pollution, caused significant oyster population declines throughout the U.S. and shifted harvest effort from the east coast further south into the Gulf of Mexico through the 19th and 20th centuries (Kirby 2004). The development of the oyster fishery in the Gulf established a significant economic resource for many coastal communities, and this long-term fishery developed economic, social, and cultural importance.

In Apalachicola Bay, Florida, harvest of the Eastern oyster (Crassostrea virginica) started in the mid-1800s, and the fishery boomed well into the 20th century. Oyster populations declined in the early 21st century culminating in a fishery collapse (Pine et al. 2015) and subsequent closure. This provides an opportunity to study the importance of oysters to the historically reliant community across social, cultural, and ecological dimensions, as impacts of the decline were felt by oyster harvesters, many seafood industry professionals, and local community members. We use semi-structured interviews to describe the impacts of the oyster population collapse on the collective community and on individuals who were directly or indirectly reliant on oysters and characterize the multidimensional foundation that oysters have provided within the Apalachicola Bay social-ecological system. More broadly, we demonstrate the need to consider social and cultural consequences of foundation species loss in future management and restoration plans.

METHODS

The Apalachicola Bay case study

Apalachicola Bay, located on the Gulf Coast of the Florida panhandle, is a 540 km² estuary known as the “Oyster Capital of the World.” The Bay once provided 90% of all oysters harvested in Florida and 10% of oysters harvested nationally (Pine et al. 2015). The commercial fishery began in the mid-1800s and flourished for over 100 years (Pine et al. 2015). At one time, the fishery supported about 1000 license holders and 40,000 fishing trips per year, with canning and processing houses lining the waterfront throughout the Bay (Havens et al. 2013). However, after decades of intense harvest, environmental stressors, and other human-related impacts, the fishery declined steeply in 2012, leading to a declaration of a Federal Fisheries Disaster in 2013 (Camp et al. 2015, Pine et al. 2015). The exact causes of this decline remain uncertain, and several potential influences have been cited, including lack of freshwater, overharvesting, and impacts from decisions related to the Gulf Horizon oil spill (Camp et al. 2015). Despite increasingly strict catch limits in the years following the declaration, the oyster population continued to decline until the state issued a five-year closure for the Bay in December 2020 (Florida Administrative Code § 68B-27.017 2021).

Apalachicola Bay is located in Franklin County, which has a population of just under 12,500, many of whom rely on seafood harvest directly or indirectly through harvest, sales, and tourism opportunities (Office of Economic and Demographic Research 2024). Much of this reliance was centralized around the oyster fishery, as this was one of the most productive and profitable fisheries in the area (Pine et al. 2015). Because oysters are an ecological foundation species, providing structure and habitat for other species, even non-oyster fishers may have had an indirect reliance on oysters (Cranfield et al. 2001). Furthermore, oyster fishers required services and places to sell their harvest, tying other local business owners to the oyster population (Moore et al. 2020). Because of the longevity and historical success of the fishery, and heavy reliance on oysters within this community, Apalachicola Bay provides an excellent case study for understanding how the collapse of an important fishery causes rippling social effects (Russell et al. 2016, Holland and Leonard 2020, Drakopulos and Poe 2023).

Participant identification

To elucidate the importance of oysters across the extent of the community of Apalachicola Bay, interviews were conducted with a range of individuals (following a protocol approved by Florida State University - IRB STUDY00003719) between 2023 and 2024. Because we were interested in understanding the extent of economic, ecological, and cultural connections to oysters within the study region, we focused on identifying participants who currently, or had previously, lived or worked in Franklin County or on the Apalachicola Bay, and had ties to the seafood industry or oyster recovery efforts. The interviewer advertised the study to local community members at community festivals and outreach events, publicized the study in the local newspaper, spoke at a Franklin County Commission meeting, and utilized local connections through existing partnerships. Publicizing the study in the local newspaper yielded both local participants and those who had since moved out of Franklin County. Most interview connections were, however, made through snowball sampling, speaking with individuals recommended by previous interviewees (Goodman 1961). Some contacted individuals declined to participate, many citing frustrations over the fishery closure or disinterest in speaking on the subject with the researcher.

Interviews were conducted in person or over the phone depending on participant availability. We conducted 28 interviews, 27 with local community members who have at one time lived or worked in the Apalachicola Bay region, and one interview with a former oyster farmer who worked in Oyster Bay located nearby and sold oysters within the Apalachicola Bay region. Three of the participants do not currently live within the area because of a variety of factors, including moving to find a more stable or better paying job as the oyster population declined. Participants ranged in their length of time living within the region, some being from multi-generational fishing families, while others recently moved or retired to the area. Although many respondents held multiple jobs over the course of their lives, we were able to categorize them into connection groups as described below.

Interview group categorization

In order to illustrate the extent of impacts and connections within the community, we categorized respondents post hoc into four connection groups for our analysis based on each participant’s job and stated connection to oysters, with relatively even representation of each group (Fig. 1, Table 1). The oyster fishers group represented 25% of participants (n = 7), supply chain group 18% (n = 5), non-seafood industry and businesses group 29% (n = 8), and community members group 29% (n = 8). We then compared these groups to ascertain commonalities in experiences among participants, and record oyster’s importance across social and ecological realms.

Oyster populations within the region fluctuated for decades but began to decline significantly in the mid-2000s and early 2010s (Havens et al. 2013, Camp et al. 2015, Pine et al. 2015, Grizzle et al. 2019). We categorized interview participants’ connection groups according to the jobs they reported holding at the time of the federal fishery disaster declaration in 2013 (Table A1.1). Although some interviewees reported changing professions or shifting the focus of their jobs prior to the declaration, citing already declining population numbers, 2013 was a significant collective turning point in the fishery when continually restrictive catch limits began, and oyster numbers remained consistently low. We also noted the job held by the participant at the time of the fishery closure in 2020, which for all but 2 participants were also the jobs they held at the time of the interview in 2023 or 2024.

Lines of questioning, data collection and thematic analysis

Interview questions were open-ended, allowing for discussion and clarification. Questioning followed several themes to understand the participants’ background, experiences, and perceptions (Appendix 1). Interviews began by collecting basic demographic information about the participant and their connection to the Bay and wild oysters. Subsets of participants who identified as fishers were asked about their participation in the oyster fishery. Participants were then questioned about the impacts of the oyster population collapse, and the importance of oysters to the Apalachicola Bay from the perspective of the ecosystem and the human community. Questioning began with broader open-ended questions about oyster importance, and then focused on specific ecosystem service loss, such as “Are there generally more or less fish in the Bay since the collapse?” These questions aimed to explore specific impacts of collapse from ecological, economic, and social standpoints. Finally, participants were asked how the collapse affected them personally (Appendix 1).

All interviews were recorded, with one exception, and recordings were transcribed verbatim for coding and analysis. Common response themes were derived from both a priori and inductive approaches (Ryan and Bernard 2003). Responses on the impacts of oyster population collapse on participants were inductively coded as individual impacts (Table 2, Table A1.2). If a participant said that they had shifted jobs prior to the fisheries disaster declaration, but indicated the shift was due to declining populations, we coded these responses in the direct impact analysis as an economic impact. Responses about impacts that happened to the community or were broadly about impacts on others were defined as collective impacts (Table 3).Collective impact themes were derived from the Millennium Ecosystem Assessment, the IPBES assessments, and categories we created that provide social and cultural context for oyster loss but do not fall into predefined categories from other assessments (MEA 2005, IPBES 2019; Table A1.3). We coded directed questions on impacts for presence/absence in relation to the ecosystem services and assigned sub-themes within these broader categories (Machado et al. 2019). Further themes and subthemes, in addition to the a priori/inductive codes, were developed deductively during analysis through observation of repetitions of common ideas and differences in responses among participants (Ryan and Bernard 2003). Thematic responses were also analyzed by connection group to understand trends among different subsections of the interview respondents. Illustrative quotations from participant interviews were edited for length and clarity.

RESULTS

Individual reliance and impacts of oyster loss

Participants reported personal experiences ranging from no impact to many impacts, and mentioned cultural, ecological, economic, and nutritional impacts (Fig. 2, Table 2). Across all participants, economic impacts were reported most often, and among the connection groups, the distribution of responses varied considerably. The Oyster Fisher group (OF group) individuals most frequently spoke of economic impacts, noting they either retired or found one or more jobs to support them after the fishery closed. The Oyster Supply Chain group (OSC group) individuals solely reported economic impacts because of the fishery collapse. The Non-Oyster Seafood Industry (NOSI group) individuals were the most diverse in their responses, with respondents citing impacts across all four categories, while two individuals did not feel there was any impact on them from the collapse. Within NOSI group responses, ecological and economic impacts were the most common with ecological impacts only reported by NOSI group individuals. Finally, half of Community Member group (CM group) participants felt no impact of the collapse, while others noted the loss of accessible free food options, loss of potential supplemental income in retirement, and loss of time spent on the water because they no longer can harvest oysters.

Economic impacts were felt by a majority of the participants (57%, n = 16), and included economic losses and economic shifts, specifically direct loss of employment, loss of alternative or reliable income sources, and indirect impacts on employment such as changes in the customer base of businesses. Loss of employment was felt by OF group individuals, the oyster fishers who could no longer go out to harvest, resulting in changing jobs or entering early retirement at the time of disaster or time of closure (18%, n = 5). NOSI and CM group individuals mentioned the loss of an alternative income source, citing that oysters were often a reliable source of supplemental income that would help pay bills, cover income when other jobs were unavailable, or supplement their retirement income. Other impacts on employment were expressed by OSC and NOSI group individuals (14%, n = 4). Two seafood dealers mentioned the direct impacts on employment of seafood house workers due to reduction in the availability of product for purchase, both oysters and other local species, resulting in fewer hours for the workers. One guide fisher mentioned how the reduction in reef-associated species and the inability to catch fish affected their clients’ perceptions of their abilities as fishers, which was classified as an indirect impact on employment because this negative perception could affect client retention. Finally, a local business owner mentioned another indirect impact on employment where they adapted their business as their clientele shifted from local seafood workers to tourists:

We went from 300 people [customers] in the morning by 9:00 to, when it stayed open at the end, we would see six or eight [customers]. To basically now no [customers], well I shouldn’t say no, because we are still selling gloves and cull irons ... to other places. We still sell some gloves to people shucking who are bringing oysters in. It’s hard to give a perspective because tourism was growing naturally anyway. Oystering was sort of doing this. [Up and down hand motion.] But basically, in the same time frame I would probably say that it directly affected at least 35 to 40% of my business in a total flip. Respondent 11, NOSI group.

Many of the experiences and reactions described by participants as economic impacts or changes from the collapse could collectively be grouped under an umbrella of economic diversification, encompassing taking on additional jobs, changing jobs within or outside of their field, and changing business models. The ability to adapt to fisheries changes, and how fishers respond, can be influenced by a variety of characteristics or conditions, and responses can occur at different levels within the overall system (Cinner et al. 2018, Barnes et al. 2020, Ojea et al. 2020, Green et al. 2021). Within OF group individuals this meant they either found other jobs, or in two cases multiple jobs, outside oyster fishing, or entered early retirement. OSC group respondents diversified within their field, either changing where they source their product, their clientele, or their business models. As one business owner put it, “we had to flip the script” (Respondent 10, OSC group). Adaptation of NOSI group respondents included guide fishers who moved offshore or changed where they fished to meet client expectations.

The loss of oysters as a food source was mentioned by eight of the interviewees (29%), across three of the four connection groups. This was one of the main impacts noted by CM group respondents, often while expressing love and enjoyment for eating oysters and frustration in no longer being able to readily harvest and consume them. Respondents who had generational ties, or had grown up fishing in the area, also expressed negative sentiments in relation to this loss, including frustration or disbelief in their inability to harvest their own food like they had done previously.

It’s a depression to know that somebody controls when you can and when you can’t go get something to eat, that - what if I go out here and I’m eating - get caught eating some oysters or get caught taking some oysters off of a pylon and I’m set up a fine, and then my name is plastered all over the paper like I’m some bad criminal or something. Respondent 16, NOSI group.

We just can’t get over the fact that we’ve got to go buy oysters to eat when we’ve done it all our lives. Respondent 22, OF group.

Oyster reefs create crucial habitat and provide food sources for many commercially and recreationally fished species (Coen et al. 1999). With the degradation of oyster reefs and thus habitat for these fished species, there were ecological changes within the bay that also affected respondents. Guide fishing captains noted changes in location and availability of target species, reporting either catching fish in new locations, or only catching fish where they know there are oysters. Additionally, five respondents, all within the NOSI group, noted that there was a reduction in reef-associated species, such as redfish or tripletail, which impacted their livelihoods. A common theme within these responses was the change in the certainty of catching fish; there was a shift from guaranteed and easy catch to greater uncertainty and effort needed to catch target species.

We were catching big redfish like that almost guaranteed; you go out there and I can catch them like that [*snaps*] all the time. That was in one trip. That was probably in 45 minutes ... As many as we wanted, all, everybody would be lined up. We haven’t done that in years. Respondent 3, NOSI group.

Three former oyster fishers discussed how the fishery closure impacted their sense of identity through the loss of their way of life, or the freedom of being their own boss, which we categorized as a cultural impact. These impacts were exclusively described by individuals who were previously oyster fishers, however they fell into different connection groups based on their jobs at the time of the fishery disaster declaration. These respondents described being an oyster fisher in relation to their roots and their connection to the area. One participant moved out of state prior to the disaster declaration, citing the inability to make a living within the seafood industry, but mentioned that, “my life-long dream is to do what I’m doing, retire and come back home, make my living in the bay, to supplement my retirement, or do whatever I wanted to, but there’s nothing there. I’ve got some family, but there is nothing there” (Respondent 19, CM group). This highlights that the loss of the oyster industry not only affected people’s present day lives, but also the futures they imagined for themselves.

Collective impacts of loss on community cultural and social identity

Evaluating responses related to community and collective impacts to ecosystem services, changes in cultural services were most frequently mentioned (100% respondents, n = 28), followed by supporting (68%, n = 19), regulating (46%, n = 13), and provisioning (43%, n = 12) services. Additional responses on the collective economic impacts of the fishery collapse were made by 79% of interviewees (n = 22).

Within non-cultural services, loss of habitat for reef-associated species was the most frequently mentioned impact (64%, n = 18), followed by changes in water quality (36%, n = 10) and changes in food availability derived from oyster reefs (32%, n = 9; Table A1.4). Within cultural services, impacts of oyster loss fell into 11 sub-themes based on participant responses (Fig. 3).

The most common impact to cultural ecosystem services mentioned was changes in community identity (54%), related to two key shifts. First, there was the physical change within the Bay municipalities along the coast, which was tied to sense of place, cultural heritage values and aesthetic values. Respondents spoke of the visual change of the landscape as the oyster fishery deteriorated through time. They mentioned the closure of oyster houses along the coastal highway and the lack of boats out on the water. Both long-time and recent residents mentioned the enjoyment they derived from seeing working boats out on the water, and noted the change in how the Bay looks now.

Down this road here there used to be gas stations, there used to be stores, and now there ain’t but one gas station. They just shut down. On this beach from here to Leavin’s [seafood processing house] there used to be 17 oyster houses. Respondent 15, OSC group.

The second shift in collective community identity was more intangible. The Apalachicola Bay region has been a working waterfront for generations, and the identity of the area was tied to the oyster through its title of “Oyster Capital of the World.” As one guide fisher put it, “the oyster was the heartbeat of Apalachicola” (Respondent 8, NOSI group). However, with the oyster declines, and subsequent closure of both the fishery and the related seafood industry businesses in the area, respondents indicated that this identity is changing along with the landscape. Some respondents were specifically concerned about the shifting community identity as it relates to tourism. As indicated by several participants, tourism seems to be filling the gap as the main industry in the area, replacing the seafood industry. Increasing numbers of guide boats, rental houses, and tourism-based businesses have changed the landscape of the Apalachicola region from a working waterfront to a tourist destination. A local business owner expressed concerns about this shift:

Even now because of the culture and atmosphere, we’re still depending on the oyster industry. Because if not, I’m afraid we’re going to see a change to where it’s not ... I’m really, tremendously dependent on the oysters for money but also for keeping the right tourism coming. People like what we are and who we are, and when you start losing the natural checkpoints, like a healthy seafood industry, it puts more demand on tourism. And our fear is that it will become somewhere like Destin, where you have too much population. Respondent 11, NOSI group.

We hypothesized that because of the branding of the region as the Oyster Capital, a decline in the species might lead to a decrease in tourism. Contrary to these expectations, only three participants indicated that they thought there was less tourism in the area than there would be if the oyster fishery were open. By contrast, 17 participants reported that tourism has grown immensely in recent years, including since the closure. This was not attributed to the fishery collapse, rather respondents suggested that the region had been recently “discovered.” A common refrain among the participants was “The Forgotten Coast ain’t forgotten anymore” (Respondent 27, CM group).

Lack of a negative impact on regional tourism was attributed to participants’ belief that visitors to the region are unaware that the fishery is closed, or that the oysters they are eating are not wild caught from the area. Four respondents mentioned changes in the type of tourist that comes to the region. Historically tourists would come for the seafood and the oysters, now tourism is driven by visiting the beach and the opportunities for recreational fishing: “People don’t come down here, bring family, because of oysters, they come for that beach. That beach and that salt” (Respondent 28, OF group). Additionally, the COVID-19 pandemic seemed to play a role in the popularity of the area, and the “discovery” of the Forgotten Coast. “[Tourism in] our county has boomed and part of that was COVID related. We are basically the perfect scenario of what people wanted. We’re low density, but open air and recreation outside” (Respondent 11, NOSI group).

Decline of social well-being within the community was the second most common theme. “The social decline and the amount of drugs and arrests, it has killed Franklin County. And it all stems back to the people that used to be - there wasn’t nothing but hard-working individuals. We didn’t have them problems, Franklin County didn’t have them problems. Now, there’s nothing for them to do” (Respondent 19, CM group). Participants cited increases in poverty, food insecurity, drug use tied to loss of employment from the oyster collapse, and lack of additional opportunities or training as contributors to overall decline in social well-being within the region. Additionally, respondents mentioned that they previously raised money for community members in need through the harvest and sale of oysters and other reef-associated seafood with local benefit events. These responses indicate that the existence of the oyster and seafood industries helped alleviate poverty and mitigate hardships.

Two cultural services were tied for the third most mentioned responses: brand and individual identity. Participants expressed pride in the Apalachicola oyster brand and highlighted the recognition of it globally. “The Apalachicola oyster is king, as far as taste and salinity ... We had the number one product, Apalachicola oysters was the pinnacle, it was the best” (Respondent 10, OSC group). Although all respondents who spoke about the branding of Apalachicola oysters were in agreement about its recognition and prestige, they were divided on whether it would, and does, continue to hold its recognition with the closure, with one participant saying: “You now have taken the best product off of the market for five years. We’ll never probably get that back. Even if oysters come back tomorrow. You probably will never get back to where you was, as far as the most desired and sought after product. Well, that was our brand. So, with that now you’ve lost your brand, you’ve lost your future of oysters” (Respondent 11, NOSI group). Whereas others believe, “Oh I guarantee you right now every one of these oyster bars that we have here, half shell bars? They’d go right back to Apalachicola oysters, that’s what everybody wants. That’s what everybody comes here and wants an Apalachicola oyster” (Respondent 27, CM group). However, restaurants in the area are still serving oysters on the half shell, harvested from other locations or sourced locally from aquaculture farms. The sale of farmed oysters, grown in the area and sold with the Apalachicola name, which have replaced wild caught oysters in restaurants and seafood markets, may help retain this branding during the closure.

Finally, a little more than a third of respondents mentioned the importance of oysters to the individual identity of people, particularly oyster fishers, describing the loss of oysters as a loss of “the way of life.” These terms and relationship to identity were cited across connection groups. Oyster fishing and the seafood industry supported the identities of community members throughout the region. One participant said, “If you were an oyster harvester, you weren’t in it for the money. It was the way of life that you had ... You wasn’t gonna get rich, but you would have a good life. And that life’s been taken away now” (Respondent 4, OF group). These identities were tied to positive sentiments, with participants using terms of love, enjoyment, and satisfaction. Additionally, this identity was tied to heritage, with terms such as “roots” and “in their blood” used to describe how interconnected oyster fishers were with their profession.

DISCUSSION

This study highlights the widespread and diverse impacts caused by the loss of an ecological foundation species, revealing how species can also act as multidimensional foundation species. Oysters within the Apalachicola Bay were intensely harvested until recently, and are still a significant part of community identity and the community’s collective memory (Cristancho and Vining 2004, Garibaldi and Turner 2004). We interviewed local community members with differing levels of connection to the historical oyster fishery to understand the importance that oysters play for the social system in Apalachicola Bay. We highlight that a variety of community members derived ecological, economic, and cultural benefits from oysters. We showcase the extent of the social, cultural, and economic connections to oysters, and illustrate the extent of impact from oyster population declines, elucidating direct impacts on oyster fishers, and direct and indirect effects on individuals throughout the community.

Respondents provided insight into the direct impacts of the oyster loss to them personally, including several individuals who were involved in the oyster industry and left their positions prior to the 2013 disaster declaration. Multiple reasons were given for this shift out of oystering, both related and unrelated to oyster population declines. By utilizing information from local experiences, we found that changes in oyster populations, or oyster harvest regulations, may have resulted in changes within the economic and social landscapes much earlier than the 2010s fishery collapse. Within ecological systems, loss of foundation species at different frequencies, severities, or time scales can impact how the system reacts in response (Orwig et al. 2013, Castorani et al. 2018). Our findings suggest a similar dynamic within the social system, including cultural and economic shifts within the community. However, in considering future studies on multi-dimensional foundation species, we recommend incorporating questions about the lifelong experiences of interview participants, and how fisheries played a role in their social and cultural experiences through time to better contextualize the impacts of species loss on the maintenance of culture (Coe and Gaoue 2020).

Interviews with different community members also provided insight into the breadth of impacts. Not every community member derives the same benefits from oysters, but our results show that there are some commonalities in the experiences of similarly connected individuals. For example, only NOSI group individuals highlighted ecological impacts relevant to them because of the fishery collapse. Participants in this group such as local guide fishing captains noted they were seeing declines in the size and availability of reef-associated species, resulting in many guides shifting fishing effort off-shore or to areas they know retain oysters. These perspectives suggest ecological changes that are affecting not just fishers, but also those reliant on the ecosystem created by the foundation species.

The aim of this work was to document the extent of species loss impacts within social, cultural, economic, and ecological dimensions, and to record the direct impacts to a range of community members who vary in their degree and types of connections to the species. We found that community members experience multiple impacts from species loss based on different connections they hold to oysters. Future research on the drivers of individual impact responses could provide better understanding of the components of a person’s identity, such as multigenerational involvement or time lived in the area, that influence their connection to the foundation species.

The interviewees represent only a small portion of the population of the region, thus limiting the viewpoints within the study. Several individuals were uninterested in speaking with the authors, citing frustration over the fishery closure and how it was communicated and handled. Although we had a diversity of respondents with differing opinions on the causes of the decline and the necessity of the fishery closure itself, we recognize that a greater sample size would have provided even more insight into the impacts of the oyster decline. Of note, we were unable to connect with commercial fishermen of oyster reef-reliant species, such as crabs and shrimp, to further understand how the decline may have impacted their catch and livelihoods directly. Greater representation from oyster farmers (n = 1, Table A1.1) could have provided further insight into any potential increase in demand for aquaculture, as farmed oysters from the region have filled some of the market gap from reduction in wild harvest.

Despite the limited number of respondents, we were able to begin to assess service losses that fell into the cultural ecosystem services category. Such services are frequently understudied because of their “intangible” nature (Milcu et al. 2013, Cabana et al. 2020). However, in recent years there has been a concerted effort to include factors of human well-being in studies of climate stress, fisheries participation, and ecosystem management (Breslow et al. 2016, Holland et al. 2020, Moore et al. 2024). Community identity tied to fishing and working waterfronts is common and may persist even in the face of fisheries collapse (Jacob et al. 2001, van Ginkel 2001, Brookfield et al. 2005). Furthermore, place character may develop and be shaped by fishing within regions, as well as the associated physical aesthetic built around and from fishing (Urquhart and Acott 2014). Studying these indicators of identity and well-being in the face of fisheries change can help identify adaptation needs and recovery opportunities (Seara et al. 2022).

We found a consensus that “the Forgotten Coast is no longer forgotten,” and tourists have discovered the region as a vacation destination. With this influx of tourism, it seems that the consumption of Apalachicola oysters is not as important as was once presumed. Globally, consumer knowledge on seafood sourcing and traceability is limited, and many people may presume they are eating local seafood without checking (Zisser et al. 2012, Ryburn et al. 2022, Hirokawa and Thompson 2023). This seems to be the case within Apalachicola, particularly among tourists who have recently discovered the area, as participants speculated that they may be unaware of the fishery closure. This region is not alone in its transition from a fishing focus to tourism (Ounanian 2016, Fabinyi et al. 2022). This transition to a tourism-centered economy can bring opportunities for livelihood diversification and high value markets (Fabinyi et al. 2022), but can also bring a changing sense of identity, increased property prices, and greater fishing intensity (Ounanian 2016, Fabinyi 2020, Kimbu et al. 2022, Miller 2022). As such, coastal policy makers focused on economic development should consider community identity and the importance of historical economies within these transitioning areas.

Finally, our study contributes to a broader literature on cultural keystone species. Cristancho and Vining (2004:162) highlighted that the applicability of the concept “largely depends on what definition of culture one takes,” which opens the door to applying these concepts to a wide array of settings. Here we built off both ecological foundation species and cultural keystone species theories. The use of our multidimensional approach may be beneficial in a number of other settings. In particular, we show the value of categorizing respondents into groups based on their varying connections with the species. Although our grouping of participants was specific to the setting, broadening the groupings may help future studies apply similar concepts in a reproducible and comparable way.

Implications for management

Our results could be useful for informing management, restoration, and coastal resilience plans for Apalachicola. With the closure of oyster harvest within the Apalachicola Bay, stakeholders have felt both an economic and cultural loss. In approaching future restoration and management endeavors, decision makers should consider the numerous benefits the community derives from oysters. Providing job training or employing community members in restoration efforts could help fishers directly impacted by loss of employment. Additionally, individuals who historically relied on oysters for a secondary or alternative income source could benefit from similar opportunities, such as access to small quota commercial harvest licenses. Nutritionally, community members currently lack access to oysters, a previously free source of fresh, healthy protein. Subsistence harvest is important for community well-being and resilience, and should be considered in management and development plans (Poe et al. 2015). For example, maintaining regulated recreational harvest on intertidal reefs, through bag limits or area closure, both of which have been used previously for commercial and recreational fishing, would allow the community to continue limited subsistence harvesting without impacting sub-tidal reef structures (https://myfwc.com/). Furthermore, the utilization of oyster reefs by other commercial and guide fishers is key for both the seafood and tourism industries within the area. Restoration of the Apalachicola Bay oyster reefs is ongoing, and the Florida Fish and Wildlife Conservation Commission is currently considering management scenarios for when the fishery closure ends, currently set for January 2026 (Apalachicola Bay System Initiative Community Advisory Board 2024). Based on the results of this study we recommend that managers account for the multidimensional importance of these habitats and consider management recommendations that address social-ecological perspectives to prioritize holistic ecosystem recovery. Finally, recognition of the cultural significance and the heritage tied to oysters and their harvest within the Apalachicola Bay should be considered when prioritizing management, local development plans, and ecosystem recovery.

These lessons can be applied more broadly to areas that are heavily reliant on harvesting a single species, particularly foundation species. Promoting opportunities for job diversification and/or multi-species harvest could help mitigate the economic impacts of species loss (Smith and McKelvey 1986, Kasperski and Holland 2013). Subsidized and supported job training can help communities facing fisheries declines (Harper and Sumaila 2019). Recognition by decision makers of the importance of seafood as a nutrition source is also crucial in supporting coastal community well-being. Remote areas that are reliant on fishing for subsistence may have limited low-cost alternative options for protein sources, and seafood subsistence can help promote food security (Loring et al. 2013, Viana et al. 2023). Understanding the reliance on seafood within a system and providing management decisions that allow for continued subsistence harvest, or affordable protein alternatives, can assist in lessening the impacts of fisheries loss. Finally, when approaching species restoration and recovery, recognizing the social and cultural importance of species, and allowing continued opportunities for fisheries harvest can help maintain the individual and community identities of an area. Balancing goals for ecosystem recovery and sustainable harvest is an important consideration when planning restoration efforts and determining management plans. However, recovering all of the many cultural, social, and economic aspects that are lost within a community when a foundation species collapses is challenging, and thus ideally proactive and adaptive management will help prevent collapse form occurring.

Building on the foundation species and cultural keystone species concepts (Dayton 1972, Cristancho and Vining 2004, Garibaldi and Turner 2004), we argue that oysters are a multidimensional foundation species within the Apalachicola Bay region. This study shows that oysters are not only important species ecologically, but their significance within the area has created a social, cultural, and economic foundation within the region. As showcased in this study, individual and collective benefits are derived from the oyster populations of Apalachicola Bay. As such, understanding the comprehensive impacts of the loss of species that serve this multidimensional foundation species role is crucial for the resilience of ecosystems and human communities.

LITERATURE CITED

Angelini, C., A. H. Altieri, B. R. Silliman, and M. D. Bertness. 2011. Interactions among foundation species and their consequences for community organization, biodiversity, and conservation. BioScience 61:782-789. https://doi.org/10.1525/bio.2011.61.10.8

Apalachicola Bay System Initiative Community Advisory Board. 2024. Apalachicola Bay System Ecosystem-Based Adaptive Restoration and Management Plan (Special Report). Florida State University Coastal & Marine Laboratory, Triumph Gulf Coast Project #69, St Teresa, Florida, USA.

Barnes, M. L., P. Wang, J. E. Cinner, N. A. J. Graham, A. M. Guerrero, L. Jasny, J. Lau, S. R. Sutcliffe, and J. Zamborain-Mason. 2020. Social determinants of adaptive and transformative responses to climate change. Nature Climate Change 10:823-828. https://doi.org/10.1038/s41558-020-0871-4

Beheshti, K. M., S. L. Williams, K. E. Boyer, C. Endris, A. Clemons, T. Grimes, K. Wasson, and B. B. Hughes. 2022. Rapid enhancement of multiple ecosystem services following the restoration of a coastal foundation species. Ecological Applications 32:e02466. https://doi.org/10.1002/eap.2466

Breslow, S. J., B. Sojka, R. Barnea, X. Basurto, C. Carothers, S. Charnley, S. Coulthard, N. Dolšak, J. Donatuto, C. García-Quijano, C. C. Hicks, A. Levine, M. B. Mascia, K. Norman, M. Poe, T. Satterfield, K. St. Martin, and P. S. Levin. 2016. Conceptualizing and operationalizing human wellbeing for ecosystem assessment and management. Environmental Science & Policy 66:250-259. https://doi.org/10.1016/j.envsci.2016.06.023

Brookfield, K., T. Gray, and J. Hatchard. 2005. The concept of fisheries-dependent communities: a comparative analysis of four UK case studies: Shetland, Peterhead, North Shields and Lowestoft. Fisheries Research 72:55-69. https://doi.org/10.1016/j.fishres.2004.10.010

Cabana, D., F. Ryfield, T. P. Crowe, and J. Brannigan. 2020. Evaluating and communicating cultural ecosystem services. Ecosystem Services 42:101085. https://doi.org/10.1016/j.ecoser.2020.101085

Camp, E. V., W. E. Pine III, K. Havens, A. S. Kane, C. J. Walters, T. Irani, A. B. Lindsey, and J. G. Morris, Jr. 2015. Collapse of a historic oyster fishery: diagnosing causes and identifying paths toward increased resilience. Ecology and Society 20(3):45. https://doi.org/10.5751/ES-07821-200345

Castorani, M. C. N., D. C. Reed, and R. J. Miller. 2018. Loss of foundation species: disturbance frequency outweighs severity in structuring kelp forest communities. Ecology 99:2442-2454. https://doi.org/10.1002/ecy.2485

Cinner, J. E., W. N. Adger, E. H. Allison, M. L. Barnes, K. Brown, P. J. Cohen, S. Gelcich, C. C. Hicks, T. P. Hughes, J. Lau, N. A. Marshall, and T. H. Morrison. 2018. Building adaptive capacity to climate change in tropical coastal communities. Nature Climate Change 8:117-123. https://doi.org/10.1038/s41558-017-0065-x

Coe, M. A., and O. G. Gaoue. 2020. Cultural keystone species revisited: are we asking the right questions? Journal of Ethnobiology and Ethnomedicine 16:70. https://doi.org/10.1186/s13002-020-00422-z

Coen, L. D., R. D. Brumbaugh, D. Bushek, R. Grizzle, M. W. Luckenbach, M. H. Posey, S. P. Powers, and S. G. Tolley. 2007. Ecosystem services related to oyster restoration. Marine Ecology Progress Series 341:303-307. https://doi.org/10.3354/meps341303

Coen, L. D., M. W. Luckenbach, and D. L. Breitburg. 1999. The role of oyster reefs as essential fish habitat: a review of current knowledge and some new perspectives. American Fisheries Society Symposium 22:438-454.

Cranfield, H. J., G. Carbines, K. P. Michael, A. Dunn, D. R. Stotter, and D. J. Smith. 2001. Promising signs of regeneration of blue cod and oyster habitat changed by dredging in Foveaux Strait, southern New Zealand. New Zealand Journal of Marine and Freshwater Research 35:897-908. https://doi.org/10.1080/00288330.2001.9517052

Cristancho, S., and J. Vining. 2004. Culturally defined keystone species. Human Ecology Review 11:153-164.

Dayton, P. K. 1972. Toward an understanding of community resilience and the potential effects of enrichments to the benthos at McMurdo Sound, Antarctica. In B. C. Parker, editor. Proceedings of the Colloquium on Conservation Problems in Antarctica. Allen Press, Lawrence, Kansas, USA.

Detmer, A. R., R. J. Miller, D. C Reed, T. W. Bell, A. C. Stier, and H. V. Moeller. 2021. Variation in disturbance to a foundation species structures the dynamics of a benthic reef community. Ecology 102:e03304. https://doi.org/10.1002/ecy.3304

Drakopulos, L., and M. Poe. 2023. Facing change: individual and institutional adaptation pathways in West Coast fishing communities. Marine Policy 147:105363. https://doi.org/10.1016/j.marpol.2022.105363

Ellison, A. M. 2019. Foundation species, non-trophic interactions, and the value of being common. iScience 13:254-268. https://doi.org/10.1016/j.isci.2019.02.020

Ellison, A. M., M. S. Bank, B. D. Clinton, E. A. Colburn, K. Elliott, C. R. Ford, D. R. Foster, B. D. Kloeppel, J. D. Knoepp, G. M. Lovett, J. Mohan, D. A. Orwig, N. L. Rodenhouse, W. V. Sobczak, K. A. Stinson, J. K. Stone, C. M. Swan, J. Thompson, B. Von Holle, and J. R. Webster. 2005. Loss of foundation species: consequences for the structure and dynamics of forested ecosystems. Frontiers in Ecology and the Environment 3:479-486. https://doi.org/10.1890/1540-9295(2005)003[0479:LOFSCF]2.0.CO;2

Fabinyi, M. 2020. The role of land tenure in livelihood transitions from fishing to tourism. Maritime Studies 19:29-39. https://doi.org/10.1007/s40152-019-00145-2

Fabinyi, M., B. Belton, W. H. Dressler, M. Knudsen, D. S. Adhuri, A. A. Aziz, Md. A. Akber, J. Kittitornkool, C. Kongkaew, M. Marschke, M. Pido, N. Stacey, D. J. Steenbergen, and P. Vandergeest. 2022. Coastal transitions: small-scale fisheries, livelihoods, and maritime zone developments in Southeast Asia. Journal of Rural Studies 91:184-194. https://doi.org/10.1016/j.jrurstud.2022.02.006

Garibaldi, A., and N. Turner. 2004. Cultural keystone species: implications for ecological conservation and restoration. Ecology and Society 9(3):1. https://doi.org/10.5751/ES-00669-090301

Goodman, L. A. 1961. Snowball sampling. Annals of Mathematical Statistics 32(1):148-170. https://doi.org/10.1214/aoms/1177705148

Grabowski, J. H., C. J. Baillie, A. Baukus, R. Carlyle, F. J. Fodrie, R. K. Gittman, A. R. Hughes, D. L. Kimbro, J. Lee, H. S. Lenihan, S. P. Powers, and K. Sullivan. 2022. Fish and invertebrate use of restored vs. natural oyster reefs in a shallow temperate latitude estuary. Ecosphere 13(5):e4035. https://doi.org/10.1002/ecs2.4035

Grabowski, J. H., R. D. Brumbaugh, R. F. Conrad, A. G. Keeler, J. J. Opaluch, C. H. Peterson, M. F. Piehler, S. P. Powers, and A. R. Smyth. 2012. Economic valuation of ecosystem services provided by oyster reefs. BioScience 62:900-909. https://doi.org/10.1525/bio.2012.62.10.10

Green, K. M., J. C. Selgrath, T. H. Frawley, W. K. Oestreich, E. J. Mansfield, J. Urteaga, S. S. Swanson, F. N. Santana, S. J. Green, J. Naggea, and L. B. Crowder. 2021. How adaptive capacity shapes the adapt, react, cope response to climate impacts: insights from small-scale fisheries. Climatic Change 164:15. https://doi.org/10.1007/s10584-021-02965-w

Grizzle, R., M. Parker, C. Snyder, M. Lamb, E. E. Dontis, and K. R. Radabaugh. 2019. Apalachicola Bay. In Oyster integrated mapping and monitoring program report for the State of Florida. Fish and Wildlife Research Institute, Florida Fish and Wildlife Conservation Commission, St. Petersburg, Florida, USA.

Harper, S., and U. R. Sumaila. 2019. Distributional impacts of fisheries subsidies and their reform: case studies of Senegal and Vietnam. IIED Working Paper. International Institute for Environment and Development, London, UK.

Havens, K., M. Allen, E. Camp, T. Irani, A. M. Lindsey, J. Glenn, A. Kane, D. Kimbro, S. Otwell, and B. Pine. 2013. Apalachicola Bay oyster situation report. Florida Sea Grant, Gainesville, Florida, USA.

Hirokawa, T., and B. S. Thompson. 2023. The influence of new sustainable fisheries policies on seafood company practices and consumer awareness in Japan. Marine Policy 157:105819. https://doi.org/10.1016/j.marpol.2023.105819

Holland, D. S., J. K. Abbott, and K. E. Norman. 2020. Fishing to live or living to fish: job satisfaction and identity of West Coast fishermen. Ambio 49(2):628–639. https://doi.org/10.1007/s13280-019-01206-w

Holland, D. S., and J. Leonard. 2020. Is a delay a disaster? Economic impacts of the delay of the California dungeness crab fishery due to a harmful algal bloom. Harmful Algae 98:101904. https://doi.org/10.1016/j.hal.2020.101904

Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES). 2019. Global assessment report on biodiversity and ecosystem services of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services. E. S. Brondizio, J. Settele, S. Díaz, and H. T. Ngo, editors. IPBES secretariat, Bonn, Germany. https://doi.org/10.5281/zenodo.3831673

Jacob, S., F. Farmer, M. Jepson, and C. Adams. 2001. Landing a definition of fishing dependent communities: potential social science contributions to meeting National Standard 8. Fisheries 26:16-22. https://doi.org/10.1577/1548-8446(2001)026%3C0016:LADOFD%3E2.0.CO;2

Kasperski, S., and D. S. Holland. 2013. Income diversification and risk for fishermen. Proceedings of the National Academy of Sciences 110:2076-2081. https://doi.org/10.1073/pnas.1212278110

Kimbu, A. N., I. Booyens, and A. Winchenbach. 2022. Livelihood diversification through tourism: identity, well-being, and potential in rural coastal communities. Tourism Review International 26:25-40. https://doi.org/10.3727/154427221X16245632411854

Kirby, M. X. 2004. Fishing down the coast: historical expansion and collapse of oyster fisheries along continental margins. Proceedings of the National Academy of Sciences 101:13096-13099. https://doi.org/10.1073/pnas.0405150101

Loring, P. A., S. C. Gerlach, and H. L. Harrison. 2013. Seafood as local food: food security and locally caught seafood on Alaska’s Kenai Peninsula. Journal of Agriculture, Food Systems, and Community Development 3:13-30. https://doi.org/10.5304/jafscd.2013.033.006

Machado, A. M. da S., F. G. Daura-Jorge, D. F. Herbst, P. C. Simões-Lopes, S. N. Ingram, P. V. de Castilho, and N. Peroni. 2019. Artisanal fishers’ perceptions of the ecosystem services derived from a dolphin-human cooperative fishing interaction in southern Brazil. Ocean & Coastal Management 173:148-156. https://doi.org/10.1016/j.ocecoaman.2019.03.003

Marshall, N. A., D. M. Fenton, P. A. Marshall, and S. G. Sutton. 2007. How resource dependency can influence social resilience within a primary resource industry. Rural Sociology 72:359-390. https://doi.org/10.1526/003601107781799254

McCay, B. J., W. Weisman, and C. Creed. 2011. Coping with environmental change: systemic responses and the roles of property and community in three fisheries. Pages 381-400 in R. E. Ommer, R. I. Perry, K. Cochrane, and P. Cury, editors. World fisheries: a social-ecological analysis. Wiley Blackwell, Oxford, UK. https://doi.org/10.1002/9781444392241.ch23

McClanahan, T., E. H. Allison, and J. E. Cinner. 2015. Managing fisheries for human and food security. Fish and Fisheries 16:78-103. https://doi.org/10.1111/faf.12045

Milcu, A. I., J. Hanspach, D. Abson, and J. Fischer. 2013. Cultural ecosystem services: a literature review and prospects for future research . Ecology and Society 18(3):44. https://doi.org/10.5751/ES-05790-180344

Millennium Ecosystem Assessment. 2005. Ecosystems and human well-being: biodiversity synthesis. World Resources Institute, Washington, D.C., USA.

Miller, K. M. 2022. Disentangling tourism impacts on small-scale fishing pressure. Marine Policy 137:104960. https://doi.org/10.1016/j.marpol.2022.104960

Moore, S. K., M. Broadwater, C. Cha, Q. Dortch, C. J. Harvey, K. C. Norman, J. Pearce, C. Pomeroy, and J. F. Samhouri. 2024. Exploring the human dimensions of harmful algal blooms through a well-being framework to increase resilience in a changing world. PLoS Climate 3:e0000411. https://doi.org/10.1371/journal.pclm.0000411

Moore, S. K., S. J. Dreyer, J. A. Ekstrom, K. Moore, K. Norman, T. Klinger, E. H. Allison, and S. L. Jardine. 2020. Harmful algal blooms and coastal communities: socioeconomic impacts and actions taken to cope with the 2015 U.S. West Coast domoic acid event. Harmful Algae 96:101799. https://doi.org/10.1016/j.hal.2020.101799

Office of Economic and Demographic Research. 2024. Franklin County - County Profiles. Florida Legislature, Tallahassee, Florida, USA.

Ojea, E., S. E. Lester, and D. Salgueiro-Otero. 2020. Adaptation of fishing communities to climate-driven shifts in target species. One Earth 2:544-556. https://doi.org/10.1016/j.oneear.2020.05.012

Orwig, D. A., A. A. Barker Plotkin, E. A. Davidson, H. Lux, K. E. Savage, and A. M. Ellison. 2013. Foundation species loss affects vegetation structure more than ecosystem function in a northeastern USA forest. PeerJ 1:e41. https://doi.org/10.7717/peerj.41

Ounanian, K. 2016. In place of fishing: coastal communities in transition. Dissertation. University of Rhode Island, Kingston, Rhode Island, USA.

Pine III, W. E., C. J. Walters, E. V. Camp, R. Bouchillon, R. Ahrens, L. Sturmer, and M. E. Berrigan. 2015. The curious case of eastern oyster Crassostrea virginica stock status in Apalachicola Bay, Florida. Ecology and Society 20(3):46. https://doi.org/10.5751/ES-07827-200346

Poe, M. R., P. S. Levin, N. Tolimieri, and K. Norman. 2015. Subsistence fishing in a 21st century capitalist society: from commodity to gift. Ecological Economics 116:241-250. https://doi.org/10.1016/j.ecolecon.2015.05.003

Reeder-Myers, L., T. J. Braje, C. A. Hofman, E. A. Elliott Smith, C. J. Garland, M. Grone, C. S. Hadden, M. Hatch, T. Hunt, A. Kelley, M. J. LeFebvre, M. Lockman, I. McKechnie, I. J. McNiven, B. Newsom, T. Pluckhahn, G. Sanchez, M. Schwadron, K. Y. Smith, T. Smith, A. Spiess, G. Tayac, V. D. Thompson, T. Vollman, E. M. Weitzel, and T. C. Rick. 2022. Indigenous oyster fisheries persisted for millennia and should inform future management. Nature Communications 13:2383. https://doi.org/10.1038/s41467-022-29818-z

Ritzman, J., A. Brodbeck, S. Brostrom, S. McGrew, S. Dreyer, T. Klinger, and S. K. Moore. 2018. Economic and sociocultural impacts of fisheries closures in two fishing-dependent communities following the massive 2015 U.S. West Coast harmful algal bloom. Harmful Algae 80:35-45. https://doi.org/10.1016/j.hal.2018.09.002

Russell, S. M., A. Arias-Arthur, K. Sparks, and A. Varney. 2016. West Coast communities and catch shares: the early years of social change. Coastal Management 44:441-451. https://doi.org/10.1080/08920753.2016.1208864

Ryan, G. W., and H. R. Bernard. 2003. Techniques to identify themes. Field Methods 15:85-109. https://doi.org/10.1177/1525822X02239569

Ryburn, S., W. Ballantine, F. Loncan, O. Manning, M. Alston, B. Steinwand, and J. Bruno. 2022. Public awareness of seafood mislabeling. PeerJ 10:e13486. https://doi.org/10.7717/peerj.13486

Seara, T., A. Owens, R. Pollnac, R. Pomeroy, and C. Dyer. 2022. Lessons learned from a natural resource disaster: the long-term impacts of the Long Island Sound lobster die-off on individuals and communities. Marine Policy 136:104943. https://doi.org/10.1016/j.marpol.2021.104943

Scyphers, S. B., J. S. Picou, and J. H. Grabowski. 2019. Chronic social disruption following a systemic fishery failure. Proceedings of the National Academy of Sciences 116:22912-22914. https://doi.org/10.1073/pnas.1913914116

Smith, C., and R. McKelvey. 1986. Specialist and generalist: roles for coping with variability. North American Journal of Fisheries Management 6:88-99. https://doi.org/10.1577/1548-8659(1986)6%3C88:SAG%3E2.0.CO;2

Smith, R. S., S. L. Cheng, and M. C. N. Castorani. 2023. Meta-analysis of ecosystem services associated with oyster restoration. Conservation Biology 37:e13966. https://doi.org/10.1111/cobi.13966

Sorte, C. J. B., V. E. Davidson, M. C. Franklin, K. M. Benes, M. M. Doellman, R. J. Etter, R. E. Hannigan, J. Lubchenco, and B. A. Menge. 2017. Long-term declines in an intertidal foundation species parallel shifts in community composition. Global Change Biology 23:341-352. https://doi.org/10.1111/gcb.13425

Springer, Y. P., C. G. Hays, M. H. Carr, and M. R. Mackey. 2010. Toward ecosystem-based management of marine macroalgae - the Bull Kelp, Nereocystis Luetkeana. Oceanography and Marine Biology 48:1-41.

Stachowicz, J. J. 2001. Mutualism, facilitation, and the structure of ecological communities: positive interactions play a critical, but underappreciated, role in ecological communities by reducing physical or biotic stresses in existing habitats and by creating new habitats on which many species depend. BioScience 51:235-246. https://doi.org/10.1641/0006-3568(2001)051[0235:MFATSO]2.0.CO;2

Steneck, R. S., T. P. Hughes, J. E. Cinner, W. N. Adger, S. N. Arnold, F. Berkes, S. A. Boudreau, K. Brown, C. Folke, L. Gunderson, P. Olsson, M. Scheffer, E. Stephenson, B. Walker, J. Wilson, and B. Worm. 2011. Creation of a gilded trap by the high economic value of the Maine lobster fishery. Conservation Biology 5:904-912. https://doi.org/10.1111/j.1523-1739.2011.01717.x

Thomas, S., K. Collins, C. Hauton, and A. Jensen. 2022. A review of the ecosystem services provided by the native oyster (Ostrea edulis): implications for restoration. IOP Conference Series: Materials Science and Engineering 1245:012010. https://doi.org/10.1088/1757-899X/1245/1/012010

Urquhart, J., T. Acott. 2014. A sense of place in cultural ecosystem services: the case of Cornish fishing communities. Society & Natural Resources 27:3-19. https://doi.org/10.1080/08941920.2013.820811

Valiente-Banuet, A., M. A. Aizen, J. M. Alcántara, J. Arroyo, A. Cocucci, M. Galetti, M. B. García, D. García, J. M. Gómez, P. Jordano, R. Medel, L. Navarro, J. R. Obeso, R. Oviedo, N. Ramírez, P. J. Rey, A. Traveset, M. Verdú, and R. Zamora. 2015. Beyond species loss: the extinction of ecological interactions in a changing world. Functional Ecology 29:299-307. https://doi.org/10.1111/1365-2435.12356

van Ginkel, R. 2001. Inshore fishermen: cultural dimensions of a maritime occupation. Pages 177-193 in D. Symes and J. Phillipson, editors. Inshore fisheries management . Springer, Dordrecht, The Netherlands. https://doi.org/10.1007/978-94-017-1892-9_10

Viana, D. F., J. Zamborain-Mason, S. D. Gaines, J. Schmidhuber, and C. D. Golden. 2023. Nutrient supply from marine small-scale fisheries. Scientific Reports 13:11357. https://doi.org/10.1038/s41598-023-37338-z

Zisser, B., D. Glaser, and I. Seggerman. 2012. Do you know where your seafood comes from? How seafood traceability stacks up against beef and produce. Oceana, Washington, D.C., USA.