Introduction

Coastal wetlands, considered as ecological ecotones between terrestrial and ocean environments, play an indispensable role in the global ecosystems (Kirwan and Megonigal 2013). While they only account for 5–8% of the global land surface (Mitsch et al. 2013), coastal wetlands are hotspots for productivity, performing critical functions including nutrient and biogeochemical cycling and providing important ecosystem services such as water quality improvement, biodiversity, and carbon sequestration (Baustian et al. 2012). Globally, tidal wetlands accumulate 53.65 gigatons of carbon (C) a year, serving as an important carbon sink (i.e., “blue” Carbon) (Wang et al. 2021) and thus considerably contributing to the global C budget (Howard et al. 2017). Many of the coastal ecosystem processes and functions are regulated or mediated by their constitutive biota such as plants and microbial communities in the soil (Emerson et al. 1999; Weiss et al. 2004; Yarwood 2018). Due to a greater focus on anerobic processes in the generally anoxic wetland soils, bacteria are often considered the primary mediators of many coastal ecosystem processes, with fungi historically being relegated to having minor importance relative to bacteria (Khan 2004). Fungal communities are thus overlooked in coastal ecosystems and studies on their diversity, function and distribution lags that of bacterial studies.

There is a growing recognition, however, that fungi can also be critical players in coastal ecosystem processes such as decomposition, largely due to their tight associations or symbiotic relationships with plant communities (Onufrak et al. 2020; Bahram and Netherway 2022; Wang et al. 2022). For example, arbuscular mycorrhizal fungi (AMF) have largely been shown to be ubiquitous and critical in wetland ecosystems (Wolfe et al. 2007; Xu et al. 2016). Other types of fungi such as endophytes (fungi that live inside plant tissues) and fungi in the rhizosphere soil zone (i.e., soil that is in contact with plant roots) are beginning to be explored as well, unmasking rich and diverse communities whose ecology and functions are critical for wetland plants and to coastal wetland ecosystems (Farrer et al. 2022). However, there is still a lack of systematic understanding of patterns related to their distribution and diversity globally and the different factors shaping these processes in coastal wetlands in space and time. Basic understanding of these dynamics is necessary in order to fully assess the magnitude of their roles in coastal wetland ecosystem functioning.

The diversity and distribution of fungi in coastal wetlands are influenced by interplay of both biotic and abiotic factors depending on the substrate, i.e., inside plants tissues or soil. For example, rhizosphere soil fungi and those living inside plant roots (e.g., mycorrhiza and endophytes) can display varying and sometimes opposite diversity patterns, likely due to different underlying mechanisms shaping their patterns (Lumibao et al. 2021). Host factors are likely to have stronger influence on endosphere fungi than on rhizosphere fungal communities, acting as selective filters for fungi colonizing their roots. However, these patterns might be contingent upon the intrinsic abiotic and biotic factors of their local environment and would thus differ when larger spatial scales, i.e., across continents, are considered. A recent review of arbuscular mycorrhizal fungi (AMF), for example, suggests that they have global distribution in coastal wetlands though they are limited by flooding, hypoxia, soil pH, salinity, and the host plant’s identity or genotype (Wang et al. 2022). Across coastal wetland habitats, pronounced variations in local environmental conditions (e.g., anoxic saltmarsh soils vs. oxygenated mangrove swamp soils) can also drive differences in fungal communities (Alzarhani et al. 2019).

In this review, we focus on belowground fungal communities that are in close associations with coastal wetland plants for a number of reasons. First, as coastal wetlands like marshes are often dominated by only a few plant species (e.g., Spartina alterniflora), functional shifts in plant-fungal associations of dominant vegetation can lead to considerable ecosystem-level outcomes. Second, plant-fungal interactions encompass a feedback mechanism. Plants influence fungal communities in the rhizosphere soil (i.e., soil in contact with their roots) and those living inside their roots (Lumibao et al. 2020). In turn, fungi can influence physiology and tolerance of the plants under an environmental stressor (Torres-Martínez et al. 2020; Lumibao et al. 2022) and help in nutrient acquisition (e.g., Schultz et al. 2001; Bowles et al. 2018; Moreau et al. 2019). The dynamics of these associations can potentially regulate coastal ecosystem processes. Third, the diversity and composition of rhizosphere soil fungi and endophytic fungal communities are subject to the competing and/or synergistic influences of both biotic, i.e., plants, and abiotic, i.e., local environment, that are likely to vary across spatial and temporal scales.

We address the patterns of fungal diversity associated with plants in naturally occurring wetland habitats across the globe. We synthesized literature that focused on rhizosphere soil and root endosphere fungal communities (including mycorrhizae) across four wetland habitat types: coastal dunes, intertidal flats, salt marshes, and tidal wetlands. Our aims were to (1) synthesize literature/data on rhizosphere and root endosphere fungi across different coastal wetland habitats, (2) determine the commonalities, if any, in fungal research across global coastal wetland habitats, and (3) identify research gaps in fungal studies in coastal wetlands.

Materials and Methods

We performed a systematic quantitative literature review by searching Scopus (Elsevier, Atlanta, U.S.A) and ISI Web of Knowledge (Core collection; Thomson Reuters, NY, U.S.A.). These databases were searched through article title, abstract, and keywords using the search string: (fung* OR microb*) AND (“saltmarsh” OR “salt marsh” OR “coastal ecosystem” OR “tidal wetland” OR “freshwater swamp” OR “tidal marsh”) AND (“rhizosphere” OR “rhizo*” OR “root” OR “endosphere” OR “endoph*”). Available literature until December 2021 was included. This search returned 382 and 145 papers in Scopus and ISI Web of Knowledge, respectively. All papers were imported into COVIDENCE (Covidence systematic review software, Veritas Health Innovation, Melbourne) for screening. Covidence removed automatically duplicates resulting in 527 papers that were included in title and abstract screening. Further screening of the 527 studies based on our criteria ultimately resulted in 51 studies that were used for the synthesis (Table S1, Online Resources 1).

We defined coastal ecosystems as types of habitats at the transition zone between terrestrial and aquatic or ocean environments. This encompasses coastal dunes, restinga forest, intertidal flats, tidal wetlands (including mangroves), forested wetlands (including freshwater wetlands), and salt marshes (Table 1). Where applicable further (i.e., more specific), categorization was carried out based on how it was defined in the papers used in this synthesis and/or Ramsar Convention (2016) as described in Tables 1 and S1 (Online Resources 1).

Table 1 Coastal wetland category description adopted from Ramsar Convention on Wetlands (2016), key representative vegetation type, the reported region they occur, and published studies used in this synthesis. The full list of host plant species/vegetation can be found in Online Resources 1
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Study Inclusion Criteria

Relevant studies were imported and screened using COVIDENCE. Selection was based on the following criteria to be included in the systematic review: study must be (1) conducted in a natural/pristine coastal ecosystem, i.e., no environmental disturbances reported; (2) conducted in field conditions without field manipulative experiments or lab experiments; and (3) focused on fungi inside roots (root endophytes or mycorrhizae) and/or rhizosphere soil. All studies not meeting the criteria were excluded in the final screening. Data extraction was conducted manually within COVIDENCE, which included information on author, publication year, geographic location of the study, habitat type, substrate, and host plant (Table S1, Online Resources 1). We did not make any distinction on whether the host plant species is native or foreign to the specific area or geographic location; thus, they are included as long as studies meet the criteria. Non-target ecosystem types that were captured during the keyword search but do not pertain to this study were removed (i.e., three seagrass studies). Mangrove studies were merged with the tidal wetland category as we did not include “mangroves” in key word searches.

Analyses

We conducted all analyses and created graphs in R (R Core Team 2020) and Microsoft Excel 365 (Version 2208, Microsoft Corporation) where applicable. To assess plant and fungal richness patterns, we counted the number of either phyla, genera, or species reported across all studies from each coastal wetland type. Functional guilds were assigned to taxa using FUNGuild (Nguyen et al. 2016). All assignments that were “highly probable” and “probable” were kept as recommended by Nguyen et al. (2016).

Results

The studies included in our review were published between 1961 and 2021 (Fig. 1a). The total mean number of studies published on root and rhizosphere communities in coastal ecosystems between 1961 and 2021 was 2 per year (Fig. 1a). Since 2019, there has been an increase in the number of studies published (6) suggesting an increasing interest in the fungal microbial communities in coastal ecosystems. Majority of studies were conducted in the USA (i.e., 21%, Fig. 1b), followed by Spain (12%), China, and India, both 8% (Fig. 1b).

Fig. 1
figure 1

a Global annual number of publications on root and rhizosphere fungal communities in coastal ecosystems. b Number of publications per country. c Number of studies per ecosystem type. Photos from left to right illustrate reported coastal ecosystems: coastal dune, tidal wetland, and saltmarsh. d Number of studies per substrate

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Fungal Communities in the Roots and Rhizosphere of Coastal Plant Communities

Our results showed that 73% of studies reported fungal communities from salt marshes, followed by 16% from tidal wetlands, 10% from coastal dunes, and other coastal ecosystems (Fig. 1c). Over half of the studies (53%) assessed fungal communities from roots, followed by rhizosphere soil (33% of studies) and 14% from both roots and rhizosphere soil (Fig. 1d). Culture and root staining methods were the most reported assessment techniques, i.e., 37% and 31%, respectively, of fungal communities in the roots and rhizosphere of coastal plant communities. High-throughput sequencing (HTS) methods were used in 8% of studies, but yielded the highest number of fungal species identified (Fig. S1, Online Resources 2). Other studies assessed fungal communities with a combination of methods, e.g., phospholipid fatty acid analysis (PLFA) and enzyme activity assessment or direct fungal spore counting (Fig. S1, Online Resources 2).

Across 51 studies we reviewed, 69 plant species’ fungal communities were reported to at least genus level. Of those, the ten most studied host species reflected saltmarsh and tidal wetland communities. They were Spartina alterniflora (12 studies), Phragmites australis (10 studies), Suaeda maritima (6 studies), Spartina patens (5 studies), Arthrocnemum macrostachyum (4 studies), Avicennia marina (4 studies), Carpobrotus edulis (3 studies), and Limonium tetragonum (3 studies). A few studies assessed fungal communities associated with succulent marsh plants (Salicornia and Batis).

Overall, salt marsh plant fungal communities have been reported the most, followed by tidal wetland and coastal dune plant communities (Fig. 2a). Some plant host species, e.g., P. australis and S. alterniflora, were reported from multiple coastal ecosystems, e.g., saltmarsh and tidal wetland, respectively (Appendix 1, Online Resources 3). Other plant species were more restricted, e.g., Calligonum polygonoides (Polygonaceae) was only reported from coastal dune, Tamarix chinensis (Tamaricaceae) from tidal wetland and Zygophylla coccineum (Zygophyllaceae) from intertidal flat. Fungal genera and species richness varied considerably between the ecosystems, with overall highest fungal species richness reported within the roots of saltmarsh plants, followed by coastal dune plants and tidal wetland plants (Fig. 2b, Online Resources 3). Root-associated fungal communities were not reported from intertidal flats (Fig. 2b). Fungal species richness in the rhizosphere was the highest around saltmarsh plants, followed by coastal dune and tidal wetland plants (Fig. 2b, Online Resources 3). At the phylum level, Ascomycota was described only from intertidal flats. Basidomycota was reported only from salt marshes while Glomeromycota was investigated in coastal dunes, salt marshes, and tidal wetlands. Mucoromycota was only reported from salt marshes. Fungal genera varied between the coastal ecosystems with 98 genera reported from salt marshes, 39 from coastal dunes, 21 from tidal wetlands, and 5 from intertidal flats. Most reported fungal genera from coastal dunes included Acaulospora, Glomus, and Funneliformis (Fig. 3). Most reported fungal genera from intertidal flats included Aspergillus, Penicillium, and Cladosporium (Fig. 3). In salt marshes, it was Glomus, Fusarium, and Aspergillus (Fig. 3). In tidal wetlands, Acaulospora, Rhizophagus, and Funneliformis (Fig. 3).

Fig. 2
figure 2

a Host plant and fungal species richness in the rhizosphere and plant roots from reviewed coastal ecosystems; b fungal species richness across the rhizosphere and the roots of coastal plants (Online Resources 3). Note that figure does not include plant and fungal richness reported in Alzarhani (2019)

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Fig. 3
figure 3

Twenty-five most abundant fungal genera across coastal dunes, intertidal flats, salt marshes, and tidal wetlands (full list found in Online Resources 3)

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In terms of functional guilds, nineteen guilds were reported across all studies reviewed here, with predominantly arbuscular mycorrhizal fungi (AMF) reported with an exception of intertidal flats and freshwater wetlands, where AMF were not reported (Fig. 4). Across all reported fungal taxa in both roots and rhizosphere soil, FUNGuild analyses revealed that 39% were AMF, 11% belong to the Plant Pathogen-Dung-Undefined-Wood Saprotroph complex, 6% plant pathogen, 2% animal pathogen, and 33% as cannot be assigned confidently to any known functional guild (Fig. 4). The rest of the assigned guilds were low in abundance and mostly occurred in salt marshes. Tidal wetlands contain primarily AMF with one identified saprotroph while intertidal flats only harbor different types of saprotrophs.

Fig. 4
figure 4

Frequency of reported fungal functional guilds across coastal dunes, intertidal flats, salt marshes, and tidal wetlands

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Discussion

It is increasingly recognized that fungi may play critical roles in coastal wetland processes (Bahram and Netherway 2022), driving soil multifunctionality (Li et al. 2022). However, fungi in these ecosystems remain underexplored; thus, a systematic understanding of their diversity and distribution is still elusive. Here, we focused on root-associated and rhizosphere soil fungal communities as they form close relationships with plants (e.g., mycorrhizal symbioses), and the dynamics of these associations can potentially influence ecosystem processes (e.g., carbon sequestration, provisioning of biodiversity).

Our synthesis provides insights into the current state of fungal research in coastal ecosystems globally, revealing an increasing number of publications over the past decade. Our analysis revealed that fungal research in coastal systems is geographically biased and skewed towards the USA with a large proportion of studies focusing on saltmarsh fungal communities. Outside the USA, the majority of studies were conducted in China, South Korea, and India. Notably, China contains Asia’s largest wetlands, accounting for about 10% of the global wetland area (Xiao et al. 2019). Tropical coastal wetlands in Southeast Asia were greatly underrepresented in this region, indicating the need for better representation of these regions globally in future coastal wetland fungal research. Moreover, our analysis revealed that coastal ecosystems have unique plant and fungal assemblages with salt marshes reported to have the highest richness of fungal taxa in their roots.

Status of Global Fungal Research in Coastal Wetlands

Coastal wetlands account for 5–8% of the global land surface (Mitsch et al. 2013; Gardner and Finlayson 2018) and are responsible for many climate-soil feedback processes including greenhouse gas fluxes, biodiversity, and carbon sequestration. Overall, the number of publications of fungal research in natural coastal wetlands showed marginal increase since 2019. Despite the wide availability of high-throughput sequencing technology over the past 10 years, which allows for in-depth and more comprehensive characterization of rhizosphere and endosphere fungi, plant root-associated and soil fungal research in coastal systems has been sporadic compared to higher number of studies focusing on bacterial or fungal diversity in terrestrial ecosystems (e.g., Tedersoo et al. 2022). Notably, our study only included studies reporting fungal communities from natural or pristine environments conducted up to 2021; hence, we did not review fungal research conducted in restored, managed, or disturbed wetlands over the past decades (e.g., Carrasco et al. 2006; Lumibao et al. 2018). For instance, there have been growing interests in the functional role of fungi in metal and oil-contaminated sites in coastal and marine environments due to their potential use for bioremediation (Zhao et al. 2022).

Fungal Diversity Across Coastal Wetlands

Coastal wetlands support diverse but unique assemblages of fungi according to wetland types. Despite salt marshes harboring naturally depauperate plant assemblages dominated by ecosystem engineers like Spartina alterniflora, they contain the highest number of root-associated and rhizosphere fungal taxa, with Glomus and Penicillium as the two most dominant genera. In general, wetland habitats with lower (reported) plant richness can still harbor a high number of root-associated and rhizosphere soil fungal taxa. For example, coastal dune habitat showed similar fungal richness (59 fungal species) despite lower plant richness (only three reported plant species) compared to tidal wetland (32 fungal species; 33 plant species) habitats. These patterns are in contrast with studies demonstrating strong correlation between high plant diversity and belowground fungal communities in terrestrial communities (e.g., Roy-Bolduc et al. 2016; Onufrak et al. 2020). Thus, our analysis revealed that low plant richness in coastal ecosystems does not translate into lower reported fungal richness in roots and/or rhizosphere.

Salt marshes also harbor diverse functional guilds—from mycorrhizae and endophytes to saprotrophs and pathogens. On the other hand, coastal dunes and tidal wetlands were almost completely composed of mycorrhizae while only saprotrophs (with a few undefined) were identified in intertidal flats. This reflects the potential influence of habitat type in promoting growth of certain fungal taxa that performs specific functions. For example, the high abundance of saprotrophs in the rhizosphere soils associated with a few plant species (e.g., Zygophyllum, Limoniastrum) in the intertidal flat as reported by El-Morsy (1999) might indicate that a large number of saprophytic fungi play a significant role in the organic matter decomposition in these habitats as found in other studies (e.g., Li et al. 2016; Lin et al. 2023). While intertidal flats are generally considered unproductive and poorly vegetated, they can still support certain plants as found in the reported study (El-Morsy 1999). Notably, as salt marshes were heavily studied in the literature, marsh fungal communities are, thus, better characterized compared to the other habitats, which explains the presence of diverse fungal guilds.

The unique assemblages—different fungal genera and species—found in each wetland habitat type could be due to different abiotic conditions, i.e., the environment supporting different plant species that in turn support unique fungal taxa. For instance, several Aspergillus species are known to thrive in the soils of semi-arid regions and Mediterranean coasts (Abdel-Azeem et al. 2020). Here, Aspergillus was reported only in salt marshes and intertidal flats (in Egypt), where it is the most common fungal genera in the latter. On the other hand, the genus Glomus was reported across all coastal wetland habitats globally except in intertidal flats and was most dominant in the salt marshes. Fungal species belonging to Glomus are exclusively mycorrhizal fungi, which are cosmopolitan and form symbiotic associations with different (host) plant species (Schwarzott et al. 2001). Thus, the ability of fungi to associate with plants might have contributed to the widespread distribution of Glomus (as well as the other mycorrhizal genera like Acaulospora) across coastal wetland habitats.

Known plant, insect, and animal pathogens such as those belonging to the genera Fusarium were reported from salt marshes (though other Fusarium species are capable of switching to necrotrophs, i.e., kill host then feed on dead host cells (Summerell 2019). Different species of Fusarium that are known plant pathogens (e.g., F. palustre) were reported in plant roots, mostly those in salt marshes across the globe (Elmer et al. 2016). F. palustre has been associated with the dieback of the saltmarsh grass, S. alterniflora, in its native North American salt marshes (Elmer 2014; Li et al. 2014).

Notably, the majority of the reported fungal species were mycorrhizal fungi (arbuscular mycorrhiza) as they were the focus of more than 40% of the studies included in our synthesis. Plant-mycorrhizal symbioses represent the oldest symbiotic relationships between plants and fungi, having co-evolved ca. 200–400 M years ago, with mycorrhiza colonizing about 85% of vascular plants (Brundrett 2002). Fungi supply nutrients to plants, and plants provide carbon substrate as food to fungi, and this symbiosis has been widely studied in terrestrial ecosystems (e.g., Kivlin et al. 2011; Soudzilovskaia et al. 2019). Based on our synthesis, mycorrhizae in natural coastal wetland plants were first reported only in 1990 (Cooke and Lefor 1990), though it became the main focus of fungal research in coastal wetlands in the following decades (e.g., D’Entremont et al. 2018; Kandalepas et al. 2010). In part, this might be driven by the potential of mycorrhiza in mediating ecosystem functions as observed in terrestrial systems. Arbuscular and ectomycorrhizal vegetation in terrestrial ecosystems store between 241 ± 15 and 100 ± 17 GT carbon, respectively, in aboveground biomass compared to non-mycorrhizal vegetation (29 ± 5.5 GT carbon) (Soudzilovskaia et al. 2019), suggesting that mycorrhizal fungi contribute significantly to carbon sequestration. High frequency of reported mycorrhizal fungi found in reviewed studies may also reflect the availability and relatively low cost of assessing mycorrhizal colonization in plant roots. Specifically, AMF characterization is a considerably less expensive approach requiring standard laboratory consumables and materials to assess root colonization—which involves root washing, staining, and counting presence of fungi under microscope—compared to more technically complex and expensive analyses such as high-throughput sequencing.

Recent studies reviewed in our synthesis have illustrated that mycorrhiza, particularly, AMF (belonging to genus Glomus), are globally ubiquitous across different coastal wetlands. For example, AMF have been reported from tidal wetlands in the US and Egypt (e.g., Bauer et al. 2003; Abd-Elgawad et al. 2020; others), mangrove-dominated habitats in India (e.g., Gaonkar and Rodrigues 2021), and from salt marshes in North America, Europe and Asia (Wang et al. 2022; Daleo et al. 2007; others). They appear to be associated with almost all the wetland plants studied, supporting previous AM findings from wetlands (e.g., Xu et al. 2016). This suggests that AMF plays an equally important functional role in coastal systems as in terrestrial systems via provisioning of critical ecosystem processes (e.g., nutrient acquisition, mediation of stress tolerance of plants), warranting further in-depth study.

Notably, our synthesis did not capture other coastal habitats such as seagrasses and mangroves. Recent studies suggest that seagrasses might also be an important reservoir of fungal diversity (Vohnik et al. 2015; Ettinger and Eisen 2019; Poli et al. 2022) as are mangrove-dominated habitats (e.g., de Souza Sebastianes et al. 2013). These findings highlight the increasing recognition of the importance of fungi in coastal ecosystems.

Our review includes studies that were published between 1961 and 2021. We acknowledge that since 2021, seminal papers have been published reviewing microbial community diversity, including fungi, in coastal ecosystems. For example, Crump and Bowen (2024) synthesized recent research on microbial habitats in estuaries. Farrer et al. (2022) reviewed the ecology of plant-microbial symbioses in coastal systems including mycorrhizae.

Key Knowledge Gaps and Future Research

While the fungal research in coastal ecosystems is slowly on the rise, there are still many research gaps that need to be addressed. Below, we outline three main research gaps resulting from our synthesis analyzing reported root-associated and rhizosphere fungal communities from coastal ecosystems.

First, increased representation of the different coastal wetland habitats, regionally and/or globally, is needed. More research is needed to better understand the diversity and functional importance of fungal communities associated with plants from restinga forests, intertidal flats, and forested wetlands. Second, the geographic bias in global fungal research can limit our inferences on the global diversity and distribution patterns of fungi in coastal wetlands. We found that the majority of studies reporting root or rhizosphere fungal communities were from the USA, Spain, and China, collectively accounting for 41% of reviewed studies. Increasing the reporting of fungal research from underrepresented countries is critical for global biodiversity assessments.

Third, we found that 68% of studies reviewed here analyzed root and rhizosphere fungal communities using exclusively culture-based or root-staining methods and only 8% of studies used sequencing methodologies. Fungal analysis using cultures is very valuable; however, only a small fraction of fungi is culturable, and thus, the true diversity of fungal communities using culture-dependent methods will be underestimated. For instance, only 10–30% of fungi are culturable by using traditional microbiological methods (Magnuson and Lasure 2002). High-throughput sequencing (HTS) offers high-quality, cutting-edge alternatives for analyzing microbiome structure and functioning from complex environmental DNA samples (Tedersoo et al. 2021). High costs associated with HTS may be a significant impediment as the total cost of sampling, extraction of DNA, PCR, and sequencing of fungal DNA at a commercial lab is AU$/US$50–120/sample with the total costs of a one-off analysis quickly escalating depending on the number of samples and thus may be inaccessible to many researchers and practitioners (Birnbaum and Trevathan-Tackett 2022). Therefore, more collaboration to advance the study of plant-associated fungal diversity in underrepresented coastal regions is warranted using the latest cutting-edge molecular tools.

We suggest that further research should focus on addressing the knowledge gaps highlighted above as well as assess the functional roles of fungal communities across coastal wetlands. Currently, majority of the reported studies were descriptive, and most are focused on single fungal guild e.g., mycorrhizae. While studies on mycorrhizae provide important insights into their ecosystem role, especially with respect to plant physiology in coastal systems, systematic community level characterization of fungi is needed—at the levels of taxonomic, functional, and metabolic diversity.

Conclusion

The need for fungal research in natural coastal wetlands cannot be understated. As increasing pressures from anthropogenic-driven changes to the coastal wetland environment impact these ecosystems, generating baseline data of fungal communities—their distribution, diversity, and functional roles—in natural, pre-disturbed environment is critical for a better predictive capacity to anticipate organismal and ecosystem-system level impacts of such disturbances. It can also provide important insights that can aid global coastal restoration and management efforts (Birnbaum and Trevathan-Tackett 2022; Farrer et al. 2022).