1 Introduction

Coral reef fish are the core biological elements that maintain the structure and function of coral reef systems. It plays an irreplaceable role in key processes such as material cycling, habitat construction and algal regulation, and these processes directly determine the resilience and productivity level of coral reef ecosystems (Xi et al., 2022; Luo et al., 2023; Huang et al., 2024; Zhao et al., 2024). Studies have confirmed that fish population density and species richness are significantly positively correlated with the health of coral bases. Changes in coral coverage and three-dimensional structure significantly affect the dynamics and ecological functions of fish communities.

Hainan Island and the South China Sea, as typical tropical sea areas, are home to coral reef ecosystems with high biodiversity. This area has been listed by the international academic community as a key hotspot for Marine biodiversity. Existing research has recorded over a thousand species of coral reef fish, covering dozens of families, genera and groups. For example, 874 species have been recorded in the Xisha Islands of the South China Sea, and more than 360 species have been investigated and confirmed around Hainan Island, fully demonstrating its species enrichment characteristics (Xi et al., 2022; Zhang et al., 2023; Zhao et al., 2024). The diversity distribution of fish species is jointly regulated by environmental factors such as the gradient of water temperature and salinity, and the complexity of coral structure (Xia et al., 2022).

Analyzing the characteristics of fish diversity and their ecological functions is the theoretical basis for formulating scientific conservation strategies. The composition characteristics of fish communities can visually represent the health status of coral reefs and their ability to maintain ecological services such as coastal protection and fishery resource supply (Shi, 2015; Zhao et al., 2019; Zhao et al., 2024). Human disturbances such as overfishing, bottom trawling operations and coastal engineering have led to coral degradation and the loss of fish diversity, triggering the unification of community structure and the decline of ecological functions (Huang et al., 2024). Currently, environmental DNA (eDNA) detection and DNA barcoding technology have been widely applied in biodiversity assessment and conservation decision support (Xi et al., 2022; Zhang et al., 2023).

This study will explore the research progress on the biodiversity characteristics and ecological functions of fish species in Hainan Island and coral reefs in the South China Sea. This study will analyze the species composition patterns, community distribution patterns and their functional roles in maintaining the homeostasis of coral reefs, and identify natural and anthropogenic driving factors such as habitat heterogeneity, water quality parameters and fishing pressure. This study aims to analyze the latest research findings, clarify the fundamental role of fish diversity conservation in the sustainable management of coral reefs, provide scientific basis for the construction of regional ecological protection networks and the restoration of degraded habitats, and through interdisciplinary data integration, guide policy-making and protection practices to achieve the long-term maintenance of coral reef ecosystem service functions.

2 Research Progress on the Biodiversity of Coral Reef Fish

2.1 A multi-functional group composed of 800 species of fish

The sea area between Hainan Island and the South China Sea is rich in coral reef fish resources. Existing data indicate that more than 800 species of coral reef fish are distributed in this area (Xi et al., 2022; Zhang et al., 2023). These species cover multiple functional groups such as herbivorous species, carnivorous species, planktonic carnivorous species and omnivorous species, and maintain the homeostasis and resilience of coral reef systems through niche differentiation (Luo et al., 2023). Taking the coastal area of Hainan Island as an example, the study has identified 363 species of fish belonging to 114 families and 24 orders. Among them, the Perciformes occupy the dominant group, confirming the taxonomic diversity characteristics of this area.

This diversity feature not only represents the regional ecological value, but also supports the complex interaction network of coral-fish communities. Coral reef fish significantly affect system health and productivity levels through key roles such as nutrient regeneration, algal regulation, and habitat construction (Xi et al., 2022; Huang et al., 2024). Empirical studies have shown that the diversity index of fish communities is significantly correlated with the complexity of coral base structures, further verifying the ecological necessity of biodiversity conservation.

2.2 Spatial heterogeneity characteristics of diverse distribution

The fish diversity of coral reefs in Hainan Island and the South China Sea shows significant spatial differentiation, which is reflected in the gradient differences between the nearshore and the open sea and between the north and south sea areas. DNA barcoding studies have shown that there are significant differences in the composition of fish communities in the northern and southern sea areas of Hainan Island, with common species accounting for only a minority (Zhang et al., 2023). Synchronous monitoring shows that the fish biomass and community abundance in the nearshore waters of southwestern Hainan are higher than those in other areas, and there are seasonal and spatial dynamic fluctuations (Luo et al., 2023).

The eDNA survey on the coral reefs of Xi Island detected 41 species of fish belonging to 16 families, revealing the regulatory effect of local habitat conditions on diversity distribution (Xi et al., 2022). This spatial pattern is driven by multiple factors such as the complexity of coral structure, water quality parameters and the intensity of human activities, resulting in the formation of characteristic fish combinations in different reef areas.

2.3 The combined effect of environmental disturbances and human activities

Marine environmental parameters (such as water temperature fluctuations, salinity gradient changes and upwelling events) have a significant regulatory effect on fish diversity. Take the Qiongdong upwelling as an example. Although the special water mass environment formed by it provides a thermal refuge for some species, the algal outbreak and the simplification of habitat structure have led to a sharp decline in fish abundance and community reconfiguration (Zhu et al., 2022). The degradation of coral skeletons caused by large-scale bleaching events further reduces habitat heterogeneity and reef-dwelling species richness (Huang et al., 2024).

Human activities such as high-intensity fishing operations, coastal engineering construction and pollution emissions are the main causes of biodiversity decline. The disturbed area shows a trend of community homogeneity, and the proportion of small and low economic value fish has increased significantly (Lu et al., 2020; Luo et al., 2023; Yu et al., 2024). It is worth noting that some areas still maintain a relatively high ecological integrity, highlighting the effectiveness of targeted conservation measures in alleviating environmental stress.

3 The Ecological Functions of Coral Reef Fish

3.1 Multiple nutritional levels synergistically support energy transfer and structural homeostasis

Coral reef fish cover multiple trophic levels from primary consumers (such as herbivorous species) to top predators, forming the basic framework of energy transfer in the ecosystem. Take herbivorous fish as an example. They constitute the main biomass components, drive biomass turnover and nutrient regeneration, and are the core elements for maintaining the fishery productivity and nutrient cascade effect of coral reefs (Robinson et al., 2023). The dynamic balance of multiple nutrient groups ensures the stability of the ecosystem. When coral coverage fluctuates or the environment is disturbed, it will trigger the biomass and structural reorganization of the groups, thereby affecting the overall health status of the system (Russ et al., 2020).

The nutritional structure characteristics of fish are closely related to the health of coral substrates. High integrity coral cover areas support higher diversity and biomass levels, while disturbances such as whiteout events or overfishing will disrupt the nutrient network, resulting in decreased energy transfer efficiency and weakened system resilience. Therefore, maintaining the diverse combination of fish at multiple trophic levels is the key to achieving the structural and functional integrity of coral reefs.

3.2 Clean symbiosis and algal regulatory functions maintain the health of corals

Coral fish directly promote coral health and enhance interspecies interactions through specialized ecological functions. The cleaning behavior is manifested as specific fish eliminating parasites, necrotic tissues and mucus on the body surface of the host. This process not only provides nutritional supplementation for the cleaners, but also improves the survival rate of the host and enhances the stability of the community (Dunkley et al., 2018). Such interactions have highly selective characteristics. Some cleaning species specifically serve specific hosts or adjust the cleaning strategy according to the environmental gradient (Wu et al., 2024).

Parrofish, spiny fish and other herbivorous fish regulate algal biomass through continuous feeding to avoid competition for light resources between corals and algae (Clements et al., 2016; Topor et al., 2019). Such grazing behaviors create substrate conditions for the attachment of coral larvae, while nest-building activities promote biodiversity through microhabitat modification. The functional synergy supports the stress resistance of corals and maintains the complex interspecies interaction network.

3.3 Nutrient cycling and carbon sink functions enhance ecological connectivity

Coral fish dominate the nutrient cycle process through metabolic activities. Its excreta are rich in macronutrients and micronutrients. The excretion characteristics of species at different trophic levels vary significantly. These nutrients play a decisive role in the physiological metabolism and systemic productivity of corals. Species diversity guarantees the heterogeneity of nutrient supply and meets the metabolic requirements of corals and other organisms (Figure 1) (Van Wert et al., 2023; Robinson et al., 2023).

Fish activities also promote carbon storage and ecological connectivity through cross-habitat migration. Its feeding and migration behaviors accelerate the transreef transport of energy and matter, while the processes of nutrition and carbon cycling directly affect the succession direction of benthic communities and the system's anti-interference ability (Pellowe et al., 2024). Therefore, scientific management of coral reef fish communities is of core value for maintaining their ecological functions.

4 Threats to the Functions of Coral Reef Fish

4.1 Ecological damage effects of illegal fishing operations

Illegal operations such as blast fishing and cyanide fishing pose a devastating threat to coral reef systems. Such methods directly damage the coral skeleton structure, dismantle the fish habitats, and lead to the acute extinction of both target and non-target species. Cyanide not only causes momentary coma or death of fish, but also the residual toxic substances have long endangered coral polyps and reef organisms, triggering a decline in diversity (Goldberg and Wilkinson, 2004; Madeira et al., 2020). Blast fishing reduces habitat heterogeneity by crushing coral substrates, hinders the recovery process of fish communities, and ultimately weakens their ecological functions (Goldberg and Wilkinson, 2004).

The ecological effects of such destructive activities are exacerbated by the coupling of multiple stress factors. For example, the toxicity of cyanide is magnified under the condition of seawater warming, resulting in a sharp increase in fish mortality and an extended recovery period, and facing a higher risk in the context of climate warming (Madeira et al., 2020). Long-term illegal fishing leads to the loss of key functional groups, reducing the resilience of coral reefs and their ability to support fishery resources (Goldberg and Wilkinson, 2004).

4.2 The chain effect of bleaching driven by climate warming

Bleaching events caused by Marine heatwaves are the core indirect threat to coral reef fish. Thermal stress leads to the large-scale death of corals and disintegrates the habitat base of reef-dwelling fish (Goldberg and Wilkinson, 2004; Cinner et al., 2009). The decline in coral coverage reduces structural heterogeneity, resulting in a decrease in fish community abundance and niche diversity (Wilson et al., 2010). Habitat degradation has a particularly significant impact on obligate species that rely on living corals, triggering changes in the composition of functional groups and the absence of key ecological processes.

The synergistic effect of climate pressure and illegal fishing has intensified its harm. Cyanide exposure combined with high temperature significantly increases the mortality rate and predation vulnerability of fish, threatening the survival of populations (Madeira et al., 2020). The continuous degradation of coral reefs has led to a significant decline in their ecological service functions such as coastal protection and fishery support (Eddy et al., 2021).

4.3 The Superimposed effect of land-based pollution and tourism development

Land-based pollutants (such as nutrient input, sediment deposition and sewage discharge) seriously threaten the health of corals. Pollutants deteriorate water quality, induce algal blooms, increase the risk of coral and fish diseases, and ultimately lead to the decline of biodiversity (Goldberg and Wilkinson, 2004). Tourism infrastructure projects have caused habitat fragmentation and physical damage. Coupled with human interference, the degradation of ecological functions has been exacerbated. Habitat fragmentation disrupts the connectivity of reef areas, hinders the migratory behavior of fish and the cross-regional transmission of energy and matter. Such disturbances form a superimposed effect with pollution and fishing pressure, resulting in the obstruction of core ecological processes such as nutrient cycling and larval replenishment (Figure 2) (El-Naggar, 2020; Ahmad et al., 2024). The cumulative effect of multiple stresses weakens the stability of coral reef systems and threatens the sustainability of their ecological service functions.

5 Coral Reef Protection and Management Strategies

5.1 Construction and optimization of marine protected area network

The scientific planning of coral reef reserve networks plays a crucial role in maintaining fish diversity and ecological functions. Studies have shown that well-managed protected areas (even if the area is limited) can significantly increase fish biomass, species richness and individual size compared with non-protected areas, verifying their supporting efficacy for coral reef health (MacNeil et al., 2015; Bayley et al., 2019). The effectiveness of protection not only depends on regional demarcation, but also requires strengthening the law enforcement supervision and ecological monitoring system to ensure the achievement of protection goals and the connection of ecological corridors (Ban et al., 2023).

Optimizing the protected area network requires a comprehensive assessment of biophysical parameters such as habitat heterogeneity and ecological connectivity. Establishing a composite management unit including core no-fishing zones and sustainable utilization zones can not only promote the recovery of fish populations but also alleviate socio-economic contradictions (Fidler et al., 2021). Strategic layout of the protected area network to enhance the connectivity between reef sections is crucial for maintaining the integrity of fish migration channels and life history, thereby improving the resilience of the system.

5.2 Implement eco-friendly fishery management

The implementation of ecologically-oriented fishery policies holds core value in alleviating the pressure on key functional species and maintaining ecological service functions. Measures such as fishing gear control, size restrictions and bans on fishing of specific species can help maintain the baseline value of fish biomass and promote resource recovery. Dynamic management strategies such as crop rotation fishing moratorium and hierarchical quotas can effectively maintain key functions such as forage regulation in areas lacking complete protected zones (Dee et al., 2014; MacNeil et al., 2015).

To adopt the ecosystem-level fishery management model, it is necessary to comprehensively assess the ecological niches of species and the cumulative effect of fishing in order to maintain the integrity of the system (Appeldoorn, 2008; Cinner et al., 2020). Promoting stakeholder participatory management can enhance policy compliance and ensure that measures are adapted to the regional ecological-socio-economic characteristics, achieving the coordinated development of protection and livelihoods (Mapstone et al., 2008).

5.3 Strengthen the scientific research support and monitoring system

Systematic scientific research and monitoring are the cornerstones of adaptive management of coral reefs. Emerging technologies such as 3D image modeling and environmental DNA (eDNA) can achieve rapid and non-destructive assessment of coral health and fish communities, providing real-time data for management decisions (Bayley et al., 2019). The dynamic monitoring system can quantify the management effectiveness, identify the ecological evolution trend, and adjust strategies in a timely manner to deal with emerging threats (Ban et al., 2023).

Long-term ecological monitoring plans are particularly crucial for tracking biomass fluctuations, functional evolution and the effects of management interventions (Bayley et al., 2019; Ban et al., 2023). The in-depth integration of scientific research and management decisions ensures that conservation actions have scientific basis and dynamic response capabilities, thereby enhancing the sustainability of the Hainan Island - South China Sea coral reef system (MacNeil et al., 2015).

6 Research Limitations and Future Prospects

6.1 Insufficiency of research on the correlation between functional diversity and ecological services

Although progress has been made in the cataloging research of coral reef fish species, there are still significant gaps in the systematic analysis of functional diversity characteristics and ecosystem services (such as nutrient cycling, habitat maintenance and interference resistance ability) (Villeger et al., 2017; Bellwood et al., 2018). Most of the existing studies focus on quantifiable parameters rather than empirically analyzing the contribution of specific functional groups to ecological processes, which restricts the prediction accuracy of the impact of fish community structure evolution under human stress on ecological services.

Although current functional diversity indicators are widely used, their relationship with ecological functions is mostly based on hypotheses rather than empirical evidence. It is necessary to break through the traditional description of morphological characteristics, quantify the ecological contributions of different functional groups, and analyze how functional redundancy and uniqueness regulate system stability (Brandl et al., 2016; Bellwood et al., 2018; Streit et al., 2019).

6.2 The lack of standardization in the monitoring system and database

The existing monitoring data of coral reef fish have problems such as dispersion, lack of standards and limited spatio-temporal coverage. The absence of a long-term standardized monitoring system hinders the ability of community dynamic assessment and ecological early warning (Yeager et al., 2017; Martin et al., 2024). The differences in monitoring methods lead to difficulties across regions, affecting the evaluation of management effectiveness (Martin et al., 2024).

The scarcity of functional trait databases has limited the construction of meta-analysis and prediction models (Villeger et al., 2017). There is an urgent need to establish a high-quality database through the application of new technologies and protocol standardization to support adaptive management decisions (Villeger et al., 2017).

6.3 Research directions of cross-domain collaboration and technology integration

In the future, regional cooperation needs to be strengthened to address the cross-border challenges of coral reef systems. The collaborative mechanism can promote data sharing, unification of monitoring standards and management linkage, and enhance the effectiveness of large-scale protection (Yeager et al., 2017; Martin et al., 2024). Multidisciplinary research integrating ecology and social sciences will deepen the understanding of functional driving mechanisms and optimize intervention strategies (Bellwood et al., 2018).

The application of technologies such as environmental DNA (eDNA) and remote sensing provides new approaches for fish community monitoring. Combining traditional ecological knowledge with community participation can enhance the dynamic monitoring and adaptive management capabilities in the Hainan Island - South China Sea region.

7 Conclusion

The biodiversity and ecological functions of coral reef fish are the core elements for maintaining the homeostasis and resilience of coral reef systems. The species abundance and functional heterogeneity of fish communities support the functional operation of the system, including biomass accumulation, climate adaptability and the maintenance of key ecological processes. These elements have dual values for biodiversity conservation and human well-being. The synergistic effect of multiple functional traits (such as grazing behavior, trophic cascade and material cycling) guarantees the health and productivity level of coral reefs.

Under multiple pressures such as overfishing, habitat degradation and climate change, it is urgent to strengthen protection management and scientific research investment. A scientifically designed network of Marine protected areas and an ecosystem-oriented fishery policy can generate significant conservation benefits and maintain fishery resources and ecological functions. However, with the intensification of human interference, biodiversity conservation is facing greater challenges, which highlights the importance of adaptive management mechanisms and continuous monitoring systems for optimizing conservation strategies.

Hainan Island and the South China Sea area, with their unique characteristics of coral reef fish diversity, have become a model area for tropical sea area protection. By strengthening the priority of classification and functional diversity protection and promoting the multi-dimensional integration of scientific research, management and policy, this region is expected to achieve balanced development of ecological protection and resource utilization, providing an innovative model reference for global coral reef management.

Acknowledgments

We would like to thank Dr Xuan continuous support throughout the development of this study.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

References

Ahmad I., Guo P., Zhao M.X., Zhong Y., Zheng X., Zhang S., Qiu J., Shi Q., Yan H., Tao S., and Xu L., 2024, Coral reefs of Pakistan: a comprehensive review of anthropogenic threats climate change and conservation status, Frontiers in Marine Science, 11: 1466834.

https://doi.org/10.3389/fmars.2024.1466834

Appeldoorn R., 2008, Transforming reef fisheries management: application of an ecosystem-based approach in the USA caribbean, Environmental Conservation, 35: 232-241.

https://doi.org/10.1017/S0376892908005018

Arai T., 2015, Diversity and conservation of coral reef fishes in the Malaysian South China Sea, Reviews in Fish Biology and Fisheries, 25: 85-101.

https://doi.org/10.1007/s11160-014-9371-9

Ban N.C., Darling E.S., Gurney G.G., Friedman W., Jupiter S.D., Lestari W.P., Yulianto I., Pardede S., Tarigan S., Prihatiningsih P., Mangubhai S., Naisilisili W., Dulunaqio S., Naggea J., Ranaivoson R., Agostini V., Ahmadia G., Blythe J., Campbell S., Claudet J., Cox C., Epstein G., Estradivari E., Fox M., Gill D., Himes-Cornell A., Jonas H., Mcleod E., Muthiga N., and McClanahan T., 2023, Effects of management objectives and rules on marine conservation outcomes, Conservation Biology, 37(6): e14156.

https://doi.org/10.1111/cobi.14156

Bayley D.T.I., Mogg A.O.M., Purvis A., and Koldewey H.J., 2019, Evaluating the efficacy of small‐scale marine protected areas for preserving reef health: a case study applying emerging monitoring technology, Aquatic Conservation: Marine and Freshwater Ecosystems, 29(12): 2026-2044.

https://doi.org/10.1002/aqc.3215

Bellwood D.R., Streit R.P., Brandl S.J., and Tebbett S.B., 2018, The meaning of the term ‘function’ in ecology: a coral reef perspective, Functional Ecology, 33(6): 948-961.

https://doi.org/10.1111/1365-2435.13265

Brandl S.J., Emslie M.J., Ceccarelli D.M., and Richards Z., 2016, Habitat degradation increases functional originality in highly diverse coral reef fish assemblages, Ecosphere, 7(11): e01557.

https://doi.org/10.1002/ECS2.1557

Cinner J., McClanahan T., Graham N., Pratchett M., Wilson S., and Raina J., 2009, Gear-based fisheries management as a potential adaptive response to climate change and coral mortality, Journal of Applied Ecology, 46: 724-732.

https://doi.org/10.1111/J.1365-2664.2009.01648.X

Cinner J., Zamborain‐Mason J., Gurney G., Graham N., Graham N., MacNeil M., Hoey A., Mora C., Villéger S., Maire E., Maire E., Maire E., McClanahan T., Maina J., Maina J., Kittinger J., Hicks C., Hicks C., D'agata S., Huchery C., Barnes M., Feary D., Williams I., Kulbicki M., Vigliola L., Wantiez L., Edgar G., Stuart‐Smith R., Sandin S., Green A., Beger M., Friedlander A., Wilson S., Brokovich E., Brooks A., Cruz‐Motta J., Booth D., Chabanet P., Tupper M., Ferse S., Sumaila U., Hardt M., Mouillot D., and Mouillot D., 2020, Meeting fisheries ecosystem function and biodiversity goals in a human-dominated world, Science, 368: 307-311.

https://doi.org/10.1126/science.aax9412

Clements K., German D., Piché J., Tribollet A., and Choat J., 2016, Integrating ecological roles and trophic diversification on coral reefs: multiple lines of evidence identify parrotfishes as microphages, Biological Journal of The Linnean Society, 120: 729-751.

https://doi.org/10.1111/BIJ.12914

Dee L., Horii S., and Thornhill D., 2014, Conservation and management of ornamental coral reef wildlife: successes shortcomings and future directions, Biological Conservation, 169: 225-237.

https://doi.org/10.1016/J.BIOCON.2013.11.025

Dunkley K., Cable J., and Perkins S.E., 2018, The selective cleaning behaviour of juvenile blue-headed wrasse (Thalassoma bifasciatum) in the Caribbean, Behavioural Processes, 147: 5-12.

https://doi.org/10.1016/j.beproc.2017.12.005

Eddy T.D., Lam V.W.Y., Reygondeau G., Cisneros-Montemayor A.M., Greer K., Palomares M.L.D., Bruno J., Ota Y., and Cheung W., 2021, Global decline in capacity of coral reefs to provide ecosystem services, One Earth, 4(9): 1278-1285.

https://doi.org/10.1016/j.oneear.2021.08.016

El-Naggar H., 2020, Human impacts on coral reef ecosystem, Natural Resources Management and Biological Sciences, 2020.

https://doi.org/10.5772/intechopen.88841

Fidler R.Y., Andradi-Brown D.A., Awaludinnoer, Pada D., Purwanto, Hidayat N., Ahmadia G., and Harborne A., 2021, The importance of biophysical context in understanding marine protected area outcomes for coral reef fish populations, Coral Reefs, 40: 791-805.

https://doi.org/10.1007/s00338-021-02085-y

Goldberg J., and Wilkinson C., 2004, Global threats to coral reefs: coral bleaching global climate change disease predator plagues and invasive species, Status of Coral Reefs of The World, 2004: 67-92.

Huang L.T., McWilliam M., Liu C.Y., Yu X.L., Jiang L., Zhang C., Luo Y., Yang J.H., Yuan X.C., Lian J.S., and Huang H., 2024, Loss of coral trait diversity and impacts on reef fish assemblages on recovering reefs, Ecology and Evolution, 14(11): e70510.

https://doi.org/10.1002/ece3.70510

Lu Y., Yang Y., Sun B., Yuan J., Yu M., Stenseth N.C., Bullock J.M., and Obersteiner M., 2020, Spatial variation in biodiversity loss across China under multiple environmental stressors, Science Advances, 6(47): eabd0952.

https://doi.org/10.1126/sciadv.abd0952

Luo Z., Yang C., Wang L., Liu Y., Shan B., Liu M., Chen C., Guo T., and Sun D., 2023, Relationships between fish community structure and environmental factors in the nearshore waters of Hainan Island South China, Diversity, 15(8): 901.

https://doi.org/10.3390/d15080901

MacNeil M., Graham N., Cinner J., Wilson S., Williams I., Maina J., Newman S., Friedlander A., Jupiter S., Polunin N., and McClanahan T., 2015, Recovery potential of the world's coral reef fishes, Nature, 520: 341-344.

https://doi.org/10.1038/nature14358

Madeira D., Andrade J., Leal M.C., Ferreira V., Rocha R.J.M., Rosa R., and Calado R., 2020, Synergistic effects of ocean warming and cyanide poisoning in an ornamental tropical reef fish, Frontiers in Marine Science, 7: 246.

https://doi.org/10.3389/fmars.2020.00246

Mapstone B., Little L., Punt A., Davies C., Smith A., Pantus F., McDonald A., Williams A., and Jones A., 2008, Management strategy evaluation for line fishing in the great barrier reef: balancing conservation and multi-sector fishery objectives, Fisheries Research, 94: 315-329.

https://doi.org/10.1016/J.FISHRES.2008.07.013

Martin B., Huveneers C., Reeves S., and Baring R., 2024, Fishing for more than shellfish: a systematic review of fish community monitoring of shellfish reefs, Restoration Ecology, 33(1): e14323.

https://doi.org/10.1111/rec.14323

McKinley S.J., Saunders B.J., Rastoin-Laplane E., Salinas-de-León P., and Harvey E., 2022, Functional diversity of reef fish assemblages in the galapagos archipelago, Journal of Experimental Marine Biology and Ecology, 549: 151695.

https://doi.org/10.1016/j.jembe.2022.151695

Pellowe K.E., Durfort A., Burkepile D.E., Mouillot D., and Lade S.J., 2024, Positive feedbacks in coastal reef social-ecological systems can maintain coral dominance, ICES Journal of Marine Science, 82(5): fsae182.

https://doi.org/10.1093/icesjms/fsae182

Robinson J.P.W., Darling E.S., Maire E., Hamilton M., Hicks C.C., Jupiter S.D., MacNeil A., Mangubhai S., McClanahan T., Nand Y., and Graham N., 2023, Trophic distribution of nutrient production in coral reef fisheries, Proceedings of the Royal Society B, 290(2008): 20231601.

https://doi.org/10.1098/rspb.2023.1601

Russ G.R., Rizzari J.R., Abesamis R.A., and Alcala A.C., 2020, Coral cover a stronger driver of reef fish trophic biomass than fishing, Ecological Applications, 31(1): e02224.

https://doi.org/10.1002/eap.2224

Streit R.P., Cumming G.S., and Bellwood D.R., 2019, Patchy delivery of functions undermines functional redundancy in a high diversity system, Functional Ecology, 33(6): 1144-1155.

https://doi.org/10.1111/1365-2435.13322

Topor Z.M., Rasher D.B., Duffy J.E., and Brandl S., 2019, Marine protected areas enhance coral reef functioning by promoting fish biodiversity, Conservation Letters, 12(4): e12638.

https://doi.org/10.1111/conl.12638

Van Wert J.C., Ezzat L., Munsterman K.S., Landfield K., Schiettekatte N., Parravicini V., Casey J., Brandl S., Burkepile D., and Eliason E., 2023, Fish feces reveal diverse nutrient sources for coral reefs, Ecology, 104(8): e4119.

https://doi.org/10.1002/ecy.4119

Villéger S., Brosse S., Mouchet M., Mouillot D., and Vanni M., 2017, Functional ecology of fish: current approaches and future challenges, Aquatic Sciences, 79: 783-801.

https://doi.org/10.1007/s00027-017-0546-z

Wang Q., Sun C.M., and Liu L.P., 2024, Ecological impacts of common carp invasions: a global perspective, International Journal of Aquaculture, 14(4): 174-183.

https://doi.org/10.5376/ija.2024.14.0018

Wilson S., Wilson S., Fisher R., Pratchett M., Graham N., Graham N., Dulvy N., Turner R., Cakacaka A., and Polunin N., 2010, Habitat degradation and fishing effects on the size structure of coral reef fish communities, Ecological Applications : A Publication of The Ecological Society of America, 20(2): 442-451.

https://doi.org/10.1890/08-2205.1

Wu J.N., Xu X.Y., and Wu L.M., 2024, The physiological and ecological effects between ocean acidification and coral reefs, International Journal of Marine Science, 14(1): 14-20.

https://doi.org/10.5376/ijms.2024.14.0003

Xi R., Gao W., Wang X., and Xing Y., 2022, Species diversity of coral reef fishes around the West Island of Sanya City South China Sea based on environmental DNA, Biodiversity Data Journal, 10: e89685.

https://doi.org/10.3897/BDJ.10.e89685

Xia J.Q., Zhu W.T., Liu X.B., Ren Y.X., Huang J.Z., Zhu M., Wu Z.Q., Wang A.M., and Li X.B., 2022, The effect of two types of grid transplantation on coral growth and the in-situ ecological restoration in a fragmented reef of the South China Sea, Ecological Engineering, 177: 106558.

https://doi.org/10.1016/j.ecoleng.2022.106558

Yeager L., Deith M., McPherson J., Williams I., and Baum J., 2017, Scale dependence of environmental controls on the functional diversity of coral reef fish communities, Global Ecology and Biogeography, 26: 1177-1189.

https://doi.org/10.1111/GEB.12628

Yu P., Luo Z., Wang L., Liu Y., Huang Y., Shan B., Yang C., and Sun D., 2024, Evaluation of ecological health of Hainan Island inshore waters in the South China Sea based on preliminary fish biotic integrity index (Actinopterygii and Elasmobranchii), Acta Ichthyologica et Piscatoria, 54: 221-233.

https://doi.org/10.3897/aiep.54.130966

Zhang J.X., Xu L., Du F.Y., Tang Q.H., Wang L.G., Ning J.J., Huang D.L., Li Y.F., Liu S.S., and Wang X.H., 2023, DNA barcoding of marine fish species in the waters surrounding Hainan Island northern South China Sea, Frontiers in Marine Science, 10: 1249073.

https://doi.org/10.3389/fmars.2023.1249073

Zhao J., Wang T., Li C., Shi J., Xie H., Luo L., Xiao Y., and Liu Y., 2024, Seven decades of transformation: evaluating the dynamics of coral reef fish communities in the Xisha Islands South China Sea, Reviews in Fish Biology and Fisheries, 34(4): 1261-1281.

https://doi.org/10.1007/s11160-024-09872-0

Zhao M.X., Zhang H.Y., Zhong Y., Jiang D.P., Liu G.H., Yan H.Q., Zhang H.Y., Guo P., Li C., Yang H.Q., Chen T.G., and Wang R., 2019, The status of coral reefs and its importance for coastal protection: a case study of Northeastern Hainan Island South China Sea, Sustainability, 11(16): 4354.

https://doi.org/10.3390/SU11164354

Zhu W.T., Ren Y.X., Liu X.B., Huang D.J., Xia J.Q., Zhu M., Yin H.Y., Chen R.W., and Li X.B., 2022, The impact of coastal upwelling on coral reef ecosystem under anthropogenic influence: coral reef community and its response to environmental factors, Frontiers in Marine Science, 9: 888888.

https://doi.org/10.3389/fmars.2022.888888