4.1 Macrobenthic Community Characteristics
Species composition is a fundamental characteristic of a community and a critical aspect of community ecology research (Zhou 2021). In this study, a total of 121 species of macrobenthos were collected, surpassing the 65 species recorded in the historical data from 2004–2005 (Zheng et al. 2007) and the 113 species reported in another study from 2006–2007 (Peng et al. 2011). This may be attributed to the multiple survey sections and the long time span as well as an increase in biodiversity in the mangrove areas (Chen et al. 2013) of Yueqing Bay due to the recent planting of additional mangroves (Chen 2016). Mangroves promote macrobenthic diversity and abundance by improving the microhabitats and food resources of macrobenthos (Demopoulos and Smith 2010; Frith 1977). Mollusca, Arthropoda, and Annelida were the dominant groups in the mangrove areas of Yueqing Bay, which is consistent with historical survey data (Peng et al. 2011; Zheng et al. 2006). The average abundance and biomass of macrobenthos were 798.13 ind./m² and 140.51 g/m², respectively, both of which were higher than historical levels (Zheng et al. 2007).
Changes in the dominant species of macrobenthos can effectively reflect variations in community structure and environmental shifts (Song et al. 2022; Yan et al. 2020). Historical studies in the mangrove areas of Yueqing Bay identified 19 dominant macrobenthic species (Peng et al. 2011), whereas 8 dominant species were observed in this study, with only 2 species (A. latericea and M. longipes) found in both datasets. Possible reasons for this discrepancy include the following: 1) There is a significant temporal gap between the studies. The previous study was conducted from 2006 to 2007, whereas the current study was conducted from 2021 to 2022, suggesting that the macrobenthic community in Yueqing Bay has changed. Yueqing Bay has undergone large-scale mangrove planting since 2005 (Chen 2016), and the study by Peng et al. coincided with a relatively early stage of mangrove growth, which may have contributed to the identification of a greater number of dominant species (Huang et al. 2017). 2) Differences in research methods. An earlier study used dominance (Y) as an evaluation index, whereas this study employed the Index of Relative Importance (IRI). The use of different methodologies could contribute to the observed discrepancy. Compared to Y, the IRI offers a more comprehensive and quantitative reflection of changes in dominant species (Han et al. 2004) and can indicate the importance of dominant species in community productivity (Smith and Knapp 2003). Among the common dominant species, A. latericea is primarily distributed in surface mud and is a frequent dominant species in mangrove ecosystems (Huang et al. 2017; Lin et al. 2021; Ma et al. 2018). M. longipes is phytophagous (Ph), and the increase in mangrove litter and organic debris provides an abundant food source (Zhang et al. 2019), which may explain why these species have maintained their dominant status over time.
4.2 Short-term responses of the Macrobenthos Community to Stock Enhancement
In this study, the number of macrobenthic species was generally greater after stock enhancement than before, with the exception of January 2022. This can be attributed to lower winter water temperatures, which can limit the survival of warm-water species (Zhou et al. 2016). When comparing October 2022 with October 2021, the number of macrobenthic species increased. Most species were common both before and after stock enhancement, with few species, typically those considered rare, uniquely present in low abundance before or after stock enhancement. For instance, Philine kinglipini and Amalda rubiginosa were observed before stock enhancement, whereas Eucrate crenata and Peronia verruculata were found after stock enhancement. Therefore, stock enhancement increased the number of macrobenthic species while largely preserving the species composition (Wang 2016).
The dominance of certain species in a biological community is closely linked to community stability. A greater degree of species dominance leads to a more unbalanced and fragile community ; a lower degree of species dominance leads to a more stable community (Ding et al. 2021; Zhou 2021). Before stock enhancement, there were three dominant species, with T. arcuata exhibiting an extremely high Index of Relative Importance (IRI) value of 6396.57. Aside from a high IRI value for P. esculenta in January 2022 due to the stocking event, the number of dominant species increased after stock enhancement, leading to more uniform IRI values. Compared with those in October 2021, the abundance and biomass of P. esculenta in October 2022 increased by 10.37% and 17.70%, respectively. However, stock enhancement caused significant changes throughout the entire macrobenthic community, resulting in more pronounced increases in species number, abundance, and biomass, which led to a decrease in the IRI value of P. esculenta from 4413.69 in October 2021 to 2475.37 in October 2022. Crabs are often considered ecosystem engineers (Lee 2008), and an increase in dominant crab species after stock enhancement can effectively promote the decomposition and transformation of mangrove litter, thereby increasing and enriching primary production in mangrove ecosystems (Xu et al. 2010).
After stock enhancement, the abundance and biomass of macrobenthos were generally greater than before enhancement, with the exception of biomass in January 2022. The lower biomass in January 2022 can be attributed to seasonal factors that limited the survival of warm-water species (Zhou et al. 2016). Despite seasonal constraints, abundance was greater in January 2022 than in October 2021, because many P. esculenta survived after stock enhancement. Notably, the emergence and subsequent dominance of S. sinensis and P. laevis in Section T2 led to the particularly high abundance observed in April 2022. These species were only abundant in Section T2 and less prevalent in other sections, possibly due to their cultivation and use as pond bait (Leng et al. 2013). by local fishermen Secondary productivity is a key indicator for assessing the state of ecosystems (Asmus 1987; Tumbiolo and Downing 1994), and an increase in secondary productivity may enhance the resistance of the macrobenthic community to organic pollution stress (Du et al. 2024). Biomass is the most significant factor influencing changes in secondary productivity, and the trends in secondary productivity and biomass are often closely aligned (Tumbiolo and Downing 1994), as observed in this study. After stock enhancement, the secondary productivity of the macrobenthic community was generally greater than before, except in January 2022.
In terms of community differences, based on the results of CLUSTER and NMDS analyses, the macrobenthic communities before stock enhancement were grouped separately at a 46% similarity level, whereas those after stock enhancement were clustered together. The difference in macrobenthic community structure before and after stock enhancement was not significant (P > 0.05), indicating that stock enhancement increased the similarity of macrobenthic communities but did not fundamentally alter the community structure. The primary contributor to the average dissimilarity in the macrobenthic community before and after stock enhancement was P. latericea. Phascolosoma esculenta, a detritivore (D), prefers to inhabit muddy or muddy-sandy mangrove areas in the mid-to-high tide zone. P. latericea is distributed mainly in the surface mud layer and is an omnivorous (O) predator (Wang et al. 2024). Omnivores primarily feed on plant litter, carrion, or small bivalves or absorb organic matter from the water through their skin or gill epithelia (Meng et al. 2023). Stock enhancement introduced many P. esculenta strains into the mangrove areas. Some P. esculenta die due to adaptability, environmental factors, and ecological carrying capacity, producing large amounts of organic matter and carrion that omnivores, such as P. latericea, can utilize. However, both omnivores and detritivores are preyed upon by carnivores (Ca) (Ge et al. 2008). The increase in P. esculenta reduced the number of P. latericea individuals preyed upon by carnivores. These factors collectively made P. latericea the main contributor to the differences in the macrobenthic community.
The MPI is based on the ABC curve. According to the MPI results, the benthic ecological quality initially declined then increased. In April 2022, the benthic ecological quality reached its lowest point and was categorized as "moderate." During this period, the emergence of numerous small individuals of S. sinensis and P. laevis, which are used as bait, caused the abundance curve to rise above the biomass curve and partially cross it. Human disturbance, the primary factor contributing to moderate pollution, negatively impacted the macrobenthic community structure (Peng et al. 2011). During the survey and sampling period of this study, fishermen were observed harvesting only the large individuals of Phascolosoma esculenta from the tidal flats. The average abundance of Phascolosoma esculenta increased from 30.13 ind./m² in October 2021 to 31.29 ind./m² in October 2022, while the average biomass decreased from 5.17 g/m² to 5.04 g/m², which reflects the impacts of the fishermen’s harvest. Therefore, post-stock enhancement management measures are crucial. According to the MPI results, the difference in benthic ecological quality between October 2021 and October 2022 was minimal, with October 2022 showing slight improvement compared with October 2021. This result is consistent with the M-AMBI findings, indicating that stock enhancement improved benthic ecological quality despite anthropogenic disturbances (Du et al. 2024; Yang et al. 2024).
4.3 Applicability of MPI
Due to the unique application scenarios and the specific applicability of benthic indices, different indices may yield varying conclusions for the same area (Brauko et al. 2016; Gao et al. 2023; Song et al. 2023). In this study, the MPI, which was previously used in the mangrove areas of Yueqing Bay (Long et al. 2008), was selected for evaluation. There are two main reasons for this choice: first, it facilitates the comparison of changes in benthic ecological quality; second, it allows for exploring the applicability of this index in the mangrove areas of Yueqing Bay, thus enriching macrobenthic monitoring techniques. The average MPI value in this study was − 12.79, indicating a high level of benthic ecological quality and a clear improvement over the period from 2004 to 2006 (Long et al. 2008). The M-AMBI was shown to be applicable to the mangrove areas of Yueqing Bay (Yang et al. 2024). The variation curves of the MPI and M-AMBI were generally similar. Both indices identified human disturbances in April 2022 and a slight improvement in benthic ecological quality, suggesting that the MPI results have high reference value.
The MPI has several limitations. It has a relatively late start and a limited dataset; at times, biological and ecological expertise may be required to refine the results (Long et al. 2008). However, its calculation method is simple and user-friendly. Additionally, compared to the results of the diversity indices and ABC curves, the MPI provides more accurate evaluations (Cai 2003). This method is accessible to nonprofessionals and has great potential for further development, such as dividing benthic ecological quality into five categories and defining specific quality ranges for different substrate types. The M-AMBI remains the preferred option for evaluating the mangrove areas of Yueqing Bay. Other benthic indices, such as the MPI, can serve as supplementary references. By considering the evaluation results from multiple indices and analyzing environmental factors, a more objective assessment can be achieved.