DNA barcoding reveals larval fish diversity and distribution along the Cibareno River (West Java, Indonesia)
Arif Wibowo A , Andi Chadijah A , Kurniawan Kurniawan A B C , Vitas Atmadi Prakoso
A B C * , Dwi Atminarso A B C , Deni Irawan A , Fathur Rochman A , Septiana Sri Astuti A , Indah Lestari Surbani D , Tri Deniansen A , Imron Rosadi E , Yohanes Yudha P. Jaya E , Sudarsono Sudarsono E , Lee J. Baumgartner B , Nicolas Hubert F and Ivor G. Stuart B
A
B
C
D
E
F
Abstract
There is a global problem with ongoing riverine infrastructure projects where, despite knowledge of potential environmental impacts, there is rapid development, often without appropriate environmental safeguards. This results in fragmentation of riverine fish communities, especially diadromous species. Understanding freshwater fish larval ecology is critical to provide insight into the likely impacts of these projects.
To assess fish larval biodiversity on the basis of DNA barcoding, abundance and its distribution pattern in the Cibareno River.
Fish larvae were collected at six locations in the Cibareno River. The larvae were identified by DNA barcoding.
A notable disparity was seen in the distribution of larval abundance in different locations. The non-native species, Poecilia reticulata, was the most abundant larval species, with an intraspecific diversity of 0.003 (99.7% similarity). The upstream area exhibited a lower level of larval species diversity than did the downstream area.
Genetic identification can reliably identify fish larvae and determine their spatial riverside distribution in the Cibareno River. The conservation of connectivity maintains fish community integrity and diversity between upstream and downstream locations in the weir building plan.
This discovery emphasises the relevance of larval identification in fish biodiversity assessment and sustainable fisheries resource monitoring.
Keywords: aquatic ecology, biodiversity, conservation, DNA barcoding, early life stages, ichthyoplankton, larval fish, species composition.
Introduction
River flow regulation is a major threat to riverine fish communities, with population declines occurring globally (Darwall and Freyhof 2015). Among the chief impacts of new infrastructure is an immediate change to upstream hydrodynamics from dynamic free-flowing to deep slow-flowing habitats, with a consequent loss of fish biodiversity (Ward and Stanford 1995; Widén et al. 2021). Fragmentation of the riverine continuum also has impacts on upstream and downstream fish migrations, with diadromous fish communities typically declining above barriers with varying degrees of isolation on freshwater fishes in upstream area (Drouineau et al. 2018). When the river flow is dammed, the resultant conditions can be more favourable to introduced species than native species (Mercado-Silva et al. 2009).
South-east Asia is emblematic of the global trend for ongoing riverine infrastructure projects where, despite strong existing knowledge of potential serious environmental impacts, especially to riverine fish communities, there is rapid development, often without appropriate environmental safeguards (King and Brown 2010; Soukhaphon et al. 2021). In some cases, fisheries resources are in serious decline (Campbell and Barlow 2020; Vu et al. 2021), highlighting the need for much greater scientific effort to quantify these changes so as to target efforts to protect key elements of riverine hydrology and fish communities (Schmutz et al. 2015; Ziegeweid et al. 2022).
Early life stages (e.g. eggs and larvae) of riverine fish are the most vulnerable to perturbation, while also playing a crucial trophic link between plankton and higher predators, as well as in recruitment and self-sustaining populations (Heimeier et al. 2010; Ardura et al. 2016). A comprehensive understanding of the biology of early fish life-history stages is necessary to improve fisheries management (Kouhanestani et al. 2020), which can help protect critical habitats and species (Lucas and Baras 2000; Cooke 2008). Specifically, knowledge of larval ecology can provide fundamental information on fish reproductive biology, such as reproduction timing, geo-locations, migration routes, and population recruitment success (Cooke et al. 2016; Wibowo et al. 2017). Such information is very important and extremely valuable for understanding the impacts of riverine infrastructure, developing management and conservation plans, establishing fishing management strategies, and contributing to the conservation of endangered and vulnerable species (Reynalte-Tataje et al. 2011; Wibowo et al. 2017).
The Cibareno River in West Java, Indonesia forms a natural border between Banten and West Java provinces in Indonesia. The upstream area is located in the Halimun National Park area of West Java. Cibareno River flows ~37 km to Pelabuhan Ratu Bay. There is no recent information available to evaluate the fish biodiversity in the river. In 2002, ~22 fish species from 10 families were recorded (Rachmatika et al. 2002). Recent assessments with DNA barcoding of the ichthyofauna of Java Island reported 159 species (Dahruddin et al. 2017; Hubert et al. 2019), suggesting that fish diversity in the Cibareno River might be underestimated. An irrigation weir project is currently under development in the middle part of the Cibareno River (Fig. 1). While conducting the survey, the dam foundation had reached a height of ~1 m, a fraction of the planned 3.25 m, stretching across the entire riverbed from left to right. This may constitute a barrier for upstream migration of fish early stages, a situation of concern for the Halimun National Park fish communities. The objective of the present study was to characterise at the species level the composition of the ichthyoplankton of the Cibareno River at various elevations and its variation during the course of 1 year. The identification of fish larvae is challenging if based on morphological characters, because of a general lack of diagnostic characters (Teletchea 2009; Ko et al. 2013). However, DNA barcoding, the use of the mitochondrial cytochrome oxidase I (COI) gene as a species tag for molecular identification, has been successfully used to identify fish larvae to the species level and characterise ichthyoplankton (Hubert et al. 2010, 2015; Wibowo et al. 2017; Hulley et al. 2018). Molecular techniques offer unprecedented insights on identification of fish larvae in aquatic environments (Chu et al. 2019; Lira et al. 2023).
Map of the study location in the Cibareno River, Sukabumi, West Java, Indonesia, with three fish sampling locations upstream and downstream of Caringin Weir. Upstream 1, 6.8157°S, 106.4361°E; Upstream 2, 6.8409°S, 106.4338°E; Upstream 3, 6.8768°S, 106.4200°E; Weir, 6.8783°S, 106.4200°E; Downstream 1, 6.8808°S, 106.4185°E; Downstream 2, 6.9161°S, 106.3985°E; and Downstream 3, 6.9743°S, 106.3946°E.
The present study makes use of the previously published DNA barcode reference library of the freshwater fishes of Java (Dahruddin et al. 2017) to document the species composition of the ichthyoplankton occurring in the Cibareno River over 1 year, by using DNA barcoding. In this study, DNA barcoding was essential for accurately identifying fish larval species in the Cibareno River because traditional methods based on morphological characteristics can be challenging. We used standardised fishing methods to examine larval community composition along the Cibareno River.
Materials and methods
Study sites
The 37-km-long Cibareno River originates in the Gunung Halimun National Park and flows south between Banten and West Java provinces through a forested and conserved catchment to the Indian Ocean, west of the Pelabuhan Ratu Bay (Rachmatika et al. 2002). The steep-gradient catchment is characterised by rocky substrates, waterfalls and swift water. Caringin Weir, in Sukabumi District, is a single weir that may disrupt fish movement, including that of high-value fish such as Tor tambra (Amanda et al. 2023). During this study, the weir was under construction and its foundation was elevated 1 m (the final height will be 3.25 m) above the riverbed, extending from one bank to the other.
Specimen collection
Larval and adult fish specimens were collected at six different locations along the Cibareno River, three of which were upstream of the location of the Caringin weir project and the other three were downstream (Fig. 1). Upstream and downstream sampling locations were 0, 5 and 10 km from the location of the weir.
The experiment was conducted under animal research and ethic approval for ‘Fish Biodiversity and Environmental Assessment of Cibareno River for Supporting Inland Fisheries and Fish Domestication’ number 60.1/BRSDM-BRPBATPP/LB.200/I/2022 granted by the Research Institute of Freshwater Aquaculture and Fisheries Extension, Ministry of Marine Affairs and Fisheries, Republic of Indonesia.
The following four types of fishing gear were consistently used to collect adult fish for each sampling event: (i) five multi-panel monofilament gillnets (each 5 m wide, with mesh stretched width of 19, 25, 37, 50 and 75 mm), (ii) ten collapsible bait traps (400 × 220 × 220 mm), (iii) ten cast nets (2-m diameter, 25-mm mesh) and (iv) two single wing fyke nets (wing, pairs of 4 m long, 0.2 m in diameter, net body 1.25 m long, 7-mm stretched mesh). All fish caught were photographed, measured (to the nearest millimetre) and weighed (to the nearest 0.1 g). All experimental fishing was conducted between 07:00 and 14:00 hours during 5 days. To capture any potential seasonal variations in fish species and relative abundance, all six sites were sampled three times during the wet season and three times during the dry season (between March and September 2022). Individuals were further identified to the species level by using monographs and field guides (Kottelat et al. 1993; Rachmatika et al. 2002; Keith et al. 2015), and further confirmed by a visual examination of specimen photographs of the Java fish species catalogued in BOLD (Dahruddin et al. 2017; Keith et al. 2017; Hubert et al. 2019).
At the same six sites, preflexion (i.e. from hatching to the onset of notochord flexion) to postflexion (i.e. from formation of the caudal fin to attainment of complete development fin rays and scales) larval fish (Leis and Carson-Ewart 2000) were sampled with a 30-cm diameter modified bongo net (0.3-mm mesh). The net was kept submerged ~5 cm below the surface for two sessions of 15 min, so as to collect replicates at each site. These sampling sessions were conducted early in the morning (from 07:00 to 08:00 hours). Larvae were kept in water after collection, and manually sorted. To reduce animal suffering, fish larvae were euthanised with anesthetic overdose (Eugenol; dose: 4 mL per 1 L of water) at the collection site. The procedure for larval storage and sorting followed Wibowo et al. (2017) with slight modifications. Each fish larva was stored in a 1.5-mL tube containing absolute ethanol. Each tube was labelled and prepared, and anesthetised larvae were photographed alongside the tubes before storing in the tubes. Larval counts were conducted by sorting similar larvae and grouping them together, facilitating the species identification and tallying the number of larvae at each station. Larvae were selected for DNA analysis from each group suspected to belong to distinct species and species composition among larval fish community was examined using multi-dimensional scaling (MDS) analysis with the vegan package (ver. 2.6-8, J. Oksanen et al., see https://cran.r-project.org/package=vegan/) in RStudio (ver. 1.2.1335, Posit Software, PBC, Boston, MA, USA, see https://posit.co/products/open-source/rstudio/). This analysis was applied to log-transformed abundance estimates to visualise variations in the composition occurring through space (sampling locations) and time (seasons).
Extraction, amplification and sequencing of DNA
Total genomic DNA was isolated using a genomic DNA extraction kit (Geneaid Tissue Genomic DNA Mini Kit GT050, Taiwan) from either muscle tissue (for adult fish) or entire larvae, following the protocol provided by the manufacturer (Wibowo et al. 2021). A 652 bp fragment of the mitochondrial cytochrome c oxidase subunit-1 gene (COI) was amplified using the primers Fish-COI-F (5′-TAA TAC GAC TCA CTA TAG GGT TCT CCA CCA ACC ACA ARG AYA TYGG-3) and COI-Fish-R (5′-ATT AAC CCT CAC TAA AGG GCA CCT CAG GGT GTC CGA ARA AYC ARAA-3′) (Ivanova et al. 2007).
The COI fragment was amplified in a 12.5-μL reaction volume that included 4.0 μL of ultrapure water, 0.625 μL of each primer (1 mM), 6.25 μL of 2× QIAGEN Multiplex PCR Master Mix (QIAGEN, Germany), and 1 μL of DNA template (~100 ng L−1). Polymerase chain reaction (PCR) cycling included an initial 15-min DNA polymerase denaturation stage at 95°C, 35 cycles of 30 s at 94°C, 90 s at 55°C, and 30 s at 72°C, and a 5-min extension at 72°C. PCR products were detected on a 1% agarose gel and purified with the ASAP PCR clean-up kit (ArcticZymes). The EZ-Seq service (Macrogen, Singapore) performed the sequencing and all sequences were submitted to GenBank (PP352528–PP 352565).
DNA barcoding and specimen identification
Chromatograms were inspected manually and trimmed to 652 bp, and a multiple-sequence alignment was performed using Geneious Prime (ver. 2024.0.1, see https://www.geneious.com; Kearse et al. 2012). Individual sequences were identified to the species level through basic local alignment search tool (BLAST) searches in NCBI GenBank database (see http://www.nlm.nih.gov/). Sequences were assigned to a known taxon when a match above 98% similarity was found (Hebert et al. 2003; Ross et al. 2008). Additional sequences available in GenBank for the taxa identified were downloaded and jointly analysed with the newly generated sequences. The final multiple sequence alignment was subsequently used to construct a phylogenetic tree for a visual inspection of the genetic divergence among sequences. We reconstructed a Maximum Likelihood (ML) tree using IQ-TREE (ver. 1.6.12, see http://iqtree.cibiv.univie.ac.at; Nguyen et al. 2015; Trifinopoulos et al. 2016) with the most-likely substitution model according to ModelFinder (see http://www.iqtree.org/ModelFinder/; Kalyaanamoorthy et al. 2017) following the Bayesian information criterion (BIC).
Results
Fish identification
In total, 800 fish (including larvae and juveniles) were collected from six locations over the 6-month period encompassing the wet and dry seasons. Among these 800 fish, 9 morphotypes were detected (Fig. 2).
Photographs of specimens representing morphotypes detected in the (a–f) fish larvae and (g–i) juveniles collected in the Cibareno River, West Java, Indonesia.
In total, 80 chromatogram (40 forward and reverse pairs) sequences were obtained from 40 individuals, including adult specimens and representative samples of the larval morphotypes identified. We did not perform molecular testing on the Oreochromis niloticus larval samples because our focus was on native fish species that had not yet been identified. In addition, the O. niloticus larvae can be clearly identified on the basis of their morphological characteristics. Sequencing failed for only 2 individuals and 38 sequences, in total, were produced, with a sequence length of 628–632 base pairs, with 1 to 8 specimens per morphotype. These 38 sequences were subsequently identified to the species level by using the BLAST procedure in GenBank (Table 1). All sequences were unambiguously identified to the species level with the 98% similarity threshold, excepting Ambassis sp., Schismatogobius sp. and Paracentropogon sp.
| Number | Status | Best match | Similarity (%) | Nearest neighbour | Similarity (%) | |
|---|---|---|---|---|---|---|
| 1 | Adult | Ambassis sp. | 99.84 | Ambassis gymnocephalus | 92.57 | |
| 2 | Adult | Ambassis sp. | 99.68 | Ambassis gymnocephalus | 92.40 | |
| 3 | Adult | Anguilla bicolor | 100 | Anguilla obscura | 96.02 | |
| 4 | Adult | Awaous grammepomus | 99.83 | Awaous ocellaris | 99.84 | |
| 5 | Adult | Awaous ocellaris | 99.82 | Awaous grammepomus | 93.16 | |
| 6 | Larvae | Awaous ocellaris | 99.82 | Awaous grammepomus | 93.72 | |
| 7 | Adult | Barbodes binotatus | 99.84 | Barbodes banksi | 99.35 | |
| 8 | Adult | Barbodes binotatus | 100 | Barbodes banksi | 99.35 | |
| 9 | Adult | Barbodes binotatus | 100 | Barbodes banksi | 99.35 | |
| 10 | Adult | Barbodes binotatus | 100 | Barbodes banksi | 99.35 | |
| 11 | Larvae | Barbodes binotatus | 100 | Barbodes banksi | 99.35 | |
| 12 | Adult | Glossogobius giuris | 99.67 | Awaous melanocephalus | 92.94 | |
| 13 | Larvae | Glyptothorax platypogon | 100 | Glyptothorax robustus | 98.53 | |
| 14 | Adult | Lamnostoma kampeni | 100 | Lamnostoma mindora | 97.07 | |
| 15 | Adult | Paracentropogon | NA | Inimicus japonicus | 87.26 | |
| 16 | Larvae | Poecilia reticulata | 100 | Poecilia wingei | 100 | |
| 17 | Larvae | Poecilia reticulata | 100 | Poecilia wingei | 100 | |
| 18 | Larvae | Poecilia reticulata | 100 | Poecilia wingei | 100 | |
| 19 | Larvae | Poecilia reticulata | 100 | Poecilia wingei | 100 | |
| 20 | Larvae | Poecilia reticulata | 100 | Poecilia wingei | 100 | |
| 21 | Larvae | Poecilia reticulata | 100 | Poecilia wingei | 100 | |
| 22 | Larvae | Poecilia reticulata | 100 | Poecilia wingei | 100 | |
| 23 | Larvae | Poecilia reticulata | 100 | Poecilia wingei | 100 | |
| 24 | Adult | Rhyacichthys aspro | 100 | Rhyacichthys gilberti | 95.22 | |
| 25 | Adult | Rhyacichthys aspro | 100 | Rhyacichthys gilberti | 95.22 | |
| 26 | Adult | Schismatogobius sp. A | NA | Schismatogobius insignus | 93.28 | |
| 27 | Adult | Sicyopterus lagocephalus | 99.02 | Sicyopterus marquesensis | 96.18 | |
| 28 | Adult | Sicyopterus microcephalus | 100 | Sicyopterus pugnans | 95.06 | |
| 29 | Adult | Sicyopterus parvei | 99.35 | Sicyopterus cynocephalus | 95.32 | |
| 30 | Adult | Sicyopterus parvei | 99.35 | Sicyopterus cynocephalus | 95.32 | |
| 31 | Larvae | Sicyopus rubicundus | 100 | Sicyopus discordipinnis | 94.67 | |
| 32 | Larvae | Sicyopus rubicundus | 100 | Sicyopus discordipinnis | 94.67 | |
| 33 | Larvae | Sicyopus zosterophorus | 100 | NA | NA | |
| 34 | Adult | Stiphodon semoni | 99.68 | Stiphodon atropurpureus | 96.24 | |
| 35 | Adult | Stiphodon semoni | 99.51 | Stiphodon atropurpureus | 96.24 | |
| 36 | Adult | Stiphodon semoni | 100 | Stiphodon atropurpureus | 96.57 | |
| 37 | Adult | Stiphodon semoni | 100 | Stiphodon atropurpureus | 96.57 | |
| 38 | Larvae | Tor tambra | 99.84 | Tor tambroides | 99.68 |
BLAST analyses resulted in the detection of 19 species (Table 1), of which 8 were detected among sampled larvae, indicating that several morphotypes identified among larvae correspond to different stages of the same species (Fig. 2). These correspond to Awaous ocellaris, Sicyopus zosterophorus, Sicyopus rubicundus, Poecilia reticulata, Glyptothorax platypogon, Tor sp. and Barbodes binotatus. The ML tree reconstructed with the TIM2 + F + I + Γ model, the most likely model according to modeltest, indicated that species represented by multiple sequences represent clusters of closely related sequences (Fig. 3). At the larval stage, morphology varies widely in some species (e.g. Poecilia reticulata), making species classification difficult by using morphological characters only. The greatest number of larvae found was for P. reticulata (eight individuals).
Maximum Likelihood (ML) tree of the 38 newly generated sequences from this study, corresponding to 19 species. Sequences were generated either with adult or larval specimens.
During the sampling period, Poecilia reticulata, Barbodes binotatus and Sicyopus rubicundus are the three species that were the most commonly captured in the sampling sites (Fig. 4).
Larvae of Poecilia reticulata dominated all larval assemblages because they were captured at a higher frequency than were the other species in both downstream and upstream locations (Fig. 5). Two more fish species, namely, Sicyopus rubicundus and Awaous ocellaris, were consistently observed in downstream areas during each sampling period.
Species composition of larvae sampled from the Cibareno River, West Java, Indonesia, by site and month.
The NMDS graph shows two clusters separating upstream and downstream locations with a value of P = 0.016 (Fig. 6). The presence of larvae of Poecilia reticulata characterised the upstream sites in April and May. Barbodes binotatus larvae were characteristic of the upstream sites in July. The presence of Tor sp. and Sicyopus zosterophorus larvae in July, Glypothorax platypogon larvae in May and Awaous ocellaris larvae in March characterised the downstream area.
Discussion
Construction of riverine infrastructure is increasing throughout Indonesia, which is typical of South-east Asia, with the immediate consequence of isolating migratory fish communities (Haryani 2021; Atminarso et al. 2024). Our case study on the Cibareno River also highlighted a strong potential for restriction of diadromous fish to the river reaches downstream of Caringin Weir and upstream isolation of a potamodromous and riverine fish community.
To understand how Cibareno River fish communities responded to construction of the Caringin Weir, we first needed to validate a technique to identify fish larvae. DNA barcoding enabled us to reliably identify eight species of fish larvae in the Cibareno River, two of which (Sicyopus rubicundus and Barbodes binotatus) had not previously been reported upstream (Rachmatika et al. 2002). Additionally, we also detected the genus Tor among the larvae sampled, the species of which are high-value species only recently documented in the Cibareno catchment (Amanda et al. 2023). For rivers where there are few historical survey data, DNA barcoding is emerging as a sound technique for characterising biodiversity and for providing a baseline library for future monitoring (Hubert et al. 2015; Wibowo et al. 2017).
The central position of the Caringin Weir in the Cibareno watershed is likely to be influential for interpreting patterns of larval diversity and abundance observed among upstream and downstream sites. Elsewhere, following weir construction, there is invariably a gross simplification of upstream free-flowing lotic habitats that become deeper, slow-flowing lentic habitats (Mueller et al. 2011; Jo et al. 2019). In these modified habitats, there are usually detrimental changes to food-webs, including phytoplankton, zooplankton and ultimately native fishes (Park et al. 2023). For the Cibareno River, evaluating a broad suite of environmental variables was beyond the scope of our study but certainly we observed a greater larval species diversity downstream of Caringin Weir where free-flowing habitat remained. This river stretch was characterised by features such as pools, aquatic vegetation, woody habitat, as well as riffle and run zones, where such a broad range of habitats can support a range of fish species (Guo et al. 2017).
In tropical rivers that experience wet and dry seasons, temporal patterns of abundance in fish larvae are usually observed (Castro et al. 2005). However, within the Cibareno River, we failed to observe variation in the larval fish assemblages between March and September, which encompassed both the dry and wet seasons. The wet season is often a time of fish reproduction, when floodplain habitats become accessible and there is greater food availability for fish larvae (i.e. the flood-pulse concept) (Junk et al. 1989). Changes in species diversity and abundance of fish larvae are therefore often related to taking advantage of flow events to access areas of improved water quality, food availability, spawning habitats and for reducing predation (Chhuoy et al. 2022). However, the Cibareno River is a high-gradient system with few adjacent floodplains; hence, the more general spawning strategies of local species appears to involve adaptions to local conditions (Godfrey et al. 2017).
In efforts to biologically control malaria carrying mosquitoes, Trinidadian guppies (P. reticulata) have been introduced to many aquatic habitats in South-east Asia (Yadav and Haq 2017). Where invasive guppies become established in new aquatic habitats with few resident species, they usually induce important perturbations of local biodiversity because of their predatory nature (Morgan et al. 2004; El-Sabaawi et al. 2016; R. Froese and D. Pauly, FishBase, ver. 04/2019, see www.fishbase.org). For the Cibareno River, habitat fragmentation and hydrological habitat simplification following construction of Caringin Weir would also appear as another trigger for exacerbating guppy abundance and impacts on resident species (Alexander et al. 2015; Suryaningsih et al. 2020). In other parts of Indonesia, guppies have outcompeted local species because of faster sexual maturity, reproduction rates and population growth (Reznick et al. 2001; Herder et al. 2022).
In conclusion, we have provided a baseline fish assemblage, from which the impacts of Caringin Weir can be monitored using DNA barcoding for future monitoring, along with other field techniques. We anticipate changes in fish communities at upstream and downstream sites, which may come to light as the full impact of the new weir manifests, despite the presence of a fishway, because upstream habitats will change from free-flowing fast-water to ponded deep-water habitats. In steep, fast-flowing catchments, the water dynamics create a constant connection along the river, making larval fish communities especially sensitive to changes in their environment. We aim for our baseline data to guide infrastructure projects in South-east Asia and support the conservation of tropical river fish populations.
Data availability
Data used in this paper are organised by the first author. Access to these data can be negotiated with him.
Declaration of funding
The project funding of the study was supported by the Food and Agriculture Organization and Global Environment Facility project ‘Inland Fisheries Practices in Freshwater Ecosystems of High Conservation Value (IFish)’ (FAO project code: GCP/INS/303/GFF, GEF ID: 5759, to FAO Indonesia with salary funding for Imron Rosadi, Yohanes Yudha P. Jaya and Sudarsono Sudarsono) and the Australian Centre for International Agricultural Research (project number FIS/2018/153, with salary and travel support for Lee Baumgartner and Arif Wibowo).
Acknowledgement
The authors are grateful to the FAO Indonesia (IFish Project Team) and the Australian Centre for International Agricultural Research (ACIAR) teams for this wonderful opportunity and collaboration. The authors gratefully acknowledge Research Center for Fisheries, the Ministry of Marine Affairs and Fisheries, Republic of Indonesia, the Government Office of Fisheries, Government office of Water Resource Management, West Java, and UPTD Water Resource Management of Sukabumi, West Java. We also extend our gratitude to Boby Bagja Pratama for refining the map of our study. Finally, we thank Fery, Daman, and undergraduate students of Aquatic Resource Management, IPB University, who significantly contributed to the fieldwork.
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