Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes

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While Betta splendens is well studied due to its importance in the ornamental fish trade, most wild Betta species remain poorly characterized, particularly at the genomic level. The Bangka Islands of Indonesia harbor several endemic Betta species threatened by peatland degradation. Methods We conducted an integrated meristic and genomic comparison of three endemic Bangka Island species—Betta burdigala, B. chloropharynx, and B. schalleri. Specimens were collected from peatland waters in Bangka, and meristic traits were examined to confirm diagnostic characteristics. High-molecular-weight DNA was extracted and sequenced using Oxford Nanopore PromethION technology, followed by de novo assembly and reference-guided scaffolding using the Betta splendens genome. Results The meristic analysis confirmed features consistent with their taxonomic placement within the coccina, waseri, and pugnax groups. Genome assembljies were highly contiguous and complete (BUSCO >97%), with B. chloropharynx showing the largest genome size, highest scaffold N50, and elevated retrotransposon content. Gene duplication analysis revealed dispersed duplications as the dominant category across all genomes, with variation in tandem and proximal duplicates. Comparative genomic analysis demonstrated high collinearity, with B. chloropharynx and B. schalleri showing the closest relationship, while B. burdigala diverged earlier. The Colony Stimulating Factor 1 Receptor A (CSF1RA) protein phylogenetic tree closely resembles the phylogenetic tree of nine fish species based on NCBI taxonomic data. We also identified two massive protein insertions in the CSF1RA of B. burdigala. Conclusions This study provides morphological and genomic evidence supporting the distinctiveness of Bangka’s endemic Betta species and delivers essential genomic resources for evolutionary research and conservation of these endangered freshwater fishes. " } { "@context": "http://schema.org", "@type": "BreadcrumbList", "itemListElement": [ { "@type": "ListItem", "position": "1", "item": { "@id": "https://f1000research.com/", "name": "Home" } }, { "@type": "ListItem", "position": "2", "item": { "@id": "https://f1000research.com/browse/articles", "name": "Browse" } }, { "@type": "ListItem", "position": "3", "item": { "@id": "https://f1000research.com/articles/15-226", "name": "Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic..." } } ] } Home Browse Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic... ALL Metrics - Views Downloads Get PDF Get XML Cite How to cite this article Helmizuryani H, Hidayat S, Nizar M et al. Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.12688/f1000research.174779.1 ) NOTE: If applicable, it is important to ensure the information in square brackets after the title is included in all citations of this article. Close Copy Citation Details Export Export Citation Sciwheel EndNote Ref. Manager Bibtex ProCite Sente EXPORT Select a format first Track Share ▬ ✚ Research Article Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] Helmizuryani Helmizuryani 1 , Saleh Hidayat 2 , Muhammad Nizar 1 , [...] Andhika Prima Prasetyo https://orcid.org/0000-0002-7412-8432 3 , Boby Muslimin https://orcid.org/0000-0002-0662-9972 3 , Mochammad Zamroni https://orcid.org/0000-0001-8029-8764 4 , Dessy Nurul Astuti 4 , Swarlanda Swarlanda 5 , Destra Ramadhanu 5 , Luthfi Nurhidayat https://orcid.org/0000-0002-0625-2984 6 Helmizuryani Helmizuryani 1 , Saleh Hidayat 2 , [...] Muhammad Nizar 1 , Andhika Prima Prasetyo https://orcid.org/0000-0002-7412-8432 3 , Boby Muslimin https://orcid.org/0000-0002-0662-9972 3 , Mochammad Zamroni https://orcid.org/0000-0001-8029-8764 4 , Dessy Nurul Astuti 4 , Swarlanda Swarlanda 5 , Destra Ramadhanu 5 , Luthfi Nurhidayat https://orcid.org/0000-0002-0625-2984 6 PUBLISHED 10 Feb 2026 Author details Author details 1 Study Program of Aquaculture, Faculty of Agriculture, Universitas Muhammadiyah Palembang, Palembang, South Sumatra, 30263, Indonesia 2 Study Program of Biology Education, Faculty of Teacher Training and Education, Universitas Muhammadiyah Palembang, Palembang, South Sumatra, 30263, Indonesia 3 Research Center for Applied Zoology, National Research and Innovation Agency, Cibinong, West Java, 16911, Indonesia 4 Research Center for Biota System, National Research and Innovation Agency, Cibinong, West Java, 16911, Indonesia 5 Yayasan Ikan Endemik Bangka Belitung, Pangkal Pinang, Bangka Belitung Island, 33123, Indonesia 6 Faculty of Biology, Universitas Gadjah Mada, Sleman, Special Region of Yogyakarta, 55281, Indonesia Helmizuryani Helmizuryani Roles: Conceptualization, Data Curation, Investigation, Methodology, Writing – Original Draft Preparation, Writing – Review & Editing Saleh Hidayat Roles: Data Curation, Investigation, Methodology, Validation Muhammad Nizar Roles: Data Curation, Investigation, Methodology, Validation Andhika Prima Prasetyo Roles: Investigation, Validation Boby Muslimin Roles: Data Curation, Formal Analysis, Investigation, Validation, Writing – Original Draft Preparation, Writing – Review & Editing Mochammad Zamroni Roles: Data Curation, Investigation, Methodology, Validation Dessy Nurul Astuti Roles: Data Curation, Investigation, Validation Swarlanda Swarlanda Roles: Data Curation, Investigation, Methodology, Validation Destra Ramadhanu Roles: Data Curation, Investigation, Methodology, Validation Luthfi Nurhidayat Roles: Data Curation, Formal Analysis, Software, Writing – Original Draft Preparation, Writing – Review & Editing OPEN PEER REVIEW DETAILS REVIEWER STATUS This article is included in the Genomics and Genetics gateway. Abstract Background The genus Betta (family Osphronemidae) comprises over 70 species, many of which are endemic to Southeast Asia and highly vulnerable to habitat loss. While Betta splendens is well studied due to its importance in the ornamental fish trade, most wild Betta species remain poorly characterized, particularly at the genomic level. The Bangka Islands of Indonesia harbor several endemic Betta species threatened by peatland degradation. Methods We conducted an integrated meristic and genomic comparison of three endemic Bangka Island species— Betta burdigala , B. chloropharynx , and B. schalleri. Specimens were collected from peatland waters in Bangka, and meristic traits were examined to confirm diagnostic characteristics. High-molecular-weight DNA was extracted and sequenced using Oxford Nanopore PromethION technology, followed by de novo assembly and reference-guided scaffolding using the Betta splendens genome. Results The meristic analysis confirmed features consistent with their taxonomic placement within the coccina , waseri , and pugnax groups. Genome assembljies were highly contiguous and complete (BUSCO >97%), with B. chloropharynx showing the largest genome size, highest scaffold N50, and elevated retrotransposon content. Gene duplication analysis revealed dispersed duplications as the dominant category across all genomes, with variation in tandem and proximal duplicates. Comparative genomic analysis demonstrated high collinearity, with B. chloropharynx and B. schalleri showing the closest relationship, while B. burdigala diverged earlier. The Colony Stimulating Factor 1 Receptor A (CSF1RA) protein phylogenetic tree closely resembles the phylogenetic tree of nine fish species based on NCBI taxonomic data. We also identified two massive protein insertions in the CSF1RA of B. burdigala. Conclusions This study provides morphological and genomic evidence supporting the distinctiveness of Bangka’s endemic Betta species and delivers essential genomic resources for evolutionary research and conservation of these endangered freshwater fishes. READ ALL READ LESS Keywords Betta burdigala, Betta chloropharynx, Betta schalleri, meristic analysis, genome assembly, synteny, conservation genomics, Bangka Island Corresponding Author(s) Luthfi Nurhidayat ( [email protected] ) Close Corresponding author: Luthfi Nurhidayat Competing interests: No competing interests were disclosed. Grant information: 123/C3/DT.05.00/PL/2025 The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Copyright: © 2026 Helmizuryani H et al . This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. How to cite: Helmizuryani H, Hidayat S, Nizar M et al. Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.12688/f1000research.174779.1 ) First published: 10 Feb 2026, 15 :226 ( https://doi.org/10.12688/f1000research.174779.1 ) Latest published: 10 Feb 2026, 15 :226 ( https://doi.org/10.12688/f1000research.174779.1 ) Introduction The genus Betta (family Osphronemidae) includes over 70 recognized species, many of which are endemic to Southeast Asia. Despite the global popularity of the domesticated Betta splendens in the ornamental fish trade, the majority of wild Betta species remain under-researched. The number of endemic wild Betta distribution and diversity of Southeast Asia has been identified for 16 years in Thailand (2 species) and Indonesia (9 species). 1 – 5 The high levels of endemism and restricted ranges of many Betta species make them particularly vulnerable to habitat loss and environmental changes. 2 , 5 The Bangka Islands, located off the east coast of Sumatra, are home to several unique Betta species adapted to specialized freshwater habitats such as peat swamps and slow-moving forest streams. Three species of Betta spp. were endemic, i.e., Betta burdigala , Betta chloropharynx , and Betta schalleri. 1 According to IUCN red list, the conservation status for them were criticaly endangered and endangered. The lack of molecular resources poses a barrier to implementing effective conservation strategies for these fishes. The genetic studies of the endemic fish Betta spp. on Bangka Island have been conducted in mitochondrial DNA analysis, eDNA metabarcoding, and comprehensive morphological examinations, which are needed to confirm the genetic relationships and taxonomic status of these species. However, the whole genome for the evolution of the endemic fish Betta spp. on Bangka Island hasn’t been revealed. 6 Genome assemblies are indeed foundational resources for advancing biological research and conservation management. They provide critical insights into genetic diversity, evolutionary biology, and species conservation strategies. High-quality genomic data could be used for Long-read sequencing and high-throughput chromosome (Hi-C) technology. Long-read sequencing technologies are essential for producing high-quality genome assemblies. These technologies help resolve complex repeats and haplotype heterozygosity, which are sources of assembly errors. 7 Hi-C confirmation capture that enables contiguous genome assemblies. 8 Genome assemblies could be informative evidence for scientific management decisions and tools for understanding the basis of genetic adaptation in various species. 9 , 10 Bangka Island presents a unique opportunity to investigate the genetic foundations of adaptation and diversity in insular freshwater fish species. The endemic Betta species of the island have likely undergone distinct evolutionary trajectories due to the island’s unique environmental conditions and geographic isolation. Producing genome assemblies for these taxa will address a significant knowledge gap and serve as a foundational step for future conservation genomics efforts in the region. This study aims to generate preliminary genome assemblies for three endemic Betta species originating from Bangka, Indonesia, utilizing whole-genome sequencing methodologies. Materials and methods Ethics statement This study received ethical approval from the Ethics Commission for Animal Care and Use, National Research and Innovation Agency Republic of Indonesia (Approval No. 220/KE.02/SK/09/2024). All animal procedures followed institutional regulations and adhered to the ARRIVE 2.0 reporting guidelines, with the corresponding checklists available at https://doi.org/10.6084/m9.figshare.31136455 . 11 Sample collection This study examines three endemic species of Betta: Betta burdigala , Betta chloropharynx , and Betta schalleri. The fish samples were collected from peatland water in South Bangka, Bangka Island, Indonesia. The map showing the origin locations of specimens on Bangka Island is presented in Figure 1 . The males of B. schalleri and B. chlorpharynx have been found in Central Bangka (2°22'00.7"S 106°10'57.9"E) and B. burdigala has been found in South Bangka (2°50'24.8"S 106°26'12.7"E). The specimens were collected using a scope net with a mesh size of 0.5 mm as active gear. In-situ sampling data for morphometric and meristic analysis were obtained from three individuals each of B. schalerii , B. chloropharynx , and B. burdigala. The ten morphometric and seven meristic characters (Suppl Fig 1) based on Nur et al. (2022) 2 were measured using a digital caliper with an accurary 0.1 mm. The sampling design in this study used purposive sampling based on mature females or males with different shapes and colors. Males have longer ventral fins than females. 12 The males also have more vibrant and diverse body colors than females. 13 The morphometric and meristic data were analyzed descriptively. Figure 1. The specimen origin locations of Betta schalleri and Betta chloropharynx are found in ST.1 (Central Bangka), while Betta burdigala is found in ST.2 (South Bangka). One male specimen from each Betta species was euthanized using the rapid chilling method, which involved transferring the fish to water maintained at approximately 2°C with crushed ice. 14 The fish was left in this condition for 20 minutes until opercular movement ceased. Subsequently, tissue samples were collected, fixed with absolute ethanol, and stored in cryotubes for whole genome sequencing at the Central Sequencing Laboratory, BRIN, Bogor, Indonesia. Genomic DNA extraction and sequencing The extraction of genomic DNA was performed using the Applied Biosystem MagMax TM DNA Multi-Sample 2.0 kit (Thermo Fisher Scientific; CAT. A36570) following the manufacturer’s instructions. Twenty mg of fish tissue is used for genome DNA extraction material for all Betta specimens. The genome concentration from the extraction was checked using Nanodrop and Qubit. Genome concentration by Nanodrop value between 227.45-289 ng/μl and Qubit value between 89.4-91.6 (ng/μl). The genome quality was checked using gel agarose with TBE agarose concentration 1%. The genomic DNA was subsequently prepared for library construction using the Ligation Sequencing DNA V14 kit (Oxford Nanopore Technology; SQK-LSK114) in accordance with the manufacturer’s instructions. Following this, the library was sequenced on a PromethION device utilizing PromethION Flow Cells Packs (Oxford Nanopore Technology; FLO-PRO114M). The sequencing parameters we employed included a run duration of 96 hours, a pre-scan interval of 1.5 hours, basecalling using the High-accuracy model at 400 bps, and a minimum Q score of 9. The sequencing software utilizes MinKNOW (25.03.7), Bream (8.4.4), configuration (6.4.10), basecalling Dorado (dna_r10.4.1_e8.2_400bps_ [email protected] ), and MinKNOW Core (6.4.8). The genome sequence data for Betta burdigala includes 16.38 million reads and 37.39 gigabases. For Betta chloropharynx , the data comprises 14.97 million reads and 57.97 gigabases. Meanwhile, Betta schalleri has 11.83 million reads and 23.75 gigabases. Genome assembly and annotation The genome assembly and annotation described in this paper were conducted in accordance with the methodology performed by Imron et al. 15 Genome assembly estimation was done using Flye 2.9.5, 16 while genome scaffolding was conducted using RagTag 2.1.0 17 guided by genome reference of Betta splendens (GCF_900634795.4). Genome size estimation was conducted using Jellyfish software version 2.3.1 18 and was further processed using GenomeScope 2.0 v2.0.1. The assembly statistics were calculated using assembly stat version 1.0.1. The completeness of the assembly was estimated using Benchmarking Universal Single-Copy Orthologous (BUSCO) version 5.8.2, utilizing miniprot. 19 – 21 Repetitive elements within the genome assembly were identified using RepeatModeler v2.0.6 in conjunction with RepeatMasker v4.1.7. ( http://www.repeatmasker.org ). Prior to annotation, these repetitive regions were soft masked to minimize interference. Structural genome annotation encompassing gene prediction was conducted using the GALBA pipeline, 22 which employs miniprot 21 and AUGUSTUS, 23 integrating protein data from closely related species as extrinsic evidence. Specifically, protein data from Betta splendens (GCF_900634795.4), Anabas testudineus (GCF_900324465.2), and Channa argus (GCF_033026475.1) were utilized. Functional annotation of the resulting gene predictions was then performed using the ‘funannotate annotate’ command from the Funannotate pipeline ( https://funannotate.readthedocs.io/en/latest/install.html ), incorporating tools such as InterProScan5, 24 Eggnog-Mapper, 25 and SignalP 5.0 26 to assign gene names and predict protein functions. Finally, the completeness of the genome annotation was evaluated using BUSCO v5.8.2. 20 Duplicate genes classification and comparative genomics The protein sequences in each genomes were aligned to eachother using BLAST+ v. 2.14.1 ( https://blast.ncbi.nlm.nih.gov/doc/blast-help/downloadblastdata.html ). The alignment results and the gff (general feature format) file containing gene position in the genome were then further analyzed using the ‘Duplicate_gene_classifier’ function of MCScanX 27 to identify gene duplications. We then performed all-against-all alignment of coding sequences (CDS) of three Betta genomes: using BLAST+ v. 2.14.1 ( https://blast.ncbi.nlm.nih.gov/doc/blast-help/downloadblastdata.html ). The alignment results were then concatenated into one file. The gff files were also concatenated. These files were then further analyzed using MCScanX 27 to detect gene synteny and collinearity. The results were then explored and visualize using SynVisio. 28 We also performed comparative analysis of Colony Stimulating Factor 1 Receptor A (CSF1RA), Melanocortin 1 Receptor (MC1R) and Paired Box 7 (PAX7) protein sequence among B. burdigala , B. chloropharynx , and B. schalleri , alongside B. splendens, Anabas testudineus, Channa argus, Oreochromis niloticus, Danio rerio, and Carassius auratus. The protein accession of CSF1RA that we used are: XP026135817.1, XP026135818.1, NP571747.1, XP021336731.1, XP003455234.1, XP013133007.1, XP067373776.1, XP067373775.1, XP026213579.1, XP055368253.1, XP055368253.1, and XP029020690.1. The protein accession of MC1R that we used are: NP_851301.1, XP_005159236.1, XP_026112973.1, XP_026112974.1, XP_005467175.1, XP_055363428.1, XP_067343026.1, XP_067343026.1, XP_026234205.1. The protein accession of PAX7 that we used are: XP_025763203.1, XP_005459058.1, XP_067361857.1, XP_029005486.1, XP_029005485.1, XP_026204402.1, XP_026204405.1, XP_025763202.1, XP_067361858.1, XP_026204404.1, XP_029005487.1, XP_026204403.1, NP_571407.2, NP_571400.1, XP_009304561.1, XP_026130952.1, XP_026130953.1. The protein sequence alignments were performed using Clustal Omega 29 via the European Bioinformatics Institute website ( https://www.ebi.ac.uk/jdispatcher/msa/clustalo ). Subsequently, these alignments were used for phylogenetic reconstruction with IQ-TREE, 30 and the results were visualized using TreeViewer. 31 Results Fish morphology The morphology photo of Betta spp. is presented in this research as shown in Figure 2 for B. burdigala (A), B. chloropharynx (B), and B. schalleri (C). Table 1 outlines the meristic characteristics that distinguish them. The meristic and morphometric characters are presented in Suppl Fig 1. Figure 2. The morphology of B. burdigala (A), B. chloropharynx (B) and B. schalleri (C). Table 1. Meristic and morphometric character of B. burdigala, B. chloropharynx and and B. schalleri. The average of morphometric characters as a percentage to standard length with standard deviation. B. burdigala B. chloropharynx B. schalleri Meristic character Dorsal fin radii D.I. 13–14 D.I.8-9 D.I.8-9 Anal fin radii A.I–II. 23–24 A.I-II. 27-28 A.I-II.22-23 Caudal radii C.XV C.XV - Standard length (cm) 2.46 ± 0.13 9.95 ± 0.91 5.21 ± 1.23 Morphometric character Post-orbital length (%) 34.71 ± 8.17 47.60 ± 1.06 45.52 ± 0.74 Dorsal fin length (%) 30.43 ± 1.68 14.00 ± 0.29 15.01 ± 0.95 Predorsal length (%) 51.96 ± 2.20 65.17 ± 0.81 63.01 ± 1.26 Postdorsal length (%) 17.33 ± 3.72 22.45 ± 1.69 21.16 ± 1.28 Orbital diameter (%) 6.52 ± 1.81 9.63 ± 0.51 9.24 ± 0.75 Body depth (%) 20.48 ± 1.28 25.78 ± 1.68 27.59 ± 0.25 Head length (%) 29.04 ± 0.60 33.97 ± 1.40 34.90 ± 1.90 Anal fin length (%) 46.15 ± 5.16 49.30 ± 0.17 47.73 ± 1.91 Preanal length (%) 47.73 ± 3.99 47.45 ± 1.16 47.91 ± 0.53 Betta burdigala Betta burdigala is classified within the Betta coccina group. 32 This species is characterized by its red coloration and a size range of 2-3 cm, with distinct morphological features illustrated in Figure 2A that differentiate it from other species within the Betta coccina group. The meristic characteristics of B. burdigala are comprehensively presented in Table 1 . In comparison to B. uberis , 33 B. burdigala exhibits fewer dorsal fin rays (14-15 vs. 14-17), subdorsal scales (11-11.5 vs. 12-13.5), predorsal scales (15-16 vs. 18-20), and postdorsal scales (8 vs. 9-11). Regarding morphometric characteristics, B. burdigala demonstrates a longer postdorsal length as a percentage of standard length (12.19-20.85% vs. 13.5-17.8%), a shorter dorsal fin base (28.12-32.03% vs. 30.0-37.2%), and a greater postorbital length (23.20-41.36% vs. 13.4-16.8%). Betta chloropharynx Betta chloropharynx , depicted in Figure 2B , is a member of the Betta waseri group. 34 This group is distinguished by a unique pattern on the ventral side of the head. Betta chloropharynx can be differentiated from B. hipposideros by several characteristics: it possesses ω-shaped black throat markings as opposed to horseshoe-shaped ones, lacks transverse lines on the caudal fin, and has fewer subdorsal scales, typically 5-6 compared to 6.5 ( Table 1 ). In comparison to B. renata , B. chloropharynx exhibits ω-shaped black throat marks instead of kidney-shaped ones, lacks transverse lines on the dorsal and caudal regions, has an unspotted operculum rather than a speckled one, and features a yellow underside on the operculum instead of a black edge, with fewer subdorsal scales, mode 5-6 versus 6.5. The distinctions between B. chloropharynx and B. spilotogena include ω-shaped black throat marks versus a central black spot, an unspotted operculum versus a spotted one, and an operculum with a yellow underside versus a posterior edge. 35 Betta schalleri In Figure 2C , Betta schalleri , when compared to other B. pugnax , 35 demonstrates distinct meristic characteristics. It possesses a greater number of anal fin rays (27 vs. 23-25), a higher count of dorsal fin rays than B. cracens and B. fusca (10-11 vs. 8-9), more subdorsal scales than B. cracens and B. fusca (6.5-7 vs. 5.5-6), a greater number of lateral scales than B. fusca (31 vs. 29), fewer lateral scales than B. cracens (31 vs. 32-33), and a reduced number of predorsal scales compared to B. cracens, B. fusca, and B. raja (17-19 vs. 20-24). In term of morphometric characteristics (expressed as a percentage of standard length; Table 1 ), B. schalleri exhibits a longer head length compared to B. pugnax , B. cracens, and B. fusca (35.5-36.5% vs. 27.5-35.2%), a shorter predorsal length compared to B. fusca (62.7-66.3% vs. 68.5-70.2%), a longer preanal length compared to B. cracens (47.16-48.30 % vs. 42.0-46.1%), a greater body depth compared to B. cracens (27.38-27.93 % vs. 21.2-24.2%), an extended dorsal fin base compared to B. cracens (14.11-16.32% vs. 10.5-11.6%), and a shorter anal fin base compared to B. cracens (45.49-50.16 % vs. 53.4-55.7%). Genome assembly The genome assemblies of B. burdigala , B. schalleri , and B. chloropharynx demonstrated high contiguity and completeness, with B. chloropharynx showing the strongest contiguity, while B. burdigala and B. schalleri also provide high-quality genomic resources suitable for comparative and evolutionary studies ( Figure 3 ). The genome assembly of B. burdigala totaled ~422 Mbp across 4,868 scaffolds and 6,904 contigs, with a scaffold N50 of 18 Mb and the longest scaffold reaching 33.2 Mb. The assembly was highly complete, with a BUSCO score of 99.1% (98.6% single-copy, 0.5% duplicated), while the GC content was 45.2% and gaps accounted for only 0.048%. The B. chloropharynx genome was slightly larger at ~474 Mbp, assembled into 3,273 scaffolds and 4,918 contigs. It showed the greatest contiguity, with a scaffold N50 of 20 Mbp, a maximum scaffold length of 35.9 Mbp, and the highest mean scaffold length (~145 kb). BUSCO completeness reached 99.3% (98.7% single-copy, 0.6% duplicated), the highest among the three species, with minimal fragmentation (0.5%) and missing genes (0.2%). The Betta schalleri assembly spanned ~433 Mbp with 8,474 scaffolds and 10,762 contigs, reflecting greater fragmentation. The scaffold N50 was 19 Mb, with the longest scaffold at 34.7 Mbp and a mean length of ~51 kb. BUSCO assessment indicated 99.1% completeness (98.8% single-copy, 0.4% duplicated), with GC content at 44.8% and low gap content (0.053%). Figure 3. Main features of the genome assemblies of Betta burdigala , Betta chloropharynx, and Betta schalleri. Genome annotation Repeats annotation Annotation of repetitive elements in the genomes of B. burdigala , B. chloropharynx , and B. schalleri revealed that transposable elements (TEs) comprise a substantial portion of each genome, with notable variation in both content and composition among species ( Table 2 ). Retroelements were a dominant repeat category, accounting for 11.3% in BBS, 11.4% in B. burdigala , and up to 15.9% in B. chloropharynx. Among these, LINEs were the most prevalent, particularly the L2/CR1/Rex and L1/CIN4 subfamilies, followed by LTR elements such as Gypsy/DIRS1 and BEL/Pao, which collectively contributed to the structural and regulatory diversification of the genome. Although SINEs and Penelope elements were detected, they represented only a minor fraction of the total retroelement content (<0.3%). Table 2. Classification of repeat elements of B. burdigala , B. chloropharynx , and B. schalleri genome assemblies. Repeat category Percentage of sequence (%) Betta burdigala Betta chloropharynx Betta schalleri Retroelements 5.98 8.39 6.18 SINEs 0.21 0.42 0.39 Penelope 0.03 0.01 0.32 LINEs 4.03 5.24 3.53 CRE/SLACS 0 0 0 L2/CR1/Rex 2.53 3.31 2.22 R1/LOA/Jockey 0.17 0.24 0.11 R2/R4/NeSL 0.25 0.37 0.36 RTE/Bov-B 0.41 0.8 0.54 L1/CIN4 0.58 0.35 0.16 BEL/Pao 0.11 0.25 0.21 Ty1/Copia 0.01 0.02 0.03 Gypsy/DIRS1 0.97 1.53 1.14 Retroviral 0.43 0.65 0.32 DNA transposons 3.85 5.54 4.02 hobo-Activator 1.18 1.86 1.5 Tc1-IS630-Pogo 0.68 1.65 1.08 En-Spm 0 0 0 MULE-MuDR 0 0 0 PiggyBac 0.13 0.2 0.1 Tourist/Harbinger 0.62 0.53 0.32 Rolling-circles 0.04 0.19 0.03 Unclassified 4.21 5.93 3.94 Total Interspersed 14.03 19.86 14.14 Small RNA 0.12 0.34 0.25 Satellites 0.06 0.07 0.05 Simple repeats 2.14 1.59 1.82 Low complexity 0.23 0.21 0.22 Total Masked Bases 16.54 22.08 16.30 DNA transposons were the second most abundant repeat class, comprising 6.5%, 9.8%, and 7.0% of the genomes of B. burdigala , B. chloropharynx , and B. schalleri , respectively ( Table 2 ). The dominant families within this group included hobo-Activator, Tc1-IS630-Pogo, and En-Spm elements. These transposons are known for their cut-and-paste mechanism, and their relative abundance suggests that DNA-mediated transposition has played an important role in shaping genome architecture, particularly in BBC where their proportion was the highest. Interestingly, MULE-MuDR and PiggyBac elements were detected at low levels or were absent, suggesting potential lineage-specific losses or underrepresentation due to assembly or annotation limitations. In addition to classified repeat categories, a significant fraction of the genome (3.9–5.9%) consisted of unclassified elements, indicating the presence of either novel transposable elements or highly diverged copies that defy current classification models ( Table 2 ). Moreover, 22–30% of repetitive content across the three genomes fell into the “Other” category, representing repeat families that could not be assigned confidently to standard classifications. The higher repeat content observed in B. chloropharynx , particularly in retrotransposons and unclassified repeats, may reflect recent TE amplification events or reduced efficacy of TE suppression mechanisms in this lineage. Structural and functional annotation and gene duplication analysis The genome annotation summary across the three Betta species— B. burdigala , B. chloropharynx , and B. schalleri —revealed a broadly similar genomic architecture, with minor differences in gene structure metrics ( Table 3 ). All three species exhibit approximately 1.3 transcripts per gene on average, and the number of predicted transcripts ranged from 31,517 in B. burdigala to 32,786 in B. chloropharynx. The mean exon size was consistent (172–174 bp), while the mean gene locus size ranged from 8,784 bp in B. burdigala to 9,013 bp in B. chloropharynx. Notably, the percentage of genes with alternative transcript variants was relatively stable across species (~23%), and most genes were multi-exonic (over 91%), indicating conserved splicing complexity in the genus. Table 3. Genome annotation summary for the genome assembly of Betta burdigala , Betta chloropharynx , and Betta schalleri. Statistic Betta burdigala (BBB) Betta chloropharynx (BBC) Betta schalleri (BBS) Max transcripts/gene 12 11 11 Mean exon size (bp) 173.2 174.1 172.7 Mean gene locus size (bp) 8784.6 9013.1 8942.5 Mean distinct exons/gene 10.4 10.2 10.4 Mean transcripts/gene 1.3 1.3 1.3 Mean transcript size (bp) 2013.8 1995.4 2016.1 Number of distinct exons 245812 251941 247092 Number of genes 23645 24597 23767 Genes with alt. transcripts 5,440 (23.0%) 5,615 (22.8%) 5,479 (23.1%) Multi-exon genes 21,835 (92.3%) 22,592 (91.8%) 21,922 (92.2%) Predicted transcripts 31517 32786 31694 Single-exon genes 1,810 (7.7%) 2,005 (8.2%) 1,845 (7.8%) Regarding genome composition, the GC content of genes and introns was consistent across species at 46% and 44%, respectively ( Figure 4A ). Exonic GC content was slightly higher, ranging from 54% in B. chloropharynx and B. schalleri to 55% in B. burdigala ( Figure 4B ). Exons accounted for 9–10% of the genome, while genes (including exons and introns) occupied approximately 47–49% ( Figure 4B ). Introns alone contributed 38–40% of the genome, reflecting their substantial role in genome size and structure ( Figure 4B ). These values suggest that, despite minor quantitative differences, the overall organization and composition of the genomes are highly conserved across the Betta species analyzed. Figure 4. Genome annotation statistics and assessment of Betta burdigala , Betta chloropharynx , and Betta schalleri. (A) GC content by component of the three betta species genome assemblies. (B) Genome composition by component of the three Betta species genome assemblies. (C) BUSCO assessment of genome annotation results for the three Betta species genome assemblies. (D) Gene duplication classification in the genome of the three Betta species. Note that a gene can be classified into more than one gene duplication type. The BUSCO analysis using the actinopterygii_odb10 dataset revealed high completeness across all three Betta species proteomes, indicating high-quality gene annotations ( Figure 4C ). Betta burdigala showed the highest completeness with 98.0% of BUSCOs identified as complete, closely followed by B. schalleri at 97.9%, and B. chloropharynx at 97.8%. The majority of these complete BUSCOs were single-copy, representing 78.5–78.8% of the total, while duplicated BUSCOs accounted for approximately 19.2–19.3%, suggesting some level of gene duplication across all three genomes. We also explored the gene duplication patterns revealing notable similarities and subtle differences in the distribution of duplication types among the three Betta species ( Figure 4D ). Across all species, dispersed duplications were the most prevalent, with B. chloropharynx showing the highest count (15,831 genes), followed closely by B. schalleri (15,269 genes) and B. burdigala (15,157 genes). This suggests that dispersed duplications represent a dominant mechanism contributing to gene expansion in these genomes, likely reflecting ongoing evolutionary pressures and functional diversification. Other duplication categories, including tandem, proximal, and whole-genome duplication (WGD)/segmental events, were represented in roughly similar proportions across species ( Figure 6 ). Tandem duplicates ranged from 2,277 in B. schalleri to 2,437 in B. chloropharynx , while proximal duplications were more variable, with B. schalleri showing the lowest count (673) compared to 924 in B. chloropharynx. WGD or segmental duplicates remained relatively consistent across species (approximately 2,270–2,370), highlighting a shared genomic history of large-scale duplication events. Singleton genes, which lack detectable paralogs, accounted for 3,114–3,178 genes depending on the species. Comparative genomic We conducted a collinearity analysis to investigate genome-level relationships among B. burdigala , B. chloropharynx , and B. schalleri. The analysis identified conserved syntenic gene blocks between each pair of species, which were subsequently quantified as collinear gene pairs ( Figure 5A ). A total of 16,948 pairs were detected between B. burdigala and B. chloropharynx (33.15%), 16,778 pairs between B. burdigala and B. schalleri (32.81%), and 17,405 pairs between B. chloropharynx and B. schalleri (34.04%). These values indicate a high degree of genome conservation across all three species, with each pair retaining roughly one-third of the total detected collinear relationships. Figure 5. Collinearity analysis of genome assembly of Betta burdigala (BBur), Betta schalleri (BSch), and Betta chloropharynx (BChl). (A) syntenic block connection of the three genomes. (B) Pairwise collinearity among Betta species. (C) Pylogenetic tree based on synteny. Among the three comparisons, the highest level of collinearity was observed between B. chloropharynx and B. schalleri , suggesting that these two species are slightly more closely related to each other than to B. burdigala ( Figure 5B ). In contrast, B. burdigala shows a similar but marginally lower level of collinearity with both B. chloropharynx and B. schalleri , implying that B. burdigala likely diverged earlier from the lineage leading to B. chloropharynx and B. schalleri. Visualization of the results using a dendrogram and a network graph further supports this interpretation. The dendrogram places B. chloropharynx and B. schalleri as sister species, with B. burdigala forming an outgroup ( Figure 5C ). The network graph illustrates strong syntenic connections among all three species, but with the thickest edge linking B. chloropharynx and B. schalleri , reflecting their higher degree of shared genome organization. We also did comparative analysis of three genes that might involve in color patterning of fishes which are Colony Stimulating Factor 1 Receptor A (CSF1RA), Melanocortin 1 Receptor (MC1R) and Paired Box 7 (PAX7). The protein sequence alignment are available in supplementary files (Supplementary file 1, Supplementary file 2, and Supplementary file 3). We found that the CSF1RA phylogenetic tree closely resembles the phylogenetic tree of the nine fish species compared based on NCBI taxonomic data ( Figure 6A and D ). On the other hand, we observed that the branching pattern of the MC1R and PAX7 phylogenetic tree does not align with the species phylogenetic tree (Suppl Fig 2). We also identified two significant protein insertions in the CSF1RA of B. burdigala ( Figure 6B and C ). The first insertion, consisting of 27 amino acids, is located at approximately alignment position 594, just before the start of the Kinase Domain (GKVLGAGAFG…). The second insertion, comprising 28 amino acids, is situated at around alignment position 701, between the two lobes of the kinase domain. Figure 6. A comparative analysis of the Colony Stimulating Factor 1 Receptor A (CSF1RA) protein sequence among B. burdigala , B. chloropharynx , and B. schalleri , alongside B. splendens , Anabas testudineus , Channa argus, Oreochromis niloticus, Danio rerio, and Carassius auratus. (A) A phylogenetic tree derived from CSFRA protein alignments across nine fish species. (B) Multiple Sequence Alignment of the CSFRA protein in these nine fish species. (C) A zoomed-in view of the sequence alignment within the red line box in (B). (D) a phylogenetic tree of nine fish species based on NCBI taxonomic data. The complete protein sequence alignment of CSF1RA is available in Supplementary file 1. Discussion Our study integrates meristic analyses and genome assemblies to provide a comparative framework for understanding divergence among three endemic Betta species from Bangka Island: B. burdigala , B. chloropharynx , and B. schalleri. Morphological comparisons revealed clear diagnostic traits that distinguish these species and align with their current taxonomic placement. B. burdigala showed reductions in dorsal and predorsal scale counts, consistent with its placement in the coccina group. 32 B. chloropharynx was distinguished by its characteristic throat markings and opercular coloration typical of the waseri group, 34 while B. schalleri exhibited greater fin ray counts and deeper body proportions, traits associated with the pugnax group. 35 These meristic patterns reinforce species boundaries and highlight morphological divergence even within closely related taxa. The genomic assemblies presented here represent the first references for these Bangka endemics. All three species demonstrated high contiguity and completeness, confirming their reliability for downstream analyses. Subtle genomic differences were also evident. B. chloropharynx displayed the largest genome size, highest scaffold N50, and elevated retrotransposon content, suggesting lineage-specific repeat dynamics. Gene duplication profiles showed dispersed duplications as the dominant category across all species, while variation in tandem and proximal duplications may indicate lineage-specific expansions. 36 – 38 These findings demonstrate that while overall genome organization and gene content remain conserved across the three species, lineage-specific signatures of divergence are detectable. The combination of high collinearity and subtle differences in repeat content and duplication patterns provides a genomic basis for understanding evolutionary relationships within the genus. 39 , 40 Moreover, the integration of meristic and genomic evidence strengthens taxonomic resolution, particularly in lineages where morphology alone may be insufficient. Together, the annotation and composition data indicate a strong conservation of gene architecture and genomic content within the Betta genus, consistent with their relatively recent divergence. Subtle differences, such as the slightly higher transcript count or gene locus size in B. chloropharynx , may hint at ongoing lineage-specific regulatory or structural adaptations. 41 , 42 The consistent duplication profiles across the three species suggest conserved duplication dynamics within the Betta genus. However, the modest variation in tandem and proximal duplication counts may point to lineage-specific gene family expansions or contraction events, potentially linked to ecological adaptations or evolutionary divergence. 37 , 43 Collinearity analysis has demonstrated significant genome conservation, with B. chloropharynx and B. schalleri exhibiting the highest degree of synteny, thereby supporting their closer evolutionary relationship compared to B. burdigala. These results further indicate that, although all three Betta species possess largely conserved genome architectures, B. chloropharynx and B. schalleri share the closest evolutionary relationship, with B. burdigala diverging earlier. This underscores the effectiveness of collinearity analysis in elucidating fine-scale relationships among recently diverged species and provides a genomic framework for further comparative and evolutionary investigations within the Betta genus. 41 , 44 , 45 The observed genomic collinearity, particularly between B. chloropharynx and B. schalleri , aligns with findings in other fish taxa, where conserved synteny is common among closely related species despite independent chromosomal fusions or rearrangements in more distant lineages. 46 , 47 Our comparative genomic analysis reveals a two-layered evolutionary dynamic underlying pigmentation diversity in Betta species: strong conservation of core pigment-cell survival pathways alongside rapid divergence in genes governing color type and pattern formation. CSF1RA, a key regulator of melanophore and color pigments (xanthophores, erythrophores, and iridophore) development, 48 – 50 follows the expected species phylogeny and shows high sequence conservation across all examined fishes, reflecting strong purifying selection on this essential signaling pathway. Notably, however, B. burdigala exhibits two large amino-acid insertions positioned adjacent to and within the kinase domain, indicating species-specific modulation rather than disruption of receptor function. While further research is necessary, such structural changes might fine-tune the color pigment pattern and density, potentially enhancing this species’ vibrant coloration. In contrast, genes associated with pigment-cell fate and color intensity—particularly PAX7 and MC1R—show phylogenetic patterns that deviate from the species tree, implying accelerated evolution or lineage-specific selection. Divergence in PAX7, a major regulator of xanthophore and erythrophore lineages, 51 – 53 is consistent with adaptive shifts affecting red–yellow pigmentation, especially relevant given the pronounced red coloration in B. burdigala. Similarly, the non-concordant evolution of MC1R, despite its variable role in teleost pigmentation, 54 – 56 suggests relaxed constraint or functional innovation within the Betta lineage. Together, these findings indicate that Betta color diversity arises from conserved developmental frameworks overlaid by rapid gene-specific evolution and structural innovation, enabling fine-scale modulation of pigment cell types and color intensity across species. 57 Although our data do not directly address ecological or paleogeographic processes, previous studies suggest that the high endemism of Bangka’s Betta species has been shaped by Pleistocene paleogeography and the persistence of specialized peat swamp habitats. 6 , 58 – 63 In this context, the divergence patterns we observed in both morphology and genomics are consistent with scenarios of historical geographic isolation and ecological filtering proposed by earlier work. Our results therefore provide the genetic and morphological evidence that complements these broader biogeographic hypotheses. These data not only support species delimitation and phylogenomic placement within the Betta genus but also establish a foundation for future work linking genomic variation with ecology, adaptation, and conservation. Given the conservation status of these taxa, the genomic resources generated here are valuable for monitoring genetic diversity and informing management strategies for Bangka’s unique freshwater biodiversity. Conclusion This study provides the first integrated meristic and genomic comparison of three endemic Betta species from Bangka Island— B. burdigala , B. chloropharynx , and B. schalleri. Meristic analyses confirmed diagnostic traits supporting their taxonomic distinction, while high-quality genome assemblies with near-complete BUSCO scores (>97%) offer reliable references for evolutionary and conservation studies. Although overall genomic architecture is conserved, subtle differences in genome size, repeat composition, and duplication profiles highlight lineage-specific divergence, with synteny analyses indicating a closer relationship between B. chloropharynx and B. schalleri. By combining morphological and genomic evidence, this study not only strengthens the resolution of species boundaries but also establishes essential resources for monitoring genetic diversity and informing conservation strategies for these endangered freshwater fishes. Data availability The data related to this project have been deposited under NCBI BioProject Accession PRJNA1328177 and are publicly available. The biosample accession numbers are SAMN51307167, SAMN51307168, and SAMN51307169 for B. burdigala , B. schalleri , and B. chloropharynx respectively. The raw genome sequence reads have been deposited in the NCBI Sequence Read Archive (SRA) under the accession numbers SRR35652347 for B. burdigala , SRR35653255 for B. schalleri , and SRR35728893 and SRR35728892 for B. chloropharynx. Additionally, the genome assemblies have been deposited in the NCBI GenBank database with accession numbers GCA_054471155.1, GCA_054471145.1, GCA_054471135.1 for B. burdigala , B. schalleri , and B. chloropharynx , respectively. The genome assemblies and their annotations have also been deposited and are available in the National Scientific Repository of the National Research and Innovation Agency (Badan Riset dan Inovasi Nasional/BRIN) of the Republic of Indonesia: Genome Betta spp. ( https://hdl.handle.net/20.500.12690/RIN/LEOQPP ). 64 Extended data All extended data have been deposited in figshare (DOI: https://doi.org/10.6084/m9.figshare.31189546 ). 65 The extended data include the following: Suppl Fig 1. The meristic (A) and morphometric (B) characters of Betta spp. Suppl Fig 2. A comparative analysis of the Melanocortin 1 Receptor (MC1R) and Paired box 7 (PAX7) protein sequences among B. burdigala , B. chloropharynx , and B. schalleri , alongside B. splendens, Anabas testudineus, Channa argus, Oreochromis niloticus, Danio rerio, and Carassius auratus. (A) A phylogenetic tree derived from MC1R protein alignments across nine fish species. (B) Multiple Sequence Alignment of the PAX7 MC1R protein in these nine fish species. (C) A phylogenetic tree derived from PAX7 protein alignments across nine fish species. (D) Multiple Sequence Alignment of the PAX7 protein in these nine fish species. The complete protein sequence alignment of MC1R and PAX7 are available in Supplementary file 2 and Supplementary file 3 respectively. Supplementary file 1. Protein alignment of CSF1R1 in 9 fishes.clustal_num Supplementary file 2. Protein alignment of MC1R in 9 fishes.clustal_num Supplementary file 3. Protein alignment of PAX7 in 9 fishes.clustal_num Reporting guidelines The complete checklists of ARRIVE 2.0 reporting guidelines is available at https://doi.org/10.6084/m9.figshare.31136455 . 11 All the underlying and extended data of this study are openly available under the terms of The CC BY 4.0 license (Creative Commons Attribution 4.0 International) . Acknowledgements We extend our sincere appreciation to the Directorate of Research and Community Service (DPPM) of Ministry of Higher Education, Science and Technology, and the Chairman of LLDIKTI II for their financial support of this research. We also express our gratitude to the Rector of Universitas Muhammadiyah Palembang, the Chairman of LPPM Universitas Muhammadiyah Palembang, and the Dean of the Faculty of Agriculture at Universitas Muhammadiyah Palembang for their support in the execution of this research. Additionally, we acknowledge PT. Timah Tbk. and Yayasan Ikan Endemik Bangka Belitung for facilitating this research. The authors are grateful for the facilities and scientific and technical support provided by the Whole Genome Sequencing Laboratory, National Research and Innovation Agency through E-Layanan Sains, Badan Riset dan Inovasi Nasional. We also wish to thank Integrated Genome Factory (IGF) Faculty of Biology UGM for granting access to computing facilities. During the preparation of this work, we used Paperpal to improve grammar and sentence structure in the manuscript. After using this tool, we reviewed and edited the content as needed and take full responsibility for the content of the publication. References 1. Helmizuryani H, et al. : Exploring freshwater fish biodiversity using eDNA metabarcoding and traditional sampling to assess floodplain waters. Journal of Water and Land Development. 2025; 100–112. Publisher Full Text 2. Nur FM, et al. : Diversity, distribution, and conservation status of Betta fish (Teleostei: Osphronemidae) in Aceh waters, Indonesia. The European Zoological Journal. 2022; 89 : 142–151. 3. Pammanasut P, et al. : Discovery of wild populations of Betta smaragdina Ladiges, 1972 (Teleostei, Osphronemidae) in a western province of Thailand. Check List. 2018; 14 : 1077–1082. 4. Tan HH: Redescription of Betta anabatoides Bleeker, and a new species of Betta from West Kalimantan, Borneo (Teleostei: Osphronemidae). Zootaxa. 2009; 2165 : 59–68. 5. Wattanadilokchatkun P, et al. : Integrative approach for landscape demography analysis of Plakad-Pa Pak-Tawan-Ok (Betta siamorientalis): Deciphering genetic and environmental factors in Eastern Thailand’s conservation efforts. Global Ecology and Conservation. 2024; 51 : e02870. Publisher Full Text 6. Valen FS, et al. : Unveiling the close relationship between Betta burdigala and Betta uberis through DNA barcoding based on COI Gene. IOP Conf. Ser. Earth Environ. Sci. 2023; 1267 : 012066. 7. Rhie A, et al. : Towards complete and error-free genome assemblies of all vertebrate species. Nature. 2021; 592 : 737–746. PubMed Abstract | Publisher Full Text 8. Totikov A, et al. : Chromosome-Level Genome Assemblies Expand Capabilities of Genomics for Conservation Biology. Genes. 2021; 12 . Publisher Full Text 9. Huang Y, et al. : Reference Genome Assembly of the Big Berry Manzanita (Arctostaphylos glauca). J. Hered. 2022; 113 : 188–196. PubMed Abstract | Publisher Full Text 10. Theissinger K, et al. : How genomics can help biodiversity conservation. Trends Genet. 2023; 39 : 545–559. PubMed Abstract | Publisher Full Text 11. Helmizuryani H, et al. : Author Checklist of The ARRIVE guidelines 2.0 for Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes. figshare. 2026. 12. Batubara JP, Laila K, Rumodang R, et al. : The Effects of Different Dosages of Pasak Bumi (Eurycoma longifolia Jack) towards The Sex of Betta Fish Embryo. IOP Conf. Ser. Earth Environ. Sci. 2022; 1118 : 012005. 13. Pattiasina BJ, Pattinasarany MM, Manuputty MMD, et al. : Masculinization of beta fish larvae Betta splendens through the different treatment immersion of honey solution and larval age. IOP Conf. Ser. Earth Environ. Sci. 2021; 797 : 012016. 14. AmericanVeterinaryMedicalAssociation: AVMA Guidelines for the euthanasia of animals.2020. Vol. Accessed 10 March 2021. 15. Imron I, et al. : Draft genome assembly of the slender walking catfish, Prophagorus nieuhofii [version 1; peer review: 2 approved with reservations, 1 not approved]. F1000Res. 2025; 14 . Publisher Full Text 16. Kolmogorov M, Yuan J, Lin Y, et al. : Assembly of long, error-prone reads using repeat graphs. Nat. Biotechnol. 2019; 37 : 540–546. PubMed Abstract | Publisher Full Text 17. Alonge M, et al. : Automated assembly scaffolding using RagTag elevates a new tomato system for high-throughput genome editing. Genome Biol. 2022; 23 : 258. 18. Marçais G, Kingsford C: A fast, lock-free approach for efficient parallel counting of occurrences of k-mers. Bioinformatics. 2011; 27 : 764–770. PubMed Abstract | Publisher Full Text 19. Simão F, Waterhouse R, Ioannidis P, et al. : Assessing genome assembly and annotation completeness with single-copy orthologs.2015; 31 : 3210–3212. Publisher Full Text 20. Manni M, Berkeley MR, Seppey M, et al. : BUSCO: Assessing Genomic Data Quality and Beyond. Current Protocols. 2021; 1 : e323. Publisher Full Text 21. Li H: Protein-to-genome alignment with miniprot. Bioinformatics. 2023; 39 . Publisher Full Text 22. Brůna T, et al. : Galba: genome annotation with miniprot and AUGUSTUS. BMC Bioinformatics. 2023; 24 : 327. Publisher Full Text 23. Hoff KJ, Stanke M: Predicting Genes in Single Genomes with AUGUSTUS. Curr. Protoc. Bioinformatics. 2019; 65 : e57. Publisher Full Text 24. Jones P, et al. : InterProScan 5: genome-scale protein function classification. Bioinformatics. 2014; 30 : 1236–1240. PubMed Abstract | Publisher Full Text 25. Cantalapiedra CP, Hernández-Plaza A, Letunic I, et al. : eggNOG-mapper v2: Functional Annotation, Orthology Assignments, and Domain Prediction at the Metagenomic Scale. Mol. Biol. Evol. 2021; 38 : 5825–5829. PubMed Abstract | Publisher Full Text 26. Almagro Armenteros JJ, et al. : SignalP 5.0 improves signal peptide predictions using deep neural networks. Nat. Biotechnol. 2019; 37 : 420–423. PubMed Abstract | Publisher Full Text 27. Wang Y, et al. : MCScanX: a toolkit for detection and evolutionary analysis of gene synteny and collinearity. Nucleic Acids Res. 2012; 40 : e49. 28. Bandi V, et al. : Visualization Tools for Genomic Conservation. Methods Mol. Biol. 2022; 2443 : 285–308. PubMed Abstract | Publisher Full Text 29. Sievers F, Higgins DG: Clustal Omega, accurate alignment of very large numbers of sequences. Methods Mol. Biol. 2014; 1079 : 105–116. PubMed Abstract | Publisher Full Text 30. Thomas KF, Wong NL-T, Ren H, et al. : IQ-TREE 3: Phylogenomic Inference Software using Complex Evolutionary Models.2025. Reference Source 31. Bianchini G, Sánchez-Baracaldo P: TreeViewer: Flexible, modular software to visualise and manipulate phylogenetic trees. Ecol. Evol. 2024; 14 : e10873. 32. Ding J, Lei W, Haryono H, et al. : Phylogenetic analysis of Betta coccina complex (Teleostei, Osphronemidae) from Peninsular Malaysia and Sumatra Island with descriptions of two new species. ZooKeys. 2025; 1238 : 161–181. PubMed Abstract | Publisher Full Text 33. Tan HH, Ng PKL: Six new species of fighting fish (Teleostei: Osphronemidae: Betta) from Borneo. Ichthyological Exploration of Freshwaters. 2006; 17 : 97–114. 34. Tan HH, Ahmad AB: Betta omega, a new species of black water fighting fish (Teleostei: Osphronemidae) from Malaysia. RAFFLES BULLETIN OF ZOOLOGY. 2018; 66 : 402–407. 35. Tan HH, Ng PKL: The labyrinth fishes (Teleostei: Anabantoidei, Channoidei) of Sumatra, Indonesia. Raffles Bull. Zool. 2005; 13 : 115–138. 36. Chen Z, et al. : De novo assembly of the goldfish (Carassius auratus) genome and the evolution of genes after whole-genome duplication. Sci. Adv. 2019; 5 : eaav0547. 37. Lu J, Peatman E, Tang H, et al. : Profiling of gene duplication patterns of sequenced teleost genomes: evidence for rapid lineage-specific genome expansion mediated by recent tandem duplications. BMC Genomics. 2012; 13 : 246. Publisher Full Text 38. Nakamura Y, et al. : Rhodopsin gene copies in Japanese eel originated in a teleost-specific genome duplication. Zoological Letters. 2017; 3 : 18. Publisher Full Text 39. Li S, et al. : Genome collinearity analysis illuminates the evolution of donkey chromosome 1 and horse chromosome 5 in perissodactyls: A comparative study. BMC Genomics. 2021; 22 : 665. Publisher Full Text 40. Wang X, et al. : Evolutionary Divergence between Toona ciliata and Toona sinensis Assayed with Their Whole Genome Sequences. Genes. 2022; 13 . Publisher Full Text 41. Kwon YM, et al. : Genomic consequences of domestication of the Siamese fighting fish. Sci. Adv. 2022; 8 : eabm4950. 42. Zhang W, et al. : The genetic architecture of phenotypic diversity in the Betta fish (Betta splendens). Sci. Adv. 2022; 8 : eabm4955. 43. Huang P, et al. : Chromosome-level genome assembly and population genetic analysis of a near-threatened rosewood species (Dalbergia cultrata Pierre Graham ex Benth) provide insights into its evolutionary and cold stress responses. Front. Plant Sci. 2023; 14 . Publisher Full Text 44. Fan G, et al. : Chromosome-level reference genome of the Siamese fighting fish Betta splendens, a model species for the study of aggression. GigaScience. 2018; 7 : giy087. Publisher Full Text 45. Srikulnath K, et al. : Overview of the betta fish genome regarding species radiation, parental care, behavioral aggression, and pigmentation model relevant to humans. Genes & Genomics. 2021; 43 : 91–104. PubMed Abstract | Publisher Full Text 46. Brasó-Vives M, et al. : Parallel evolution of amphioxus and vertebrate small-scale gene duplications. Genome Biol. 2022; 23 : 243. 47. Shen Q, et al. : Chromosome-level genome assembly of the butterfly hillstream loach Beaufortia pingi. Sci. Data. 2024; 11 : 1260. Publisher Full Text 48. Patterson LB, Bain EJ, Parichy DM: Pigment cell interactions and differential xanthophore recruitment underlying zebrafish stripe reiteration and Danio pattern evolution. Nat. Commun. 2014; 5 : 5299. 49. Braasch I, Salzburger W, Meyer A: Asymmetric Evolution in Two Fish-Specifically Duplicated Receptor Tyrosine Kinase Paralogons Involved in Teleost Coloration. Mol. Biol. Evol. 2006; 23 : 1192–1202. PubMed Abstract 50. Stanley ER, et al. : Biology and action of colony-stimulating factor-1. Mol. Reprod. Dev. 1997; 46 : 4–10. PubMed Abstract 51. Nord H, Dennhag N, Muck J, et al. : Pax7 is required for establishment of the xanthophore lineage in zebrafish embryos. Mol. Biol. Cell. 2016; 27 : 1853–1862. PubMed Abstract | Publisher Full Text 52. Roberts RB, Moore EC, Kocher TD: An allelic series at pax7a is associated with colour polymorphism diversity in Lake Malawi cichlid fish. Mol. Ecol. 2017; 26 : 2625–2639. PubMed Abstract | Publisher Full Text 53. Pokrajac NT, Singh AP: Spotting the band. Proc. Natl. Acad. Sci. 2024; 121 : e2412259121. 54. Tezuka A, Yamamoto H, Yokoyama J, et al. : The MC1R gene in the guppy (Poecilia reticulata): Genotypic and phenotypic polymorphisms. BMC. Res. Notes. 2011; 4 : 31. Publisher Full Text 55. Cal L, Suarez-Bregua P, Cerdá-Reverter JM, et al. : Fish pigmentation and the melanocortin system. Comp. Biochem. Physiol. A Mol. Integr. Physiol. 2017; 211 : 26–33. PubMed Abstract | Publisher Full Text 56. Mundy NI: Conservation and convergence of colour genetics: MC1R mutations in brown cavefish. PLoS Genet. 2009; 5 : e1000388. 57. Maan ME, Sefc KM: Colour variation in cichlid fish: Developmental mechanisms, selective pressures and evolutionary consequences. Semin. Cell Dev. Biol. 2013; 24 : 516–528. PubMed Abstract | Publisher Full Text 58. Ng PKL, Tan HH: Freshwater fishes of Southeast Asia: potential for the aquarium fish trade and conservation issues. Aquar. Sci. Conserv. 1997; 1 : 79–90. 59. Voris HK: Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations. J. Biogeogr. 2000; 27 : 1153–1167. 60. Kottelat M: The fishes of the inland waters of Southeast Asia: a catalogue and core bibliography of the fishes known to occur in freshwaters, mangroves and estuaries. Raffles Bull. Zool. 2013; 27 : 1–663. 61. Sathiamurthy E, Voris H: Maps of Holocene Sea Level Transgression and Submerged Lakes on the Sunda Shelf. Tropical Nat. Hist. 2006; 1–44. Publisher Full Text 62. Ramadhanu D, Prananda M, Wulandari UA, et al. : Perbandingan Pola Rasio Morfometrik dan Karakteristik Habitat Dua Spesies Ikan Cupang Alam (Betta spp.) Endemik Pulau Bangka. Bioscientist: Jurnal Ilmiah Biologi. 2023; 11 : 1501–1508. 63. Khanati O, Lista D, Lindiatika L, et al. : Bioecology of Wild Betta Fish (Betta schalleri) of Bangka Island Waters. Jurnal Lahan Suboptimal: Journal of Suboptimal Lands. 2024; 13 . Publisher Full Text 64. Helmizuryani H, et al. : Genome Betta spp. V2 edn (RIN Dataverse).2025. 65. Helmizuryani H, et al. : [Extended data] Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes.2026. Comments on this article Comments (0) Version 1 VERSION 1 PUBLISHED 10 Feb 2026 ADD YOUR COMMENT Comment Author details Author details 1 Study Program of Aquaculture, Faculty of Agriculture, Universitas Muhammadiyah Palembang, Palembang, South Sumatra, 30263, Indonesia 2 Study Program of Biology Education, Faculty of Teacher Training and Education, Universitas Muhammadiyah Palembang, Palembang, South Sumatra, 30263, Indonesia 3 Research Center for Applied Zoology, National Research and Innovation Agency, Cibinong, West Java, 16911, Indonesia 4 Research Center for Biota System, National Research and Innovation Agency, Cibinong, West Java, 16911, Indonesia 5 Yayasan Ikan Endemik Bangka Belitung, Pangkal Pinang, Bangka Belitung Island, 33123, Indonesia 6 Faculty of Biology, Universitas Gadjah Mada, Sleman, Special Region of Yogyakarta, 55281, Indonesia Helmizuryani Helmizuryani Roles: Conceptualization, Data Curation, Investigation, Methodology, Writing – Original Draft Preparation, Writing – Review & Editing Saleh Hidayat Roles: Data Curation, Investigation, Methodology, Validation Muhammad Nizar Roles: Data Curation, Investigation, Methodology, Validation Andhika Prima Prasetyo Roles: Investigation, Validation Boby Muslimin Roles: Data Curation, Formal Analysis, Investigation, Validation, Writing – Original Draft Preparation, Writing – Review & Editing Mochammad Zamroni Roles: Data Curation, Investigation, Methodology, Validation Dessy Nurul Astuti Roles: Data Curation, Investigation, Validation Swarlanda Swarlanda Roles: Data Curation, Investigation, Methodology, Validation Destra Ramadhanu Roles: Data Curation, Investigation, Methodology, Validation Luthfi Nurhidayat Roles: Data Curation, Formal Analysis, Software, Writing – Original Draft Preparation, Writing – Review & Editing Competing interests No competing interests were disclosed. Grant information 123/C3/DT.05.00/PL/2025 The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Article Versions (1) version 1 Published: 10 Feb 2026, 15:226 https://doi.org/10.12688/f1000research.174779.1 Copyright © 2026 Helmizuryani H et al . This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Download Export To Sciwheel Bibtex EndNote ProCite Ref. Manager (RIS) Sente metrics Views Downloads F1000Research - - PubMed Central info_outline Data from PMC are received and updated monthly. - - Citations open_in_new 0 open_in_new 0 open_in_new SEE MORE DETAILS CITE how to cite this article Helmizuryani H, Hidayat S, Nizar M et al. Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.12688/f1000research.174779.1 ) NOTE: If applicable, it is important to ensure the information in square brackets after the title is included in all citations of this article. COPY CITATION DETAILS track receive updates on this article Track an article to receive email alerts on any updates to this article. TRACK THIS ARTICLE Share Open Peer Review Current Reviewer Status: ? Key to Reviewer Statuses VIEW HIDE Approved The paper is scientifically sound in its current form and only minor, if any, improvements are suggested Approved with reservations A number of small changes, sometimes more significant revisions are required to address specific details and improve the papers academic merit. Not approved Fundamental flaws in the paper seriously undermine the findings and conclusions Version 1 VERSION 1 PUBLISHED 10 Feb 2026 Views 0 Cite How to cite this report: Kron NS. Reviewer Report For: Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.5256/f1000research.192703.r457789 ) The direct URL for this report is: https://f1000research.com/articles/15-226/v1#referee-response-457789 NOTE: it is important to ensure the information in square brackets after the title is included in this citation. Close Copy Citation Details Reviewer Report 05 Mar 2026 Nicholas S Kron , University of Miami, Rickenbacker Cswy, Miami, USA; Genevia Technologies, Finland, Finland Approved with Reservations VIEWS 0 https://doi.org/10.5256/f1000research.192703.r457789 Summary In “Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes”, Helmizuryani et al. describe the collection, sampling, genome assembly and annotation, and comparative genomic analysis of three species of the genus Betta from the Bangka ... Continue reading READ ALL Summary In “Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes”, Helmizuryani et al. describe the collection, sampling, genome assembly and annotation, and comparative genomic analysis of three species of the genus Betta from the Bangka islands of Indonesia. The authors demonstrate high quality, sub-chromosome level genome assemblies using Oxford Nanopore sequencing and established long read assembly and annotation workflows. Based on genome annotation results, the authors describe the genomes of the three species as highly colinear and consistent, with minor variation among them possibly relating to evolutionary divergence among the three, with special attention paid to pigmentation associated genes. Comments The study employs commonly used open-source tools and standard methodologies for assembly and annotation. The genomic resources generated by this study are novel and valuable contributions and will surely be useful for future studies in these species and for teleost and vertebrate genomics more broadly. The manuscript would benefit from proof reading for spelling, grammar, consistency, etc.: Consistency of italicization for species names throughout. Introduction: “The number of endemic wild Betta distribution and diversity of Southeast Asia has been identified for 16 years in Thailand (2 species) and Indonesia (9 species)” Introduction: “Three species of Betta spp. were endemic, i.e., Betta burdigala , Betta chloropharynx , and Betta schalleri.” Introduction: “The genetic studies of the endemic fish Betta spp. on Bangka Island have…” Introduction: “However, the whole genome for the evolution of the endemic fish Betta spp. on Bangka Island hasn’t been revealed.” Introduction: “High-quality genomic data could be used for Long-read sequencing and high-throughput chromosome (Hi-C) technology.” The authors reference the benefits of Hi-C in their introduction but do not use Hi-C in their analysis. Are sufficient details of methods and analysis provided to allow replication by others? (partly) The authors provide the software and version numbers to software used which is appreciated and facilitates reproducibility. However, the authors do not specify if software was used with default parameters or parameters deviating from default. These can be critical for reproducing or evaluating the analyses done, especially for tools like BLAST. Nor do the authors provide links to repositories (e.g. github, zenodo) of scripts used for verification and/or reproduction of methods. I would strongly recommend a remark in the methods stating that unless otherwise specified, default parameters are used, and explicit description when non-standard parameters are used in text or a supplement. If possible, provide access to analysis scripts in their entirety. Are the conclusions drawn adequately supported by the results? (Partly) The assembly BUSCO assessments have a very low duplicate BUSCO count but the gene model level BUSCO assessments are much higher (~20%). This is not unexpected as isoform redundancy, pseudo-genes, etc do crop up and can impact BUSCO analysis. Have the authors run the annotation level BUSCO analysis using only longest isoforms to verify? It is not clear from the methods how exactly BUSCO was run on the annotation results. The authors do not mention any secondary verification on the gene models / predicted duplicates to verify they are not artifacts/pseudogenes/etc. While automated pipelines like funannotate are powerful and convenient, secondary validation of gene models is often needed especially if proceeding to duplication analysis with tools like MCScanX/OrthoFinder/Café. The authors do not describe the assembly or annotation of the mitochondrial genome of each species. While not strictly necessary per se, these data are readily isolated from the long reads and would provide a more complete picture of the genomes and further context to the syntenty-based phylogenetic analysis. The authors use RagTag for reference-based scaffolding of assembled contigs. In the absence of Hi-C/OmniC data, this is a suitable alternative, especially given the authors have access to a closely related congener Betta splendens . This success of this approach is determined by the completeness/continuity/scaffold size of the input assemblies and the reference. Given this, comparisons of the assembly and annotation statistics of the new assemblies to the reference would provide important context for these assemblies. This would improve the ability of readers to gauge the quality of the assemblies and the success of the method. For example, a table produced by a tool like QUAST that compares the major assembly statistics would be suitable. The RagTag method used can cause scaffolded assemblies to resemble the reference genome in terms of structure, especially for more fragmented/low coverage assemblies. The authors need to clearly discuss the potential impact of these potential artefacts on their collinearity results and conclusions. Given the authors are using gene models for their comparative analysis (not sequences directly verified with molecular methods), the authors should report support for identified protein insertions with primary data (e.g. insertion spanning long reads, depth of coverage at the insertion site, etc.). In the discussion, the authors make claims about purifying selection, rate of gene evolution, lineage specific selection, etc.. These are inferences that can only be drawn with population genetic data. These claims are improper for the data presented (e.g. assemblies of single individuals) and should be removed or qualified as such. Additional comments The manuscript would be strengthened by reporting metrics from initial k-mer profiling with GenomeScope2. These could be reported in the form of the annotated k-mer distribution plot generated by GenomeScope2. The authors do not describe any methods for decontamination (either technical or biological). If these were not performed, this should be explicitly noted. If they were, what methods were employed and if they were pre-assembly or post assembly should be specified. The authors do make publicly available the raw data, genome assemblies, and annotations. I would also recommend additionally submitting the repeats to public databases such as Dfam or FishTEDB. Is the work clearly and accurately presented and does it cite the current literature? Yes Is the study design appropriate and is the work technically sound? Yes Are sufficient details of methods and analysis provided to allow replication by others? Partly If applicable, is the statistical analysis and its interpretation appropriate? I cannot comment. A qualified statistician is required. Are all the source data underlying the results available to ensure full reproducibility? Yes Are the conclusions drawn adequately supported by the results? Partly Competing Interests: No competing interests were disclosed. Reviewer Expertise: bioinformatics, marine biology I confirm that I have read this submission and believe that I have an appropriate level of expertise to confirm that it is of an acceptable scientific standard, however I have significant reservations, as outlined above. Close READ LESS CITE CITE HOW TO CITE THIS REPORT Kron NS. Reviewer Report For: Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.5256/f1000research.192703.r457789 ) The direct URL for this report is: https://f1000research.com/articles/15-226/v1#referee-response-457789 NOTE: it is important to ensure the information in square brackets after the title is included in all citations of this article. COPY CITATION DETAILS Report a concern Respond or Comment COMMENT ON THIS REPORT Views 0 Cite How to cite this report: Xing B. Reviewer Report For: Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.5256/f1000research.192703.r461103 ) The direct URL for this report is: https://f1000research.com/articles/15-226/v1#referee-response-461103 NOTE: it is important to ensure the information in square brackets after the title is included in this citation. Close Copy Citation Details Reviewer Report 26 Feb 2026 Bingpeng Xing , Third Institute of Oceanography Ministry of Natural Resources, Xiamen, China Approved with Reservations VIEWS 0 https://doi.org/10.5256/f1000research.192703.r461103 This manuscript focuses on three endemic Betta fishes from Bangka Island— Betta burdigala, B. chloropharynx, and B. schalleri— conducting an integrated study of meristic analysis and whole-genome assembly, and it makes the related sequencing data and assembly results publicly available. For endemic ... Continue reading READ ALL This manuscript focuses on three endemic Betta fishes from Bangka Island— Betta burdigala, B. chloropharynx, and B. schalleri— conducting an integrated study of meristic analysis and whole-genome assembly, and it makes the related sequencing data and assembly results publicly available. For endemic fishes of Bangka Island peatland wetlands, this study provides the first genomic resources and has potential value for conservation genomics and species evolution research. Overall, the data release is relatively complete, and the assembly contiguity and BUSCO completeness are high, meeting the basic requirements of a resource-type genome paper. However, the manuscript still needs strengthening in the following aspects: 1. Reference genome–guided scaffolding may affect structural and collinearity conclusions. The authors used RagTag together with the Betta splendens reference genome for scaffold construction. This approach is feasible for generating chromosome-scale assemblies, but it may also introduce structural bias to some extent, making the assembly closer to the reference genome structure. The conclusions in the current manuscript such as “high collinearity,” “closer phylogenetic relationships,” and “highly conserved structure” are all based on this scaffolding result; therefore: it is recommended to clearly state the potential limitations of reference-guided scaffolding in the Methods and Discussion; and clarify whether the collinearity analysis was based on the original contigs or the reference-guided scaffolds. 2. The sample size is limited, and over-inference at the species level should be avoided. The morphological analysis includes only three individuals per species, and genome sequencing includes only one individual per species. This is acceptable for a resource-type paper, but inferences in the following aspects should be more cautious: patterns of interspecific differences; gene duplication dynamics; specific gene insertion events; and inference of evolutionary divergence time. 3. The two “large insertions” in CSF1RA require stronger supporting evidence. The manuscript reports that CSF1RA in B. burdigala contains two large amino-acid insertions and speculates that they may affect pigment regulation. This result has potential novelty but currently lacks the following validation: long-read coverage; read-level support at the insertion boundaries; whether there are low-complexity or repetitive-sequence artifacts; and whether assembly or annotation errors have been excluded. 4. Evidence from a single-gene phylogenetic analysis is limited. The authors constructed a phylogenetic tree using the single gene CSF1RA and compared it with the NCBI taxonomy tree. A single-gene tree can only serve as a consistency check and cannot provide strong support for phylogenetic relationships. 5. Genome quality assessment metrics can still be strengthened. The manuscript reports N50, scaffold number, BUSCO, GC content, and gap percentage; these metrics are relatively basic. It is recommended to add sequencing depth (estimated coverage); read length N50 or mean read length; k-mer completeness analysis; assembly quality value (QV); and contamination detection methods (if any). Adding 2–3 of the above metrics can significantly improve confidence in the assembly. 6. The background statements in the Introduction needs revision. There are unclear and illogical sentences in the Introduction, such as the description of “16 years” and “2 species in Thailand and 9 species in Indonesia,” which is confusing. Rewrite the relevant paragraphs to make the logic clearer; describe more accurately recent progress in taxonomy and diversity research of the genus Betta; and standardize IUCN status as “Critically Endangered (CR)” or “Endangered (EN)” with clear citations. 7. The Latin name format is not standardized. The manuscript should be unified: genus and species names in italics, such as Betta burdigala; abbreviated form B. burdigala; in “Betta spp.” only Betta in italics and “spp.” not in italics; and the family name Osphronemidae not in italics. There are many cases of missing italics or inconsistency, and a full-text unified check is recommended. 8. There are many spelling and language errors, including but not limited to: criticaly → critically, Genome assembljies → assemblies, inconsistent spelling such as chlorpharynx / chloropharynx, etc. A comprehensive language check is recommended. Is the work clearly and accurately presented and does it cite the current literature? Partly Is the study design appropriate and is the work technically sound? Partly Are sufficient details of methods and analysis provided to allow replication by others? Partly If applicable, is the statistical analysis and its interpretation appropriate? Partly Are all the source data underlying the results available to ensure full reproducibility? Yes Are the conclusions drawn adequately supported by the results? Partly Competing Interests: No competing interests were disclosed. Reviewer Expertise: marine biology，genome，fish，genetic，DNA，eDNA I confirm that I have read this submission and believe that I have an appropriate level of expertise to confirm that it is of an acceptable scientific standard, however I have significant reservations, as outlined above. Close READ LESS CITE CITE HOW TO CITE THIS REPORT Xing B. Reviewer Report For: Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.5256/f1000research.192703.r461103 ) The direct URL for this report is: https://f1000research.com/articles/15-226/v1#referee-response-461103 NOTE: it is important to ensure the information in square brackets after the title is included in all citations of this article. COPY CITATION DETAILS Report a concern Respond or Comment COMMENT ON THIS REPORT Comments on this article Comments (0) Version 1 VERSION 1 PUBLISHED 10 Feb 2026 ADD YOUR COMMENT Comment keyboard_arrow_left keyboard_arrow_right Open Peer Review Reviewer Status info_outline Alongside their report, reviewers assign a status to the article: Approved The paper is scientifically sound in its current form and only minor, if any, improvements are suggested Approved with reservations A number of small changes, sometimes more significant revisions are required to address specific details and improve the papers academic merit. Not approved Fundamental flaws in the paper seriously undermine the findings and conclusions Reviewer Reports Invited Reviewers 1 2 Version 1 10 Feb 26 read read Bingpeng Xing , Third Institute of Oceanography Ministry of Natural Resources, Xiamen, China Nicholas S Kron , University of Miami, Rickenbacker Cswy, USA; Genevia Technologies, Finland, Finland Comments on this article All Comments (0) Add a comment Sign up for content alerts Sign Up You are now signed up to receive this alert Browse by related subjects keyboard_arrow_left Back to all reports Reviewer Report 0 Views copyright © 2026 Kron N. This is an open access peer review report distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. 05 Mar 2026 | for Version 1 Nicholas S Kron , University of Miami, Rickenbacker Cswy, Miami, USA; Genevia Technologies, Finland, Finland 0 Views copyright © 2026 Kron N. This is an open access peer review report distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. format_quote Cite this report speaker_notes Responses (0) Approved With Reservations info_outline Alongside their report, reviewers assign a status to the article: Approved The paper is scientifically sound in its current form and only minor, if any, improvements are suggested Approved with reservations A number of small changes, sometimes more significant revisions are required to address specific details and improve the papers academic merit. Not approved Fundamental flaws in the paper seriously undermine the findings and conclusions Summary In “Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes”, Helmizuryani et al. describe the collection, sampling, genome assembly and annotation, and comparative genomic analysis of three species of the genus Betta from the Bangka islands of Indonesia. The authors demonstrate high quality, sub-chromosome level genome assemblies using Oxford Nanopore sequencing and established long read assembly and annotation workflows. Based on genome annotation results, the authors describe the genomes of the three species as highly colinear and consistent, with minor variation among them possibly relating to evolutionary divergence among the three, with special attention paid to pigmentation associated genes. Comments The study employs commonly used open-source tools and standard methodologies for assembly and annotation. The genomic resources generated by this study are novel and valuable contributions and will surely be useful for future studies in these species and for teleost and vertebrate genomics more broadly. The manuscript would benefit from proof reading for spelling, grammar, consistency, etc.: Consistency of italicization for species names throughout. Introduction: “The number of endemic wild Betta distribution and diversity of Southeast Asia has been identified for 16 years in Thailand (2 species) and Indonesia (9 species)” Introduction: “Three species of Betta spp. were endemic, i.e., Betta burdigala , Betta chloropharynx , and Betta schalleri.” Introduction: “The genetic studies of the endemic fish Betta spp. on Bangka Island have…” Introduction: “However, the whole genome for the evolution of the endemic fish Betta spp. on Bangka Island hasn’t been revealed.” Introduction: “High-quality genomic data could be used for Long-read sequencing and high-throughput chromosome (Hi-C) technology.” The authors reference the benefits of Hi-C in their introduction but do not use Hi-C in their analysis. Are sufficient details of methods and analysis provided to allow replication by others? (partly) The authors provide the software and version numbers to software used which is appreciated and facilitates reproducibility. However, the authors do not specify if software was used with default parameters or parameters deviating from default. These can be critical for reproducing or evaluating the analyses done, especially for tools like BLAST. Nor do the authors provide links to repositories (e.g. github, zenodo) of scripts used for verification and/or reproduction of methods. I would strongly recommend a remark in the methods stating that unless otherwise specified, default parameters are used, and explicit description when non-standard parameters are used in text or a supplement. If possible, provide access to analysis scripts in their entirety. Are the conclusions drawn adequately supported by the results? (Partly) The assembly BUSCO assessments have a very low duplicate BUSCO count but the gene model level BUSCO assessments are much higher (~20%). This is not unexpected as isoform redundancy, pseudo-genes, etc do crop up and can impact BUSCO analysis. Have the authors run the annotation level BUSCO analysis using only longest isoforms to verify? It is not clear from the methods how exactly BUSCO was run on the annotation results. The authors do not mention any secondary verification on the gene models / predicted duplicates to verify they are not artifacts/pseudogenes/etc. While automated pipelines like funannotate are powerful and convenient, secondary validation of gene models is often needed especially if proceeding to duplication analysis with tools like MCScanX/OrthoFinder/Café. The authors do not describe the assembly or annotation of the mitochondrial genome of each species. While not strictly necessary per se, these data are readily isolated from the long reads and would provide a more complete picture of the genomes and further context to the syntenty-based phylogenetic analysis. The authors use RagTag for reference-based scaffolding of assembled contigs. In the absence of Hi-C/OmniC data, this is a suitable alternative, especially given the authors have access to a closely related congener Betta splendens . This success of this approach is determined by the completeness/continuity/scaffold size of the input assemblies and the reference. Given this, comparisons of the assembly and annotation statistics of the new assemblies to the reference would provide important context for these assemblies. This would improve the ability of readers to gauge the quality of the assemblies and the success of the method. For example, a table produced by a tool like QUAST that compares the major assembly statistics would be suitable. The RagTag method used can cause scaffolded assemblies to resemble the reference genome in terms of structure, especially for more fragmented/low coverage assemblies. The authors need to clearly discuss the potential impact of these potential artefacts on their collinearity results and conclusions. Given the authors are using gene models for their comparative analysis (not sequences directly verified with molecular methods), the authors should report support for identified protein insertions with primary data (e.g. insertion spanning long reads, depth of coverage at the insertion site, etc.). In the discussion, the authors make claims about purifying selection, rate of gene evolution, lineage specific selection, etc.. These are inferences that can only be drawn with population genetic data. These claims are improper for the data presented (e.g. assemblies of single individuals) and should be removed or qualified as such. Additional comments The manuscript would be strengthened by reporting metrics from initial k-mer profiling with GenomeScope2. These could be reported in the form of the annotated k-mer distribution plot generated by GenomeScope2. The authors do not describe any methods for decontamination (either technical or biological). If these were not performed, this should be explicitly noted. If they were, what methods were employed and if they were pre-assembly or post assembly should be specified. The authors do make publicly available the raw data, genome assemblies, and annotations. I would also recommend additionally submitting the repeats to public databases such as Dfam or FishTEDB. Is the work clearly and accurately presented and does it cite the current literature? Yes Is the study design appropriate and is the work technically sound? Yes Are sufficient details of methods and analysis provided to allow replication by others? Partly If applicable, is the statistical analysis and its interpretation appropriate? I cannot comment. A qualified statistician is required. Are all the source data underlying the results available to ensure full reproducibility? Yes Are the conclusions drawn adequately supported by the results? Partly Competing Interests No competing interests were disclosed. Reviewer Expertise bioinformatics, marine biology I confirm that I have read this submission and believe that I have an appropriate level of expertise to confirm that it is of an acceptable scientific standard, however I have significant reservations, as outlined above. reply Respond to this report Responses (0) Kron NS. Peer Review Report For: Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.5256/f1000research.192703.r457789) NOTE: it is important to ensure the information in square brackets after the title is included in this citation. The direct URL for this report is: https://f1000research.com/articles/15-226/v1#referee-response-457789 keyboard_arrow_left Back to all reports Reviewer Report 0 Views copyright © 2026 Xing B. This is an open access peer review report distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. 26 Feb 2026 | for Version 1 Bingpeng Xing , Third Institute of Oceanography Ministry of Natural Resources, Xiamen, China 0 Views copyright © 2026 Xing B. This is an open access peer review report distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. format_quote Cite this report speaker_notes Responses (0) Approved With Reservations info_outline Alongside their report, reviewers assign a status to the article: Approved The paper is scientifically sound in its current form and only minor, if any, improvements are suggested Approved with reservations A number of small changes, sometimes more significant revisions are required to address specific details and improve the papers academic merit. Not approved Fundamental flaws in the paper seriously undermine the findings and conclusions This manuscript focuses on three endemic Betta fishes from Bangka Island— Betta burdigala, B. chloropharynx, and B. schalleri— conducting an integrated study of meristic analysis and whole-genome assembly, and it makes the related sequencing data and assembly results publicly available. For endemic fishes of Bangka Island peatland wetlands, this study provides the first genomic resources and has potential value for conservation genomics and species evolution research. Overall, the data release is relatively complete, and the assembly contiguity and BUSCO completeness are high, meeting the basic requirements of a resource-type genome paper. However, the manuscript still needs strengthening in the following aspects: 1. Reference genome–guided scaffolding may affect structural and collinearity conclusions. The authors used RagTag together with the Betta splendens reference genome for scaffold construction. This approach is feasible for generating chromosome-scale assemblies, but it may also introduce structural bias to some extent, making the assembly closer to the reference genome structure. The conclusions in the current manuscript such as “high collinearity,” “closer phylogenetic relationships,” and “highly conserved structure” are all based on this scaffolding result; therefore: it is recommended to clearly state the potential limitations of reference-guided scaffolding in the Methods and Discussion; and clarify whether the collinearity analysis was based on the original contigs or the reference-guided scaffolds. 2. The sample size is limited, and over-inference at the species level should be avoided. The morphological analysis includes only three individuals per species, and genome sequencing includes only one individual per species. This is acceptable for a resource-type paper, but inferences in the following aspects should be more cautious: patterns of interspecific differences; gene duplication dynamics; specific gene insertion events; and inference of evolutionary divergence time. 3. The two “large insertions” in CSF1RA require stronger supporting evidence. The manuscript reports that CSF1RA in B. burdigala contains two large amino-acid insertions and speculates that they may affect pigment regulation. This result has potential novelty but currently lacks the following validation: long-read coverage; read-level support at the insertion boundaries; whether there are low-complexity or repetitive-sequence artifacts; and whether assembly or annotation errors have been excluded. 4. Evidence from a single-gene phylogenetic analysis is limited. The authors constructed a phylogenetic tree using the single gene CSF1RA and compared it with the NCBI taxonomy tree. A single-gene tree can only serve as a consistency check and cannot provide strong support for phylogenetic relationships. 5. Genome quality assessment metrics can still be strengthened. The manuscript reports N50, scaffold number, BUSCO, GC content, and gap percentage; these metrics are relatively basic. It is recommended to add sequencing depth (estimated coverage); read length N50 or mean read length; k-mer completeness analysis; assembly quality value (QV); and contamination detection methods (if any). Adding 2–3 of the above metrics can significantly improve confidence in the assembly. 6. The background statements in the Introduction needs revision. There are unclear and illogical sentences in the Introduction, such as the description of “16 years” and “2 species in Thailand and 9 species in Indonesia,” which is confusing. Rewrite the relevant paragraphs to make the logic clearer; describe more accurately recent progress in taxonomy and diversity research of the genus Betta; and standardize IUCN status as “Critically Endangered (CR)” or “Endangered (EN)” with clear citations. 7. The Latin name format is not standardized. The manuscript should be unified: genus and species names in italics, such as Betta burdigala; abbreviated form B. burdigala; in “Betta spp.” only Betta in italics and “spp.” not in italics; and the family name Osphronemidae not in italics. There are many cases of missing italics or inconsistency, and a full-text unified check is recommended. 8. There are many spelling and language errors, including but not limited to: criticaly → critically, Genome assembljies → assemblies, inconsistent spelling such as chlorpharynx / chloropharynx, etc. A comprehensive language check is recommended. Is the work clearly and accurately presented and does it cite the current literature? Partly Is the study design appropriate and is the work technically sound? Partly Are sufficient details of methods and analysis provided to allow replication by others? Partly If applicable, is the statistical analysis and its interpretation appropriate? Partly Are all the source data underlying the results available to ensure full reproducibility? Yes Are the conclusions drawn adequately supported by the results? Partly Competing Interests No competing interests were disclosed. Reviewer Expertise marine biology，genome，fish，genetic，DNA，eDNA I confirm that I have read this submission and believe that I have an appropriate level of expertise to confirm that it is of an acceptable scientific standard, however I have significant reservations, as outlined above. reply Respond to this report Responses (0) Xing B. Peer Review Report For: Beyond the Beauty: Meristic and Genomic Signatures of Bangka’s Endemic Betta Fishes [version 1; peer review: 2 approved with reservations] . F1000Research 2026, 15 :226 ( https://doi.org/10.5256/f1000research.192703.r461103) NOTE: it is important to ensure the information in square brackets after the title is included in this citation. The direct URL for this report is: https://f1000research.com/articles/15-226/v1#referee-response-461103 Alongside their report, reviewers assign a status to the article: Approved - the paper is scientifically sound in its current form and only minor, if any, improvements are suggested Approved with reservations - A number of small changes, sometimes more significant revisions are required to address specific details and improve the papers academic merit. 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