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Nusantara Bioscience
ISSN : 20873948     EISSN : 20873956     DOI : https://doi.org/10.13057/nusbiosci
Core Subject : Education,
Aims and Scope Nusantara Bioscience (Nusantara Biosci) encourages submission of manuscripts dealing with all aspects of biological sciences that emphasize issues germane to biological and nature conservation.
Arjuna Subject : Umum - Umum
Articles 550 Documents
Phylogeny and recombination of papaya begomoviruses in Northern and Central-East India AARSHI SRIVASTAVA; VINEETA PANDEY; RAKESH KUMAR VERMA; AVINASH MARWAL; RAMWANT GUPTA; MUHAMMAD SHAFIQ SHAHID; R. K. GAUR
Nusantara Bioscience Vol. 17 No. 2 (2025)
Publisher : Smujo International

Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n170212

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Abstract. Srivastava A, Pandey V, Verma RK, Marwal A, Gupta R, Shahid MS, Gaur RK. 2025. Phylogeny and recombination of papaya begomoviruses in Northern and Central-East India. Nusantara Bioscience 17: 298-312. Papaya plants exhibiting characteristic leaf curl symptoms were surveyed in 2022 across Uttar Pradesh, Delhi, and Chhattisgarh, India. Begomovirus infection was suspected based on symptomatology and confirmed in 11 of 47 collected samples using PCR with Begomovirus-specific primers. This study aimed to elucidate the diversity and distribution of begomoviruses infecting papaya in these regions. Full-length viral genomes were amplified through Rolling Circle Amplification (RCA), and associated alphasatellites and betasatellites were detected using PCR with universal primers. The amplified viral genomes (~2.7 kb), betasatellites (~1.4 kb), and alphasatellites (~1.3 kb) were cloned and sequenced. Sequence analysis revealed 94.57-99.46% nucleotide identity of DNA-A with known isolates of Papaya Leaf Curl Virus (PaLCuV), Cotton Leaf Curl Virus (CLCuV), Croton Yellow Vein Mosaic Virus (CYVMV), Tomato Leaf Curl New Delhi Virus (ToLCNDV), and Cotton Leaf Curl Multan Virus (CLCuMuV). The DNA-B component exhibited 98.33% identity with ToLCNDV. Ten betasatellites shared 86.81-99.71% identity with related species, whereas two alphasatellites showed approximately 98.5% identity with PaLCuA and PaLCVSA. One betasatellite (PL36) displayed 86.81% identity and was identified as a novel Papaya Leaf Curl Raipur Betasatellite (PaLCuRPRB). Furthermore, six PaLCuV isolates showing <91% identity was classified as new PaLCuV variants.
Invasion dynamics and elevational range expansion of insects in tropical agricultural landscapes of Wonosobo, Central Java, Indonesia SAKINA ENOVA RAHMADHANI; SIBRINA SALSABILA; ROI ANDRIANTO; SYARIFAH HASNA ROSYIDA; QUROTUL AINIA; ARU DEWANGGA; AHMAD DWI SETYAWAN
Nusantara Bioscience Vol. 17 No. 2 (2025)
Publisher : Smujo International

Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n170216

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Abstract. Rahmadhani SE, Salsabila S, Andrianto R, Rosyida SH, Ainia Q, Dewangga A, Setyawan D. 2025. Invasion dynamics and elevational range expansion of insects in tropical agricultural landscapes of Wonosobo, Central Java, Indonesia. Nusantara Bioscience 17: 355-374. Biological invasions in tropical mountain agroecosystems are increasingly reported, yet the processes driving elevational range expansion of invasive insects remain poorly understood. This study examined invasion dynamics and elevational range expansion of insects across a gradient of lowland to highland agricultural landscapes in Wonosobo District, Central Java, Indonesia. Using a targeted sampling approach, we quantified invasion intensity with the Relative Invasiveness Index (RII) and elevational expansion with the Altitudinal Expansion Index (AEI). Functional feeding group composition and environmental drivers were further analyzed to evaluate invasion-related community reorganization. A total of 692 insect individuals were recorded, of which 295 individuals (42.6%) belonged to nine invasive or potentially invasive taxa. RII values were lowest at lowland (13.2%), highest at mid-elevation (53.6%), and moderately high at highland sites (48.7%). In contrast, AEI was only positive in highland systems (mean AEI = 45.7%; range: 38.7-59.6%), where upslope expansion was detected in saprophagous Diptera, with Leucostoma simplex showing the highest elevational shift (up to 856 m above historically documented limits). Functional composition shifted from herbivore-dominated assemblages at low elevation to predator-dominated at mid-elevation and saprophage-detritivore-dominated at high elevation, indicating functional homogenization under increasing invasion pressure. Canonical Correspondence Analysis revealed that elevation and light intensity were the primary drivers of upslope expansion, while temperature and wind exposure influenced invasion dominance. These findings demonstrate that elevational gradients in tropical agricultural landscapes function as invasion filters rather than biodiversity gradients, with mid-elevation systems acting as transitional invasion hotspots and highland systems representing high-risk zones for invasion-driven functional simplification. The study highlights the need for elevation-specific invasion risk zoning and early intervention strategies to mitigate emerging invasion threats in tropical mountain agroecosystems.
Prediction of potential climate change impacts on the geographic distribution shift of Selaginella kraussiana and S. uncinata in East, South and Southeast Asia AHMAD DWI SETYAWAN; SUTARNO SUTARNO; SUGIYARTO SUGIYARTO; SUNARTO SUNARTO; ILYAS NURSAMSI; MUHAMMAD NUR SULTON; GILANG DWI NUGROHO
Nusantara Bioscience Vol. 18 No. 1 (2026)
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Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n180105

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Abstract. Setyawan AD, Sutarno, Sugiyarto, Sunarto, Nursamsi I, Sulton MN, Nugroho GD. 2026. Prediction of potential climate change impacts on the geographic distribution shift of Selaginella kraussiana and S. uncinata in East, South and Southeast Asia. Nusantara Bioscience 18 (1): n180105. https://doi.org/10.13057/nusbiosci/n180105. Climate change is increasingly altering the geographic distribution of invasive plant species, yet comparative assessments of closely related invasive taxa remain limited. This study evaluated the potential impacts of climate change on the future distribution of two invasive lycophytes, Selaginella kraussiana and S. uncinata, across East, South, and Southeast Asia. Species distribution models were developed using the Maximum Entropy (MaxEnt) algorithm based on occurrence records compiled from GBIF, field observations, and published literature. The models incorporated 15 environmental variables representing bioclimatic, edaphic, UVB-radiation, and topographic factors. Future habitat suitability was projected for 2030, 2050, and 2080 under four Representative Concentration Pathway (RCP) scenarios (2.6, 4.5, 6.0, and 8.5). Model performance was high for both species, with AUC values of 0.935 for S. kraussiana and 0.966 for S. uncinata, indicating excellent predictive accuracy. Environmental controls differed markedly between the species. Selaginella kraussiana was primarily associated with temperature-related variables, particularly the minimum temperature of the coldest month (bio_6), whereas S. uncinata was more strongly associated with annual precipitation (bio_12) and other moisture-related variables. Future projections indicated substantial habitat expansion for S. kraussiana, with total suitable habitat increasing from 11.47 × 10⁶ km² under current conditions to 16.43 × 10⁶ km² under RCP 8.5 by 2080. Expansion was projected mainly into higher-latitude and higher-elevation subtropical and temperate regions. In contrast, S. uncinata exhibited relatively stable distribution patterns, with total suitable habitat changing only slightly from 2.95 × 10⁶ km² to 3.04 × 10⁶ km² across future climate scenarios. These findings suggest that S. kraussiana may experience a greater climate-driven increase in invasion potential than S. uncinata and demonstrate the value of integrating species distribution modeling with ecological interpretation to support invasion-risk assessment, early detection, and climate-adaptive management of invasive lycophytes in Asia.
Climatic and elevational drivers of Selaginella species richness in Java, Indonesia AHMAD DWI SETYAWAN; SUTARNO SUTARNO; SUGIYARTO SUGIYARTO; SUNARTO SUNARTO; MUHAMMAD NUR SULTON
Nusantara Bioscience Vol. 18 No. 1 (2026)
Publisher : Smujo International

Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n180106

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Abstract. Setyawan AD, Sutarno, Sugiyarto, Sunarto, Sulton MN. 2026. Climatic and elevational drivers of Selaginella species richness in Java, Indonesia. Nusantara Bioscience 18 (1): n180106. https://doi.org/10.13057/nusbiosci/n180106. Understanding environmental factors that shape species richness patterns, yet the determinants of Selaginella diversity in Java, Indonesia, remain poorly understood. Selaginella, an ancient lycophyte genus widely distributed in tropical regions, is highly sensitive to environmental conditions, particularly moisture availability and temperature. This study examined the relationships between Selaginella species richness and elevational and climatic gradients across Java, Indonesia. A total of 1,962 occurrence records representing 21 identified species and one unidentified taxon were compiled from field surveys, herbarium collections, and verified biodiversity databases. Species richness was estimated using a 0.25° × 0.25° grid system and spatially visualized through Inverse Distance Weighting (IDW) interpolation. The influence of elevation, Annual Mean Temperature, Annual Mean Precipitation, Annual Mean Solar Radiation, and Annual Mean Moisture Index on species richness was evaluated using polynomial regression, Pearson correlation analysis, and Principal Component Analysis (PCA). Species richness showed pronounced spatial heterogeneity, with hotspots concentrated in the humid montane regions of West Java. Richness exhibited nonlinear relationships with all environmental variables and peaked at intermediate elevations. Annual Mean Moisture Index (R² = 0.616) and Annual Mean Precipitation (R² = 0.527) were the variables most strongly associated with species richness, whereas Annual Mean Temperature and solar radiation showed negative relationships. Correlation and PCA results further indicated that moisture availability is closely linked to richness patterns, while elevation influences richness indirectly through its effects on local climatic conditions. These findings demonstrate that humid submontane and montane environments are critical for maintaining Selaginella diversity in Java and highlight the importance of conserving climatically suitable habitats under future environmental change.
Short Communication: Exploratory AFLP-based genetic structuring among selected Selaginella species from Indonesia AHMAD DWI SETYAWAN; TATIK CHIKMAWATI; MIFTAHUDIN MIFTAHUDIN; SUTARNO SUTARNO; SUGIYARTO SUGIYARTO; SUNARTO SUNARTO; YULITA KUSUMADEWI
Nusantara Bioscience Vol. 17 No. 2 (2025)
Publisher : Smujo International

Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n170217

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Abstract. Setyawan AD, Chikmawati T, Miftahudin, Sutarno, Sugiyarto, Sunarto. 2025. Short Communication: Exploratory AFLP-based genetic structuring among selected Selaginella species from Indonesia. Nusantara Bioscience 17 (2): 375-386. Selaginella is one of the oldest groups of living vascular plants and helps us understand how early land plants evolved and diversified. Despite its ecological and morphological diversity, molecular data remain limited for many tropical species. This study presents an exploratory assessment of multilocus genetic structuring among selected Selaginella species using Amplified Fragment Length Polymorphism (AFLP) markers. AFLP profiles were generated using four selective primer combinations, producing a total of 248 polymorphic fragments. Genetic similarity among species was quantified using Jaccard’s coefficient, and patterns of multilocus resemblance were examined through UPGMA clustering and principal coordinate analysis (PCoA). Pairwise similarity values ranged from 0.59 to 1.00, indicating a continuum of genetic differentiation rather than sharply delimited genetic groups. UPGMA clustering and PCoA revealed two broad species assemblages and substantial variation in multilocus genetic similarity among the analyzed taxa. The clustering pattern showed partial correspondence with major growth-form categories, with predominantly creeping to ascending species grouped separately from taxa exhibiting a cypress-like habit together with S. ornata. At the same time, several morphologically distinct species exhibited relatively high AFLP similarity, highlighting both congruence and incongruence between molecular and morphological patterns. Because the final comparative analyses were based on representative AFLP profiles for each species, the results should be regarded as an exploratory overview of interspecific multilocus differentiation rather than an assessment of intraspecific variation. Although AFLP markers do not permit direct phylogenetic inference, the findings provide a useful molecular baseline for future studies employing broader sampling and sequence-based or genomic approaches. Our findings demonstrate the value of integrating multilocus AFLP data with morphological evidence to better understand genetic structuring and species relationships in Selaginella. Although exploratory in nature, this study provides a useful molecular baseline for future sequence-based and genomic investigations.
Climate-associated shifts in potential habitat of the vulnerable tree Eusideroxylon zwageri in Southeast Asia using an exploratory MaxEnt assessment ARDITA AYU WULANDARI; AULIA TSALATSA AGUSTINA; AZKA DITA AULIA; BIAN RAFI SAFARAZ; ARUM NUR MUKARROMAH; ASFI DZIHNI; ATIKAH KHOIRIYAH AZZAM; AULIA SYAH HAFIFFAH; MUHAMMAD NUR SULTON; AHMAD DWI SETYAWAN
Nusantara Bioscience Vol. 18 No. 1 (2026)
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Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n180107

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Abstract. Wulandari AA, Agustina AT, Aulia AD, Safaraz BR, Mukarromah AN, Dzihni A, Azzam AK, Hafiffah AS, Sulton MN, Setyawan AD. 2026. Climate-associated shifts in potential habitat of the vulnerable tree Eusideroxylon zwageri in Southeast Asia using an exploratory MaxEnt assessment. Nusantara Bioscience 18 (1): n180107. https://doi.org/10.13057/nusbiosci/n180107. Eusideroxylon zwageri (Bornean ironwood) is a slow-regenerating, high-value rainforest tree threatened by timber extraction, habitat conversion, and climate change. We examined its current and future potential habitat in Southeast Asia using an archived MaxEnt workflow based on 57 GBIF occurrence records and 18 climatic, topographic, edaphic, vegetation, radiation, and anthropogenic predictors. Baseline climate was represented by WorldClim variables, whereas projections used HadGEM2-ES conditions for 2030, 2050, and 2080 under RCP 2.6, 4.5, 6.0, and 8.5. The baseline model had an AUC of 0.91. Precipitation of the driest quarter contributed 57.40%, followed by human population density (9.85%), minimum temperature of the coldest month (9.43%), and annual temperature range (7.78%). Under baseline conditions, high-suitability habitat occupied 0.22×106 km2 (5.16% of the modeled region), principally in Borneo, Sumatra, Brunei Darussalam, and Peninsular Malaysia. Across future scenarios, total modeled area remained approximately constant (4.25-4.28×106 km2), but the distribution among suitability classes changed; high suitability ranged from 4.94% to 6.82%. Within Borneo, high suitability ranged from 22.35% to 38.06%, and the 2080 suitability centroid shifted 7.81-66.54 km depending on the RCP. These outputs indicate redistribution of modeled environmental suitability, not demonstrated adaptation, occupancy, or population persistence. The maps are therefore appropriate for regional screening but require confirmation through bias-corrected occurrence data, spatially independent validation, multiple climate models, model-tuning and extrapolation diagnostics, dynamic land-cover scenarios, and field assessment before site-level conservation decisions.
Implications of irrigation regimes and nitrogen timing on rice yield and quality under different production systems SHAHID NAZEER; NADEEM AKBAR; SHAKEEL AHMAD ANJUM; MUHAMMAD ANWAR-UL-HAQ
Nusantara Bioscience Vol. 18 No. 1 (2026)
Publisher : Smujo International

Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n180108

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Abstract. Nazeer S, Akbar N, Anjum SA, Anwar-Ul-Haq M. 2026. Implications of irrigation regimes and nitrogen timing on rice yield and quality under different production systems. Nusantara Bioscience 18 (1): n180108. https://doi.org/10.13057/nusbiosci/n180108. Increasing production costs, shortage of water, and labor are posing threats to the sustainability of rice-wheat cropping system in Pakistan. The current study assessed the impact of production practices, irrigation strategies, and nitrogen placement on rice yield and grain quality. Field experiment was carried out for two consecutive seasons from 2022-2023 at Water Management Research Centre, PARS, University of Agriculture, Faisalabad, Pakistan, through a randomized complete block design with factorial combination of two production practices (transplanted rice, TPR; direct-seeded rice, DSR), two irrigation management systems (Alternate Wetting and Drying, AWD; aerobic irrigation, AI) and nitrogen application timing before irrigation and after irrigation. Over the years, DSR performed better than TPR in terms of higher plant stand, productive tillers, and grain yield with an average grain yield of 5.75 t ha-¹ for DSR and 5.21 t ha-¹ for TPR. On the other hand, TPR proved superior to DSR in selected grain quality attributes. The AWD irrigation had a positive effect on the plant growth and yield attributes as well as grain quality, resulting in higher grain yield (6.5 t ha-¹) and higher protein content (7.81%) compared to the AI system. Application of the nitrogen fertilizer prior to the irrigation had more pronounced effect on growth, yield, and quality parameters than application after the irrigation. Interactions among the production systems, irrigation methods, and nitrogen fertilization showed that the best yield results could be obtained from the combination of the DSR technique (5.75 t ha-1), AWD (5.95 t ha-1), and nitrogen fertilizer application prior to the irrigation (5.68 t ha-1). Moreover, this combination led to better grain quality parameters, such as protein content and milling qualities.
Field-based assessment of rhizosphere bacterial communities in banana and Goeppertia allouia in response to a banana fiber-derived bioproduct VAN LONG NGUYEN; XUAN HOAN NGUYEN; XUAN CUONG DANG; THI THANH NGAN TRUONG; VIET THE HO
Nusantara Bioscience Vol. 18 No. 1 (2026)
Publisher : Smujo International

Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n180109

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Abstract. Nguyen VL, Nguyen XH, Dang XC, Truong TTN, Ho VT. 2026. Field-based assessment of rhizosphere bacterial communities in banana and Goeppertia allouia in response to a banana fiber-derived bioproduct. Nusantara Bioscience 18 (1): n180109. https://doi.org/10.13057/nusbiosci/n180109. Banana fiber processing generates substantial residual biomass that can potentially be converted into value-added agricultural inputs. This field-based study provides a comparative assessment of rhizosphere bacterial communities associated with banana (Musa spp.) and Goeppertia allouia observed under local management practices in a farmer-managed agricultural system. Utilizing three composite biological replicates per treatment, rhizosphere soil samples were collected from fertilized and unfertilized plots, and microbial communities were analyzed using 16S rRNA gene sequencing. Amplicon Sequence Variants (ASVs) were generated using the DADA2 pipeline and classified against the SILVA database. No statistically significant differences in alpha diversity were observed, although ASV richness ranged from 6,045 to 10,270 across samples. In contrast, beta diversity analysis revealed differences in microbial community composition. Bray-Curtis dissimilarity values indicated separation between the two crop species (> 80), while within-treatment variation ranged from 38.30 to 66.34. PERMANOVA analysis showed that plant species were significantly associated with variation in microbial community composition (R² = 0.41, p = 0.001), whereas fertilizer treatment showed a weaker but detectable association (p < 0.05). At the phylum level, Proteobacteria (35-45%) and Firmicutes (5-10%) were relatively more abundant in fertilized soils, while Acidobacteriota (10-15%) and Chloroflexi (12-18%) were more represented in unfertilized soils. These observed shifts reflect crop-specific and management-associated bacterial profiles, with plant species showing a stronger association than fertilizer treatment. Overall, the findings provide preliminary insights into rhizosphere microbial variation under field conditions. Given the farmer-managed design, lack of baseline soil characterization, and observational nature of the study, the results should be interpreted as indicative of community-level patterns rather than definitive evidence of treatment effects. From a practical perspective, the findings support cautious use of banana fiber-derived bioproducts as a complementary soil input, while emphasizing that crop identity and field heterogeneity must be considered when interpreting microbiome responses.
Taxonomic and landscape-use composition of planted vegetation across a tropical university campus in Central Java, Indonesia FADIA AULIANISSA AINAYA; NUR ROHMAYANI ANGGELIKA PUTRI; DANASTRI NUR ATHAYA RADYA PUTRI; AIKO YHOVIERA FARRAZ MU&#039;ALI; SESILIA RETNO AYU NINGTYAS; HERLINA NOFITASARI; AHMAD DWI SETYAWAN
Nusantara Bioscience Vol. 18 No. 1 (2026)
Publisher : Smujo International

Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n180110

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Abstract. Ainaya FA, Putri NRA, Putri DNAR, Mu'ali AYF, Ningtyas SRA, Nofitasari H, Setyawan AD. 2026. Taxonomic and landscape-use composition of planted vegetation across a tropical university campus in Central Java, Indonesia. Nusantara Bioscience 18 (1): n180110. https://doi.org/10.13057/nusbiosci/n180110. Urban university campuses are intensively managed landscapes that can retain diverse planted floras and support long-term biodiversity planning. This study produced a traceable inventory of planted vegetation at the main campus of Universitas Sebelas Maret, Surakarta, Central Java, Indonesia. Seventeen sampling stations were distributed purposively among ten campus landscape zones; each station comprised four nested component plots assigned to tree, pole, sapling, and seedling/low-stature classes, giving 68 component plots rather than 68 independent sampling units. The archived checklist was audited at the taxon level, and every entry was assigned to one mutually exclusive primary landscape-use category. The inventory contained 99 operational taxonomic entries representing 98 named species because two cultivated forms of Psidium guajava were retained separately; entries belonged to 39 families. Fabaceae was the richest family (17 entries), followed by Arecaceae and Myrtaceae (seven each). Ornamental plants were the largest use category (37 entries; 37.4%), followed by fruit trees (27; 27.3%), shade trees (19; 19.2%), timber species (6; 6.1%), multipurpose plants (6; 6.1%), and medicinal plants (4; 4.0%). Across structural classes, 61 taxa occurred in the seedling/low-stature class, 28 in sapling, 28 in pole, and 32 in tree classes; entries could occur in multiple classes. Sixty-five entries (65.7%) were represented in only one class, whereas five (5.1%) occurred in all four; ornamental entries were concentrated in the seedling/low-stature class. Because plot-level abundance, DBH data, vouchers, and repeated surveys were unavailable, abundance-based diversity, regeneration, sector comparisons, and ecosystem-service estimates could not be validated. The audited inventory provides a reproducible baseline for future georeferenced, abundance-based, and temporal campus vegetation monitoring.
Abundance-diversity decoupling in tropical seagrass meadows of Mati City, Philippines ANTHONY AMORES; SOPHIA NADENN AGUILAR; ERROLE MAXEY
Nusantara Bioscience Vol. 18 No. 2 (2026)
Publisher : Smujo International

Show Abstract | Download Original | Original Source | Check in Google Scholar | DOI: 10.13057/nusbiosci/n180201

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Abstract. Amores A, Aguilar SN, Maxey E. 2026. Abundance-diversity decoupling in tropical seagrass meadows of Mati City, Philippines. Nusantara Bioscience 18 (2): n180201. https://doi.org/10.13057/nusbiosci/n180201. Seagrass ecosystems provide important ecological functions but are increasingly exposed to environmental and human pressures. Understanding changes in these ecosystems requires assessment of both abundance and diversity over time. This study examined temporal variation in seagrass communities in two tropical meadows, Lawigan and Tamisan, in Mati City, Davao Oriental, Philippines, using quarterly sampling from March 2024 to December 2025. Seagrass shoots were quantified using transect-quadrat methods, and community structure was evaluated through diversity indices and individual-based rarefaction analyses. Eight seagrass species were recorded during the study period. Total shoot abundance increased by approximately 350%, from 2,098 individuals in 2024 to 9,443 in 2025, whereas pooled annual species richness increased only slightly from six to seven species. Despite higher abundance, diversity indices declined, indicating greater dominance by a few species. Species-level analyses showed marked increases in Enhalus acoroides and Cymodocea serrulata, while other species remained stable, declined, or occurred intermittently. Rarefaction analyses revealed similar expected richness among samples in 2024 but greater variability in 2025 under standardized sampling effort. The results show that increases in seagrass abundance do not necessarily correspond to increases in diversity. Instead, higher abundance was associated with greater species dominance and uneven community structure. These findings highlight the importance of combining abundance, diversity, and standardized richness measures in long-term monitoring to improve assessments of tropical seagrass ecosystem condition.