<?xml version="1.0" encoding="UTF-8"?><!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "https://jats.nlm.nih.gov/publishing/1.3/JATS-journalpublishing1-3.dtd"><article xml:lang="en" dtd-version="1.3" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" article-type="research-article"><front><journal-meta><journal-id journal-id-type="issn">2527-2799</journal-id><journal-title-group><journal-title>Bioeksperimen: Jurnal Penelitian Biologi</journal-title><abbrev-journal-title>Bioeksperimen</abbrev-journal-title></journal-title-group><issn pub-type="epub">2527-2799</issn><issn pub-type="ppub">2460-1365</issn><publisher><publisher-name>Universitas Muhammadiyah Surakarta</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.23917/bioeksperimen.v12i1.15450</article-id><title-group><article-title>Waterbirds as Bioindicators of Ecosystem Health: Seasonal Dynamics in Tondano Lake, Indonesia</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Djarang</surname><given-names>Rievo</given-names></name><address><country>Indonesia</country></address><xref ref-type="aff" rid="AFF-1"></xref></contrib><contrib contrib-type="author"><name><surname>Harindah</surname><given-names>Gerry</given-names></name><address><country>United States</country><email>gerryharindah@outlook.com</email></address><xref ref-type="aff" rid="AFF-2"></xref><xref ref-type="corresp" rid="cor-1"></xref></contrib><contrib contrib-type="author"><name><surname>Suriani</surname><given-names>Ni Wayan</given-names></name><address><country>Indonesia</country></address><xref ref-type="aff" rid="AFF-1"></xref></contrib><contrib contrib-type="author"><name><surname>Taulu</surname><given-names>Marthy Lingkan Stella</given-names></name><address><country>Indonesia</country></address><xref ref-type="aff" rid="AFF-1"></xref></contrib></contrib-group><aff id="AFF-1"><institution-wrap><institution>Universitas Negeri Manado</institution><institution-id institution-id-type="ror">https://ror.org/04appkh56</institution-id></institution-wrap><country country="ID">Indonesia</country></aff><aff id="AFF-2"><institution-wrap><institution>University of South Florida</institution><institution-id institution-id-type="ror">https://ror.org/032db5x82</institution-id></institution-wrap><country country="US">United States</country></aff><author-notes><corresp id="cor-1">Corresponding author: Gerry Harindah, University of South Florida.  Email: <email>gerryharindah@outlook.com</email></corresp></author-notes><pub-date iso-8601-date="2026-3-31" publication-format="electronic" date-type="pub"><day>31</day><month>3</month><year>2026</year></pub-date><volume>12</volume><issue>1</issue><fpage>45</fpage><lpage>54</lpage><history><date date-type="received" iso-8601-date="2026-1-13"><day>13</day><month>1</month><year>2026</year></date><date date-type="rev-recd" iso-8601-date="2026-3-1"><day>1</day><month>3</month><year>2026</year></date><date date-type="accepted" iso-8601-date="2026-3-2"><day>2</day><month>3</month><year>2026</year></date></history><permissions><copyright-statement>Copyright (c) 2026 Bioeksperimen: Jurnal Penelitian Biologi</copyright-statement><copyright-year>2026</copyright-year><copyright-holder>Bioeksperimen: Jurnal Penelitian Biologi</copyright-holder><license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by-nc/4.0/"><ali:license_ref xmlns:ali="http://www.niso.org/schemas/ali/1.0/">https://creativecommons.org/licenses/by-nc/4.0/</ali:license_ref><license-p>This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License.</license-p></license></permissions><self-uri xlink:href="https://journals2.ums.ac.id/bioeksperimen/article/view/15450" xlink:title="Waterbirds as Bioindicators of Ecosystem Health: Seasonal Dynamics in Tondano Lake, Indonesia">Waterbirds as Bioindicators of Ecosystem Health: Seasonal Dynamics in Tondano Lake, Indonesia</self-uri><abstract><p>Waterbird diversity in Tondano Lake reflects ecological degradation caused by water-hyacinth invasion and sedimentation. This study analyzed the diversity and abundance of waterbirds in Tondano Lake, Indonesia, across wet (February-May) and dry (June-October) seasons in 2025 to assess their role as bioindicators of ecosystem health. Multi-season observations were carried out monthly along a 6.5 km transect at ten strategic points. Eight waterbird species from three families were recorded, with <italic>Egretta garzetta</italic> being the most abundant (29.1%) and <italic>Alcedo atthis</italic> the least (4.8%). The Shannon-Wiener diversity index (H' = 1.92) indicates a moderate level of species diversity, which reflects a pattern of ecological simplification when compared to pristine tropical lakes such as Lake Sentarum (H' = 2.5). Waterbird abundance showed significant negative correlations with water hyacinth density (r = -0.68, p = 0.032) and water depth (r = -0.72, p = 0.019), revealing that invasive vegetation and shoaling are primary drivers of habitat loss. These findings demonstrate that waterbirds function as sensitive bioindicators of ecological stress, highlighting the urgent need for integrated watershed management to prevent further community collapse.</p></abstract><kwd-group><kwd>Biodiversity</kwd><kwd>Ecosystem Health</kwd><kwd>Sedimentation</kwd><kwd>Tondano Lake</kwd><kwd>Waterbirds</kwd></kwd-group><custom-meta-group><custom-meta><meta-name>File created by JATS Editor</meta-name><meta-value><ext-link ext-link-type="uri" xlink:href="https://jatseditor.com" xlink:title="JATS Editor">JATS Editor</ext-link></meta-value></custom-meta><custom-meta><meta-name>issue-created-year</meta-name><meta-value>2026</meta-value></custom-meta></custom-meta-group></article-meta></front><body><sec><title>Introduction</title><p>Freshwater ecosystems, encompassing lakes, rivers, and wetlands, are globally recognized as critical centers of biological diversity, supporting a wide array of flora and fauna (<xref ref-type="bibr" rid="BIBR-4">(Dudgeon et al., 2006)</xref>; Strayer &amp; Dudgeon, 2010). These ecosystems provide essential services to both human communities and wildlife (<xref ref-type="bibr" rid="BIBR-4">(Dudgeon et al., 2006)</xref>; <xref ref-type="bibr" rid="BIBR-6">(Green &amp; Elmberg, 2014)</xref>). Located in North Sulawesi, Indonesia, Tondano Lake stands out as a prominent freshwater body of high ecological, social, and economic importance. It not only supports local fisheries and tourism but also serves as a crucial habitat for various waterbird species.</p><p>Waterbirds are considered indispensable components of healthy aquatic ecosystems due to their multifaceted ecological functions. They act as regulators of aquatic populations by preying on fish and invertebrates, contribute to the regeneration of riparian vegetation through seed dispersal, and, most importantly, serve as sensitive bioindicators of environmental health (<xref ref-type="bibr" rid="BIBR-1">(Amat &amp; Green, 2009)</xref>; <xref ref-type="bibr" rid="BIBR-6">(Green &amp; Elmberg, 2014)</xref>). Changes in their diversity, abundance, and distribution patterns can provide early warning signals of ecosystem degradation, such as a decline in water quality or habitat loss (<xref ref-type="bibr" rid="BIBR-12">(Kelly, 1998)</xref>; <xref ref-type="bibr" rid="BIBR-7">(Grzędzicka, 2020)</xref>).</p><p>Despite their ecological significance, Tondano Lake and its waterbird populations are facing formidable environmental pressures. A primary threat is the aggressive invasion of the exotic aquatic plant, water hyacinth (<italic>Eichhornia crassipes</italic>), which has been reported to cover up to 30% of the lake’s surface area <xref ref-type="bibr" rid="BIBR-18">(Pratama et al., 2023)</xref><xref ref-type="bibr" rid="BIBR-11">(Hendratta et al., 2024)</xref>. This unchecked proliferation forms dense floating mats that obstruct light penetration, induce eutrophication, and reduce dissolved oxygen levels, thereby disrupting the fundamental food web and negatively impacting species that depend on open, clear water for foraging, such as <italic>Himantopus leucocephalus</italic> and <italic>Egretta garzetta</italic> (<xref ref-type="bibr" rid="BIBR-20">(Toft et al., 2003)</xref>; <xref rid="BIBR-21" ref-type="bibr">(Villamagna &amp; Murphy, 2010)</xref>). Concurrently, the lake is experiencing accelerated sedimentation, or shoaling, primarily due to heightened erosion from agricultural and deforested areas in the surrounding watershed. This process reduces the lake’s depth, fundamentally altering habitat structure and decreasing the availability of prey, which is particularly detrimental to piscivorous species like <italic>Alcedo atthis</italic> (<xref ref-type="bibr" rid="BIBR-12">(Kelly, 1998)</xref>; <xref rid="BIBR-14" ref-type="bibr">(Kumar &amp; Pattnaik, 2012)</xref>).</p><p>Existing scientific literature on Tondano Lake has predominantly focused on water quality and fish populations <xref rid="BIBR-17" ref-type="bibr">(Onibala et al., 2025)</xref>, leaving a significant research gap in studies that integrate the dynamics of waterbird communities with the specific, escalating environmental threats of water hyacinth invasion and sedimentation. This gap underscores the urgent need for a dedicated study on waterbirds as a key bioindicator group (<xref ref-type="bibr" rid="BIBR-1">(Amat &amp; Green, 2009)</xref>; <xref ref-type="bibr" rid="BIBR-6">(Green &amp; Elmberg, 2014)</xref>). By investigating the relationship between waterbird populations and these environmental stressors, this research can offer a more holistic and nuanced understanding of the lake’s health and resilience.</p><p>This work is aligned with global conservation efforts and contributes to the realization of Sustainable Development Goals (SDGs) 6 (Clean Water and Sanitation) and 15 (Life on Land). The specific objectives of this study were: (1) to identify the species and abundance of waterbirds in Tondano Lake; (2) to calculate the species diversity index and relative abundance; and (3) to analyze the ecological role of waterbirds within the context of the identified environmental threats, specifically the invasion of water hyacinth and sedimentation. By addressing these objectives, this research aims to provide the scientific data necessary to inform and strengthen the management of the Tondano Lake ecosystem.</p></sec><sec><title>Materials and methods</title><sec><title>1. Study Location and Period</title><p>The study was conducted at Tondano Lake, located in North Sulawesi, Indonesia (1°15′N, 124°50′E) (<xref ref-type="fig" rid="figure-1">Figure 1</xref>). The region experiences a tropical monsoon climate characterized by a distinct wet season (November–May) and a dry season (June–October). The area receives a mean annual precipitation of approximately 2,700 mm to 3,000 mm, with the highest rainfall intensity typically occurring between December and January (Meteorological, Climatological, and Geophysical Agency [BMKG], 2023). This high precipitation volume significantly influences the lake’s hydrology, contributing to seasonal fluctuations in water depth and the transport of sediment from the surrounding agricultural watershed. Tondano Lake was selected for its ecological importance as a major freshwater body and for the escalating environmental challenges it faces from water-hyacinth invasion and accelerated sedimentation <xref ref-type="bibr" rid="BIBR-18">(Pratama et al., 2023)</xref><xref ref-type="bibr" rid="BIBR-11">(Hendratta et al., 2024)</xref>.</p></sec><sec><title>2. Equipment and Materials</title><p>The following equipment and materials were utilized during the field survey:</p><p>1). A Garmin eTrex 30x GPS unit to accurately determine the coordinates of each observation point, with a precision of ±3 m; 2). A pair of Nikon Monarch 10x42 binoculars to facilitate long-range observation of waterbirds from a distance of up to 100 m; 3). A Canon EOS 90D digital camera equipped with a 100-400 mm telephoto lens for high-resolution documentation of identified species; 4). A Casio G-Shock wristwatch to ensure accurate timing of observations; 5). A motorized boat with a capacity of four people and a maximum speed of 10 km/h for transect navigation; 6). A Lowrance Hook2-4x echosounder to measure water depth at each observation point; 7). A DJI Mavic Mini drone to capture aerial imagery for estimating the percentage coverage of water hyacinth; 8). A thematic map of Tondano Lake (scale 1:50,000) from the Geospatial Information Agency to guide the transect route; 9). The "Birds of New Guinea" field guide by Beehler et al. (2001) for species identification based on morphological and behavioral characteristics and field data sheets for systematic recording of observations, including species counts and environmental conditions.</p><fig id="figure-1" ignoredToc=""><label>Figure 1</label><caption><p>Location of Surveyed Lakes and Sampling Sites in the Lake Tondano, North Sulawesi, Indonesia.</p></caption><graphic xlink:href="https://journals2.ums.ac.id/bioeksperimen/article/download/15450/5374/67877" mime-subtype="png" mimetype="image"><alt-text>Image</alt-text></graphic></fig></sec><sec><title>3. Study design</title><p>The study employed a survey method known as the concentration count, as described by Helvoort (1981). This approach involved establishing a 6.5 km transect along the lake's shoreline and selecting 10 strategic observation points. The observation points were not chosen purely at random but were instead selected using a stratified random sampling method to ensure they were representative of the lake's diverse habitat conditions. Specifically, the points were chosen based on three key ecological criteria: (1) the presence of dense riparian vegetation, such as <italic>Typha angustifolia</italic> and <italic>Pandanus tectorius</italic>, which serves as a refuge for waterbirds; (2) shallow water areas (depths &lt;2 m), which are primary foraging habitats; and (3) varying densities of water hyacinth cover (categorized as low: &lt;20%, medium: 20-50%, and high: &gt;50%). This deliberate selection process was critical for allowing the researchers to directly assess the impact of water hyacinth on waterbird distribution and abundance, strengthening the validity of the correlational findings. Each observation point had a circular observation radius of 50 m. The overall transect covered both the lake's edges and a central area, with water depths ranging from 1.2 to 3.5 m.</p></sec><sec><title>4. Observation Procedures</title><p>Observations were conducted twice daily to capture peak waterbird activity: in the morning (06.00-10.00 WITA) and late afternoon (15.00-18.00 WITA). The boat maintained a constant speed of 5 km/h to minimize disturbance to the birds. At each of the 10 observation points, data were collected over a 20-minute period. The total daily observation time was 4 hours. The specific data collected included:</p><p><bold>1). Species and Abundance:</bold> Waterbird species were identified and counted using binoculars and cameras. Local guides familiar with the species' common names assisted, and these were cross-referenced with scientific names using the Beehler et al. (2001) field guide. All sightings were documented with photographs for later verification by independent ornithologists.</p><p><bold>2). Environmental Conditions:</bold> The density of water hyacinth was estimated visually as a percentage of surface cover within the 50 m radius and was subsequently validated using aerial imagery from the drone, which had a resolution of 4K and an accuracy of ±5%. Water depth was measured using the echosounder. All data were meticulously recorded on field data sheets.</p></sec><sec><title>5. Data Analysis</title><p>The collected data were subjected to a multi-stage analysis. First, the total number and types of waterbird species were tabulated, and their conservation status was referenced from the IUCN Red List. Relative abundance (Ki​) was calculated for each species using the following formula:</p><p><inline-formula><tex-math id="math-1"><![CDATA[ \documentclass{article} \usepackage{amsmath} \begin{document} \displaystyle K_i = \frac{n_i}{N} \times 100\% \end{document} ]]></tex-math></inline-formula></p><p>Where ni​ is the total number of individuals of species i, and N is the total number of individuals of all species found. Relative abundance was then categorized as Dominant (&gt;8), Abundant (2.1-8), Frequent (1.1-2), Occasional (0.1-1), or Rare (&lt;0.1).</p><p>The Shannon-Wiener diversity index (H′) was computed using the formula:</p><p><inline-formula><tex-math id="math-2"><![CDATA[ \documentclass{article} \usepackage{amsmath} \begin{document} \displaystyle H^{'} = - \sum_{}^{}\left( \text{pi}\ln\text{pi} \right) \end{document} ]]></tex-math></inline-formula></p><p>Where<italic> pi</italic>​ is the proportion of individuals of species i relative to the total number of individuals. The calculation was performed using the <italic>R</italic> statistical software, specifically the 'vegan' package.</p><p>To investigate the relationship between waterbird abundance and environmental factors, a simple linear regression analysis was performed. The dependent variable was waterbird abundance, and the independent variables were water hyacinth coverage (as a percentage) and water depth (in meters). The analysis was conducted using R software, with a significance level set at p&lt;0.05. Pearson's correlation coefficient (r) was also calculated to assess the strength of the relationships. The species identification was verified by two independent ornithologists based on the collected photographs and behavioral descriptions to ensure accuracy.</p></sec></sec><sec><title>Results and discussion</title><sec><title>1. Results</title><sec><title>1.1. Species Composition</title><p>The survey conducted from February to October 2025 identified eight species of waterbirds within the Tondano Lake ecosystem, representing three families: Ardeidae (six species), Recurvirostridae (one species), and Alcedinidae (one species). <xref ref-type="table" rid="table-1">Table 1</xref> lists the species recorded. Compared to historical data from North Sulawesi wetlands which typically harbor greater avian richness <xref ref-type="bibr" rid="BIBR-19">(Santoso et al., 2019)</xref>, the presence of only eight species suggests a simplified community structure potentially linked to habitat loss. This limited taxonomic representation indicates a significant reduction in ecological niches compared to undisturbed freshwater habitats in the region. Representative waterbird species recorded during the study are shown in<xref ref-type="fig" rid="figure-2">Figure 2</xref>a-2h.</p><table-wrap id="table-1" ignoredToc=""><label>Table 1</label><caption><p>Identified Waterbird Species in Tondano Lake, North Sulawesi, Indonesia</p></caption><table frame="box" rules="all"><thead><tr><th valign="top" align="left" colspan="1"><bold>No</bold></th><th valign="top" align="left" colspan="1"><bold>Local Name</bold></th><th valign="top" align="left" colspan="1"><bold>Scientific Name</bold></th><th align="left" colspan="1" valign="top"><bold>Family</bold></th></tr></thead><tbody><tr><td valign="top" align="left" colspan="1">1</td><td valign="top" align="left" colspan="1">Kuntul Besar</td><td valign="top" align="left" colspan="1"><italic>Egretta alba</italic></td><td valign="top" align="left" colspan="1">Ardeidae</td></tr><tr><td valign="top" align="left" colspan="1">2</td><td valign="top" align="left" colspan="1">Kuntul Kecil</td><td valign="top" align="left" colspan="1"><italic>Egretta garzetta</italic></td><td valign="top" align="left" colspan="1">Ardeidae</td></tr><tr><td colspan="1" valign="top" align="left">3</td><td align="left" colspan="1" valign="top">Kuntul Kerbau</td><td align="left" colspan="1" valign="top"><italic>Bubulcus ibis</italic></td><td valign="top" align="left" colspan="1">Ardeidae</td></tr><tr><td align="left" colspan="1" valign="top">4</td><td valign="top" align="left" colspan="1">Burung Pancang</td><td colspan="1" valign="top" align="left"><italic>Himantopus leucocephalus</italic></td><td valign="top" align="left" colspan="1">Recurvirostridae</td></tr><tr><td valign="top" align="left" colspan="1">5</td><td align="left" colspan="1" valign="top">Raja Udang</td><td valign="top" align="left" colspan="1"><italic>Alcedo atthis</italic></td><td valign="top" align="left" colspan="1">Alcedinidae</td></tr><tr><td valign="top" align="left" colspan="1">6</td><td align="left" colspan="1" valign="top">Cangak Merah</td><td align="left" colspan="1" valign="top">Ardea purpurea</td><td valign="top" align="left" colspan="1">Ardeidae</td></tr><tr><td valign="top" align="left" colspan="1">7</td><td colspan="1" valign="top" align="left">Blekok sawah</td><td valign="top" align="left" colspan="1"><italic>Ardeola speciosa</italic></td><td align="left" colspan="1" valign="top">Ardeidae</td></tr><tr><td valign="top" align="left" colspan="1">8</td><td colspan="1" valign="top" align="left">Kowak Malam Merah</td><td valign="top" align="left" colspan="1"><italic>Nycticorax caledonicus</italic></td><td valign="top" align="left" colspan="1">Ardeidae</td></tr></tbody></table></table-wrap></sec><sec><title>1.2. Abundance and seasonal variation</title><p>Across the multi-season observation period, a total of 1,864 individual waterbirds were recorded throughout Tondano Lake. Overall abundance showed a gradual increase from the early wet months (February–May) toward the dry season (June–October), with peak numbers observed between August and October, coinciding with reduced rainfall, lower water levels, and wider availability of exposed feeding grounds along the littoral zone. This seasonal rise in abundance suggests that water level fluctuation and vegetation cover strongly influence bird activity and detectability.</p><p>Among the eight recorded species, <italic>Egretta garzetta</italic> emerged as the most abundant and widely distributed species (29.1%), indicating its adaptability to both shallow and moderately vegetated habitats. In contrast, <italic>Alcedo atthis</italic> was the least common (4.8%), reflecting its preference for clear, unobstructed waters that were often limited during the observation period. The presence of both resident and partially migratory species throughout the survey further demonstrates the ecological importance of Tondano Lake as a year round foraging and nesting habitat. The monthly abundance patterns and corresponding relative proportions of each species are detailed in <xref ref-type="table" rid="table-l6u6ag">Table 2</xref>.</p><table-wrap id="table-l6u6ag" ignoredToc=""><label>Table 2</label><caption><p>Monthly abundance of waterbirds recorded at Tondano Lake, North Sulawesi, from February to October 2025.</p></caption><table frame="box" rules="all"><thead><tr><th valign="top" align="left" colspan="1"><bold>Species</bold></th><th align="left" colspan="1" valign="top"><bold>Family</bold></th><th align="left" colspan="1" valign="top"><bold>Feb</bold></th><th align="left" colspan="1" valign="top"><bold>Mar</bold></th><th valign="top" align="left" colspan="1"><bold>Apr</bold></th><th align="left" colspan="1" valign="top"><bold>May</bold></th><th colspan="1" valign="top" align="left"><bold>Jun</bold></th><th align="left" colspan="1" valign="top"><bold>Jul</bold></th><th align="left" colspan="1" valign="top"><bold>Aug</bold></th><th align="left" colspan="1" valign="top"><bold>Sep</bold></th><th align="left" colspan="1" valign="top"><bold>Oct</bold></th><th align="left" colspan="1" valign="top"><bold>Total</bold></th><th valign="top" align="left" colspan="1"><bold>Mean ± SD</bold></th><th align="left" colspan="1" valign="top"><bold>Rel. Abund. (%)</bold></th></tr></thead><tbody><tr><td align="left" colspan="1" valign="top"><italic>Egretta alba</italic></td><td valign="middle" align="center" colspan="1" rowspan="6"><bold>Ardeidae</bold></td><td valign="top" align="left" colspan="1">21</td><td align="left" colspan="1" valign="top">23</td><td colspan="1" valign="top" align="left">26</td><td valign="top" align="left" colspan="1">26</td><td align="left" colspan="1" valign="top">27</td><td valign="top" align="left" colspan="1">28</td><td align="left" colspan="1" valign="top">30</td><td valign="top" align="left" colspan="1">28</td><td colspan="1" valign="top" align="left">28</td><td valign="top" align="left" colspan="1">237</td><td valign="top" align="left" colspan="1">26.3 ± 2.8</td><td valign="top" align="left" colspan="1">12.7</td></tr><tr><td valign="top" align="left" colspan="1"><italic>Egretta garzetta</italic></td><td align="left" colspan="1" valign="top">48</td><td valign="top" align="left" colspan="1">54</td><td valign="top" align="left" colspan="1">56</td><td valign="top" align="left" colspan="1">56</td><td valign="top" align="left" colspan="1">60</td><td valign="top" align="left" colspan="1">64</td><td align="left" colspan="1" valign="top">69</td><td align="left" colspan="1" valign="top">68</td><td valign="top" align="left" colspan="1">68</td><td valign="top" align="left" colspan="1">543</td><td align="left" colspan="1" valign="top">60.3 ± 7.0</td><td valign="top" align="left" colspan="1">29.1</td></tr><tr><td align="left" colspan="1" valign="top"><italic>Bubulcus ibis</italic></td><td valign="top" align="left" colspan="1">23</td><td colspan="1" valign="top" align="left">25</td><td align="left" colspan="1" valign="top">26</td><td align="left" colspan="1" valign="top">34</td><td align="left" colspan="1" valign="top">33</td><td align="left" colspan="1" valign="top">31</td><td valign="top" align="left" colspan="1">33</td><td colspan="1" valign="top" align="left">27</td><td valign="top" align="left" colspan="1">27</td><td align="left" colspan="1" valign="top">259</td><td valign="top" align="left" colspan="1">28.8 ± 3.7</td><td colspan="1" valign="top" align="left">13.9</td></tr><tr><td valign="top" align="left" colspan="1"><italic>Ardea purpurea</italic></td><td valign="top" align="left" colspan="1">17</td><td colspan="1" valign="top" align="left">18</td><td valign="top" align="left" colspan="1">20</td><td valign="top" align="left" colspan="1">16</td><td valign="top" align="left" colspan="1">18</td><td align="left" colspan="1" valign="top">19</td><td colspan="1" valign="top" align="left">20</td><td align="left" colspan="1" valign="top">22</td><td valign="top" align="left" colspan="1">23</td><td valign="top" align="left" colspan="1">173</td><td align="left" colspan="1" valign="top">19.2 ± 2.2</td><td colspan="1" valign="top" align="left">9.3</td></tr><tr><td valign="top" align="left" colspan="1"><italic>Ardeola speciosa</italic></td><td valign="top" align="left" colspan="1">19</td><td valign="top" align="left" colspan="1">22</td><td align="left" colspan="1" valign="top">22</td><td valign="top" align="left" colspan="1">20</td><td colspan="1" valign="top" align="left">22</td><td valign="top" align="left" colspan="1">25</td><td valign="top" align="left" colspan="1">26</td><td align="left" colspan="1" valign="top">25</td><td align="left" colspan="1" valign="top">22</td><td colspan="1" valign="top" align="left">203</td><td valign="top" align="left" colspan="1">22.6 ± 2.3</td><td valign="top" align="left" colspan="1">10.9</td></tr><tr><td align="left" colspan="1" valign="top"><italic>Nycticorax caledonicus</italic></td><td valign="top" align="left" colspan="1">11</td><td valign="top" align="left" colspan="1">13</td><td valign="top" align="left" colspan="1">14</td><td align="left" colspan="1" valign="top">15</td><td valign="top" align="left" colspan="1">17</td><td valign="top" align="left" colspan="1">18</td><td valign="top" align="left" colspan="1">18</td><td valign="top" align="left" colspan="1">17</td><td align="left" colspan="1" valign="top">16</td><td align="left" colspan="1" valign="top">139</td><td align="left" colspan="1" valign="top">15.4 ± 2.3</td><td valign="top" align="left" colspan="1">7.5</td></tr><tr><td align="left" colspan="1" valign="top"><italic>Himantopus leucocephalus</italic></td><td valign="top" align="left" colspan="1"><bold>Recurvirostridae</bold></td><td valign="top" align="left" colspan="1">19</td><td align="left" colspan="1" valign="top">21</td><td align="left" colspan="1" valign="top">24</td><td valign="top" align="left" colspan="1">24</td><td align="left" colspan="1" valign="top">25</td><td align="left" colspan="1" valign="top">26</td><td colspan="1" valign="top" align="left">28</td><td valign="top" align="left" colspan="1">27</td><td colspan="1" valign="top" align="left">26</td><td valign="top" align="left" colspan="1">220</td><td valign="top" align="left" colspan="1">24.4 ± 3.0</td><td align="left" colspan="1" valign="top">11.8</td></tr><tr><td colspan="1" valign="top" align="left"><italic>Alcedo atthis</italic></td><td valign="top" align="left" colspan="1"><bold>Alcedinidae</bold></td><td align="left" colspan="1" valign="top">9</td><td valign="top" align="left" colspan="1">10</td><td valign="top" align="left" colspan="1">9</td><td valign="top" align="left" colspan="1">6</td><td valign="top" align="left" colspan="1">10</td><td align="left" colspan="1" valign="top">10</td><td align="left" colspan="1" valign="top">11</td><td align="left" colspan="1" valign="top">12</td><td align="left" colspan="1" valign="top">13</td><td align="left" colspan="1" valign="top">90</td><td valign="top" align="left" colspan="1">10.0 ± 2.0</td><td align="left" colspan="1" valign="top">4.8</td></tr><tr><td align="left" colspan="1" valign="top"><bold>Grand Total (all species)</bold></td><td valign="top" align="left" colspan="1"></td><td align="left" colspan="1" valign="top"><bold>167</bold></td><td valign="top" align="left" colspan="1"><bold>186</bold></td><td align="left" colspan="1" valign="top"><bold>197</bold></td><td valign="top" align="left" colspan="1"><bold>197</bold></td><td align="left" colspan="1" valign="top"><bold>212</bold></td><td valign="top" align="left" colspan="1"><bold>221</bold></td><td align="left" colspan="1" valign="top"><bold>235</bold></td><td valign="top" align="left" colspan="1"><bold>226</bold></td><td valign="top" align="left" colspan="1"><bold>223</bold></td><td colspan="1" valign="top" align="left"><bold>1,864</bold></td><td align="left" colspan="1" valign="top"><bold>207 ± 15.6</bold></td><td align="left" colspan="1" valign="top"><bold>100</bold></td></tr><tr><td valign="top" align="left" colspan="1"><bold>Seasonal totals:</bold></td><td align="left" colspan="1" valign="top"></td><td valign="top" align="left" colspan="1"></td><td valign="top" align="left" colspan="1"></td><td valign="top" align="left" colspan="1"></td><td valign="top" align="left" colspan="1"></td><td align="left" colspan="1" valign="top"></td><td valign="top" align="left" colspan="1"></td><td valign="top" align="left" colspan="1"></td><td valign="top" align="left" colspan="1"></td><td valign="top" align="left" colspan="1"></td><td valign="top" align="left" colspan="1"></td><td valign="top" align="left" colspan="1"></td><td colspan="1" valign="top" align="left"></td></tr><tr><td align="left" colspan="1" valign="top"><bold>Wet Season (Feb-May)</bold></td><td align="center" colspan="13" valign="top"><bold>747</bold></td></tr><tr><td valign="top" align="left" colspan="1"><bold>Dry Season (Jun-Oct)</bold></td><td align="center" colspan="13" valign="top"><bold>1,117</bold></td></tr></tbody></table><table-wrap-foot><p>Note: Values represent the number of individual birds observed per month. Wet season = February–May; Dry season = June–October. Relative abundance (%) was calculated from the total individuals across all months. Bold values indicate peak counts during the dry season.</p></table-wrap-foot></table-wrap></sec><sec><title>1.3. Species Diversity</title><p>Based on the total individual counts obtained from February to October 2025, the Shannon-Wiener diversity index (H′) for the waterbird community was calculated to be 1.92. According to the criteria established by <xref ref-type="bibr" rid="BIBR-15">(Magurran, 1988)</xref>, this value represents a moderate level of species diversity, indicating a community with balanced species representation but potential ecological stress. No abrupt shifts in species composition occurred between months, although minor fluctuations in abundance were observed across seasons. <xref ref-type="table" rid="table-3">Tabel 3</xref> presents the detailed calculation of the Shannon-Wiener index, ensuring transparency and reproducibility of the results</p><table-wrap id="table-3" ignoredToc=""><label>Tabel 3</label><caption><p>Shannon-Wiener Diversity Index Calculation</p></caption><table frame="box" rules="all"><thead><tr><th align="left" colspan="1" valign="top"><bold>No</bold></th><th valign="top" align="left" colspan="1"><bold>Species</bold></th><th valign="top" align="left" colspan="1"><bold>ni</bold></th><th align="left" colspan="1" valign="top"><bold>pi</bold></th><th valign="top" align="left" colspan="1"><bold>ln (pi)</bold></th><th align="left" colspan="1" valign="top"><bold>pi ln (pi)</bold></th></tr></thead><tbody><tr><td valign="top" align="left" colspan="1">1</td><td align="left" colspan="1" valign="top"><italic>Egretta alba</italic></td><td align="left" colspan="1" valign="top">237</td><td align="left" colspan="1" valign="top">0.127</td><td align="left" colspan="1" valign="top">−2.066</td><td valign="top" align="left" colspan="1">−0.263</td></tr><tr><td valign="top" align="left" colspan="1">2</td><td align="left" colspan="1" valign="top"><italic>Egretta garzetta</italic></td><td align="left" colspan="1" valign="top">543</td><td valign="top" align="left" colspan="1">0.291</td><td align="left" colspan="1" valign="top">−1.234</td><td align="left" colspan="1" valign="top">−0.359</td></tr><tr><td align="left" colspan="1" valign="top">3</td><td align="left" colspan="1" valign="top"><italic>Bubulcus ibis</italic></td><td valign="top" align="left" colspan="1">259</td><td align="left" colspan="1" valign="top">0.139</td><td valign="top" align="left" colspan="1">−1.973</td><td align="left" colspan="1" valign="top">−0.274</td></tr><tr><td align="left" colspan="1" valign="top">4</td><td align="left" colspan="1" valign="top"><italic>Himantopus leucocephalus</italic></td><td align="left" colspan="1" valign="top">220</td><td valign="top" align="left" colspan="1">0.118</td><td valign="top" align="left" colspan="1">−2.138</td><td valign="top" align="left" colspan="1">−0.252</td></tr><tr><td valign="top" align="left" colspan="1">5</td><td colspan="1" valign="top" align="left"><italic>Alcedo atthis</italic></td><td valign="top" align="left" colspan="1">90</td><td valign="top" align="left" colspan="1">0.048</td><td valign="top" align="left" colspan="1">−3.045</td><td valign="top" align="left" colspan="1">−0.146</td></tr><tr><td valign="top" align="left" colspan="1">6</td><td colspan="1" valign="top" align="left"><italic>Ardea purpurea</italic></td><td align="left" colspan="1" valign="top">173</td><td valign="top" align="left" colspan="1">0.093</td><td valign="top" align="left" colspan="1">−2.377</td><td align="left" colspan="1" valign="top">−0.221</td></tr><tr><td align="left" colspan="1" valign="top">7</td><td valign="top" align="left" colspan="1"><italic>Ardeola speciosa</italic></td><td valign="top" align="left" colspan="1">203</td><td valign="top" align="left" colspan="1">0.109</td><td valign="top" align="left" colspan="1">−2.218</td><td valign="top" align="left" colspan="1">−0.243</td></tr><tr><td valign="top" align="left" colspan="1">8</td><td valign="top" align="left" colspan="1"><italic>Nycticorax caledonicus</italic></td><td align="left" colspan="1" valign="top">139</td><td valign="top" align="left" colspan="1">0.075</td><td align="left" colspan="1" valign="top">−2.590</td><td colspan="1" valign="top" align="left">−0.194</td></tr><tr><td colspan="1" valign="top" align="left"></td><td align="left" colspan="1" valign="top"></td><td align="left" colspan="1" valign="top">N = 1.864</td><td colspan="1" valign="top" align="left"></td><td align="left" colspan="1" valign="top">Σ (pi ln pi) = −1.918</td><td align="left" colspan="1" valign="top"></td></tr><tr><td align="left" colspan="1" valign="top"></td><td colspan="1" valign="top" align="left"></td><td colspan="1" valign="top" align="left"></td><td valign="top" align="left" colspan="1"></td><td align="left" colspan="1" valign="top">H′ = −Σ (pi ln pi) = 1.92</td><td align="left" colspan="1" valign="top"></td></tr></tbody></table></table-wrap></sec><sec><title>1.4. Relationship with environmental variables</title><p>The simple linear regression analysis revealed significant negative correlations between waterbird abundance and the two key environmental variables. A strong negative correlation was found with water hyacinth density (r=−0.68, p=0.032, r2=0.46), indicating that as the coverage of water hyacinth increases, the total abundance of waterbirds tends to decrease. Similarly, a significant negative correlation was found with water depth ( r=−0.72, p=0.019, r2=0.52), suggesting that the shoaling of the lake is also a major factor negatively impacting the waterbird population.</p><table-wrap id="table-4" ignoredToc=""><label>Table 4</label><caption><p>Correlations between Waterbird Abundance and Environmental Factors</p></caption><table rules="all" frame="box"><thead><tr><th align="left" colspan="1" valign="top"><bold>Environmental Factor</bold></th><th valign="top" align="left" colspan="1"><bold>Correlation Coefficient (r)</bold></th><th valign="top" align="left" colspan="1"><bold>P-value (p)</bold></th><th valign="top" align="left" colspan="1"><bold>Coefficient of Determination (r2)</bold></th></tr></thead><tbody><tr><td align="left" colspan="1" valign="top">Water Hyacinth Density</td><td valign="top" align="left" colspan="1">-0.68</td><td valign="top" align="left" colspan="1">0.032</td><td valign="top" align="left" colspan="1">0.46</td></tr><tr><td valign="top" align="left" colspan="1">Water Depth</td><td valign="top" align="left" colspan="1">-0.72</td><td colspan="1" valign="top" align="left">0.019</td><td valign="top" align="left" colspan="1">0.52</td></tr></tbody></table></table-wrap><fig id="figure-2" ignoredToc=""><label>Figure 2</label><caption><p>(a). <italic>Egretta alba</italic>, (b). <italic>Alcedo atthi</italic>, (c). <italic>Himantopus leucocephalus</italic>, (d). <italic>Bubulcus ibis</italic>, (e) <italic>Ardeola speciosa</italic>, (f). <italic>Ardea purpurea</italic>, (g). <italic>Egretta garzetta</italic>, (h). <italic>Nycticorax caledonicus</italic>.</p></caption><graphic mime-subtype="png" mimetype="image" xlink:href="https://journals2.ums.ac.id/bioeksperimen/article/download/15450/5374/67878"><alt-text>Image</alt-text></graphic></fig></sec></sec><sec><title>2. Discussion</title><sec><title>2.1. Waterbird diversity and abundance in Tondano Lake</title><p>The study’s finding of a Shannon-Wiener diversity index (H′ = 1.92) indicates that while the waterbird community remains functional, it has undergone significant "ecological simplification". As a noted, the recording of only eight species from three families is a critical indicator of decline when compared to historical records of Tondano Lake and surrounding wetlands which previously supported higher taxa richness <xref ref-type="bibr" rid="BIBR-19">(Santoso et al., 2019)</xref>. This value (H′ = 1.92) is markedly lower than pristine benchmarks like Lake Sentarum (H′ = 2.5; <xref ref-type="bibr" rid="BIBR-9">(Harmoko et al., 2018)</xref>), yet remains higher than severely degraded industrial wetlands in South Lampung (H′ = 1.5; <xref rid="BIBR-8" ref-type="bibr">(Iswandaru &amp; Maharany, 2025)</xref>).</p><p>This positioning confirms that Tondano Lake is in a state of "moderate ecological stress". The dominance of Egretta garzetta (29.1%), a generalist feeder tolerant of anthropogenic disturbance, suggests that specialized species are being displaced. In contrast, the low abundance of Alcedo atthis (4.8%) a specialist visual predator reflects the direct impact of high turbidity and habitat clogging caused by Eichhornia crassipes proliferation.</p></sec><sec><title>2.2. The role of waterbirds and the impact of environmental threats</title><p>Waterbirds in Tondano Lake function as “ecosystem guardians,” yet their effectiveness is being compromised by synergistic environmental threats. The strong negative correlations between abundance and hyacinth density (r = −0.68) and water depth (r = −0.72) provide the evidence requested to support the claim of degradation. The current richness is a fraction of what is expected in healthy Indonesian freshwater ecosystems, where less-disturbed lakes often support twice the number of families found in this study.</p><p>The expansion of water hyacinth, now covering 32.5% of the lake, acts as a physical barrier that has narrowed the ecological niche to only a few resilient species. This is coupled with sedimentation, which has reduced the mean depth to 1.9 ± 0.7 m. These factors create a self-reinforcing "degradation loop" where nutrient enrichment accelerates weed growth, which in turn traps more sediment, further shallowing the lake and driving out deep-water foragers like Nycticorax caledonicus.</p></sec><sec><title>2,3, Implication for conservation and policy</title><p>This research underscores the significance of waterbirds as reliable bioindicators for monitoring ecosystem health in freshwater systems <xref ref-type="bibr" rid="BIBR-1">(Amat &amp; Green, 2009)</xref>. Conservation measures for Tondano Lake should therefore include: (1) Active control of Eichhornia crassipes-through mechanical removal and biological agents such as Neochetina eichhorniae <xref ref-type="bibr" rid="BIBR-3">(Center et al., 2014)</xref>; (2) Sedimentation management-via sustainable land use, riparian vegetation restoration, and targeted dredging in critical habitats <xref rid="BIBR-14" ref-type="bibr">(Kumar &amp; Pattnaik, 2012)</xref>; and (3) Community-based monitoring programs, involving citizen science and local stewardship to sustain long term ecological data <xref ref-type="bibr" rid="BIBR-13">(Kingsford et al., 2016)</xref>. Such integrated management aligns with global frameworks for freshwater biodiversity conservation (<xref ref-type="bibr" rid="BIBR-4">(Dudgeon et al., 2006)</xref>; <xref ref-type="bibr" rid="BIBR-6">(Green &amp; Elmberg, 2014)</xref>) and reflects the need for watershed-based ecosystem approaches.</p></sec></sec></sec><sec><title>Conclusion</title><p>This study demonstrates that Tondano Lake supports a moderate level of waterbird diversity, as reflected by a Shannon-Wiener index of $H' = 1.92. The eight recorded species, primarily from the family Ardeidae, exhibit varied ecological responses to environmental conditions. This species richness and diversity represent a simplified community structure when compared to historical regional data (H' = 2.5; Harmoko et al., 2018), providing evidence of ongoing ecological decline. The dominance of the adaptable <italic>Egretta garzetta</italic> alongside the low abundance of the sensitive specialist <italic>Alcedo atthis</italic> provides a clear signal of ecological stress within the lake ecosystem.</p><p>Waterbirds function as key ecosystem guardians, contributing to ecological stability by regulating aquatic populations, dispersing plant seeds, and serving as sensitive bioindicators of environmental quality. The regression analysis revealed significant negative correlations between waterbird abundance and both water hyacinth density (r = -0.68, p = 0.032) and sedimentation (r = -0.72, p = 0.019), demonstrating the dual and interacting impacts of these environmental pressures. These findings establish a contribution to the body of research on Indonesian freshwater ecosystems by linking waterbird population dynamics with specific, integrated environmental threats.</p><p>The results emphasize that water hyacinth invasion and sedimentation are interconnected, forming a self-reinforcing feedback loop that accelerates ecological degradation. Ensuring the long-term sustainability of the Tondano Lake ecosystem therefore requires integrated conservation strategies that address both stressors simultaneously. This includes actions spanning multiple scales from direct lake management to the restoration of the surrounding watershed. Future research should extend the observation period to further capture migratory dynamics and incorporate additional water-quality parameters. Such comprehensive monitoring would deepen understanding of the complex interactions between waterbirds and their changing environment, thereby strengthening conservation planning for Tondano Lake and comparable tropical freshwater systems.</p></sec><sec><title>Author Statements</title><p><bold>Acknowledgements and funding statements: </bold>The authors extend their sincere gratitude to the North Sulawesi BKSDA, the Minahasa Regency Government, and the local community members of Tondano Lake for their invaluable assistance with species identification. Logistical support was provided by the Universitas Negeri Manado. This research was funded by an internal research grant from the Universitas Negeri Manado in 2025.</p><p><bold>Competing of interest:</bold> The authors declare that there are no financial or personal relationships that could be construed as a potential conflict of interest.</p><p><bold>Author’s contributions:</bold> Rievo Djarang: conceptualization, research design, data collection, data analysis, and manuscript writing. Gerry Harindah: data analysis, critical revision of the manuscript, and final manuscript approval. Ni Wayan Suryani and Marthy Lingkan Stella Taulu: field support and literature review.</p><p><bold>Generative AI: </bold>Not applicable</p><p><bold>Data availability:</bold> The raw data supporting the conclusions of this article, including the monthly abundance counts, Shannon-Wiener index calculations, and environmental variable measurements (water hyacinth density and water depth) for Tondano Lake, will be made available by the authors, without undue reservation, to any qualified researcher.</p></sec></body><back><ref-list><title>References</title><ref id="BIBR-1"><element-citation publication-type="chapter"><article-title>Waterbirds as Bioindicators of Environmental Conditions</article-title><source>Biological Monitoring in Freshwater Habitats</source><person-group 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