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<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title-group>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
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<issn pub-type="epub">2296-701X</issn>
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<publisher-name>Frontiers Media S.A.</publisher-name>
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<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2025.1644458</article-id>
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<subj-group subj-group-type="heading">
<subject>Brief Research Report</subject>
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<title-group>
<article-title>Optimal control of <italic>Lantana camara</italic>: an entropy-based sustainable strategy</article-title>
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<contrib-group>
<contrib contrib-type="author" equal-contrib="yes">
<name><surname>Kumar</surname><given-names>Shyam</given-names></name>
<xref ref-type="author-notes" rid="fn003"><sup>&#x2020;</sup></xref>
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<name><surname>Mishra</surname><given-names>Preet</given-names></name>
<xref ref-type="author-notes" rid="fn003"><sup>&#x2020;</sup></xref>
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<name><surname>Singh</surname><given-names>R. K. Brojen</given-names></name>
<xref ref-type="corresp" rid="c001"><sup>*</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/340915/overview"/>
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<aff id="aff1"><institution>School of Computational and Integrative Sciences, Jawaharlal Nehru University</institution>, <city>New Delhi</city>,&#xa0;<country country="in">India</country></aff>
<author-notes>
<corresp id="c001"><label>*</label>Correspondence: R. K. Brojen Singh, <email xlink:href="mailto:brojen@jnu.ac.in">brojen@jnu.ac.in</email></corresp>
<fn fn-type="equal" id="fn003">
<label>&#x2020;</label>
<p>These authors have contributed equally to this work</p></fn>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2026-01-22">
<day>22</day>
<month>01</month>
<year>2026</year>
</pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2025</year>
</pub-date>
<volume>13</volume>
<elocation-id>1644458</elocation-id>
<history>
<date date-type="received">
<day>10</day>
<month>06</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>17</day>
<month>12</month>
<year>2025</year>
</date>
<date date-type="rev-recd">
<day>19</day>
<month>11</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2026 Kumar, Mishra and Singh.</copyright-statement>
<copyright-year>2026</copyright-year>
<copyright-holder>Kumar, Mishra and Singh</copyright-holder>
<license>
<ali:license_ref start_date="2026-01-22">https://creativecommons.org/licenses/by/4.0/</ali:license_ref>
<license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License (CC BY)</ext-link>. The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</license-p>
</license>
</permissions>
<abstract>
<p>Framing control policies to mitigate the impact of invasive plants on indigenous biodiversity within the Sustainable Development Goals (SDG) framework is the primary objective of this work. Using reported ecological dynamics of the invasive species <italic>Lantana camara</italic>, we develop a minimal three-species network model, where each node follows generalized Lotka-Volterra (GLV) dynamical equations. Employing Lie algebra and network control theory, we establish the model&#x2019;s controllability and accessibility criteria. Through nonlinear optimization programming, we derive sustainable policies for controlling abundances of <italic>Lantana camara</italic>. We have also used Shannon entropy as an indicator to assess the sustainability of these optimal policies. The analysis of the sensitivity measured using this technique reveals that the control strategy is critically dependent on the ratio of the intrinsic growth rates of the <italic>Lantana camara</italic> and the control plant. Thus, we get a modular algorithmic decision support mechanism for designing control policies to manage <italic>Lantana camara</italic> abundances.</p>
</abstract>
<kwd-group>
<kwd><italic>Lantana camara</italic></kwd>
<kwd>generalized Lotka-Volterra</kwd>
<kwd>Shannon entropy</kwd>
<kwd>sustainable development goals</kwd>
<kwd>sensitivity analysis</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declared that financial support was received for this work and/or its publication. PM would like to acknowledge CSIR Direct-SRF(via File no. 09/0263(18990)/2024-EMR-I), UGC and JNU for financial support and SK would like to acknowledge ICMR-SRF(via File No: 3/1/3/9/M/2022-NCD-II), JNU and UGC for financial support during the work.</funding-statement>
</funding-group>
<counts>
<fig-count count="5"/>
<table-count count="0"/>
<equation-count count="7"/>
<ref-count count="34"/>
<page-count count="10"/>
<word-count count="6372"/>
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<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Models in Ecology and Evolution</meta-value>
</custom-meta>
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</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Strategic control of invasive plants in an ecosystem is a global issue, as they are one of the major causes of biodiversity loss, environmental damage, and a threat to endangered native species. In this work, we focus on controlling the invasive species <italic>Lantana camara</italic>, which is one of the primary drivers of imbalance between local and global plant diversity (<xref ref-type="bibr" rid="B14">Kohli et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B31">Venkataraman, 2015</xref>; <xref ref-type="bibr" rid="B10">Hiremath et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B28">Sivakumar et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B15">Kumar et&#xa0;al., 2021</xref>). This complex process of invasion leads to a state of dominance by the invasive species in the ecosystem, which we have termed as <italic>diseased state</italic>. We also know that <italic>Lantana camara</italic> promotes the growth of deleterious microbes, which decrease the population of beneficial microbes in the soil (<xref ref-type="bibr" rid="B7">Fan et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B15">Kumar et&#xa0;al., 2021</xref>) and implicitly hinder the growth of other plants. The main goal of this work is to propose a model and analyze it to guide the diseased ecosystem to a possible desired healthy state (<xref ref-type="bibr" rid="B21">Montoya et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B11">Jones et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B34">Weidlich et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B27">Singh et&#xa0;al., 2022</xref>) in a sustainable manner as suggested in the United Nations Sustainable Development Goals (UN-SDG) (<xref ref-type="bibr" rid="B30">United Nations General Assembly, 2015</xref>) using suitable external inputs. We have thus defined two-fold objectives: first, to manage the abundance of the <italic>Lantana camara</italic> population, and second, to provide a possible mathematical framework for addressing important sustainability issues related to controlling invasive plants in general. We derive our theoretical model systematically by following some of the basic ecosystem management rules.</p>
<p>The basic ecological observations incorporated into our model framework are as follows. For our control objectives, we introduce a native plant into the ecosystem, which helps control the abundance of <italic>Lantana camara</italic> and also increases soil microbe abundance. Some studies demonstrate how to utilize plant-plant interactions to regulate the abundances of plants within a spatial distance threshold from each other in the ecosystem (<xref ref-type="bibr" rid="B24">Sakai and Suzuki, 1954</xref>; <xref ref-type="bibr" rid="B12">Joshi, 1991</xref>). This interaction also has its own benefits as certain plants are known to improve soil quality by increasing and promoting the growth of good microbes in the soil (<xref ref-type="bibr" rid="B9">Hamilton and Frank, 2001</xref>; <xref ref-type="bibr" rid="B33">Wardle et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B26">Singh et&#xa0;al., 2014</xref>). It has been found in certain field studies (<xref ref-type="bibr" rid="B12">Joshi, 1991</xref>) that some non-invasive plants like <italic>Ipomoea staphylina</italic>, <italic>Mimosa pudica</italic> (Mimosa has also added benefits of being a leguminous plant) can act as being control plants, which further provides a practical basis to our theoretical modeling and analysis. From these basic important observations, we propose a minimal model, which consists of these two fundamental interactions i.e. plant-plant interactions and the plant-microbe interactions, and analyze this model for possible sustainable control strategies of invasive species Lantana camara. The dynamics of each nodal variable is defined by generalized Lotka-Volterra (GLV) model. The control strategies in the model are incorporated using nonlinear control theory (<xref ref-type="bibr" rid="B17">Liu and Barab&#xe1;si, 2016</xref>; <xref ref-type="bibr" rid="B2">Angulo et&#xa0;al., 2019</xref>). This is a basic minimal model specially designed to incorporate control theoretical technique and can be extended to more general interactions.</p>
<p>The behavior of such ecological systems can be well studied within the framework of population dynamics where the growth of the population are represented by dynamical systems. In these models, we study how the population states of species of the system change over time due to interactions within individuals of the same species and with individuals of other species present in the ecosystem. Thus we set up an optimal control problem whereby we want to find the suitable optimal control inputs that can steer the population state, evolving under the proposed GLV dynamical system, to a desired state (<xref ref-type="bibr" rid="B13">Kalman and Soc,1963</xref>; <xref ref-type="bibr" rid="B3">Brockett, 1973</xref>; <xref ref-type="bibr" rid="B22">Nijmeijer and van der Schaft, 1990</xref>; <xref ref-type="bibr" rid="B6">Conte et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B18">Liu et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B4">Brockett, 2015</xref>; <xref ref-type="bibr" rid="B17">Liu and Barab&#xe1;si, 2016</xref>; <xref ref-type="bibr" rid="B2">Angulo et&#xa0;al., 2019</xref>).</p>
</sec>
<sec id="s2">
<label>2</label>
<title>Model description</title>
<p>Components of an ecosystem such as plants, animals, soil microbes, etc. interact in a complex manner as shown in <xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1a</bold></xref> and can be represented by a network as shown in <xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1b</bold></xref>. In such networks defined by a graph <inline-formula>
<mml:math display="inline" id="im1"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi>V</mml:mi><mml:mo>,</mml:mo><mml:mi>E</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, ecological agents are represented by the nodes (<bold>V</bold>) and interactions by edges (<bold>E</bold>) among them (<xref ref-type="bibr" rid="B5">Cohen, 1968</xref>; <xref ref-type="bibr" rid="B23">Roberts, 1978</xref>; <xref ref-type="bibr" rid="B2">Angulo et&#xa0;al., 2019</xref>). In our study, we represent the dynamics of the individual nodes by GLV model (<xref ref-type="bibr" rid="B20">May and Leonard, 1975</xref>). Once we construct the network dynamical system, we then ask how we can steer the system in a finite amount of time to achieve a certain desired state by suitable input signals to the chosen nodes (ecological agents) of the network as shown schematically in <xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1b</bold></xref>. For the analysis and solution to this situation, network theory-based controllability (<xref ref-type="bibr" rid="B18">Liu et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B17">Liu and Barab&#xe1;si, 2016</xref>; <xref ref-type="bibr" rid="B2">Angulo et&#xa0;al., 2019</xref>) predictions have been systematically implemented in this work.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Schematic of the ecological system, its modeling. <bold>(a)</bold> Shows the eco-system of the three interacting species: <italic>Lantana camara</italic>, control plants, and microbes. <bold>(b)</bold> A network representation of GLV-type interactions among ecological agents. The abundances of <italic>Lantana camara x</italic><sub>1</sub>(<italic>t</italic>), control plants <italic>x</italic><sub>2</sub>(<italic>t</italic>), and microbes <italic>x</italic><sub>3</sub>(<italic>t</italic>) as nodes in the network representing the dynamical system. Control inputs <italic>u</italic><sub>1</sub>(<italic>t</italic>) (reducing <italic>Lantana camara</italic>) and <italic>u</italic><sub>2</sub>(<italic>t</italic>) (enhancing control plants) act on <italic>x</italic><sub>1</sub> and <italic>x</italic><sub>2</sub> respectively, with arrows indicating the nodes at which the control is applied.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1644458-g001.tif">
<alt-text content-type="machine-generated">Diagram showing two parts: (a) illustrates an ecosystem with Lantana camara labeled as \(x_1\), a native plant as \(x_2\), both growing in soil, and microbes labeled \(x_3\) in the soil. (b) depicts a network model with nodes \(x_1\), \(x_2\), \(x_3\) connected by arrows, influenced by inputs \(u_1\) and \(u_2\).</alt-text>
</graphic></fig>
<p>The minimal ecological model we propose in this work is a three-node network, whose individual dynamics is governed by nonlinear GLV (<xref ref-type="bibr" rid="B8">Goel et&#xa0;al., 1971</xref>; <xref ref-type="bibr" rid="B20">May and Leonard, 1975</xref>; <xref ref-type="bibr" rid="B19">Mambuca et&#xa0;al., 2022</xref>), with the interacting edges built from experimental observations (<xref ref-type="bibr" rid="B31">Venkataraman, 2015</xref>; <xref ref-type="bibr" rid="B28">Sivakumar et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B15">Kumar et&#xa0;al., 2021</xref>). We define the state vector <inline-formula>
<mml:math display="inline" id="im2"><mml:mrow><mml:mi>x</mml:mi><mml:mo>=</mml:mo><mml:msup><mml:mrow><mml:mo>[</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub><mml:mo>]</mml:mo></mml:mrow><mml:mi>T</mml:mi></mml:msup></mml:mrow></mml:math></inline-formula> representing the abundances of <italic>Lantana camara</italic> (<inline-formula>
<mml:math display="inline" id="im3"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>), control-plants (<inline-formula>
<mml:math display="inline" id="im4"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>), and soil microbes (<inline-formula>
<mml:math display="inline" id="im5"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>). The interaction strength matrix (the adjacency matrix) <inline-formula>
<mml:math display="inline" id="im6"><mml:mrow><mml:mi>A</mml:mi><mml:mo>=</mml:mo><mml:msub><mml:mrow><mml:mo>[</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>]</mml:mo></mml:mrow><mml:mrow><mml:mn>3</mml:mn><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> with <inline-formula>
<mml:math display="inline" id="im7"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> quantifies the relative strength of interactions between the nodes (the participating ecological agents) in the GLV model. The uncontrolled dynamical equations are given by</p>
<disp-formula id="eq1"><label>(1)</label>
<mml:math display="block" id="M1"><mml:mrow><mml:mtable><mml:mtr columnalign="left"><mml:mtd columnalign="left"><mml:mfrac><mml:mrow><mml:mi>d</mml:mi><mml:mi>x</mml:mi></mml:mrow><mml:mrow><mml:mi>d</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:mfrac><mml:mo>=</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mtext>diag</mml:mtext><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>[</mml:mo><mml:mi>A</mml:mi><mml:mi>x</mml:mi><mml:mo>+</mml:mo><mml:mi>r</mml:mi><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:mtext>&#xa0;&#xa0;</mml:mtext><mml:mi>A</mml:mi><mml:mo>=</mml:mo><mml:mo>[</mml:mo><mml:mtable equalrows="true" equalcolumns="true"><mml:mtr><mml:mtd><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>11</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>12</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>13</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>21</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>22</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>23</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>31</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>32</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>33</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mtr></mml:mtable><mml:mo>]</mml:mo><mml:mo>,</mml:mo></mml:mtd></mml:mtr><mml:mtr columnalign="left"><mml:mtd columnalign="left"><mml:mtext>&#xa0;&#xa0;diag</mml:mtext><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mo>[</mml:mo><mml:mtable equalrows="true" equalcolumns="true"><mml:mtr><mml:mtd><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:mtd><mml:mtd><mml:mn>0</mml:mn></mml:mtd><mml:mtd><mml:mn>0</mml:mn></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mn>0</mml:mn></mml:mtd><mml:mtd><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mtd><mml:mtd><mml:mn>0</mml:mn></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mn>0</mml:mn></mml:mtd><mml:mtd><mml:mn>0</mml:mn></mml:mtd><mml:mtd><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub></mml:mrow></mml:mtd></mml:mtr></mml:mtable><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:mtext>&#xa0;&#xa0;</mml:mtext><mml:mi>r</mml:mi><mml:mo>=</mml:mo><mml:mo>[</mml:mo><mml:mtable equalrows="true" equalcolumns="true"><mml:mtr><mml:mtd><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>3</mml:mn></mml:msub></mml:mrow></mml:mtd></mml:mtr></mml:mtable><mml:mo>]</mml:mo></mml:mtd></mml:mtr></mml:mtable></mml:mrow></mml:math>
</disp-formula>
<p>where <inline-formula>
<mml:math display="inline" id="im8"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> (<inline-formula>
<mml:math display="inline" id="im9"><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn>1</mml:mn><mml:mo>,</mml:mo><mml:mn>2</mml:mn><mml:mo>,</mml:mo><mml:mn>3</mml:mn></mml:mrow></mml:math></inline-formula>) represent the intrinsic growth rates adjusted for carrying capacities of the ecological agents and where <inline-formula>
<mml:math display="inline" id="im10"><mml:mrow><mml:mtext>diag&#xa0;</mml:mtext><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> denotes the diagonal matrix with diagonal entries from vector <inline-formula>
<mml:math display="inline" id="im11"><mml:mi>x</mml:mi></mml:math></inline-formula>. The state variables <inline-formula>
<mml:math display="inline" id="im12"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im13"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, and <inline-formula>
<mml:math display="inline" id="im14"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> represent normalized population densities (biomass fractions) and are dimensionless quantities satisfying <inline-formula>
<mml:math display="inline" id="im15"><mml:mrow><mml:mn>0</mml:mn><mml:mo>&#x2264;</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>&#x2264;</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:math></inline-formula>. The biomass proportions are normalized such that the total biomass sums to unity, with plants contributing 85% and microbes 15% to the total, ensuring the model accounts for relative biomass contributions while maintaining ecological relevance.</p>
<p>The time variable <inline-formula>
<mml:math display="inline" id="im16"><mml:mi>t</mml:mi></mml:math></inline-formula> is measured in months. The intrinsic growth rates <inline-formula>
<mml:math display="inline" id="im17"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> have units of inverse time (<inline-formula>
<mml:math display="inline" id="im18"><mml:mrow><mml:msup><mml:mrow><mml:mtext>time</mml:mtext></mml:mrow><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>), representing per capita growth rates. The interaction coefficients <inline-formula>
<mml:math display="inline" id="im19"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> are dimensionless parameters in the interval <inline-formula>
<mml:math display="inline" id="im20"><mml:mrow><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, quantifying the relative strength of interspecific interactions. The control inputs <inline-formula>
<mml:math display="inline" id="im21"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im22"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> have units of <inline-formula>
<mml:math display="inline" id="im23"><mml:mrow><mml:msup><mml:mrow><mml:mtext>time</mml:mtext></mml:mrow><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, representing the rate of change in population density induced by external interventions such as removal/addition of biomass, habitat modification, or resource allocation. The control vector fields <inline-formula>
<mml:math display="inline" id="im24"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im25"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> are dimensionless unit vectors indicating which state variables are directly affected by the controls. This dimensional consistency ensures that all terms in the dynamical equations have units of <inline-formula>
<mml:math display="inline" id="im26"><mml:mrow><mml:mo>&#xa0;</mml:mo><mml:msup><mml:mrow><mml:mtext>time</mml:mtext></mml:mrow><mml:mrow><mml:mo>&#x2212;</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, representing rates of change of the dimensionless population densities.</p>
<p>In our model, we impose two control signals <inline-formula>
<mml:math display="inline" id="im27"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im28"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> on the abundance variables of <italic>Lantana camara</italic> (<inline-formula>
<mml:math display="inline" id="im29"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) and control-plant (<inline-formula>
<mml:math display="inline" id="im30"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>), respectively. The choice of these driver nodes is based on the maximum matching algorithm applied to our system (<xref ref-type="bibr" rid="B3">Brockett, 1973</xref>; <xref ref-type="bibr" rid="B2">Angulo et&#xa0;al., 2019</xref>), which identifies suitable nodes at which control inputs can be effectively applied. After incorporating the control input signals into <xref ref-type="disp-formula" rid="eq1">Equation 1</xref>, we have the following control equation:</p>
<disp-formula id="eq2"><label>(2)</label>
<mml:math display="block" id="M2"><mml:mrow><mml:mtable><mml:mtr columnalign="left"><mml:mtd columnalign="left"><mml:mfrac><mml:mrow><mml:mi>d</mml:mi><mml:mi>x</mml:mi></mml:mrow><mml:mrow><mml:mi>d</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:mfrac><mml:mo>=</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>+</mml:mo><mml:munderover><mml:mo>&#x2211;</mml:mo><mml:mrow><mml:mi>j</mml:mi><mml:mo>=</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mn>2</mml:mn></mml:munderover><mml:msub><mml:mi>u</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:msub><mml:mi>f</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>;</mml:mo></mml:mtd></mml:mtr><mml:mtr columnalign="left"><mml:mtd columnalign="left"><mml:mtext>&#x2003;</mml:mtext><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mo>[</mml:mo><mml:mtable equalrows="true" equalcolumns="true"><mml:mtr><mml:mtd><mml:mn>1</mml:mn></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mn>0</mml:mn></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mn>0</mml:mn></mml:mtd></mml:mtr></mml:mtable><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:mtext>&#xa0;&#xa0;</mml:mtext><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mo>[</mml:mo><mml:mtable equalrows="true" equalcolumns="true"><mml:mtr><mml:mtd><mml:mn>0</mml:mn></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mn>1</mml:mn></mml:mtd></mml:mtr><mml:mtr><mml:mtd><mml:mn>0</mml:mn></mml:mtd></mml:mtr></mml:mtable><mml:mo>]</mml:mo></mml:mtd></mml:mtr></mml:mtable></mml:mrow></mml:math>
</disp-formula>
<p>where, it was assumed that the susceptibilities (<inline-formula>
<mml:math display="inline" id="im31"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mi>j</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) of the species to input are constant and equal to one.</p>
</sec>
<sec id="s3">
<label>3</label>
<title>Methods</title>
<p>The general controllability problem for our ecological system can be defined as steering the system to reach a desired target state <inline-formula>
<mml:math display="inline" id="im32"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>D</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> from a given initial state <inline-formula>
<mml:math display="inline" id="im33"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>0</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> in finite time using appropriate control input signals <inline-formula>
<mml:math display="inline" id="im34"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>. To determine whether such steering is possible, we must analyze the system&#x2019;s accessibility and controllability properties (<xref ref-type="bibr" rid="B3">Brockett, 1973</xref>; <xref ref-type="bibr" rid="B29">Sussmann, 1987</xref>; <xref ref-type="bibr" rid="B22">Nijmeijer and van der Schaft, 1990</xref>; <xref ref-type="bibr" rid="B6">Conte et&#xa0;al., 1999</xref>). For the control-affine system given in <xref ref-type="disp-formula" rid="eq2">Equation 2</xref>, where <inline-formula>
<mml:math display="inline" id="im35"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> represents the drift dynamics and <inline-formula>
<mml:math display="inline" id="im36"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> are the control vector fields, accessibility is determined through the Lie algebra rank condition (LARC). The accessibility distribution at a point <inline-formula>
<mml:math display="inline" id="im37"><mml:mi>x</mml:mi></mml:math></inline-formula> is constructed from the vector fields and their iterated Lie brackets as shown in <xref ref-type="disp-formula" rid="eq3">Equation 3</xref>:</p>
<disp-formula id="eq3"><label>(3)</label>
<mml:math display="block" id="M3"><mml:mrow><mml:mtable><mml:mtr columnalign="left"><mml:mtd columnalign="left"><mml:mi mathvariant="script">A</mml:mi><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mtext>span</mml:mtext><mml:mo>{</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>,</mml:mo></mml:mtd></mml:mtr><mml:mtr columnalign="left"><mml:mtd columnalign="left"><mml:mo>[</mml:mo><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:mo>&#x2026;</mml:mo><mml:mo>}</mml:mo><mml:mo>,</mml:mo></mml:mtd></mml:mtr></mml:mtable></mml:mrow></mml:math>
</disp-formula>
<p>where the Lie bracket of two vector fields <italic>X</italic> and <italic>Y</italic> is defined as in <xref ref-type="disp-formula" rid="eq4">Equation 4</xref>:</p>
<disp-formula id="eq4"><label>(4)</label>
<mml:math display="block" id="M4"><mml:mrow><mml:mo>[</mml:mo><mml:mi>X</mml:mi><mml:mo>,</mml:mo><mml:mi>Y</mml:mi><mml:mo>]</mml:mo><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mfrac><mml:mrow><mml:mtext mathvariant="italic">&#x2202;Y</mml:mtext></mml:mrow><mml:mrow><mml:mtext mathvariant="italic">&#x2202;x</mml:mtext></mml:mrow></mml:mfrac><mml:mi>X</mml:mi><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>&#x2212;</mml:mo><mml:mfrac><mml:mrow><mml:mtext mathvariant="italic">&#x2202;X</mml:mtext></mml:mrow><mml:mrow><mml:mtext mathvariant="italic">&#x2202;x</mml:mtext></mml:mrow></mml:mfrac><mml:mi>Y</mml:mi><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>.</mml:mo></mml:mrow></mml:math>
</disp-formula>
<p>The system is accessible at <italic>x</italic> if the accessibility distribution spans the entire state space, i.e., <inline-formula>
<mml:math display="inline" id="im38"><mml:mrow><mml:mi>dim</mml:mi><mml:mtext>&#xa0;</mml:mtext><mml:mo>(</mml:mo><mml:mi mathvariant="script">A</mml:mi><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mi>n</mml:mi></mml:mrow></mml:math></inline-formula>, where <italic>n</italic> is the dimension of the state space. This is verified by constructing the accessibility distribution matrix given in <xref ref-type="disp-formula" rid="eq5">Equation 5</xref>:</p>
<disp-formula id="eq5"><label>(5)</label>
<mml:math display="block" id="M5"><mml:mrow><mml:mi mathvariant="script">C</mml:mi><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:mo>[</mml:mo><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:mo>[</mml:mo><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>]</mml:mo><mml:mo>,</mml:mo><mml:mo>&#x2026;</mml:mo><mml:mo>]</mml:mo><mml:mo>,</mml:mo></mml:mrow></mml:math>
</disp-formula>
<p>and checking the rank condition given in <xref ref-type="disp-formula" rid="eq6">Equation 6</xref>:</p>
<disp-formula id="eq6"><label>(6)</label>
<mml:math display="block" id="M6"><mml:mrow><mml:mtext>rank</mml:mtext><mml:mo>[</mml:mo><mml:mi mathvariant="script">C</mml:mi><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo><mml:mo>]</mml:mo><mml:mo>=</mml:mo><mml:mi>n</mml:mi><mml:mo>.</mml:mo></mml:mrow></mml:math>
</disp-formula>
<p>is crucial to distinguish between accessibility and controllability (<xref ref-type="bibr" rid="B29">Sussmann, 1987</xref>). Accessibility ensures that the reachable set from any initial state has a non-empty interior, meaning a neighborhood of any target state can be reached. Controllability, on the other hand, guarantees that any target state can be reached exactly in finite time. For control-affine systems with drift, accessibility is a necessary but not always sufficient condition for local controllability (<xref ref-type="bibr" rid="B22">Nijmeijer and van der Schaft, 1990</xref>; <xref ref-type="bibr" rid="B6">Conte et&#xa0;al., 1999</xref>). Additional conditions, such as Sussmann&#x2019;s condition for small-time local controllability, may need to be verified (<xref ref-type="bibr" rid="B29">Sussmann, 1987</xref>).</p>
<p>For our three-species ecological network dynamics with two control inputs, we analyze the accessibility and controllability properties under the constraint that all species interaction coefficients <inline-formula>
<mml:math display="inline" id="im39"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> lie in the open interval <inline-formula>
<mml:math display="inline" id="im40"><mml:mrow><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, representing non-negligible ecological interactions. The system also possesses invariant manifolds at coordinate planes where any species is absent, which restricts global controllability. A comprehensive mathematical analysis, including detailed computation of Lie brackets, rank verification, and examination of invariant sets, is provided in the <xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref> in the <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Information (SI)</bold></xref>. The biological interpretation of these mathematical properties and their implications for control design in ecological management are discussed in the Results section.</p>
<sec id="s3_1">
<label>3.1</label>
<title>Entropy as sustainability criteria</title>
<p>We identify sustainability with the idea that any action we take must not cause further depletion of resources. For example, if an action plan is designed such that one type of action is taken over a long period, it results in a cost-effective but not sustainable policy. Thus, we used Shannon entropy as an indicator of the degree of uniformity in the actions taken, i.e., we need to classify actions that use the maximum possible combinations in the control input space.</p>
<p>We have applied Shannon entropy to evaluate the degree of the sustainability of the control policies of the model. Shannon entropy is a fundamental information measure used to quantify the diversity in a distribution (<xref ref-type="bibr" rid="B16">Lin, 1991</xref>). It is mathematically defined as: <inline-formula>
<mml:math display="inline" id="im41"><mml:mrow><mml:mi>H</mml:mi><mml:mo>=</mml:mo><mml:mo>&#x2212;</mml:mo><mml:msubsup><mml:mo>&#x2211;</mml:mo><mml:mrow><mml:mi>i</mml:mi><mml:mo>=</mml:mo><mml:mn>1</mml:mn></mml:mrow><mml:mi>n</mml:mi></mml:msubsup><mml:msub><mml:mi>p</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mi>log</mml:mi><mml:mtext>&#xa0;</mml:mtext><mml:mo>(</mml:mo><mml:msub><mml:mi>p</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>)</mml:mo><mml:mo>,</mml:mo></mml:mrow></mml:math></inline-formula> where <inline-formula>
<mml:math display="inline" id="im42"><mml:mrow><mml:msub><mml:mi>p</mml:mi><mml:mi>i</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> represents the proportion of data in the <inline-formula>
<mml:math display="inline" id="im43"><mml:mi>i</mml:mi></mml:math></inline-formula>-th bin, and <inline-formula>
<mml:math display="inline" id="im44"><mml:mi>n</mml:mi></mml:math></inline-formula> is the total number of bins (<xref ref-type="bibr" rid="B25">Shannon, 1948</xref>). The entropy reaches its minimum value, <inline-formula>
<mml:math display="inline" id="im45"><mml:mrow><mml:mi>H</mml:mi><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula>, when all the data is concentrated in a single bin (<inline-formula>
<mml:math display="inline" id="im46"><mml:mrow><mml:msub><mml:mi>p</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:math></inline-formula> for one bin and <inline-formula>
<mml:math display="inline" id="im47"><mml:mrow><mml:msub><mml:mi>p</mml:mi><mml:mi>j</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula> for all <inline-formula>
<mml:math display="inline" id="im48"><mml:mrow><mml:mi>j</mml:mi><mml:mo>&#x2260;</mml:mo><mml:mi>i</mml:mi></mml:mrow></mml:math></inline-formula>), indicating no diversity or uncertainty. Conversely, the entropy is maximized when the data is evenly distributed across all bins (<inline-formula>
<mml:math display="inline" id="im49"><mml:mrow><mml:msub><mml:mi>p</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mfrac><mml:mn>1</mml:mn><mml:mi>n</mml:mi></mml:mfrac></mml:mrow></mml:math></inline-formula> for all <inline-formula>
<mml:math display="inline" id="im50"><mml:mi>i</mml:mi></mml:math></inline-formula>), in which case <inline-formula>
<mml:math display="inline" id="im51"><mml:mrow><mml:mi>H</mml:mi><mml:mo>=</mml:mo><mml:mi>log</mml:mi><mml:mo>(</mml:mo><mml:mi>n</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, representing maximum diversity. For example, with six bins, the maximum entropy is <inline-formula>
<mml:math display="inline" id="im52"><mml:mrow><mml:mi>H</mml:mi><mml:mo>=</mml:mo><mml:mi>log</mml:mi><mml:mo>(</mml:mo><mml:mn>6</mml:mn><mml:mo>)</mml:mo><mml:mo>&#x2248;</mml:mo><mml:mn>1.7918</mml:mn></mml:mrow></mml:math></inline-formula>. Policies with higher entropy indicate a more robust application of interventions, such as non-monotone cutting of Lantana (<inline-formula>
<mml:math display="inline" id="im53"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) and planting of control plant species (<inline-formula>
<mml:math display="inline" id="im54"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>), which ensures ecological stability and sustainability.</p>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Numerical methods</title>
<p>We employ a model predictive control (MPC) framework to address the optimal control problem for managing the Lantana population dynamics defined by our three-species ecological system. The control objective is formulated as a variational problem that seeks to minimize a cost function while satisfying system dynamics and constraints. The cost function, given by</p>
<disp-formula id="eq7"><label>(7)</label>
<mml:math display="block" id="M7"><mml:mrow><mml:mtable><mml:mtr columnalign="left"><mml:mtd columnalign="left"><mml:mi>J</mml:mi><mml:mo>=</mml:mo><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mtext>control</mml:mtext></mml:mrow></mml:msub><mml:mo>&#xb7;</mml:mo><mml:mo>(</mml:mo><mml:msubsup><mml:mi>u</mml:mi><mml:mn>1</mml:mn><mml:mrow><mml:mn>2</mml:mn></mml:mrow></mml:msubsup><mml:mo>+</mml:mo><mml:msubsup><mml:mi>u</mml:mi><mml:mn>2</mml:mn><mml:mrow><mml:mn>2</mml:mn></mml:mrow></mml:msubsup><mml:mo>)</mml:mo><mml:mo>+</mml:mo><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mtext>biodiversity</mml:mtext></mml:mrow></mml:msub></mml:mtd></mml:mtr><mml:mtr columnalign="left"><mml:mtd columnalign="left"><mml:mo>&#xb7;</mml:mo><mml:mo>(</mml:mo><mml:msup><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mrow><mml:mn>1</mml:mn><mml:mi>d</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo></mml:mrow><mml:mn>2</mml:mn></mml:msup><mml:mtext>&#xa0;</mml:mtext><mml:mo>+</mml:mo><mml:mtext>&#xa0;</mml:mtext><mml:msup><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mrow><mml:mn>2</mml:mn><mml:mi>d</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo></mml:mrow><mml:mn>2</mml:mn></mml:msup><mml:mtext>&#xa0;</mml:mtext><mml:mo>+</mml:mo><mml:mtext>&#xa0;</mml:mtext><mml:msup><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub><mml:mo>&#x2212;</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mrow><mml:mn>3</mml:mn><mml:mi>d</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo></mml:mrow><mml:mn>2</mml:mn></mml:msup><mml:mo>)</mml:mo><mml:mo>,</mml:mo></mml:mtd></mml:mtr></mml:mtable></mml:mrow></mml:math>
</disp-formula>
<p>balances two competing objectives: <inline-formula>
<mml:math display="inline" id="im55"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mtext>control</mml:mtext></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>1000</mml:mn></mml:mrow></mml:math></inline-formula> penalizes aggressive control efforts (harvesting <inline-formula>
<mml:math display="inline" id="im56"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> and augmentation <inline-formula>
<mml:math display="inline" id="im57"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>), while <inline-formula>
<mml:math display="inline" id="im58"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mtext>biodiversity</mml:mtext></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>10</mml:mn></mml:mrow></mml:math></inline-formula> penalizes deviations from the desired ecological equilibrium state <inline-formula>
<mml:math display="inline" id="im59"><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mrow><mml:mn>1</mml:mn><mml:mi>d</mml:mi></mml:mrow></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mrow><mml:mn>2</mml:mn><mml:mi>d</mml:mi></mml:mrow></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mrow><mml:mn>3</mml:mn><mml:mi>d</mml:mi></mml:mrow></mml:msub><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mo>(</mml:mo><mml:mn>0.1</mml:mn><mml:mo>,</mml:mo><mml:mn>0.75</mml:mn><mml:mo>,</mml:mo><mml:mn>0.15</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>. The optimal control problem was implemented in Python using the CasADi optimization framework (<xref ref-type="bibr" rid="B1">Andersson et&#xa0;al., 2019</xref>). The nonlinear dynamics were discretized over a time horizon of <inline-formula>
<mml:math display="inline" id="im60"><mml:mrow><mml:mi>T</mml:mi><mml:mo>=</mml:mo><mml:mn>25</mml:mn></mml:mrow></mml:math></inline-formula> units using the fourth-order Runge-Kutta (RK4) method with step size <inline-formula>
<mml:math display="inline" id="im61"><mml:mrow><mml:mtext>&#x394;</mml:mtext><mml:mi>t</mml:mi><mml:mo>=</mml:mo><mml:mn>0.03125</mml:mn></mml:mrow></mml:math></inline-formula>, divided into <inline-formula>
<mml:math display="inline" id="im62"><mml:mrow><mml:mi>N</mml:mi><mml:mo>=</mml:mo><mml:mn>40</mml:mn></mml:mrow></mml:math></inline-formula> control intervals. The discretized problem was formulated as a nonlinear programming (NLP) problem with state constraints maintaining populations within <inline-formula>
<mml:math display="inline" id="im63"><mml:mrow><mml:mo>[</mml:mo><mml:mn>0.0</mml:mn><mml:mo>,</mml:mo><mml:mn>1.0</mml:mn><mml:mo>]</mml:mo></mml:mrow></mml:math></inline-formula> and terminal constraints ensuring convergence to the target equilibrium. The NLP was solved using IPOPT (<xref ref-type="bibr" rid="B32">W&#xe4;chter and Biegler, 2006</xref>). Control bounds were imposed as <inline-formula>
<mml:math display="inline" id="im64"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>[</mml:mo><mml:mo>&#x2212;</mml:mo><mml:mn>5</mml:mn><mml:mo>,</mml:mo><mml:mn>0</mml:mn><mml:mo>]</mml:mo></mml:mrow></mml:math></inline-formula> for harvesting the invasive species and <inline-formula>
<mml:math display="inline" id="im65"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>[</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>5</mml:mn><mml:mo>]</mml:mo></mml:mrow></mml:math></inline-formula> for augmenting native species, with two scenarios investigated: simultaneous application of both controls and selective harvesting only (<inline-formula>
<mml:math display="inline" id="im66"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula>). The optimization was performed for various initial conditions, confirming successful convergence in all cases as shown in the results section.</p>
<p>We would like to point out that the form of the cost function chosen here is not exhaustive and one of the choice can be incorporating the entropy term which would require further detailed investigation.</p>
</sec>
</sec>
<sec id="s4" sec-type="results">
<label>4</label>
<title>Results</title>
<p>We provide solutions to the optimal control problem of the management of abundances of the invasive plant <italic>Lantana camara</italic>. We also analyzed the qualitative and quantitative behavior of these calculated optimal control policies, which can drive the system from any initial condition to a given desired state. We first present the results of analyzing the GLV network-based dynamical control systems model using Lie algebraic approaches. Secondly, by incorporating the principles of information theory into the model, we present Shannon entropy as a measure of the degree of sustainability of the control policies aligned with the Sustainable Development Goals (SDGs) paradigms.</p>
<sec id="s4_1">
<label>4.1</label>
<title>Accessibility and controllability conditions</title>
<p>The controllability analysis of the three-species system (2) establishes the theoretical foundation for our optimal control framework and clarifies the extent to which desired system states can be reached. Under the ecological constraint that all interaction coefficients satisfy <inline-formula>
<mml:math display="inline" id="im67"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> for all <inline-formula>
<mml:math display="inline" id="im68"><mml:mrow><mml:mi>i</mml:mi><mml:mo>,</mml:mo><mml:mi>j</mml:mi><mml:mo>&#x2208;</mml:mo><mml:mo>{</mml:mo><mml:mn>1</mml:mn><mml:mo>,</mml:mo><mml:mn>2</mml:mn><mml:mo>,</mml:mo><mml:mn>3</mml:mn><mml:mo>}</mml:mo></mml:mrow></mml:math></inline-formula> (<xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref>, Assumption 1), representing non-negligible but bounded species interactions, the system is locally accessible at every point in the positive orthant <inline-formula>
<mml:math display="inline" id="im69"><mml:mrow><mml:msubsup><mml:mi>&#x211d;</mml:mi><mml:mrow><mml:mo>+</mml:mo><mml:mo>+</mml:mo></mml:mrow><mml:mn>3</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula> where all three species coexist (<xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref>, Theorem 1). Importantly, this accessibility property holds uniformly across the entire parameter regime <inline-formula>
<mml:math display="inline" id="im70"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>&#x2014;it is not confined to specific parameter configurations but applies universally to all biologically realistic interaction strengths.</p>
<p>The proof involves computing Lie brackets [<italic>f</italic><sub>0</sub>, <italic>f</italic><sub>1</sub>] and [<italic>f</italic><sub>0</sub>, <italic>f</italic><sub>2</sub>] between the drift vector field (natural ecosystem dynamics) and control vector fields (management interventions), which reveal additional directions in state space accessible by combining natural dynamics with control actions. The accessibility distribution matrix <inline-formula>
<mml:math display="inline" id="im71"><mml:mrow><mml:mi mathvariant="script">C</mml:mi><mml:mo>(</mml:mo><mml:mi>x</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> (<xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref>, <xref ref-type="disp-formula" rid="eq1">Equation 12</xref>) has full rank equal to three throughout <inline-formula>
<mml:math display="inline" id="im72"><mml:mrow><mml:msubsup><mml:mi>&#x211d;</mml:mi><mml:mrow><mml:mo>+</mml:mo><mml:mo>+</mml:mo></mml:mrow><mml:mn>3</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula>, satisfying the LARC. This accessibility depends critically on non-zero interaction coefficients <inline-formula>
<mml:math display="inline" id="im73"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>31</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im74"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>32</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, representing how <italic>Lantana camara</italic> and native plants influence soil microbes (<xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref>, Remark 1). These strictly positive coefficients provide the indirect control pathway: the control parameters <inline-formula>
<mml:math display="inline" id="im75"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im76"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> acting directly on plant populations can indirectly influence microbes through ecological coupling. If either of the coefficients <inline-formula>
<mml:math display="inline" id="im77"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>31</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> or <inline-formula>
<mml:math display="inline" id="im78"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>32</mml:mn></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> were zero, the accessibility matrix rank would reduce to 2, destroying controllability. Local accessibility guarantees that from any initial state <inline-formula>
<mml:math display="inline" id="im79"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>&#x2208;</mml:mo><mml:msubsup><mml:mi>&#x211d;</mml:mi><mml:mrow><mml:mo>+</mml:mo><mml:mo>+</mml:mo></mml:mrow><mml:mn>3</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula>, the reachable set has non-empty interior (<xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref>, Theorem 2), meaning we can reach a neighborhood of any target state through appropriate control trajectories.</p>
<p>However, the system is not globally controllable. Forward-invariant manifolds exist at coordinate planes where a species has zero population (<xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref>, Lemma 1, Theorem 4), and this limitation persists uniformly across all parameter values <inline-formula>
<mml:math display="inline" id="im80"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>. This reveals fundamentally different dependencies on parameters versus initial conditions.</p>
<p>Control efficacy is universal across the parameter regime, and accessibility holds for all values in <inline-formula>
<mml:math display="inline" id="im81"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> without exception. However, control efficacy exhibits a sharp dichotomy with respect to initial conditions: if all species are initially present (<inline-formula>
<mml:math display="inline" id="im82"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>&#x2208;</mml:mo><mml:msubsup><mml:mi>&#x211d;</mml:mi><mml:mrow><mml:mo>+</mml:mo><mml:mo>+</mml:mo></mml:mrow><mml:mn>3</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula>), any target state can be approached.</p>
<p>We also investigated small-time local controllability (STLC) using Sussmann&#x2019;s sufficient condition. The system fails this condition uniformly across <inline-formula>
<mml:math display="inline" id="im83"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> (<xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref>, Theorem 3): the Lie brackets <inline-formula>
<mml:math display="inline" id="im84"><mml:mrow><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>]</mml:mo></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im85"><mml:mrow><mml:mo>[</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>]</mml:mo></mml:mrow></mml:math></inline-formula>, essential for controlling microbes, cannot be expressed as combinations of control vector fields <inline-formula>
<mml:math display="inline" id="im86"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im87"><mml:mrow><mml:msub><mml:mi>f</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, which span only the plant population plane. This failure is a structural feature persisting across all parameter values. While the condition is sufficient but not necessary, the failure suggests potential time-scale limitations on restoration speed (<xref ref-type="supplementary-material" rid="SM1"><bold>Appendix</bold></xref>, Remark 3). Target states are eventually reachable from <inline-formula>
<mml:math display="inline" id="im88"><mml:mrow><mml:msubsup><mml:mi>&#x211d;</mml:mi><mml:mrow><mml:mo>+</mml:mo><mml:mo>+</mml:mo></mml:mrow><mml:mn>3</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula>, but arbitrarily rapid transitions may not be achievable, aligning with ecological reality where restoration unfolds over extended periods. The optimal trajectories (<xref ref-type="fig" rid="f2"><bold>Figures&#xa0;2</bold></xref>&#x2013;<xref ref-type="fig" rid="f4"><bold>4</bold></xref>) reflect these natural time-scale constraints.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Shannon entropy <italic>H</italic> for evaluating the sustainability of control policies. <bold>(a)</bold> Box plot of <italic>H</italic> against log(<italic>c</italic><sub>control</sub><italic>/c</italic><sub>biodiversity</sub>) for the ratio of species abundances <inline-formula>
<mml:math display="inline" id="im89"><mml:mrow><mml:mo>&#xa0;</mml:mo><mml:mfrac><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mfrac></mml:mrow></mml:math></inline-formula>, this gives the information about the distribution, in time, of the values of <inline-formula>
<mml:math display="inline" id="im90"><mml:mrow><mml:mfrac><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mfrac><mml:mo>&#xa0;</mml:mo></mml:mrow></mml:math></inline-formula>This indicates how this ratio behaves in time when initially it is <inline-formula>
<mml:math display="inline" id="im91"><mml:mrow><mml:mo>&#xa0;</mml:mo><mml:mfrac><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mfrac></mml:mrow></mml:math></inline-formula> &gt; 1 and crosses over to <inline-formula>
<mml:math display="inline" id="im92"><mml:mrow><mml:mfrac><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mfrac></mml:mrow></mml:math></inline-formula> &lt; 1, across a set of varied initial conditions. <bold>(b)</bold><italic>H<sub>u</sub></italic><sub>1</sub> and <italic>H<sub>u</sub></italic><sub>2</sub> from the distribution of the control signals in time for the same set of initial conditions as in <bold>(a)</bold>. <bold>(c)</bold><italic>H<sub>x</sub></italic><sub>1</sub> and <italic>H<sub>x</sub></italic><sub>2</sub> for species abundance distribution in time for the same set of initial conditions as in <bold>(a)</bold>, reflecting control effectiveness and biodiversity outcomes.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1644458-g002.tif">
<alt-text content-type="machine-generated">Three-panel figure comparing Shannon entropy in different conditions. Panel a shows box plots of Shannon entropy \( H \) across \( \log_{10} \left( \frac{C_{\text{control}}}{C_{\text{biodiversity}}} \right) \), ranging from 0.5 to 1.5. Panel b and c display violin plots for \( H_{u1}, H_{u2}, H_{x1}, \) and \( H_{x2} \) with different distributions and medians, labeled with respective colors and legends. Data points are plotted throughout to indicate variance.</alt-text>
</graphic></fig>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Sensitivity analysis: Mean control costs show an increasing trend with an increase in the growth rate of <italic>Lantana</italic>. <bold>(a, b)</bold> correspond to the initial condition population condition as <inline-formula>
<mml:math display="inline" id="im93">
<mml:mrow>
<mml:mo>(</mml:mo>
<mml:msub>
<mml:mi>x</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mo>(</mml:mo>
<mml:mn>0</mml:mn>
<mml:mo>)</mml:mo>
<mml:mo>,</mml:mo>
<mml:msub>
<mml:mi>x</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:mo>(</mml:mo>
<mml:mn>0</mml:mn>
<mml:mo>)</mml:mo>
<mml:mo>,</mml:mo>
<mml:msub>
<mml:mi>x</mml:mi>
<mml:mn>3</mml:mn>
</mml:msub>
<mml:mo>(</mml:mo>
<mml:mn>0</mml:mn>
<mml:mo>)</mml:mo>
<mml:mo>)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mo>(</mml:mo>
<mml:mn>0.85</mml:mn>
<mml:mo>,</mml:mo>
<mml:mn>0.1</mml:mn>
<mml:mo>,</mml:mo>
<mml:mn>0.05</mml:mn>
<mml:mo>)</mml:mo>
</mml:mrow>
</mml:math>
</inline-formula> where we have a high abundance of Lantana. Figures <bold>(c, d)</bold> show results for a moderate abundance of Lantana with the initial condition <inline-formula>
<mml:math display="inline" id="im94">
<mml:mrow>
<mml:mo>(</mml:mo>
<mml:msub>
<mml:mi>x</mml:mi>
<mml:mn>1</mml:mn>
</mml:msub>
<mml:mo>(</mml:mo>
<mml:mn>0</mml:mn>
<mml:mo>)</mml:mo>
<mml:mo>,</mml:mo>
<mml:msub>
<mml:mi>x</mml:mi>
<mml:mn>2</mml:mn>
</mml:msub>
<mml:mo>(</mml:mo>
<mml:mn>0</mml:mn>
<mml:mo>)</mml:mo>
<mml:mo>,</mml:mo>
<mml:msub>
<mml:mi>x</mml:mi>
<mml:mn>3</mml:mn>
</mml:msub>
<mml:mo>(</mml:mo>
<mml:mn>0</mml:mn>
<mml:mo>)</mml:mo>
<mml:mo>)</mml:mo>
<mml:mo>=</mml:mo>
<mml:mo>(</mml:mo>
<mml:mn>0.6</mml:mn>
<mml:mo>,</mml:mo>
<mml:mn>0.3</mml:mn>
<mml:mo>,</mml:mo>
<mml:mn>0.1</mml:mn>
<mml:mo>)</mml:mo>
</mml:mrow>
</mml:math>
</inline-formula>. The interaction matrix is fixed as: <italic>a</italic><sub>11</sub> = 0.3, <italic>a</italic><sub>12</sub> = 0.1, <italic>a</italic><sub>13</sub> = 0.4, <italic>a</italic><sub>21</sub> = 0.8, <italic>a</italic><sub>22</sub> = 0.2, <italic>a</italic><sub>23</sub> = 0.5, <italic>a</italic><sub>31</sub> = 0.6, <italic>a</italic><sub>32</sub> = 0.7, <italic>a</italic><sub>33</sub> = 0.7. The mean is taken over for different values of the parameter <italic>r</italic><sub>2</sub>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1644458-g003.tif">
<alt-text content-type="machine-generated">Four line graphs showing the relationship between different variables related to mean cost, with lines representing different values of r&#x2083; (0.20, 0.35, 0.50, 0.65, 0.80). Graphs (a) and (c) display log&#x2081;&#x2080;(mean cost) versus r&#x2081;/r&#x2082;, with two regimes shaded differently. Graphs (b) and (d) plot mean cost against r&#x2081; within a defined range. The lines vary in color corresponding to r&#x2083; values, reflecting diverse patterns in cost behavior across regimes.</alt-text>
</graphic></fig>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Control dynamics of the system across various initial species abundances. Growth rates: <inline-formula>
<mml:math display="inline" id="im97"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0.85</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im98"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0.4</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im99"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>3</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0.3</mml:mn></mml:mrow></mml:math></inline-formula>. Desired state: <inline-formula>
<mml:math display="inline" id="im100"><mml:mrow><mml:mo>(</mml:mo><mml:msubsup><mml:mi>x</mml:mi><mml:mn>1</mml:mn><mml:mi>d</mml:mi></mml:msubsup><mml:mo>,</mml:mo><mml:msubsup><mml:mi>x</mml:mi><mml:mn>2</mml:mn><mml:mi>d</mml:mi></mml:msubsup><mml:mo>,</mml:mo><mml:msubsup><mml:mi>x</mml:mi><mml:mn>3</mml:mn><mml:mi>d</mml:mi></mml:msubsup><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mo>(</mml:mo><mml:mn>0.1</mml:mn><mml:mo>,</mml:mo><mml:mn>0.75</mml:mn><mml:mo>,</mml:mo><mml:mn>0.15</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>. Interaction matrix: <inline-formula>
<mml:math display="inline" id="im101"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>11</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.6</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im102"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>12</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.6</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im103"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>13</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.5</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im104"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>21</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.6</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im105"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>22</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.3</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im106"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>23</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.5</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im107"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>31</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.7</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im108"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>32</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.6</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im109"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mn>33</mml:mn></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>0.7</mml:mn></mml:mrow></mml:math></inline-formula>. Cost parameters: <bold>(a, c)</bold>: Control cost <inline-formula>
<mml:math display="inline" id="im110"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mstyle mathvariant="bold" mathsize="normal"><mml:mi>c</mml:mi><mml:mi>o</mml:mi><mml:mi>n</mml:mi><mml:mi>t</mml:mi><mml:mi>r</mml:mi><mml:mi>o</mml:mi><mml:mi>l</mml:mi></mml:mstyle></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>1000.0</mml:mn></mml:mrow></mml:math></inline-formula>, biodiversity penalty <inline-formula>
<mml:math display="inline" id="im111"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mstyle mathvariant="bold" mathsize="normal"><mml:mi>b</mml:mi><mml:mi>i</mml:mi><mml:mi>o</mml:mi><mml:mi>d</mml:mi><mml:mi>i</mml:mi><mml:mi>v</mml:mi><mml:mi>e</mml:mi><mml:mi>r</mml:mi><mml:mi>s</mml:mi><mml:mi>i</mml:mi><mml:mi>t</mml:mi><mml:mi>y</mml:mi></mml:mstyle></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>1.0</mml:mn></mml:mrow></mml:math></inline-formula>. <bold>(b, d)</bold>: Control cost <inline-formula>
<mml:math display="inline" id="im112"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mstyle mathvariant="bold" mathsize="normal"><mml:mi>c</mml:mi><mml:mi>o</mml:mi><mml:mi>n</mml:mi><mml:mi>t</mml:mi><mml:mi>r</mml:mi><mml:mi>o</mml:mi><mml:mi>l</mml:mi></mml:mstyle></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>1000.0</mml:mn></mml:mrow></mml:math></inline-formula>, biodiversity penalty <inline-formula>
<mml:math display="inline" id="im113"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mstyle mathvariant="bold" mathsize="normal"><mml:mi>b</mml:mi><mml:mi>i</mml:mi><mml:mi>o</mml:mi><mml:mi>d</mml:mi><mml:mi>i</mml:mi><mml:mi>v</mml:mi><mml:mi>e</mml:mi><mml:mi>r</mml:mi><mml:mi>s</mml:mi><mml:mi>i</mml:mi><mml:mi>t</mml:mi><mml:mi>y</mml:mi></mml:mstyle></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:mn>1000.0</mml:mn></mml:mrow></mml:math></inline-formula>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1644458-g004.tif">
<alt-text content-type="machine-generated">Four graphs visualize ecological control models. Graphs (a) and (b) are 3D plots showing initial, final, and desired states of ecosystems with variables: Lantana camara, Native plants, and Microbes. Graphs (c) and (d) are line charts displaying control variables over time, differentiated by red and green dashed lines.</alt-text>
</graphic></fig>
<p>Hence, control efficacy is universal across the biologically realistic parameter regime <inline-formula>
<mml:math display="inline" id="im114"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> and accessibility holds for all parameter values within this range, and controllability is not confined to specific parameter configurations. However, control efficacy is fundamentally constrained by initial conditions, i.e., desired states are reachable if and only if the initial state contains all three species at positive densities (<inline-formula>
<mml:math display="inline" id="im115"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>0</mml:mn></mml:msub><mml:mo>&#x2208;</mml:mo><mml:msubsup><mml:mi>&#x211d;</mml:mi><mml:mrow><mml:mo>+</mml:mo><mml:mo>+</mml:mo></mml:mrow><mml:mn>3</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula>). These results provide rigorous justification that computed optimal trajectories are theoretically achievable across diverse ecosystems with varying ecological characteristics, provided management begins early to maintain all species at positive densities throughout restoration. Given this analysis, we proceed to solve numerically the control problem and present the results in subsequent sections.</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Comparative analysis of control strategies</title>
<p><xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5</bold></xref> presents a comprehensive comparison of system dynamics under different control scenarios. <xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5a</bold></xref> illustrates the uncontrolled state-space trajectories in the <inline-formula>
<mml:math display="inline" id="im126"><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> for multiple initial conditions, showing the dismal state of Lantana invasion if no control is applied. <xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5b</bold></xref> depicts the controlled trajectories under two distinct control strategies: single control with <inline-formula>
<mml:math display="inline" id="im127"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula> (red curves) and dual control with <inline-formula>
<mml:math display="inline" id="im128"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula> (blue curves). Our simulations reveal that the case where only <inline-formula>
<mml:math display="inline" id="im129"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> is active, while <inline-formula>
<mml:math display="inline" id="im130"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula>, fails to drive the system to the desired state within the specified time horizon, indicating that the control input <inline-formula>
<mml:math display="inline" id="im131"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> alone is insufficient for effective system stabilization. Hence, we have omitted this scenario. We observe that both the single-control strategy (<inline-formula>
<mml:math display="inline" id="im132"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula>) and the dual-control strategy (<inline-formula>
<mml:math display="inline" id="im133"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula>) successfully guide the system trajectories to the target state <inline-formula>
<mml:math display="inline" id="im134"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>d</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> for all initial conditions examined, but they demonstrate different control input signal structures. <xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5c</bold></xref> provides a quantitative comparison of the total costs associated with these two successful strategies, where the cost function balances state deviation from the target and control effort. The box plots reveal a striking difference: the single-control strategy incurs approximately double the cost (median <inline-formula>
<mml:math display="inline" id="im135"><mml:mo>&#x2248;</mml:mo></mml:math></inline-formula> 1300) compared to the dual-control strategy (median <inline-formula>
<mml:math display="inline" id="im136"><mml:mo>&#x2248;</mml:mo></mml:math></inline-formula> 700), demonstrating a substantial cost reduction of nearly 50%. The insets display representative time profiles of the control inputs, where inset (i) shows that in the single-control case, <inline-formula>
<mml:math display="inline" id="im137"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> must exhibit large variations to compensate for the absence of <inline-formula>
<mml:math display="inline" id="im138"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, while inset (ii) illustrates that in the dual-control case, the control effort is distributed between <inline-formula>
<mml:math display="inline" id="im139"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im140"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>. Collectively, these results establish that while the single-control strategy can stabilize the system, the dual-control policy provides superior performance by reducing intervention costs by approximately 50% and also producing smoother control actions that are more practical to implement in ecological management contexts, where resource constraints and minimized ecosystem disruption are critical considerations.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p><bold>(a)</bold> Uncontrolled state-space trajectories in the <inline-formula>
<mml:math display="inline" id="im116"><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> phase space converge away from the desired state (red cross), illustrating the prevalence of invasion dynamics in the absence of interventions. <bold>(b)</bold> Controlled trajectories under two strategies: red curves denote the case <inline-formula>
<mml:math display="inline" id="im117"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mtext>&#x2009;</mml:mtext><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula>, while blue curves denote <inline-formula>
<mml:math display="inline" id="im118"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mtext>&#x2009;</mml:mtext><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula>. Both control strategies steer the system closer to the desired state <inline-formula>
<mml:math display="inline" id="im119"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mi>d</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> (red cross), though with different transient dynamics. <bold>(c)</bold> Comparison of cumulative control costs between the two strategies (box plots), showing that simultaneous application of <inline-formula>
<mml:math display="inline" id="im120"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im121"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> yields substantially lower costs than the single-control case. Insets (i) and (ii) display the control input signals <inline-formula>
<mml:math display="inline" id="im122"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> (solid) and <inline-formula>
<mml:math display="inline" id="im123"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>t</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> (dashed) for two scenarios <inline-formula>
<mml:math display="inline" id="im124"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mtext>&#x2009;</mml:mtext><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula> and <inline-formula>
<mml:math display="inline" id="im125"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mtext>&#x2009;</mml:mtext><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>&#x2260;</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula>, respectively, highlighting distinct control effort distributions over time.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1644458-g005.tif">
<alt-text content-type="machine-generated">Panel (a) shows a 3D plot with trajectories from an initial state to a final state, indicating desired states with a green dot. Panel (b) illustrates trajectories with red and blue lines under different control settings. Panel (c) includes box plots for total cost and graphs showing control variables \(u_1\) and \(u_2\) over time, where (i) deals with \(u_1 \neq 0, u_2 = 0\) and (ii) with both controls active.</alt-text>
</graphic></fig>
<p>Since we find that the case of double controls incurs significantly lower cost than the single control one, all subsequent analyses are performed for the case of double control only.</p>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Sustainability analysis using Shannon entropy</title>
<p>Selection of a sustainable policy requires a sieve, which we now elucidate. When designing control strategies for managing the invasive species <italic>Lantana camara</italic>, optimizing policies solely based on a fixed cost function is insufficient to ensure ecological sustainability. To address this limitation, we introduced Shannon entropy as an additional metric to evaluate and select control policies. Shannon entropy quantifies the diversity and balance of control efforts, providing insights into how evenly the management inputs are distributed over time. We show in <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;4</bold></xref> how on changing the two parameters of the cost function, i.e., the control cost penalty parameter (<inline-formula>
<mml:math display="inline" id="im141"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mi>C</mml:mi><mml:mi>o</mml:mi><mml:mi>n</mml:mi><mml:mi>t</mml:mi><mml:mi>r</mml:mi><mml:mi>o</mml:mi><mml:mi>l</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) and the Bio-Diversity Penalty parameter (<inline-formula>
<mml:math display="inline" id="im142"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mi>b</mml:mi><mml:mi>i</mml:mi><mml:mi>o</mml:mi><mml:mi>d</mml:mi><mml:mi>i</mml:mi><mml:mi>v</mml:mi><mml:mi>e</mml:mi><mml:mi>r</mml:mi><mml:mi>s</mml:mi><mml:mi>i</mml:mi><mml:mi>t</mml:mi><mml:mi>y</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) used in the optimization technique, we observed the different trajectories available in <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;4</bold></xref>. This also illustrates various optimal control policies, each with distinct ecological implications. For example, the policy in <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;4</bold></xref> employs a non-monotone application of control inputs&#x2014;cutting <italic>Lantana camara</italic> (<inline-formula>
<mml:math display="inline" id="im143"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) and planting control plant species (<inline-formula>
<mml:math display="inline" id="im144"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>)&#x2014;throughout the management period. This approach promotes a gradual replacement of <italic>Lantana camara</italic> with control plants, fostering ecological balance while minimizing disruptions to soil health, microbial communities, and the broader environment. In contrast, the policy in <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;4</bold></xref> adopts a more aggressive strategy, concentrating intensive cutting and planting efforts at the beginning and end of the management period, with prolonged intervals of inaction in between. The abrupt removal of <italic>Lantana</italic> without consistent planting risks leaving the land vulnerable to degradation, which can lead to soil erosion, nutrient depletion, microbial imbalance, and other cascading ecological issues. While this strategy may seem cost-effective, it compromises long-term stability and sustainability.</p>
<p>We report the impact of changes in the parameters of the cost function on the sustainability characterization of the feasible optimal control policies. Fixing the initial conditions as shown in <xref ref-type="fig" rid="f5"><bold>Figures&#xa0;5a, b</bold></xref> and then using the defined cost function <inline-formula>
<mml:math display="inline" id="im145"><mml:mi>J</mml:mi></mml:math></inline-formula> in (<xref ref-type="disp-formula" rid="eq7">Equation 7</xref>), and keeping the interaction matrix fixed(as given in the caption of <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;4</bold></xref>), we computed the optimal control trajectories for various parameter pairs: <inline-formula>
<mml:math display="inline" id="im146"><mml:mrow><mml:mo>(</mml:mo><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mtext>control</mml:mtext></mml:mrow></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mtext>biodiversity</mml:mtext></mml:mrow></mml:msub><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mo>(</mml:mo><mml:msup><mml:mrow><mml:mn>10</mml:mn></mml:mrow><mml:mn>3</mml:mn></mml:msup><mml:mo>,</mml:mo><mml:msup><mml:mrow><mml:mn>10</mml:mn></mml:mrow><mml:mn>3</mml:mn></mml:msup><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:mo>(</mml:mo><mml:msup><mml:mrow><mml:mn>10</mml:mn></mml:mrow><mml:mn>3</mml:mn></mml:msup><mml:mo>,</mml:mo><mml:msup><mml:mrow><mml:mn>10</mml:mn></mml:mrow><mml:mn>2</mml:mn></mml:msup><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:mo>(</mml:mo><mml:msup><mml:mrow><mml:mn>10</mml:mn></mml:mrow><mml:mn>3</mml:mn></mml:msup><mml:mo>,</mml:mo><mml:mn>10</mml:mn><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:mo>(</mml:mo><mml:msup><mml:mrow><mml:mn>10</mml:mn></mml:mrow><mml:mn>3</mml:mn></mml:msup><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, across multiple initial conditions. For each trajectory, we have used six bins for discretization (We also found that on increasing the number of bins, the distribution did not change in shape). The Shannon entropy (<inline-formula>
<mml:math display="inline" id="im147"><mml:mi>H</mml:mi></mml:math></inline-formula>) was calculated for the time series of <inline-formula>
<mml:math display="inline" id="im148"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im149"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im150"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, <inline-formula>
<mml:math display="inline" id="im151"><mml:mrow><mml:msub><mml:mi>u</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, and the ratio of the Lantana and control plant abundances <inline-formula>
<mml:math display="inline" id="im152"><mml:mrow><mml:mfrac><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mfrac></mml:mrow></mml:math></inline-formula>. As can be seen in the distribution of the control inputs shown in <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2b</bold></xref>, in the case of the example trajectory in <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;4</bold></xref> corresponds to the ratio of <inline-formula>
<mml:math display="inline" id="im153"><mml:mrow><mml:mi>l</mml:mi><mml:mi>o</mml:mi><mml:msub><mml:mi>g</mml:mi><mml:mrow><mml:mn>10</mml:mn></mml:mrow></mml:msub><mml:mo>(</mml:mo><mml:mfrac><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mi>c</mml:mi><mml:mi>o</mml:mi><mml:mi>n</mml:mi><mml:mi>t</mml:mi><mml:mi>r</mml:mi><mml:mi>o</mml:mi><mml:mi>l</mml:mi></mml:mrow></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mi>b</mml:mi><mml:mi>i</mml:mi><mml:mi>o</mml:mi><mml:mi>d</mml:mi><mml:mi>i</mml:mi><mml:mi>v</mml:mi><mml:mi>e</mml:mi><mml:mi>r</mml:mi><mml:mi>s</mml:mi><mml:mi>i</mml:mi><mml:mi>t</mml:mi><mml:mi>y</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:mfrac><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mn>0</mml:mn></mml:mrow></mml:math></inline-formula> having a lower value of Shannon entropy as compared to the trajectory in <xref ref-type="fig" rid="f2"><bold>Figure&#xa0;4</bold></xref> corresponding to the ratio of <inline-formula>
<mml:math display="inline" id="im154"><mml:mrow><mml:mi>l</mml:mi><mml:mi>o</mml:mi><mml:msub><mml:mi>g</mml:mi><mml:mrow><mml:mn>10</mml:mn></mml:mrow></mml:msub><mml:mo>(</mml:mo><mml:mfrac><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mi>c</mml:mi><mml:mi>o</mml:mi><mml:mi>n</mml:mi><mml:mi>t</mml:mi><mml:mi>r</mml:mi><mml:mi>o</mml:mi><mml:mi>l</mml:mi></mml:mrow></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mrow><mml:mi>b</mml:mi><mml:mi>i</mml:mi><mml:mi>o</mml:mi><mml:mi>d</mml:mi><mml:mi>i</mml:mi><mml:mi>v</mml:mi><mml:mi>e</mml:mi><mml:mi>r</mml:mi><mml:mi>s</mml:mi><mml:mi>i</mml:mi><mml:mi>t</mml:mi><mml:mi>y</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:mfrac><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mn>3</mml:mn></mml:mrow></mml:math></inline-formula> which has a higher entropy value. Based on these results and analysis, we report that there is a meaningful way to design a sustainable policy for a desirable ecological state by maximizing the Shannon entropy as the basis for selecting the action plan to control abundances. This trend promotes a more uniform distribution of control efforts over time, as indicated by higher entropy values. Policies with higher entropy condition align with ecological sustainability principles by ensuring continuous and evenly distributed management efforts, ultimately promoting a balanced and desirable resilient ecosystem.</p>
</sec>
<sec id="s4_4">
<label>4.4</label>
<title>Sensitivity analysis</title>
<p>We study sequentially how sensitive the proposed control policies are to the changes in model parameter values. There are two types of parameters regulating our proposed model: firstly, we have the dynamical systems parameters, and secondly, we have the parameters in the optimization of the control cost function. We checked and systematically analyzed the sensitivity of the control cost to both types of parameters, as shown in the previous sections. Now for the dynamical systems parameters, since the precise values of model parameters are unknown, we analyzed the trends of control costs by uniformly varying intrinsic growth rates (<inline-formula>
<mml:math display="inline" id="im155"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>r</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>,</mml:mo><mml:msub><mml:mi>r</mml:mi><mml:mn>3</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) across the range <inline-formula>
<mml:math display="inline" id="im156"><mml:mrow><mml:mo>[</mml:mo><mml:mn>0.2</mml:mn><mml:mo>,</mml:mo><mml:mn>0.8</mml:mn><mml:mo>]</mml:mo></mml:mrow></mml:math></inline-formula>, generating 125 tuples while keeping the values of the elements of the interaction matrix fixed. Simulations were conducted for two initial conditions: <italic>Case I</italic> as shown in <xref ref-type="fig" rid="f3"><bold>Figures&#xa0;3a, b</bold></xref>, where, the abundance of the Lantana camara is high over the landscape (<inline-formula>
<mml:math display="inline" id="im157"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mn>0.85</mml:mn><mml:mo>,</mml:mo><mml:mn>0.1</mml:mn><mml:mo>,</mml:mo><mml:mn>0.05</mml:mn></mml:mrow></mml:math></inline-formula>), and <italic>Case II</italic>, where its abundance is moderate (<inline-formula>
<mml:math display="inline" id="im158"><mml:mrow><mml:msub><mml:mi>x</mml:mi><mml:mn>1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:msub><mml:mi>x</mml:mi><mml:mn>3</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>)</mml:mo><mml:mo>=</mml:mo><mml:mn>0.6</mml:mn><mml:mo>,</mml:mo><mml:mn>0.3</mml:mn><mml:mo>,</mml:mo><mml:mn>0.1</mml:mn></mml:mrow></mml:math></inline-formula>) as shown in the <xref ref-type="fig" rid="f3"><bold>Figures&#xa0;3c, d</bold></xref>. The analysis revealed two distinct regimes based on the ratio of Lantana camara&#x2019;s growth rate (<inline-formula>
<mml:math display="inline" id="im159"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) to that of the control plants (<inline-formula>
<mml:math display="inline" id="im160"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>), which matches the intuitive idea that we can use only certain plants to outgrow Lantana camara. In <italic>Regime I</italic>(<inline-formula>
<mml:math display="inline" id="im161"><mml:mrow><mml:mfrac><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mfrac><mml:mo>&lt;</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:math></inline-formula>), where the growth rate of Lantana camara is lower than that of control plants, control costs are relatively low. In contrast, <italic>Regime II</italic> ((<inline-formula>
<mml:math display="inline" id="im162"><mml:mrow><mml:mfrac><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:mfrac><mml:mo>&gt;</mml:mo><mml:mn>1</mml:mn></mml:mrow></mml:math></inline-formula>)), where Lantana camara outcompetes control plants with higher growth rates, incurs significantly higher management costs, as evident in <xref ref-type="fig" rid="f3"><bold>Figures&#xa0;3a, c</bold></xref>. These trends highlight the crucial role of competition dynamics, as higher costs emerge when Lantana camara dominates resource consumption and growth. <xref ref-type="fig" rid="f3"><bold>Figures&#xa0;3b, d</bold></xref> further illustrate that the behavior of the mean (averaged over various values of <inline-formula>
<mml:math display="inline" id="im163"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) control costs for scenarios of high and moderate Lantana camara abundances. The results clearly show that as <inline-formula>
<mml:math display="inline" id="im164"><mml:mrow><mml:msub><mml:mi>r</mml:mi><mml:mn>1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> increases, the management costs consistently rise. While the system shows limited sensitivity to the small variations within each regime, the absolute cost strongly depends on the relative growth rates and the initial abundance of the Lantana camara. These findings significantly highlight that the effective management of invasive species, such as Lantana camara, requires addressing growth dynamics and competition, as higher growth rates and competitive dominance significantly escalate management efforts to restore ecological balance.</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusion and discussion</title>
<p>Our study on optimal control strategies for managing the invasive plant <italic>Lantana camara</italic> highlights the crucial role of <italic>Lantana camara</italic>&#x2019;s growth rate and competition with control plants in determining control costs, with two regimes identified: low costs when <italic>Lantana camara</italic>&#x2019;s growth rate is lower than control plants, and high costs when <italic>Lantana</italic> outcompetes them. By integrating Shannon entropy as a metric for sustainability, we ensure that control policies are evenly distributed over time, promoting long-term ecological stability and minimizing risks like soil erosion and microbial imbalance. Sensitivity analysis reveals that control costs are influenced by both intrinsic growth rates and initial conditions, underscoring the necessity for adaptive, balanced management strategies.</p>
<p>The controllability analysis establishes that while the system is accessible at every point in <inline-formula>
<mml:math display="inline" id="im165"><mml:mrow><mml:msubsup><mml:mi>&#x211d;</mml:mi><mml:mrow><mml:mo>+</mml:mo><mml:mo>+</mml:mo></mml:mrow><mml:mn>3</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula> where all species coexist (uniformly across the parameter regime <inline-formula>
<mml:math display="inline" id="im166"><mml:mrow><mml:msub><mml:mi>a</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub><mml:mo>&#x2208;</mml:mo><mml:mo>(</mml:mo><mml:mn>0</mml:mn><mml:mo>,</mml:mo><mml:mn>1</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>), it lacks global controllability due to forward-invariant manifolds at coordinate planes. The key finding is that control efficacy depends fundamentally on initial conditions rather than parameter regimes: our optimal control strategies are effective across diverse ecosystems with varying ecological characteristics, but only when applied before any species reaches local extinction. This emphasizes the critical importance of early intervention in invasive species management and maintaining minimal viable populations throughout restoration efforts.</p>
<p>The failure of Sussmann&#x2019;s sufficient condition for small-time local controllability suggests inherent time-scale limitations on restoration speed. While target states are eventually reachable from initial conditions in <inline-formula>
<mml:math display="inline" id="im167"><mml:mrow><mml:msubsup><mml:mi>&#x211d;</mml:mi><mml:mrow><mml:mo>+</mml:mo><mml:mo>+</mml:mo></mml:mrow><mml:mn>3</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula>, arbitrarily rapid transitions may not be achievable, aligning with ecological reality where restoration unfolds over extended periods. The gradual optimal control trajectories (<xref ref-type="fig" rid="f2"><bold>Figures&#xa0;2</bold></xref>-<xref ref-type="fig" rid="f4"><bold>4</bold></xref>) reflect these fundamental dynamical constraints rather than numerical artifacts. Additionally, the dramatic cost increase when <italic>Lantana</italic> growth substantially exceeds control plant growth (<xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5</bold></xref>) indicates the system approaching the controllability boundary where control plants become critically low, underscoring the practical significance of our entropy-based sustainability index in distributing control efforts temporally to prevent unintentional population collapse.</p>
<p>These findings suggest important extensions for future work. Integrating entropy directly into the objective functional represents a profound shift in perspective, moving the optimization framework from pure cost minimization (analogous to potential energy). This &#x2018;entropic type&#x2019; functional, allows the formal control policy to balance the drive for efficiency and low cost against the necessity for action diversity and resource sustainability. The rigorous analysis of optimality conditions within this new formulation remains a significant and promising avenue for future theoretical research. Hybrid control strategies combining optimization with discrete species reintroduction events would address severely degraded sites where species are already absent. Incorporating stochasticity would account for parameter uncertainty and environmental noise, with chance constraints maintaining populations safely above zero while respecting invariant manifold limitations. Multi-timescale analysis could reveal whether exploiting fast microbial responses to steer slower plant dynamics improves controllability or reduces restoration time. Our controllability methodology also provides systematic tools for network expansion: Lie bracket calculations can predict whether adding species requires additional control variables, guiding experimental design for more complex multi-species management.</p>
<p>In conclusion, our integrated approach combining optimal control theory, entropy-based sustainability metrics, and rigorous controllability analysis ensures that control policies minimize costs while respecting fundamental dynamical constraints. The accessibility guarantees provide confidence that computed trajectories are achievable across diverse ecosystems provided management maintains all species at positive densities, while the identified limitations establish realistic boundaries for applicability and highlight where complementary approaches are necessary. This framework opens avenues for managing invasive species through plant-based interactions and provides tools for assessing control feasibility in expanded ecological networks.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Material</bold></xref>. Further inquiries can be directed to the corresponding author.</p></sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>SK: Data curation, Visualization, Methodology, Conceptualization, Validation, Writing &#x2013; original draft, Software, Formal analysis, Investigation, Writing &#x2013; review &amp; editing, Resources. PM: Resources, Methodology, Conceptualization, Validation, Visualization, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Formal analysis, Data curation, Investigation, Software. RS: Project administration, Validation, Methodology, Writing &#x2013; review &amp; editing, Investigation, Supervision, Writing &#x2013; original draft, Conceptualization, Funding acquisition.</p></sec>
<ack>
<title>Acknowledgments</title>
<p>We would like to thank the reviewers for their valuable comments and suggestions. The authors would like to thank Dr. A. Langlen Chanu, Dr. Jyoti Bhadana, S.C.C. Vyom, and Dr. Rubi Jain for valuable discussions.</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p></sec>
<sec id="s10" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declared that generative AI was not used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p></sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p></sec>
<sec id="s12" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fevo.2025.1644458/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fevo.2025.1644458/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Supplementaryfile1.pdf" id="SM1" mimetype="application/pdf"/></sec>
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