Article Type : Research Article
Authors : Pascal E, Vasquez-Pascal H, Rodriguez P and Alvarado Y
Keywords : Biomathematics, Mosquito control, Mangroves, Differential equations, Eco-epidemiological model
This research develops a biomathematical model based on nonlinear differential equations to analyze the population dynamics of Culicidae in mangrove ecosystems and their impact on controlling mosqu ito vectors. The system models the interactions between humans (H), mosquitoes (M), natural predators (P), and mangroves (B), integrating field observations conducted in Parque La Marina, Maracaibo. Stability analysis and numerical simulations demonstrate that natural predation is the most effective control mechanism, while mangroves exert a dual influence that can either amplify or suppress mosquito populations. The results highlight that the conservation of predators such as Poecilia reticulata and dragonflies, alongside appropriate mangrove management, constitutes a sustainable strategy to reduce the risk of vector-borne diseases. The study provides a quantitative framework for designing integrated control policies that balance public health and ecological conservation.
Vector-borne diseases, particularly those transmitted by mosquitoes, constitute one of the greatest threats to global public health in the 21st century. Organisms such as Aedes aegypti and Anopheles spp. are competent vectors for pathogens causing dengue, malaria, chikungunya, Zika, and yellow fever, which collectively result in hundreds of millions of cases annually and hundreds of thousands of deaths, predominantly in tropical and subtropical regions [1]. Controlling these mosquito populations is therefore a cornerstone of public health strategies to mitigate these diseases. However, controlling mosquitoes as pests and vectors faces complex challenges. Traditional interventions, such as the application of chemical insecticides, are often compromised by the development of resistance in mosquito populations, negative environmental impacts on non-target species, and high implementation costs [2]. This situation has driven the search for more sustainable, effective, and environmentally friendly integrated control strategies. In this context, mangrove ecosystems emerge as a critical ecological factor with a dual and often paradoxical influence on vector dynamics. On one hand, mangroves provide invaluable ecosystem services, such as coastal protection, carbon sequestration, and biodiversity maintenance [3]. On the other hand, the favorable conditions of these wetlands (brackish water, decomposing organic matter, and dense vegetation) can create ideal habitats for the oviposition and larval development of certain mosquito species, potentially increasing the risk of disease transmission in adjacent human communities [4]. Understanding this interaction is fundamental for designing interventions that maximize the benefits of mangroves while mitigating their potential epidemiological disadvantages. It is here that biomathematics presents itself as an indispensable tool. The construction of mathematical models, particularly through systems of differential equations, allows for the quantification and simulation of the complex nonlinear interactions between the components of an ecosystem. As noted by Smith [5] and Murray [6], these models translate qualitative ecological concepts into a quantitative framework, enabling the exploration of scenarios, the prediction of outcomes, and the evaluation of intervention strategies in silico, before their costly field implementation. A robust model can integrate the population growth rates of humans, mosquitoes, and their natural predators, along with the modulatory influence of mangroves, to identify optimal intervention points. For example, predator-prey models, widely studied in the literature [7], can be adapted to include natural mosquito predators (such as larvivorous fish, dragonflies, and insectivorous birds) that inhabit mangroves. Furthermore, the influence of the mangrove (B) can be modeled as a parameter that directly affects the growth rates (r_M) or mortality of the different species. This approach allows crucial questions to be answered: under what conditions does the mangrove act as a disease risk amplifier? How can the conservation or restoration of mangroves, coupled with the promotion of natural predators, lead to sustainable and self-regulating control of mosquito populations?
The
present article seeks to fill this gap by formulating and conducting a
preliminary analysis of a nonlinear differential equations model that captures
the ecological and epidemiological dynamics among humans, mosquitoes, their
predators, and the mangrove. The ultimate goal is to provide a mathematical
framework that serves as a basis for designing ecologically informed and
quantitatively grounded vector control strategies in these critical ecosystems.
This study employs a mixed-methods research approach, combining field observation in a specific ecosystem with the development and analysis of a theoretical mathematical model. The methodology was structured into three sequential phases: 1) Characterization of the study area and field observations, 2) Formulation of the mathematical model based on the identified ecological interactions, and 3) Theoretical analysis of the model.
Study Area and Field
Observations
Fieldwork was conducted in Parque La Marina, located in the city of Maracaibo, Zulia state, Venezuela (coordinates: 10°39'39.0"N 71°37'13.0" W). This park harbors a representative mangrove ecosystem of the Lake Maracaibo coastline, dominated primarily by species such as the red mangrove (Rhizophora mangle). This site was selected due to its proximity to urban areas, which creates a critical interface between the natural ecosystem and the human population, potentially elevating the risk of vector-borne disease transmission. Observations were conducted during periodic visits in the months of the rainy season of 2023, a period of peak mosquito activity. The objective was to identify and document the key faunal components of the local area, with special emphasis on the mosquito species present and their natural predators.
Species Collection and Identification
These empirical observations provided the fundamental ecological basis for including a predator variable (P) in the mathematical model, validating the biological relevance of the proposed interaction terms.
Formulation of the Mathematical Model
Based on the ecological interactions documented in the literature and validated by our field observations, we formulated a model of nonlinear ordinary differential equations. The model comprises four population compartments:
Dynamics of the Human Population (H)
Where: T: represents time.
The equations governing the system dynamics are
Dynamics of the Mosquito Population (M)
Dynamics of the Predator Population (P)
Dynamics of the Mangrove Biomass (B)
Theoretical Analysis of the Model
The analysis of the equation system will be carried out in the following stages:
The stability of each equilibrium will be determined by analyzing the signs of the real parts of the eigenvalues of J evaluated at that point (Routh-Hurwitz criterion) (3).
Results and Analysis
To analyze the long-term behavior of the system, the equilibrium points were calculated by solving the system of equations when
Two types of relevant equilibria were identified:
E0= (H=0, M=0, P=0, B=0), This point represents the extinction of all populations, a scenario that is ecologically catastrophic.
By solving the nonlinear algebraic system, solutions of the form E1= (H?, M?, P?, B?) were obtained, where each population persists in a steady state. The explicit solution is complex; however, numerical analysis has revealed that coexistence is possible under specific parameter conditions. For example, a representative solution is:
E1? (H?=95.2, M?=10.1, P?=22.3, B?=185.6)
These
values, obtained with a set of reference parameters, indicate a state where
mosquito and predator populations are maintained at moderate levels due to
predation pressure and the influence of the mangrove.
Jacobian Matrix and Local Stability Analysis
The Jacobian matrix of the system, essential for determining the local stability of the equilibrium points, was calculated as:
Analysis of the Trivial Point E?
Evaluating J (0,0,0,0) yields a diagonal matrix:
The eigenvalues are ?1=rH, ?2=rM, ?3=rP, and ?4=rB. Given that all intrinsic growth rates (ri) are positive by biological definition, all eigenvalues are positive.
Conclusion: Since all the real parts of the eigenvalues are negative, the equilibrium point E1 is locally asymptotically stable. The presence of complex conjugate eigenvalues suggests that the dynamics approaching equilibrium exhibit damped oscillations.
Numerical Simulations and Scenarios
To
visualize the system's dynamics and test the model's robustness, numerical
simulations were performed by integrating the differential equations using the
4th-order Runge-Kutta method [7].
Figure 1: Predators observed in Parque La Marina. Libellulidae nymph. (Archival photographs from the research team).
Table 1: Model parameters, reference values for numerical simulation, and their biological justification. The values presented constitute the baseline scenario of the model, derived from the scientific literature and calibrated with field observations in the mangrove ecosystem of Parque La Marina, Maracaibo. Each parameter includes its corresponding dimensional unit, ensuring the mathematical consistency of the system.
|
Parameter |
Worth |
Description
and Basis |
|
rH? |
0.01 |
Low human growth rate (Smith, 2018) |
|
aM? |
0.02 |
High rate of mosquito-human interaction (dengue) |
|
?H? |
0.05 |
Slightly positive influence of the mangrove |
|
rM? |
0.5 |
High intrinsic rate of mosquito growth |
|
aP? |
0.1 |
Effective predation rate (Li & Cui, 2019) |
|
?M? |
-0.01 |
Mangrove as a slightly unfavorable habitat |
|
rP? |
0.05 |
Moderate growth rate for predators |
|
?P? |
0.02 |
Mangrove is beneficial to predators |
|
rB? |
0.1 |
Mangrove growth rate |
|
? |
0.001 |
Low impact of mosquitoes on the mangrove |
|
? |
0.001 |
Low human impact (protected area) |
Figure
2: Temporal
dynamics of the ecological model under baseline conditions.
Figure
3: Scenario 1 simulation with mangrove as mosquito
amplifier (?_M = +0.02).
Figure
4: Scenario 2 simulation with 50% reduced predation
rate (aP = 0.05).
Figure
5: Scenario 3 simulation with high anthropogenic impact
on mangroves (? = 0.01, 10× Base Case).
Base Case (Figure 2)
Under the parameters from Table 1, the system converges to the equilibrium point E?, confirming the stability analysis. The mosquito population (M) is maintained at a low level (~10 individuals) due to the effective pressure from predators (P), which are sustained at a stable level (~22 individuals). Humans (H) and mangroves (B) coexist at stable levels [8]. Temporal dynamics of the ecological model under baseline conditions. Population trajectories for humans (H), mosquitoes (M), predators (P), and mangrove biomass (B) over a 100-day simulation. Horizontal dashed lines indicate the stable equilibrium values (E?: H?95.2, M?10.1, P?22.3, B?185.6). The system exhibits damped oscillations converging to equilibrium, demonstrating asymptotic stability. Key observations: (A) Human population recovers and stabilizes with mangrove support; (B) Mosquito population is suppressed by effective predation; (C) Predator population grows moderately due to prey consumption and mangrove habitat; (D) Mangrove biomass remains resilient under low anthropogenic pressure. Parameter values correspond to Table 1 (Base Case).
Scenario 1: Mangrove as Mosquito Amplifier (?_M > 0) (Figure 3)
When
the mangrove provides ideal habitats for mosquitoes (?_M = 0.02), the mosquito
population (M) reaches a significantly higher equilibrium level (~45
individuals). This leads to greater morbidity in the human population,
reflected in a slightly lower equilibrium level of H. Predators (P) also
increase due to greater prey availability, but are insufficient to completely
suppress the mosquito outbreak [9]. Population dynamics showing the
consequences of mangroves providing optimal mosquito habitats. (A) Human
population equilibrium decreases to ~87.3 due to higher disease morbidity. (B)
Mosquito population increases dramatically to ~45.8 (353% higher than Base
Case). (C) Predator population grows to ~34.2 in response to increased prey
availability, but remains insufficient for effective biological control. (D)
Mangrove biomass equilibrium reduces to ~172.1 under increased system stress.
This scenario demonstrates the epidemiological risk when mangroves function as
mosquito breeding amplifiers rather than ecological barriers.
Scenario 2: Predation
Effectiveness (Figure 4)
A
50% reduction in the predation rate (aP=0.05) was simulated, representing, for
example, a decline in P. reticulata populations due to pollution. The result
was a drastic increase in the equilibrium mosquito population (~65
individuals), demonstrating the critical importance of this biological control
mechanism. The human population is seriously affected [10]. This scenario
simulates the loss of predator effectiveness, e.g., due to pollution affecting
P. reticulata populations. (A) Human population equilibrium dramatically
decreases to ~71.5 (24.9% reduction from Base Case), indicating a severe public
health impact. (B) Mosquito population explodes to ~65.4 (547% increase from
Base Case), demonstrating the critical importance of biological control. (C) Predator
population shows limited growth to ~24.6 despite increased prey availability,
highlighting the functional limitation of reduced predation rates. (D) Mangrove
biomass declines to ~154.8, reflecting ecosystem stress from trophic imbalance.
This scenario provides the strongest evidence for protecting predator
populations as a frontline defense against mosquito-borne diseases.
Scenario 3: High
Anthropogenic Impact (? > 0) (Figure 5)
An increase in the rate of mangrove degradation by humans (? = 0.01) leads to a decrease in mangrove biomass (B). Since, in the base case, the mangrove has a net positive effect on predators (?_P > 0), its decline results in a reduction of the predator population (P), which indirectly allows for an increase in mosquitoes (M). This illustrates an ecosystem-mediated "cascade effect" [11]. This scenario demonstrates a clear ecosystem cascade effect: (D) Mangrove biomass decreases dramatically to ~81.3 due to human degradation (-56.2% from Base). (C) Predator population declines to ~15.2 (-31.8% from Base) due to loss of mangrove habitat and resources (?P>0). (B) Mosquito population increases to ~38.7 (+283% from Base) as biological control weakens. (A) Human population equilibrium decreases to ~82.4 (-13.4% from Base) due to increased mosquito-borne disease morbidity. The cascade (Human impact ? Mangrove loss ? Predator decline ? Mosquito increase ? Human health impact) illustrates the interconnectedness of ecosystem health and public health, showing that environmental degradation ultimately rebounds negatively on human populations.
A local sensitivity analysis was conducted using partial derivatives to identify the parameters with the greatest influence on the equilibrium mosquito population (M*). The results indicate that M* is most sensitive to:
The
present study developed and analyzed a nonlinear mathematical model that
integrates the population dynamics of humans, mosquitoes, their predators, and
mangroves in a coastal ecosystem. The results obtained provide profound
theoretical insights into the forces governing vector proliferation and offer a
quantitative framework for evaluating control strategies.
Interpretation of Key
Results and Control Mechanisms
The
central finding of this work is the demonstration that a stable coexistence
state (E?) is achievable under plausible ecological conditions. The stability
of this equilibrium, characterized by damped oscillations, suggests that the
ecosystem possesses an inherent resilience capable of buffering minor
perturbations. However, the key to public health lies in the levels at which
populations stabilize, particularly that of mosquitoes [12]. The sensitivity analysis identified the predation
rate (a_P) as the most critical factor for suppressing the mosquito population.
This empirically validates the field observations conducted in Parque La
Marina, where predators such as Poecilia reticulata and Libellulidae nymphs
were present. Our numerical models show that a decrease in a_P (Scenario 2)
leads to a loss of control over the mosquito population. This has a direct
implication for management: strategies that protect and promote natural
predators (such as banning broad-spectrum pesticides that affect them or the controlled
introduction of larvivorous fish into water bodies) are likely among the most
effective for sustainable long-term control [7, 13]. Secondly, the parameter ?_M, which quantifies the
effect of the mangrove on mosquitoes, proved to be a powerful modulator of the
system. The model effectively captures the dual nature of the mangrove. When
?_M > 0 (Scenario 1), the mangrove acts as a vector amplifier, increasing
epidemiological risk. Conversely, when ?_M < 0 (as in our Base Case), the
mangrove can contribute to mosquito suppression, possibly by promoting
predators or creating unfavorable breeding conditions. This underscores that
mangrove management cannot be homogeneous; it must be based on a specific
diagnosis of how the structure and health of the local ecosystem influence
vector ecology [4]. Finally, the model reveals
a vitally important indirect cascade effect (Scenario 3). Anthropogenic
degradation of the mangrove (high ?) not only negatively impacts the biomass of
B but also, by weakening its supportive role for predators (?_P > 0),
indirectly leads to an increase in the mosquito population. This highlights
that mangrove conservation is not merely an ecological goal but a potent public
health strategy, as emphasized by (3).
Relationship with
Existing Literature and Model Contribution
Our work
aligns with and extends previous literature. On one hand, it agrees with
classical predator-prey models that highlight top-down control as a fundamental
regulator of insect populations (7). On the other hand, the explicit inclusion
of the mangrove (B) as a dynamic compartment influencing all species goes a
step beyond traditional epidemiological models, which often treat the
environment as a static background (6). The novelty of our approach lies in integrating the
ecological-epidemiological interface into a single mathematical framework.
While the WHO (2022) advocates for "integrated vector management," it
often lacks quantitative tools to predict the consequences of specific
interventions in complex systems. This model provides a "virtual
laboratory" for testing such interventions, such as predator conservation
programs or mangrove habitat management, before their field implementation.
Study Limitations and
Future Directions
It is crucial to recognize
the limitations of this model as a simplification of reality. First, the model
assumes spatial homogeneity, ignoring the patchy distribution of mosquitoes and
predators within the mangrove. Future iterations could benefit from a
metapopulation or agent-based modeling approach to capture this heterogeneity.
Second, the current model aggregates all predators
into a single variable (P). A promising future research direction would be to
disaggregate this variable to separately model the dynamics of aquatic
predators (such as P. reticulata) and terrestrial predators (such as spiders or
birds), each with their own vital rates and modes of interaction.
Third, the model does not explicitly differentiate
between susceptible and infected mosquitoes. Incorporating epidemiological
sub-compartments (for example, using an SIR framework for humans and an SI
framework for mosquitoes) would allow for the direct study of disease
prevalence and the impact of different strategies on the pathogen's basic
reproduction number R? (6). Finally, the fine-tuning of
parameters with real longitudinal data from Parque La Marina is an essential
step to transform this theoretical model into a robust predictive tool for
public health and environmental managers in the city of Maracaibo, Venezuela
[14].
This study has developed
and analyzed an innovative biomathematical model that integrates the population
dynamics of humans, mosquitoes, predators, and mangroves in a coastal
ecosystem. Through the analysis of nonlinear differential equations, numerical simulations,
and field observations in Parque La Marina, Maracaibo, significant conclusions
have been drawn that contribute to both theoretical knowledge and practical
applications in the control of epidemiological vectors.
The model demonstrates the existence of a locally
asymptotically stable non-trivial equilibrium point E?, where all populations
coexist. This equilibrium is characterized by damped oscillations, indicating
the system's capacity to recover from minor perturbations, provided key
ecological interactions are maintained. The sensitivity analysis revealed that the
equilibrium mosquito population (M*) is particularly sensitive to the predation
rate (a_P*). This scientifically validates the field observations that
identified Poecilia reticulata and Libellulidae nymphs as crucial
biological control agents in the mangroves of Parque La Marina.
The parameter ?_M proved to be a determining
factor: when positive, the mangrove amplifies mosquito populations; when
negative, it contributes to their control. This duality underscores the need
for specific evaluations of each mangrove ecosystem.
The model identified that human impact on mangroves
(?) not only reduces their biomass but also indirectly increases mosquito
populations by weakening the ecological support for natural predators.
This study demonstrates how mathematical models can
serve as virtual laboratories to evaluate intervention strategies, allowing for
the optimization of resources and the prediction of non-intuitive consequences
of management policies.