Nurfadillah Nurfadillah
| Nur Fadli* | Zulkarnain Jalil | Noor Adelyna Mohammed Akib
© 2025 The authors. This article is published by IIETA and is licensed under the CC BY 4.0 license (http://creativecommons.org/licenses/by/4.0/).
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Feeding habits and growth patterns of fish in lake ecosystems are influenced by food availability, competition, predation, and environmental conditions. This study aimed to analyze the feeding habits and length-weight relationships of native and invasive fish species in Aneuk Laot Lake, Sabang, Indonesia. Fish were sampled using traps and gill nets with mesh sizes of 0.5, 0.75, and 1 inch. A total of 483 fish were collected: 98 Rasbora sp., 104 Barbodes sp., and 218 Amphilophus trimaculatus. Feeding habits were examined using the digestive tract dissection method, and the data were analyzed based on the volumetric index, frequency of occurrence, index of preponderance, niche breadth, and niche overlap. Growth patterns were assessed through length-weight relationship analysis using linear regression and t-test analysis. Results indicated that Rasbora sp. is herbivorous, primarily consuming Cosmarium sp. (84.4%), whereas Barbodes sp. preferred Daphnia sp. (45.1%). A. trimaculatus primarily preys on small fish (62.5%). Both Barbodes sp. and A. trimaculatus are omnivorous. There was a high dietary overlap (0.96) between Rasbora sp. and Barbodes sp., indicating strong interspecific competition. A. trimaculatus preys on Rasbora sp., demonstrating its impact as an invasive predator. All species exhibited negative allometric growth (b < 3), suggesting environmental stress or food limitations. These findings highlight the need for managing invasive species to conserve native fish populations and maintain ecological balance.
aquatic resources management, indigenous fish, length-weight relationships (LWRs), niche breadth, trophic ecology
Food is a critical biological factor that determines the survival of a fish [1, 2]. The feeding habits of fish serve as a basis for assessing their role and position in the food chain [3-5]. Additionally, information on diet is essential for managing sustainable fisheries and plays a vital role in selecting suitable species for aquaculture [6, 7]. The utilization of identical food sources by fish in the wild leads to direct competition [8-11].
Each species has a unique ecological niche, encompassing how it utilizes resources and interacts with its environment. This niche includes aspects such as diet composition, feeding time, habitat, and interactions with other species [12-14]. These complex interactions shape a species’ realized niche, which is often narrower than its fundamental niche due to competition and other ecological constraints [15, 16]. If two species occupy highly similar niches, intense competition for limited resources is likely to occur. This competition can reduce the availability of resources for native fish, ultimately leading to a decline in population size and local fish species diversity. Additionally, invasive fish that act as efficient predators and prey on native fish, including juveniles and eggs, can cause a drastic decline in native fish populations.
The presence of invasive fish species can disrupt the existence of native fish species [17-23]. Similarly, concerns arise regarding the presence of non-native fish species, which affect competition for food and habitat [22, 24-26]. As one of the main doctrines in ecological theory, this principle states that two species competing for the same limited resources cannot coexist stably [15]. This principle, first formally formulated by G.F. Gause, highlights the instability of ecological systems in which multiple species compete for the same resources [27, 28]. The Competitive Exclusion Principle states that the species that utilizes resources more efficiently will outcompete and eventually eliminate the other species from its habitat [27].
This situation occurs in Aneuk Laot Lake. Aneuk Laot Lake is the second-largest lake in Aceh Province [29]. This lake provides significant benefits to the community in Sabang City, including its use as a fishing area for residents around the lake [30, 31]. The supply of fish for consumption in Aneuk Laot Lake has primarily relied on the catches from local fishermen operating within its waters. Intensive fishing by the community is believed to cause high fishing pressure, which could lead to a decline in fish resources. The decline in fish species composition is suspected to be caused by several issues, including increasing activities along the lake’s riparian zone each year, contributing to the pollution load entering the lake’s water body. The rise in anthropogenic waste loads disrupts the fish habitats in the lake. Environmental degradation, in turn, affects fish production [30, 32-34].
Research on the feeding habits of fish in Aneuk Laot Lake is particularly important, as the lake serves as a vital habitat for various species, including some that may be endemic [35]. Research on the feeding habits of fish in the waters of Aneuk Laot Lake has been previously conducted, but it was limited to invasive fish species that were caught. In contrast, the feeding habits of native fish in this lake have never been studied. This report represents the first documented study on the feeding habits and growth patterns of native fish species in Aneuk Laot Lake. This study is an essential strategic step for ensuring the survival of species and the sustainability of their ecosystems. Understanding habitats as feeding grounds is crucial for gaining insights into the ecological processes within the lake and how different species interact within the ecosystem [36-38].
Continuous fluctuations in the Aneuk Laot Lake ecosystem pose a serious threat to the survival of aquatic organisms. These threats include potential habitat loss and biodiversity loss within the lake environment. Therefore, although fish resources are fundamentally renewable, they are limited, necessitating proper utilization, protection, and management to ensure their long-term availability and ecological balance [39, 40]. As a result, empirical research consistently demonstrates that coexisting species tend to differentiate in at least one key aspect of their ecological niche, thereby reducing direct competition and enabling resource partitioning. This mechanism allows multiple species to exploit different resources or utilize the same resources in slightly different ways, thereby minimizing interspecific competition.
In this context, species can avoid direct competition by utilizing resources differently or occupying different spatial and temporal niches within the ecosystem [27]. Therefore, this principle highlights the dynamic interaction between competition and niche differentiation in shaping community structure and biodiversity [41]. Quantifying a species' niche is a complex task that often requires an in-depth analysis of resource use patterns, habitat preferences, and activity over a given time frame [15]. A comprehensive understanding of species niches is crucial for predicting community assembly dynamics, forecasting species responses to environmental disturbances such as climate change and habitat loss, and anticipating the ecological consequences of species introductions and removals. All these aspects play a vital role in effective conservation management and ecosystem restoration. Therefore, our hypothesis in this study is that the non-native fish Amphilophus trimaculatus has a broad niche breadth, which may disrupt the presence of native fish species in Lake Aneuk Laot, specifically Rasbora sp. and Barbodes sp.
This research can provide the necessary data to formulate adaptation and mitigation strategies and serve as a basis for developing conservation and ecosystem management policies. Accurate and up-to-date data are essential for effective policies and appropriate interventions [42, 43]. One of the approaches to support the conservation and sustainability of native fish resources in Aneuk Laot Lake is through studies on feeding habits, fish growth, and the factors influencing habitat availability for aquatic biota. These studies are crucial as a foundation for sustainable management, particularly for Aneuk Laot Lake, which is classified as a productive water body. The presence of native fish species is crucial for the sustainability of fisheries in this lake and serves as a biological indicator of the aquatic ecosystem through the analysis of fish length and weight. This provides a foundation for implementing protective measures to mitigate the impacts of invasive fish species [44-47].
This study was conducted from March to November 2024 in Aneuk Laot Lake, Sabang City, Weh Island, Indonesia. Fish sampling was carried out every two months over nine months at three observation stations selected based on habitat characteristics. Aneuk Laot Lake is a unique lake as it has no inlet or outlet [29, 48]. It is relatively small, covering an area of approximately ±39.86 hectares, with a maximum length of 1,419 meters, an average width of 250 meters, and a maximum depth of 22 meters [29].
Fish collection was conducted through direct capture using traps and gill nets. The gill nets used varied in size, with mesh sizes of 0.5-inch, 0.75 inch, and 1 inch. The deployment of traps and nets was performed twice daily: the first deployment occurred at 07:00 WIB and was retrieved at 17:00 WIB, while the second deployment occurred at 18:00 WIB and was retrieved at 06:00 WIB. The research location map is presented in Figure 1.
Figure 1. The research location map of Aneuk Laot Lake
A total of 483 fish individuals were sampled for analysis of feeding habits and growth patterns, consisting of 98 Rasbora sp., 104 Barbodes sp., and 218 Amphilophus trimaculatus. Observations indicated that Rasbora sp. was more frequently captured in calm areas with greater depths (Station 1). In contrast, Barbodes sp. and Amphilophus trimaculatus were predominantly captured in areas densely populated with aquatic vegetation (Stations 2 and 3).
Fish length was measured as the total length (from the tip of the mouth to the tip of the caudal fin) using a caliper with a precision of 0.1 cm. Total fish weight was measured using a digital scale with a precision of 0.01 grams.
The analysis of fish feeding habits involved dissection using surgical scissors, starting from the anus toward the upper abdomen below the lateral line and along the lateral line to the posterior of the operculum, then ventrally toward the abdominal cavity. After dissection, the fish's digestive organs (stomach and intestines) were preserved in 4% formalin [8, 49]. Subsequent observations of the digestive tract contents were conducted at the Marine Biology Laboratory, Faculty of Marine and Fisheries, Syiah Kuala University, Banda Aceh.
The observations of fish digestive tract contents were analyzed for feeding habits using several parameters, including the volumetric method, index of preponderance, frequency of feeding occurrence [50], niche breadth, and dietary overlap analysis. The volumetric method was applied using the formula referenced from [50].
$V=\frac{V i}{V t} \times 100 \%$ (1)
where, V represents the percentage of a specific type of food (%), vi is the volume of a specific type of food (ml), and vt is the total volume of all types of food (ml).
The calculation of the index of preponderance is conducted to determine the types of food consumed by fish. The index of preponderance combines two methods: the frequency of occurrence method and the volumetric method, using the formula equation proposed by study [49].
$I P=\frac{V i \times F O}{\Sigma V i \times F O} \times 100 \%$ (2)
where, IP represents the index of preponderance (%), Vi is the percentage volume of a specific food type (%), FO is the percentage frequency of occurrence of a specific food type (%), and ΣVi × FO is the total of Vi × FO for all food types.
Subsequently, the identification of all stomach contents was conducted using the frequency of occurrence method, calculated with the following formula [49].
$F O(\%)=\frac{The\,\, total\,\, occurrance\,\, of\,\, a\,\, specific\,\, food\,\, type}{The\,\, total\,\, number\,\, of\,\, stomachs\,\, containing\,\, food} \times 100$ (3)
Testing the difference in feeding habits among fish groups using the Chi-Square ($\chi^2$) statistical test. The food niche breadth indicates the diversity of food types consumed by fish. The determination of niche breadth, according to Levin’s Niche Breadth index, is calculated using the following formula:
$B i=\frac{1}{\Sigma P i^2}$ (4)
where, Bi represents the food niche breadth, and Pi is the proportion of fish species associated with a specific food type.
In this calculation, standardization is required to ensure that the resulting niche breadth values range between 0 and 1, with intervals that are neither too large nor insignificant. Therefore, a formula based on study [51] is applied.
$B A=\frac{B i-1}{n-1}$ (5)
where, BA is the standardized niche breadth (range 0–1), Bi represents the niche breadth, and ‘n’ is the total number of food organisms utilized.
Subsequently, to calculate the overlap value between fish, the formula proposed by index Pianka [52] can be used. This approach examines the dietary competition based on the contents of the fish intestines, allowing for the identification of similarities or differences in the types of food utilized by the fish.
$O_{i j}=\frac{\Sigma\left(p_i p_j\right)}{\sqrt{\Sigma\left(p_i^2\right) \Sigma\left(p_j^2\right)}}$ (6)
where, $O_{i j}$ represents the overlap between species i and species j, $p_i p_j$ are the proportions of food resources utilized by species i and j, respectively, and $\sum\left(p_i p_j\right)$ is the sum of the products of the proportions of the same food resources consumed by both species.
The relative length of the gut (RLG) can be calculated using the formula provided by study [50].
$R L G(\%)=\frac{Gut\,\, length\,\,(mm)}{Total\,\, body\,\, length\,\, (mm)}$ (7)
Gut length refers to the total length of the intestine of the fish, typically measured from the beginning of the digestive tract to the end. It is used in calculating the RLG to assess the feeding habits and dietary classification of the fish species.
Additionally, growth patterns can be analyzed through the calculation of the length-weight relationship of fish, referring to the formula by [53].
$W=a L^b$ (8)
where, W represents the total weight of the fish (g), and L is the total length of the fish (cm). The parameters a and b are constants. To test whether b = 3 or b ≠ 3, a t-test (partial test) is conducted with the following hypotheses: H0: b=3, indicating that the relationship between length and weight is isometric, and H1: b ≠ 3, indicating that the relationship between length and weight is allometric. The decision is made by comparing the p-value to the significance level (α=0.05), If the p-value < 0.05, reject the null hypothesis (H0), meaning the difference is statistically significant. If p-value ≥ 0.05, fail to reject the null hypothesis, meaning there is no significant difference.
The value of b characterizes the growth pattern with the following criteria: If b = 3, the growth pattern is isometric, meaning that the fish's length and weight grow at the same rate. If b > 3, the growth pattern is positive allometric, indicating that the weight grows faster than the length. and if b < 3, the growth pattern is negative allometric, meaning that the length grows faster than the weight. Isometric growth signifies balanced growth in length and weight, while negative allometric growth indicates faster length growth compared to weight. Conversely, positive allometric growth means that weight growth outpaces length growth [54, 55].
3.1 Food habits of native and invasive fish
The feeding habits of native and invasive fish often differ significantly due to their ecological roles and adaptability. Native fish typically feed on food resources available within their natural habitat, maintaining a balanced ecosystem [19, 25]. In contrast, invasive fish often exhibit broader feeding plasticity, allowing them to exploit various food sources, which can lead to competition with native species [56, 57].
The focus of this study is on the feeding habits and growth patterns of three fish species captured in the waters of Aneuk Laot Lake, consisting of two native species (Rasbora sp. and Barbodes sp.) and one invasive species (Amphilophus trimaculatus). The analysis of the feeding habits of the three fish species is presented in Table 1.
Table 1. Feeding habits of Barbodes sp., Rasbora sp., and Amphilophus trimaculatus based on the volumetric index (V %), frequency of feeding occurrence (FO %), and index of preponderance (IP %)
|
No. |
Species of Fish |
Organism of Food |
V (%) |
IP (%) |
FO (%) |
|
1 |
Barbodes sp. |
Daphnia sp. Botryococcus sp. Cosmarium sp. Synedra sp. Frustulia sp. Aquatic plant Ant Detritus |
29.45 12.71 19.99 2.98 2.71 11.43 15.98 4.76 |
45.14 8.85 16.70 0.83 0.75 6.372 0.03 1.33 |
61.1 27.7 33.3 11.1 11.1 22.2 50.0 11.1 |
|
2 |
Rasbora sp. |
Scenedesmus sp. Synedra sp. Cosmarium sp Frustulia sp. Skeletonema sp. Tabellaria sp. Botryococcus sp. Pedistrum sp. |
4.61 2.71 63.68 9.16 5.08 2.01 12.74 0.66 |
0.96 0.94 84.4 4.47 2.48 0.42 6.22 0.09 |
15.0 25.0 95.0 35.0 35.0 15.0 35.0 10.0 |
|
3 |
Amphilophus trimaculatus |
Fish Euphausiacea Detritus Cosmarium sp. Daphnia sp. Synedra sp. Coscinodiscus sp. Skeletonema sp. Frustulia sp. Botryococcus sp. |
35.39 4.18 5.07 19.22 7.39 7.12 2.25 1.06 4.36 10.26 |
62.59 2.28 1.38 18.30 5.03 2.90 0.61 0.14 1.19 5.58 |
48.1 14.8 7.40 25.9 18.5 11.1 7.40 3.71 7.40 14.8 |
Based on the digestive tract contents of the three fish species, various organisms were identified. However, plankton was found in all the fish studied. Differences were observed in the primary food preferences, leading to distinct feeding patterns. According to the index of preponderance, the main food of Rasbora sp. was Cosmarium sp. at 84.4%, while the primary food of Barbodes sp. was Daphnia sp. at 45.14%. For Amphilophus trimaculatus, the main food identified was fish, accounting for 62.59% (Table 1). This is consistent with the statement by study [6], which classifies an IP% value of ≥45% as a primary food, an IP% of > 4 – 25% as supplementary food, and an IP% of ≤4% as complementary food.
Based on the composition of food types found in the digestive tracts of the fish, it was determined that Rasbora sp. belongs to the herbivorous group. The study results show that, according to the index of preponderance (IP), the primary food for Rasbora sp. is Cosmarium sp. (84%), while Botryococcus sp. (6%) and Frustulia sp. (4%) serve as complementary foods. Meanwhile, Scenedesmus sp., Synedra sp., Skeletonema sp., Tabellaria sp., and Pediastrum sp. are supplementary foods.
In contrast, Barbodes sp. and Amphilophus trimaculatus are classified as omnivorous fish, meaning their digestive tracts contain both animal-based and plant-based food types [6]. For Barbodes sp., Daphnia sp. (45%) was identified as the primary food, while Botryococcus sp., Cosmarium sp., insects, and aquatic plants serve as complementary foods, with IP values ranging from 4% to 25%. On the other hand, detritus, Synedra sp., and Frustulia sp. are only supplementary foods for Barbodes sp., with IP values of less than 4%.
For Amphilophus trimaculatus, the primary food is fish fry, accounting for 62.59%. Complementary foods include Cosmarium sp. (18%), Botryococcus sp. (5%), and Daphnia sp. (5%), while supplementary foods are Euphausiacea, Synedra sp., Coscinodiscus sp., Skeletonema sp., Frustulia sp., and detritus, all with IP values of less than 4%. Amphilophus trimaculatus is an opportunistic feeder, meaning it utilizes any food resources available in its environment. The Amphilophus trimaculatus, commonly known as the three-spot cichlid, has a varied diet that mainly includes small fishes, macro-invertebrates, and aquatic and terrestrial insects [58-60].
Understanding these feeding habits is essential for assessing ecological interactions, resource competition, and the impact of invasive species on native populations and the overall ecosystem [22, 26]. Observations of feeding habits revealed that Amphilophus trimaculatus (Three-spot cichlid) was found to prey on small fish, suspected to be Rasbora sp. (Figure 2). This conclusion is based on the discovery of cycloid scales in the stomach contents of Amphilophus trimaculatus. A frequency of occurrence of 48% indicates that nearly half of the analyzed Amphilophus trimaculatus samples contained cycloid scales in their stomachs. These findings suggest that Rasbora sp. constitutes a significant part of the diet of Amphilophus trimaculatus in this study, highlighting predatory patterns and interspecies interactions within their habitat. This predation likely influences the distribution and abundance of Rasbora sp., a native fish species of Aneuk Laot Lake. Therefore, it is crucial to implement control measures for the invasive species (Amphilophus trimaculatus), which damages habitats and disrupts native fish populations. Native species tend to be more sensitive to environmental changes compared to invasive species, which generally have a higher tolerance. In freshwater ecosystems, some top predators exhibit significant overlap in their trophic niches, reflecting similarities in their feeding patterns [61]. The trophic niche space, about the structure of the food web, can change over time. Understanding these changes is crucial for assessing resource utilization and trophic interactions [61]. A species’ niche encompasses various relationships it maintains within its ecosystem, including its spatial and temporal positioning, adaptations to the environment, and resource utilization patterns [61].
(a)
(b)
(c)
Figure 2. Feeding strategies of (a) Barbodes sp., (b) Rasbora sp., and (c) Amphilophus trimaculatus based on prey frequency of occurrence (FO, %) plotted against prey-specific abundance (PSA, %)
The invasive species often cause major changes in ecosystems because they dominate resources and outcompete native species. The introduction of non-native fish species has significantly altered freshwater ecosystems, frequently leading to a decline in native species richness and overall functional diversity [62]. The rapid shifts driven by invasive fish highlight the importance of understanding the interaction between environmental characteristics and the ability for native and non-native species to coexist [63]. Invasive fish are especially hard to control because the aquatic environment presents challenges for detection, monitoring, and removal efforts [64]. The findings of this study serve as a fundamental dataset for the management of Aneuk Laot Lake, with a focus on controlling invasive fish populations, conserving native fish habitats, and regulating fisheries to maintain ecosystem balance and ensure optimal fish population growth.
Contrarily, Rasbora sp. preys entirely on plankton, with a frequency of occurrence of 100%, indicating that it is a pure planktivore (Figure 2). This means the feeding habits of this fish are exclusively composed of plankton, including phytoplankton and zooplankton. This behavior signifies a specific ecological adaptation to its habitat, making it an important component of the food chain. The availability of plankton directly affects the survival of Rasbora sp., thereby supporting its growth and reproduction (Figure 2).
Meanwhile, Barbodes sp. utilizes a variety of food resources, including plankton, insects, aquatic plants, and detritus. The highest frequency of food occurrence was recorded for Daphnia sp. at 61.1%, followed by insects at 50%. This indicates that the feeding habits of this fish are highly diverse. The strong tendency to consume Daphnia sp. highlights its preference for this type of plankton, while the high consumption of insects reflects the fish's adaptation to the available food resources in its habitat. Such feeding patterns contribute to maintaining ecological balance by regulating the populations of their prey species and supporting the integrity of trophic structures within their ecosystem.
3.2 Niche breadth and dietary overlap
The food niche refers to the range of food resources utilized by a species within its ecosystem, reflecting its role in the trophic structure. Dietary overlap occurs when different species exploit similar food resources, potentially leading to competition [65, 66]. Analyzing food niche breadth and dietary overlap helps in understanding interspecific interactions, resource partitioning, and the potential impact of invasive species on native populations [67, 68].
Niche breadth describes the extent of food utilization by a specific fish species and helps determine dietary selectivity within its group [69]. A small niche breadth value indicates that a fish species is selective toward food resources in its environment, whereas a large value suggests a diverse range of consumed food types [70]. Based on fish size, the food niche breadth value for Barbodes sp. was 3.64 with a standardized value of 0.66, Rasbora sp. had a niche breadth of 1.39 with a standardized value of 0.02, and Amphilophus trimaculatus exhibited a niche breadth of 2.31 with a standardized value of 0.03 (Table 2).
Table 2. Food niche breadth of native and invasive fish in Aneuk Laot Lake
|
Species of Fish |
Niche Breadth |
Standardized |
|
Barbodes sp. |
3.64 |
0.66 |
|
Rasbora sp. |
1.39 |
0.02 |
|
Amphilophus trimaculatus |
2.31 |
0.05 |
Based on the food niche breadth values presented in Table 2, Barbodes sp. exhibits a more specific dietary adaptation. The range of values indicates that this species has a more diversified feeding pattern compared to Rasbora sp., suggesting that Rasbora sp. is more dependent on certain prey types that are consistently available. However, Amphilophus trimaculatus demonstrates a higher adaptability to changes in prey availability, indicating that this species has greater dietary flexibility.
Native fish species (Barbodes sp. and Rasbora sp.) generally exhibit specialized adaptations to utilize locally available food resources. They can consume a variety of food types, including plankton, aquatic invertebrates, aquatic plants, and even other fish, depending on the species and ecological conditions of the lake. The availability of these food resources plays a crucial role in supporting the health and population of native fish while maintaining the ecological balance of the lake. However, the presence of invasive species or environmental changes can disrupt food availability and negatively impact native fish populations.
Invasive fish species, such as Amphilophus trimaculatus, often have a broader niche breadth compared to native species, allowing them to compete more effectively and dominate food resources [19, 26]. A large niche breadth in invasive fish enables them to adapt rapidly to new habitats, contributing to their success as invasive species while impacting trophic structure and biodiversity within the ecosystems they inhabit.
Non-native fish species typically adopt a generalist feeding strategy with adaptable dietary patterns, enabling them to establish themselves and thrive as invasive species. In this study, the invasive fish Amphilophus trimaculatus exhibited a mixed and generalized feeding strategy, utilizing the available food resources within the Aneuk Laot Lake habitat. Table 3 illustrates the shared utilization of food resources between Rasbora sp. and Barbodes sp., with a dietary overlap value of 0.96, which is close to 1. In contrast, the invasive fish Amphilophus trimaculatus showed a lower dietary overlap with native fish species, ranging from 0.47 to 0.49. According to study [71], invasive species influence the demographics of native species through predation and competition for food resources. In this study, Amphilophus trimaculatus was found to have the potential to impact the population of the native fish species Rasbora sp. through predation, as evidenced by the presence of Rasbora sp. scales in its stomach contents. This impact extends beyond economic consequences, affecting the balance of the food chain and potentially triggering cascading effects within the ecosystem. In general, invasive fish tend to adopt an opportunistic feeding strategy, meaning they do not rely on specific food types but instead adjust their diet based on the availability of food in their environment. Fish species with this strategy exhibit flexibility in prey selection and can utilize a wide range of food resources depending on habitat conditions and seasonal variations.
Table 3. Dietary overlap values of Barbodes sp., Rasbora sp., and Amphilophus trimaculatus captured in Aneuk Laot Lake, Sabang City, Aceh
|
|
Rasbora sp. |
Barbodes sp. |
Amphilophus trimaculatus |
|
Rasbora sp. |
- |
0.96 |
0.49 |
|
Barbodes sp. |
0.96 |
- |
0.47 |
|
Amphilophus trimaculatus |
0.49 |
0.47 |
- |
Figure 3 illustrates the shared utilization of food resources between native and invasive fish species. Native fish in Aneuk Laot Lake exhibit a specialist-stenophagic feeding strategy, whereas invasive fish demonstrate a generalist-opportunistic feeding behavior. Native fish populations with a specialist and stenophagic-monophagic feeding strategy are less likely to expand, making them more vulnerable to environmental changes and competition. A similar situation is observed with the presence of flowerhorn fish (Amphilophus trimaculatus) in Aneuk Laot Lake, where its population continues to grow, posing a threat to the declining native fish populations. The presence of flowerhorn fish was first reported by researcher [8], whereas previous research [72] did not identify Amphilophus trimaculatus in the lake, suggesting that its introduction occurred after that period.
Figure 3. Variations in food types of native and invasive fish in Aneuk Laot Lake, with Barbodes sp. represented by the blue label, Rasbora sp. by the green label, and Amphilophus trimaculatus by the red label
The interplay between niche overlap and competitive differences significantly shapes the evolutionary trajectory of ecological communities [73]. While an increase in the number of competing species doesn't necessarily decrease tolerable niche overlap due to environmental variability, it does highlight the relationship between overlap and diffuse competition [52]. The degree of niche overlap and the magnitude of competitive differences are intricately governed by a network of complex ecological processes, consequently opening diverse avenues for species to coexist within a shared environment [73, 74]. Species can coexist by differentiating resource use, thereby minimizing direct interactions, or by maintaining partially overlapping resource use alongside near-equivalent average competitive abilities. The initial conditions of a community, including factors such as resource availability, environmental stability, and the existing species composition, play a crucial role in determining whether evolutionary shifts in niche overlap or competitive abilities will foster or hinder species coexistence [74].
Study [50] stated that fish species capable of adapting their feeding habits can significantly influence their population dynamics in natural habitats. A similar phenomenon was observed in Lake Matano, where the flowerhorn fish exhibited strong adaptability, leading to a population increase [75]. A precautionary approach is necessary when introducing new fish species into Aneuk Laot Lake, as the establishment of new species can alter the food chain structure within the fish community. High ecological pressure in the form of niche competition from introduced fish populations may threaten the existence of native fish species [76].
3.3 Growth patterns of native and invasive fish
The length-weight relationship of fish is a crucial biological parameter for understanding growth, body condition, and the ecology of a species. Fish feeding habits play a significant role in determining growth patterns [77, 78]. In this study, Barbodes sp. exhibited a negative allometric growth pattern, meaning that the proportional increase in length was faster than in weight. This finding aligns with the study [79], which reported that the growth pattern of Barbodes sp. identified in Krueng Simpoe also followed a negative allometric pattern. Similarly, study [80] stated that the growth pattern of Barbodes lateristriga in the Tebudak River, Pisak Village, Bengkayang Regency, was negative allometric for both male and female fish. In contrast, a study conducted in a national park in northern Vietnam found that Barbodes semifasciolatus exhibited a positive allometric growth pattern [81]. Study [35] stated that fish growth and fluctuations in food resources in aquatic environments are influenced by seasonal variations.
A similar pattern was observed in Rasbora sp. from Aneuk Laot Lake, which exhibited a negative allometric growth pattern with a b value of 2.77 (Figure 4). Referring to [53], fish growth patterns are determined based on the b value, where growth is classified as isometric (b = 3), negative allometric (b<3), or positive allometric (b>3). The length-weight relationship of fish is an important indicator of their overall health and condition [82, 83].
(a)
(b)
(c)
Figure 4. Length-weight relationship of native and invasive fish in Aneuk Laot Lake, Sabang City, Aceh (a) growth pattern of Barbodes sp., (b) growth pattern of Rasbora sp., and (c) growth pattern of Amphilophus trimaculatus
In fish species experiencing high predation pressure, individuals tend to grow longer before increasing their body weight. This phenomenon is evident in the native fish of Aneuk Laot Lake (Rasbora sp. and Barbodes sp.), which serve as prey for Amphilophus trimaculatus. The growth pattern of Amphilophus trimaculatus also follows a negative allometric trend. This is likely due to numerous predators in a shared habitat, leading to competition among these fish for the same prey resources. As a result, individual predators may not obtain sufficient food to adequately increase their body mass.
Interestingly, this study found a similar growth pattern for both native and invasive fish, with a b value of less than 3, indicating a negative allometric growth pattern (Figure 4). This suggests that environmental degradation in the waters of Aneuk Laot Lake, such as pollution, sedimentation, and temperature fluctuations, may be causing stress in these fish, leading to inhibited growth. Study [84] reported that Lake Aneuk Laot is classified as heavily polluted, with several water quality parameters exceeding environmental standards, including total suspended solids (TSS), biochemical oxygen demand (BOD), chemical oxygen demand (COD), dissolved oxygen (DO), and total phosphate. Furthermore, study [84] explained that sedimentation disrupts the photosynthesis process and leads to the mortality of aquatic organisms. The TSS concentration in Aneuk Laot Lake was recorded at 102.25 mg/L, exceeding the permissible water quality standards for lakes.
In contrast, studies conducted in Lake Matano, Sulawesi, found that flowerhorn fish in those waters exhibited a positive allometric growth pattern [85]. Meanwhile, other studies reported that flowerhorn fish in the P.B. Soedirman Reservoir, Central Java, demonstrated an isometric growth pattern [86]. These differences indicate that fish growth patterns can vary depending on environmental conditions and other influencing factors. Furthermore, slope values (b) can fluctuate primarily due to diverse environmental, human-induced, and biological factors, including gonadal maturation, dietary intake, stomach fill level, and growth stage [81]. Trophic interactions are crucial in aquatic environments and dictate the flow of energy and nutrients within an ecosystem [87]. Understanding the feeding ecology of fish offers valuable insights into population dynamics, resource partitioning, habitat preferences, and energy transfer, all of which have implications for fisheries management [87].
This study confirms that Rasbora sp. and Barbodes sp. exhibit significant dietary overlap, leading to competition for food resources, with Pianka’s overlap index value of 0.96. Predation by Amphilophus trimaculatus further threatens native fish populations. The growth pattern of native fish in Aneuk Laot Lake follows a negative allometric trend (b < 3), with a b-value of 2.77, indicating environmental pressure. These findings provide critical insights for ecological management strategies aimed at mitigating the impact of invasive species and preserving biodiversity in Aneuk Laot Lake. The study serves as baseline data for lake management, emphasizing invasive fish population control, native fish habitat conservation, and the implementation of fisheries regulations to maintain ecosystem balance and optimize fish population growth.
The authors would like to thank Universitas Syiah Kuala which has funded this research on the Lector Research Scheme in 2024 with contract No.: 353/UN11.2.1/PG.01.03/SPK/PTNBH/2024 on May 3rd, 2024.
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