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Article

Trait Composition and Assemblage Structure Analyses of Lacustrine Fishes: Synthesizing a Proposal for Better Fishing Practices

by
Gustavo Díaz
1,2,*,
Evelyn Habit
1,
Roberto Urrutia
1,2,
Aliro Manosalva
1,
Ricardo O. Barra
1,2 and
Ricardo Figueroa
1,2
1
Departamento de Sistemas Acuáticos, Facultad de Ciencias Ambientales y Centro EULA-Chile, Universidad de Concepción, Concepción 4070386, Chile
2
Centro de Recursos Hídricos para la Agricultura y la Minería, CRHIAM (ANID/FONDAP/1523A0001), Concepción 4030000, Chile
*
Author to whom correspondence should be addressed.
Water 2024, 16(16), 2333; https://doi.org/10.3390/w16162333
Submission received: 26 June 2024 / Revised: 22 July 2024 / Accepted: 24 July 2024 / Published: 20 August 2024
(This article belongs to the Section Biodiversity and Functionality of Aquatic Ecosystems)
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Abstract

:
Fish provide ecosystem services and contribute to human well-being through fishing. In Chile, subsistence fishing provides food to local communities, whereas recreational fishing contributes to economic income via tourism. In the Lanalhue coastal lake basin, unregulated subsistence fishing and formal recreational fishing primarily target large fish species to ensure satisfactory catches and food provision. However, the development of unplanned fishing activities and lack of scientific knowledge can lead to socioecological conflicts and tension between users. To address this issue, this study focuses on improving fishing practices through the analysis of fish assemblages of the Lanalhue lake basin. The life cycle traits of each fish species were analyzed, as well as their abundance and biomass in each habitat. Twelve fish species were identified, and their distribution and abundance explain the differences in fish assemblage composition and structure among habitats. To promote better fishing practices, we propose four target species and specific management actions. These include defining fishing seasons, delineating fishing sites, and establishing fishing quotas. This approach, considering both community structure and functional aspects of fish assemblages, can serve as a valuable guide for developing sustainable fishing practices in the Lanalhue coastal lake basin.

1. Introduction

Benefits obtained by the human population from the functioning of nature are defined as ecosystem services [1], which are the result of interactions between ecological, geophysical, and social components that support human well-being [2]. Freshwater ecosystems offer various goods and services, such as climate regulation and recreation, but food provisioning through fishing is probably the most important [3,4,5]. Fish are the main animal protein source worldwide [3,6], with large fisheries developed in rivers and lakes. Thus, the economic importance of some fish species negatively affects the conservation of fish diversity and the supply of services. The economic interest in target fish species leads to overexploitation of fish, resulting in unsustainable fishing and social conflicts among users of aquatic ecosystems [7,8]. In addition, in aquatic ecosystems that lack economically important fish, there is generally less knowledge of and interest in fish fauna [9].
In Chile, commercial fisheries have not been developed in freshwater ecosystems. Recreational fishing is regulated, with an emphasis on fly fishing, with the catch and release of fish of all sizes and a focus on salmonid species because they are good fighters and reach large body sizes [10]. Unlike recreational fishing, subsistence fishing is unregulated, practiced by local people, and focuses on both non-native and native species that provide an acceptable biomass for the catch effort (e.g., trout, carp, silversides, perch, puye) [11]. Moreover, indigenous people practice unregulated fishing for subsistence and as a traditional activity focused on large-bodied species (e.g., trout, carp, perch), disregarding knowledge about the biology, ecology, and conservation of the species [12,13]. Thus, fishing is highly focused on introduced fish (mainly salmonids; [10]), and native Chilean fish are often undesirable due to their generally small body sizes (<15 cm as adults; [14,15]). This leads to the protection of introduced salmonid species by the agency that manages freshwater resources in Chile (SERNAPESCA), with disregard for the adverse effects on native species (e.g., [16,17]). Furthermore, there is a dearth of biological and ecological information concerning native species across various aquatic ecosystems over time, and social aspects related to fishermen remain insufficiently studied. This situation highlights a critical issue: generalist fishing policies often overlook crucial factors. These policies fail to account for the local social and economic contexts of users, as well as the ecological conditions affecting local fish populations (such as abundance and conservation status). Consequently, the current management of inland water fisheries in Chile falls short of adhering strictly to existing inland fishing standards (as exemplified by Cooke et al. 2016; 2021 [18,19]).
The Chilean lakes situated in the coastal Nahuelbuta Range (between 36°51′ S, 73°05′ W and 38°09′ S, 73°19′ W) share similar environmental characteristics and are collectively referred to as the “Nahuelbutan Lakes” [20]. Among these lakes, the Lanalhue Lake basin (including the lake itself and its tributaries) has been significantly affected by the rapid expansion of forestry plantations and urban development driven by the timber industry [21,22,23].
To revitalize the local economy, regional government agencies have promoted Lanalhue Lake as a recreational fishing destination. As a result, tourism has surged, leading to a population increase relative to the number of year-round residents [21,24,25]. However, this shift has also sparked socioecological conflicts. Local residents rely on the lake for water and sustenance, while the indigenous people (Mapuche) attribute spiritual significance to it [26,27].
Resolving these conflicts is challenging due to the lack of fundamental information about the resources that support traditional and subsistence activities. This knowledge gap hinders sustainable territorial planning that is necessary to safeguard the ecosystem services provided by the lake basin. Nevertheless, solutions to these socioecological conflicts arising from fishing activities in Lanalhue Lake are feasible through the application of scientific insights about fish species in lacustrine habitats. This knowledge can pave the way for sustainable fishing development [28,29]. Unfortunately, the fish fauna of Lanalhue Lake remains poorly studied [20].
This study aims to determine a set of target species for fishing and analyze the spatial distribution of species biomass and the environmental factors influencing their presence to develop planned fishing activities that contribute to resolve socioecological conflicts arising from regulated and unregulated fishing practices in the Lanalhue coastal lake basin. In a context of limited data availability, this study represents the first attempt in Chile to propose planned fishing activities for both tourism and local communities. As such, this approach integrates various factors and can potentially be applied in other data-limited contexts.

2. Materials and Methods

2.1. Study Area

The study was conducted in the Lanalhue coastal lake basin, which includes Lanalhue Lake and its tributaries. It is located in the Nahuelbuta Range, part of the Coastal Range, south-central Chile (37°50′ S, 73°22′ W) (Figure 1). The lake is located 12 m.a.s.l. and has a total area of 31.9 km2, with an elongated shape with high sinuosity. The maximum depth is 25 m (southern area), and the minimum is 6 m (northern area). Neither zone presents considerable water level fluctuations among seasons.
The southern part of the Lanalhue basin is the site of the main town, Contulmo (6031 inhabitants), while tourism development is concentrated in the northern area. Mapuche (indigenous people) settlements are present all around the lake [25,30]. Lanalhue Lake provides recreational services such as water sports, recreational fishing, and birdwatching, which increase during the summer [20,31]. Tributary rivers and the lake provide opportunities for recreational and subsistence fishing but are affected by significant human stressors [30].
So far, only three fish native species have been recorded: Creole perch (Percichthys trucha), pocha del sur (Cheirodon kiliani), and puye (Galaxias maculatus). Two non-native species have also been observed: common carp (Cyprinus carpio) and Argentinean silverside (Odontesthes bonariensis) [20,32,33,34]. The fish fauna of the tributaries has not yet been described.

2.2. Field Surveys

A total of 20 sites were sampled during the austral summer in two consecutive years (2019–2020), with a selective sampling design employed. Fish surveys in the summer season offer a high probability of capturing the maximum diversity because Chilean coastal aquatic systems exhibit their minimal annual water flow. The best locations of sampling sites and appropriate fishing gear were chosen to increase the efficiency of fish catches and record maximum diversity based on environmental conditions in each habitat type.
Ten sites were located in tributaries and 10 in the lake (Figure 1). One sampling site was located in each tributary river, near their confluence with the lake (Figure 1), where coastal rivers exhibit the maximum diversity and abundance of fish [34]. The tributaries were all wadable and were characterized by riffle habitats with variable mean water velocities between 0.14 and 0.46 m/s and gravel and cobble substrates. Tributaries were sampled using a Halltech HT-2000 (Halltech, ON, Canada) electrofishing backpack, with a single pass covering 30 to 50 m in an upstream direction over 30 min. The lake was sampled in both littoral (5 sites) and pelagic (5 sites) habitats, with records obtained from different beaches along the perimeter of the lake, as well as limnetic zones with different depths and geomorphological features (Figure 1). Beach seines (5 m long, 1.5 m high, and 10 mm mesh size) were used to sample shallow littoral habitats (<1 m depth) characterized by gravel, cobbles, and grass patches. The extended net was swept parallel to the shore twice, covering a total of 40 m2. Based on the bathymetry of the lake, pelagic habitats were sampled at different depths (3 to 25 m) using multi-mesh gillnets (20 m long, 1.5 m high; 10, 15, 20, 30, 50, 60, 70, and 120 mm bar mesh size). Midwater gillnets were set for 12 h (set at sunset and withdrawn at dawn). The collected fish were sedated with the anesthetic Benzocaine 20% (BZ-20), identified in situ at the species level according to the available identification keys [14,35,36], counted, measured, weighed, and returned to their original habitats.
At each sampling site, a set of environmental variables was recorded from a single sample in the middle of the fish-sampled area in littoral and riverine habitats and a midwater sample in pelagic habitats. Water quality variables were measured based on standard methods (APHA-AWWA-WPCF, 1995). Temperature, pH, conductivity, total dissolved solids, dissolved oxygen and oxygen saturation, and turbidity were measured in situ (Hanna, HI-9828 and HI-98703, Hanna Instruments, Woonsocket, RI, USA). Total N and total P were analyzed by the certified laboratory of the EULA-Chile Center (Universidad de Concepción, Chile). Depth was measured as the mean depth in tributaries and littoral sampling sites based on three measurements along the fish-sampled area, and as total depth in pelagic habitats. Finally, from the pelagic sampling point to its nearest coastline, the slope and distance were recorded to calculate a rate that reflects geographic variability in pelagic habitats.

2.3. Statistical Analyses

A three-step set of statistical analyses was used to formulate a proposal for better fishing practices and move toward sustainable fishing activities in this poorly studied basin: (1) characterization of the fish assemblage composition and structure, (2) target species selection, and (3) a fishing proposal based on the biomass distribution, habitat characterization of the target species, and fishing regulations (Figure 2). Following the conceptual model and arguments of Fenichel et al. for fishing science, our three-step method was used to generate a data framework to apply filters to build a regulatory instrument for suitable policies and recommendations [37].
Characterization of the fish assemblage composition and structure: The aim is to ascertain differences among fish assemblages in different habitats in terms of species composition and abundance. To explore fish assemblage composition dissimilarities among habitats (tributaries, littoral areas, and pelagic areas), a non-metric multidimensional scaling analysis was conducted using the entire presence–absence dataset (all species) [38] (Figure 2). The correspondence of assemblage composition to habitats and statistical significance were tested using a canonical analysis of principal coordinates (CAP; [39,40]). This analysis is based on Sorensen dissimilarity matrices for fish presence–absence data and provides correspondence values that commonly indicate how unique a fish community composition is to each habitat (Figure 2). The contribution of fish species to the assemblage structure of each of the different habitats was (separately) calculated through a similarity percentage analysis (SIMPER), using a Bray–Curtis dissimilarity matrix based on the square-root-transformed fish capture per unit of effort (CPUE) (Figure 2).
Target species selection: For the second step, a set of target species was determined based on the combined analysis of proxies (functional life history traits and sustainability indicators), assemblage structure, Chilean conservation status, and records of prior catches (desirability). To do so, a set of proxies was selected to elucidate a theoretical set of ideal species (target species) for subsistence and recreational fishing based on the exploration of proxy variability and affinity with fish species (Figure 2). Functional life history traits were determined from the literature (Supplementary Materials, S.M.1) [41,42,43,44,45,46,47,48,49,50,51,52], and sustainability indicators were calculated from fish length–weight data (Table 1). Next, proxies that lacked accurate data (intervals or ranges) were transformed into categorical variables (Table 1 and Supplementary Materials, Table S1), and the relationships among them were evaluated for subsequent statistical analyses, which resulted in the deletion of the maximum weight observed. Next, a principal component analysis (PCA) was performed to determine proxy variation and select theoretical target fish species. A priori, the ideal set of target species was expected to meet the following criteria: high meat amount per catch (large body size and weight), persistence over time (high fecundity and adult percentage), no conservation concerns (no endangered native species), and satisfaction of local users’ fishing demands. Consequently, native fish classified as endangered according to the species conservation classification of the Chilean Environmental Ministry were excluded from this analysis, since better fishing practices imply conservation [53]. The use of this fish species classification was useful for this purpose because it is focused on a local scale. In addition, to support these findings and determine a better set of target species for recreational and subsistence fishing, we contrasted the results with prior catch records of those fish species (e.g., desirability). The species conservation classification and prior catch records for each species are presented in Table 2 (see results).
The chosen proxies must reflect important issues for fisheries management and theoretically will contribute to better fishing practices according to the following arguments: fish length proxies such as maximum total length, mean total length, and maximum weight indicate a high meat provision per subsistence fishing catch [3,54]. Specifically, species mean total length (in the catch) is also correlated with fishing mortality and suggests the exploitation level [55]. Furthermore, these proxies are useful for identifying large fish, which reflect angler preferences and catch satisfaction in recreational fishing [56,57]. Fecundity indicators such as number of eggs per spawn event and the percentage of mature fish in catches indicate the probability of species successfully maintaining their populations, ideally in suitable habitats [29,58,59]. In addition, Froese proposes this ratio (percentage of mature fish) as an indicator of overfishing because it serves as an estimate of the size of fish selected for fishing [60,61]. Meanwhile, regarding the available information and in situ observations, we propose shoaling behavior and habitat use as potential proxies of catch feasibility because desirable common native species in littoral habitats often exhibit shoaling and open-water swimming behavior [14,62,63]. Therefore, a high total biomass catch is possible per single effort, facilitating subsistence fishing.
Fishing proposal based on the spatial distribution of target species biomass and habitat conditions: Finally, biomass boxplots based on individual weights (excluding extreme outliers) of the target species that presented more than two individuals at a minimum of two sites per habitat were used to explore the differences among sampling sites in each habitat type. As appropriate, analysis of variance (ANOVA; parametric data) and Kruskal–Wallis (non-parametric data) analysis were used to test significant differences in species biomass among habitats. Furthermore, Tukey and Dunn tests were used to determine pairwise differences and sites that determined significant differences in species biomass. The use of these statistical tools allows better sites for catching the target species based on biomass to be identified. In addition, general linear models (GLMs) were used to determine the environmental variables that explain the significant differences in the fish biomass of the target species for preservation of the habitat conditions at these sites in order to safeguard fish resources (biomass) (Figure 2). Prior to the use of GLMs, highly correlated environmental variables were removed (R > 0.80) [64]. Consequently, total dissolved solids was not included for any habitat. Additionally, pH was not included for littoral habitats, and oxygen saturation and total N were not considered for pelagic habitats. Thus, GLMs were employed assuming a Gaussian distribution of the raw biomass data of each target species, as response variables to the debugged set of environmental variables. Finally, to propose fishing restrictions related to fishing seasons and quotas, we analyzed the current regulations for each target species and suggest modifications according to the ecological status of the target species observed in this study (see Cooke and Cowx, 2006 [65]).
Spatial statistical differences in target species biomass and their relationships with environmental variables, as well as, additional statistical analyses were performed in R software (v.4.0.2) [66]. Specifically, N-MDS, SIMPER, and PCA analyses were performed with the vegan package (v.2.6-6.1), while CAP was executed using the BiodiversityR package (v.2.16-1) [38,67].

3. Results

A total of 1918 individuals were caught, 594 in tributary rivers and 1324 in the lake, in littoral and pelagic habitats. Individuals corresponded to 12 fish species, nine of which were recorded for the first time. Of the 12 fish species, 10 were native and two were introduced (Table 2). There are records of fishing activity for six of the native fish species, as they are classified as desirable species for fishing (Table 2). Cheirodon galusdae was the only native species found in all three habitats. Among the introduced species, Oncorhynchus mykiss was found in all habitats, and C. carpio was found only in pelagic habitats (Table 2). Fish assemblage compositions corresponded to their respective habitats (100% in each case; p= 0.001), with a high dissimilarity among them (Figure 3). Assemblages in tributaries were characterized by an abundance of rainbow trout (Oncorhynchus mykiss) and native catfish (Trichomycterus areolatus) (Table 3). In the lacustrine habitats, the littoral zone was characterized by the native species Galaxias maculatus and Basilichthys microlepidotus, and the pelagic zone was characterized by Creole perch (P. trucha) and common carp (C. carpio) (Table 3).
Among the life history traits, the percentage of mature fish and total mean length explained a large proportion of the variance expressed in PC1, which accounted for 71.9% of the total variance. Meanwhile, maximum total length was the life history trait with the greatest proportion of variance in PC2 (21.9%) (Figure 4). The species with the highest proportion of variability associated with these proxies are P. trucha, O. mikyss, and C. carpio, which are proposed as target species for fishing activities (Figure 4). In contrast, seven species did not present relationships with these proxies, among them those with shoaling behavior (C. galusdae, and G. maculatus). Nonetheless, despite having little association with the proxies related to large sizes and a high proportion of mature fish, G. maculatus should be included in the target species set because juvenile individuals are desirable for subsistence fishing (Figure 4). Thus, the target group consists of O. mikyss for tributaries and G. maculatus for littoral habitats, as well as P. trucha and C. carpio for pelagic habitats. This species set is composed of the desirable species for fishing and presents the records of previous catches for subsistence and recreational fishing.
Of the target species, Galaxias maculatus was abundant only in littoral habitats (Table 3 and Table 4), with significant biomass differences among sampling sites (Figure 5a). This species presented the highest average biomass values at sampling sites L3 and L5, at which the largest average body total length (4.8 cm) and highest abundance (n = 131) were observed (Table 4). Percichthys trucha, which was present only in pelagic habitats, presented significant biomass differences among the five sampling sites determined by site P3 (Figure 5b; Supplementary Materials, Table S2). The highest average biomass was found at the central sampling site, P3, with an average total length of 30.8 cm (n = 11) and a maximum of 33.3 cm (Table 4). The non-native species C. carpio also occurred only in pelagic habitats, but there were no statistical differences among sampling sites (Figure 5c). Although O. mykiss was present in the three habitats, it was notably abundant in tributaries (Table 3 and Table 4), with significant biomass differences explained mainly by sites R1, R5, and R9 (Figure 5d; Supplementary Materials, Table S2). Sampling sites R1, R2, R6, R7, and R8 presented the highest biomass, as a consequence of both the high abundance and large size of individuals (Table 4).
GLM analyses revealed significant relationships only for O. mykiss and G. maculatus. Indeed, the pelagic species, common carp and creole perch, did not present any significant relationships with the measured environmental variables. This suggests that water quality may not be a significant factor for planning fishing areas of pelagic target species. The biomass of O. mykiss presented significant relationships with depth, temperature, dissolved oxygen, oxygen saturation, turbidity, total phosphorous, total nitrogen, and pH (Table 5). Therefore, as expected, the distribution and biomass of this species depends on water quality variables. Meanwhile, G. maculatus biomass in littoral habitats presented significant relationships with depth, dissolved oxygen, oxygen saturation, and turbidity (Table 5), demonstrating the influence of oxygen availability and littoral habitat depth on this species.
In accordance with the specific results for each species, the detailed fishing proposal (Supplementary Materials, S.M.2) includes the following guidelines: G. maculatus could be caught in the central littoral zone (L2, L3, and L4) of Lanalhue Lake due to its high abundance and desirable size for subsistence fishing from August to November (Figure 6 and Figure 7). P. trucha exhibits good size and high biomass per individual in the central area (P2, P3 and P4; Figure 7) of the lake, and we recommend catches in this zone, with a maximum extraction of three fish or 15 kg per fisherman (over 25 cm in length) (for more details, see Supplementary Materials, S.M.2). Because C. carpio and O. mykiss are introduced species and due to their adverse effects on native species, we recommend their removal in any location and season (Figure 6 and Figure 7) (for more details, see Supplementary Materials, S.M.2) [48,68,69,70].
In addition, previous information indicates a high feasibility of capture and amount of biomass (one or a group of individuals) per fishing effort (hooks or nets), which makes them suitable for subsistence fishing in all habitats. However, only P. trucha in limnetic habitats and O. mykiss in limnetic and riverine habitats present a high biomass per individual and are recognized as good fighters, making them a priority for recreational fishing (Figure 8).

4. Discussion

In this study, we analyzed fish assemblages across diverse habitats, including tributaries, littoral zones, and pelagic areas. Our primary objectives were twofold: first, to identify a set of target species for fishing based on life history traits and sustainability indicators, and second, to analyze the spatial distribution of species biomass and the environmental factors influencing their presence. By doing so, we aimed to pinpoint optimal locations for fishing activities.
Additionally, we crafted a comprehensive proposal that aligns with scientific regulations, emphasizing responsible fishing practices. This proposal encompasses several key considerations:
  • Target species selection: We recommend specific species whose biology and ecology could support different fishing activities in the study area.
  • Fishing seasons: Timing matters! We propose well-defined fishing seasons to ensure minimal impact on fish populations during critical life stages.
  • Best fishing areas: Based on our analysis, we highlight areas with abundant fish biomass and ecological significance.
  • Fishing quotas: Our proposal suggests appropriate catch restrictions to prevent overexploitation.
The outcomes of our study significantly contribute to the understanding of freshwater fish ecology and effective fishing management in Chile. By promoting sustainable resource use, we address socioecological conflicts while minimizing the need for extensive data collection. We encourage the adoption of our approach in lesser-known ecosystems, to characterize fish as a resource, to understand the local drivers that regulate fisheries, and to promote responsible fishing practices [71].

4.1. Characterization of Fish Assemblages

We documented the presence of nine fish species previously unrecorded in the Lanalhue Lake basin. This discovery underscores the limited exploration of fish biodiversity in coastal basins, as highlighted by previous research [20]. Among these newly recorded species, the presence of the migratory lamprey (Geotria australis) and the endangered species peladilla (Aplochiton zebra) stand out. Their occurrence in small tributaries reveals the significance of these highly connected systems for spawning and nursery activities. This update represents a significant advance in the ecology of fish species inhabiting the Nahuelbutan lakes. Moreover, it contributes to the understanding and appreciation of these often-overlooked aquatic ecosystems, contributing to informed fishing management decisions. Further insights emerged from our examination of different habitats:
(1)
Riverine (tributaries) habitats: Streams dominated by native catfish (Trichomycterus areolatus) and rainbow trout (Oncorhynchus mykiss), as occurs in Andean and other coastal rivers [34,72,73]. Notably, the abundance and biomass of rainbow trout in tributaries suggest its competitive success over native fish in terms of food and shelter [74] (see Supplementary Materials, Table S3).
(2)
Lacustrine (littoral and pelagic) habitats: While rainbow trout presented a lower abundance, they were represented by larger-bodied sizes (Supplementary Materials, Table S3). This coincides with findings from other Chilean coastal lakes but diverges from Andean oligotrophic lakes [75,76]. Mesotrophic coastal lakes are predominantly dominated by the native Creole perch (Percichthys trucha) and common carp (Cyprinus carpio), consistent with our results [20]. Interestingly, only large adult specimens of Creole perch were captured in pelagic habitats. This suggests that the species is as equally abundant and successful as in southern Andean lakes, as documented by Macchi et al. and Ortiz et al. [77,78]. Furthermore, the absence of P. trucha in rivers likely reflects their habitat preferences; juvenile P. trucha inhabit shallow habitats in larger rivers during summer [44,48]. Consequently, while they may be present in the outlet river of Lanalhue Lake (Paicaví River), their absence in small tributaries is consistent with their habitat choices.
The results showed variation in fish composition across different habitats, influenced by environmental filtering and by fishing gear selection [79]. Despite the potential biases introduced by gear selectivity, our result coincides with the known species in coastal Chilean rivers and Nahuelbutan lakes [20,34,76,80].

4.2. Target Species for Subsistence and Recreational Fishing

The combined analysis of fish assemblage structure and species proxies (traits and indicators), together with the evaluation of the qualitative characterization of species (conservation status and desirability), resulted in a useful tool to propose a group of target species for fishing activities. This simple approach, based on limited and non-complex data, including knowledge about the current fishing practices and species preferences of fishermen, gives support for a planned scientific proposal. Therefore, this proposal contributes to strengthening the weak connection between scientific information and public stakeholders to enhance fishing policies [81]. In addition, this approach is an advance because it does not only account for individual fish meat quantity and quality as other studies have previously [57,82,83]. For example, Chilean cauque (O. mauleanum) can reach a large body size and have tasty meat and thus could be considered a priori for subsistence fishing (Supplementary Materials, Table S3). However, our results suggested that they should be dismissed as a target species. This species did not show a large body size in this lake, and probably hosts harmful parasites as in Andean lakes [84,85]. Additionally, although the target species were primarily of large body size and with no conservation concerns, G. maculatus (with opposite attributes) also was included as a target species. Indigenous and local people capture whitebait and juveniles of G. maculatus for sale and food [63,86]. Such catches for subsistence fishing cannot be disregarded if progress is to be made in reducing conflicts among users, and they confirm the need to consider the differing motivations among fishermen to generate better fishing practices [56]. While G. maculatus catches could trigger overexploitation, the effects of subsistence fishing by small populations are completely unknown. Moreover, it is preferable to exploit this resource for traditional activities rather than to promote the feeding of wild trout populations, as suggested by the Ministry of the Economy (1981) [63].
Unlike Galaxias maculatus, P. trucha, O. mykiss, and C. carpio were not only abundant species but also provide a large quantity of meat per individual and present a very high proportion of adults in the Lanalhue lake basin. These parameters are expected to be stable throughout the year, and there are only slight decreases in the number of juvenile G. maculatus during the first month of our fishing calendar (Figure 6), as well as of adult O. mykiss in the winter season. Therefore, the data appear to be reliable for our purpose. Percichthys trucha is considered a game species in Chile, but there are no specific population studies that support this view; therefore, its extraction does not seem to follow sustainability principles [81]. Cyprinus carpio and O. mykiss were introduced in Chile for food and sport fishing [87]. These species are widespread, and their capture would support population control, decreasing their multiple negative impacts on native fish [10,17]. For example, large individuals of O. mykiss prey mainly on native species like G. maculatus and C. galusdae [17,88]. Due to their habits and behavior, carp disperse sediment into the water column, destroying the nesting sites of native fish and occasionally gleaning eggs [14]. Thus, according to users’ preferences, the extraction of these species should be promoted as a way to provide food for local people.

4.3. Environmental Conditions That Explain Species Biomass

Based on our results, water quality was particularly relevant to O. mykiss and G. maculatus biomass in Lanalhue Lake and its tributaries. As expected, the higher biomass of rainbow trout in tributaries presented significant relationships with nutrients, temperature, and oxygen because it is a good bioindicator of good water quality [89,90]. For G. maculatus, turbidity and oxygen—variables that affect its feeding and metabolism—presented a relationship with biomass. Turbidity affects the size and biomass of planktivorous species because they are visual predators [91,92]. In addition, G. maculatus juveniles present high breathing rates, which may explain the negative relationships between biomass and dissolved oxygen and oxygen saturation [93,94]. The absence of significant relationships between the measured environmental variables and both P. trucha and C. carpio biomass could suggest homogenous characteristics of the pelagic zone of the lake (Supplementary Materials, Table S4). However, Percichthys trucha is a species that requires good visibility to move and catch its prey, meaning that high turbidity does not favor high abundance [44,95]. By contrast, the systems with high abundances of C. carpio are often characterized by high turbidity, as it feeds by sucking sediment, which increases the amount of suspended matter [96]. The results suggest that the maintenance of or even reduction in the nutrient load in tributaries and lakes is important for supporting fish abundance and biomass, and consequently, fishing over time [54,97,98]. In addition, we encourage additional management measures to ensure the sustainability of recreational fishing in tributaries, as it is the most threatened in these settings due to anthropogenic pressures and the strict dependence on O. mykiss.

5. Conclusions

Lanalhue Lake lacks management measures specific to its watershed and hydrobiological species. Proposing such measures is essential, especially considering that regional government agencies have assigned the lake as a recreational fishing site [24]. To ensure better practices, regulations, and restrictions for the Lanalhue Lake basin, they should align with resource management, economic growth, and the preservation of traditional activities [65].
To this end, we craft a fishing proposal, based on limited and non-complex fish data, including knowledge about the current fishing practices and species preferences of fishermen. The result is a planned scientific proposal that recommends several key actions: (a) Revising recreational fishing laws: Government agencies should consider amending the Recreational Fishing Law (Law 20.256, Ministry of the Economy) to encourage responsible recreational and subsistence fishing. For instance, increasing the number of trout caught can help maintain smaller fish populations; (b) Promoting carp as a subsistence fishing target: Indigenous communities in the Lanalhue Lake basin can benefit from subsistence fishing. Promoting the catching of carp, a non-native species, could provide sustenance while also managing invasive species in Chilean aquatic ecosystems (Supplementary Materials, S.M.2); (c) Encouraging recreational fishing in riverine habitats: Anglers should focus on non-native rainbow trout in riverine habitats (tributary rivers). These trout are abundant, provide a good fight, and grow to substantial sizes; (d) Emphasizing native creole perch in pelagic habitats (lake): Native creole perch, found in pelagic habitats (the lake), is another excellent option. They offer abundant catches, are strong fighters, and contribute to the overall biomass (Figure 6); (e) Balancing feasibility and biomass: When developing subsistence fishing practices, native and non-native fish species that yield a high meat-to-effort ratio (feasibility and/or biomass) must be prioritized. These species must not be classified as endangered or vulnerable (Figure 6).
By implementing these strategies, we can strike a balance between conservation, economic benefits, and cultural practices, ensuring a sustainable future for the Lanalhue Lake basin.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/w16162333/s1, Table S1: Proxy values for no-conservation-concern fish species, Table S2: Significant pairwise test of target species that showed differences in biomass among sites, Table S3: Minimum, maximum, and mean length (cm) and weight (g) for each fish species captured in this study, Table S4: Environmental variables of tributaries, littoral, and pelagic habitats., S.M.1: Literature used to determine life history traits for each fish species, S.M.2: Detailed fishing planning proposal for each target species.

Author Contributions

Conceptualization, G.D., E.H., R.O.B. and R.F.; methodology, G.D. and A.M.; software, G.D. and A.M.; validation, G.D., E.H., R.U., R.O.B. and R.F.; formal analysis, G.D., E.H. and A.M.; investigation, G.D., E.H. and R.F.; resources, E.H. and R.U.; data curation, G.D. and A.M.; writing—original draft preparation, G.D., E.H., R.U., A.M., R.O.B. and R.F.; writing—review and editing, G.D., E.H., R.U., A.M., R.O.B. and R.F.; visualization, R.O.B. and R.F.; supervision, G.D., E.H. and R.F.; project administration, E.H. and R.U.; funding acquisition, E.H. and R.U. All authors have read and agreed to the published version of the manuscript.

Funding

This study was funded by the Programa de Recuperación de los Servicios Ambientales de los Ecosistemas Lacustres de la Provincia de Arauco (PRELA) of the Chilean Ministry of Environment, which provided resources to R. Urrutia. G. Díaz, R. Urrutia, R. O. Barra and R. Figueroa are grateful for the financial support of ANID/FONDAP/1523A0001. Also, the authors thank to FONDECYT project 1190647, awarded to E. Habit.

Data Availability Statement

Data available on request due to restrictions.

Acknowledgments

The authors thank the Programa de Recuperación de los Servicios Ambientales de los Ecosistemas Lacustres de la Provincia de Arauco (PRELA). Also, thank the Centro de Recursos Hídricos para la Agricultura y la Minería CRHIAM ANID/FONDAP/1523A0001. This research was partially funded by FONDECYT project 1190647, awarded to E.H. The authors also wish to express their gratitude to Alejandra Zurita for mapping support and to Waldo San Martin, Alessandra Perfetti, Laura Habit, Joaquín Cárcamo, and Diego Moraga for their help during the fieldwork.

Conflicts of Interest

The authors declare no conflicts of interest. In addition, the funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Figure 1. Lanalhue Lake basin and fish sampling sites in riverine (tributaries) and lacustrine habitats (littoral and pelagic).
Figure 1. Lanalhue Lake basin and fish sampling sites in riverine (tributaries) and lacustrine habitats (littoral and pelagic).
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Figure 2. Schematic representation of statistical methods. Analyses were grouped (dotted line) following the connected specific aims (bottom boxes) proposed in this study.
Figure 2. Schematic representation of statistical methods. Analyses were grouped (dotted line) following the connected specific aims (bottom boxes) proposed in this study.
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Figure 3. Ordination plot based on the fish composition in the three different habitats for two consecutive sampling years.
Figure 3. Ordination plot based on the fish composition in the three different habitats for two consecutive sampling years.
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Figure 4. Principal component analysis based on proxies (life history traits and sustainability indicators) of the non-endangered native fish species present in the study area. Proxies correspond to the red-colored text. The green color indicates target non-native species, while the blue color points out target native species. See Table 2 for fish name abbreviations.
Figure 4. Principal component analysis based on proxies (life history traits and sustainability indicators) of the non-endangered native fish species present in the study area. Proxies correspond to the red-colored text. The green color indicates target non-native species, while the blue color points out target native species. See Table 2 for fish name abbreviations.
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Figure 5. Target species biomass and p-values based on the significant differences among sites. Native species are shown in blue (a,b) and non-native in red (c,d). Extreme outliers of O. mykiss (>80 g (n = 4)) were not included. p-values at the top right show the differences of fish species biomass among sites. Circles up to the boxplots represent outliers. Asterisks show sites that determined significant differences in biomass.
Figure 5. Target species biomass and p-values based on the significant differences among sites. Native species are shown in blue (a,b) and non-native in red (c,d). Extreme outliers of O. mykiss (>80 g (n = 4)) were not included. p-values at the top right show the differences of fish species biomass among sites. Circles up to the boxplots represent outliers. Asterisks show sites that determined significant differences in biomass.
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Figure 6. Proposed fishing sites for native (a,b) and non-native (c,d) target fish species of the Lanalhue basin. Light green circles show the mentioned sites in the proposal for each species, and large dark green circles show the optimal sites for fishing. For more details, see Supplementary Materials, S.M.2.
Figure 6. Proposed fishing sites for native (a,b) and non-native (c,d) target fish species of the Lanalhue basin. Light green circles show the mentioned sites in the proposal for each species, and large dark green circles show the optimal sites for fishing. For more details, see Supplementary Materials, S.M.2.
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Figure 7. Annual fishing calendar proposed for target fish species of the Lanalhue basin and their restrictions (fishing quotas). Green squares show fishing season months, and red squares show closed months.
Figure 7. Annual fishing calendar proposed for target fish species of the Lanalhue basin and their restrictions (fishing quotas). Green squares show fishing season months, and red squares show closed months.
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Figure 8. Decision tree scheme for sustainable fishing activities. Diagram shows target species and suitable habitats according to fishing options for users. Different types of dashed lines show target species groups associated with fishing indicators.
Figure 8. Decision tree scheme for sustainable fishing activities. Diagram shows target species and suitable habitats according to fishing options for users. Different types of dashed lines show target species groups associated with fishing indicators.
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Table 1. List of proxies used to determine ideally the set of suitable species for subsistence and recreational fishing. Proxies are fish life history traits and sustainability indicators. Proxy values documented as ranges were transformed to numbers (numerical categories) to execute statistical analyses.
Table 1. List of proxies used to determine ideally the set of suitable species for subsistence and recreational fishing. Proxies are fish life history traits and sustainability indicators. Proxy values documented as ranges were transformed to numbers (numerical categories) to execute statistical analyses.
Numeral
Categories		Type of VariableArgument or Biological Feature
Life history traitsMaximum total length recordNumerical Maximum length of each fish species recorded in the literature
Maximum weight observedNumerical Maximum weight of each fish species registered during fieldwork
FecundityCategorical1 (Low)Less than 100 eggs for each fish per spawn event
2 (Medium)Between 100 and 999 eggs for each fish per spawn event
3 (High)Between 1000 and 9999 eggs for each fish per spawn event
4 (Very high)More than 100,000 eggs for each fish per spawn event
Habitat useCategorical1 (Benthic)Associated with substrate (boulders, pebbles, gravel, or sand).
2 (Benthic–pelagic)Associated both with substrate and water column
3 (Pelagic)Uses only the water column
Shoaling behaviorCategorical0 (No shoaling behavior)No shoaling behavior
1 (Exhibit shoaling behavior)Fish form a part of shoals
Sustainability indicatorsMean total length observedNumerical Mean total length of each fish species registered during fieldwork
Percentage of mature fish (ratio)Numerical Proportion of adults for each fish species
Table 2. List of native and non-native fish species captured in riverine and lacustrine habitats and their conservation status. “x” indicates presence by habitat.
Table 2. List of native and non-native fish species captured in riverine and lacustrine habitats and their conservation status. “x” indicates presence by habitat.
SpeciesRiverine LacustrineConservation StatusRecords of Prior Catches
Family Scientific name (abbreviation)TributariesLittoralPelagic
Native
GeotriidaeGeotria australis (Ga)x VulnerableSubsistence fishing
CharacidaeCheirodon galusdae (Cg)xxxVulnerableNo
TrichomycteridaeTrichomycterus areolatus (Ta)x VulnerableNo
GalaxiidaeGalaxias maculatus (Gm)xx Less concernSubsistence fishing
Brachygalaxias bullocki (Bb)x VulnerableNo
Aplochiton zebra (Az)x EndangeredSubsistence and recreational fishing
AtheriniidaeBasilichthys microlepidotus (Bm) x Near-threatened Subsistence and recreational fishing
Odontesthes mauleanum (Oma) xxVulnerableSubsistence and recreational fishing
PercychthidaePercichthys trucha (Pt) xLess concernSubsistence and recreational fishing
PerciliidaePercilia gillissi (Pg)xx EndangeredNo
Non-native
SalmonidaeOncorhynchus mykiss (Omy)xxx-Subsistence and recreational fishing
CyprinidaeCyprinus carpio (Cc) x-Subsistence and recreational fishing
Table 3. Values of fish species average abundance (CPUE) and contribution to similarity within habitats. Results show species contribution at a 90 cut-off from the SIMPER analysis.
Table 3. Values of fish species average abundance (CPUE) and contribution to similarity within habitats. Results show species contribution at a 90 cut-off from the SIMPER analysis.
SpeciesAv. AbundanceContribution %
RiverineTributariesOncorhynchus mykiss0.7165.91
Trichomycterus areolatus0.5018.62
Geotria australis0.2910.30
LacustrineLittoralGalaxias maculatus0.9256.91
Basilichthys microlepidotus0.6342.18
PelagicPercichthys trucha0.1259.23
Cyprinus carpio0.0720.04
Odonthestes mauleanum0.0714.33
Table 4. Total number of individuals captured during the two sampling seasons and the maximum and average body size (cm) of the target species in each sampling site. Empty cells represent no capture.
Table 4. Total number of individuals captured during the two sampling seasons and the maximum and average body size (cm) of the target species in each sampling site. Empty cells represent no capture.
SpeciesRiverine HabitatLacustrine Habitats
TributariesLittoralPelagic
R1R2R3R4R5R6R7R8R9R10L1L2L3L4L5P1P2P3P4P5
O. mykissTotal abundance6511223133861238 1 1 1
Maximum length17.725.96.616.85.925.42118.213.3 6.5 40.5 24.7
Mean length9.07.76.26.14.17.810.19.54.7 6.5 40.5 24.7
G. maculatusTotal abundance 11 607737109131
Maximum length 6.15.7 5.45.55.75.35.3
Mean length 6.15.7 4.14.54.84.44.6
P. truchaTotal abundance 92011242
Maximum length 29.533.233.333.828.0
Mean length 25.926.630.827.826.7
C. carpioTotal abundance 17114
Maximum length 44.442.040.041.5
Mean length 33.142.040.034.7
Table 5. Significant explanatory environmental variables of O. mykiss and G. maculatus biomass, based on GLM.
Table 5. Significant explanatory environmental variables of O. mykiss and G. maculatus biomass, based on GLM.
SpeciesEnvironmental Variables in the ModelEstimateStd. Errorzp
RiverineO. mykissDepth−15.3530.747−20.553<0.001
Tributaries Temperature−0.2270.025−9.039<0.001
Dissolved Oxygen−0.3400.071−4.785<0.001
Oxygen saturation0.0340.0056.066<0.001
Turbidity−0.2550.012−20.535<0.001
Total P35.9262.60913.766<0.001
Total N−8.5050.699−12.155<0.001
pH1.1920.06318.812<0.001
LacustrineG. maculatusDepth−7.2092.159−3.339<0.001
Littoral Dissolved Oxygen−0.6780.280−2.4190.015
Oxygen saturation−0.0700.030−2.3090.020
Turbidity−0.4370.069−6.325<0.001
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Díaz, G.; Habit, E.; Urrutia, R.; Manosalva, A.; Barra, R.O.; Figueroa, R. Trait Composition and Assemblage Structure Analyses of Lacustrine Fishes: Synthesizing a Proposal for Better Fishing Practices. Water 2024, 16, 2333. https://doi.org/10.3390/w16162333

AMA Style

Díaz G, Habit E, Urrutia R, Manosalva A, Barra RO, Figueroa R. Trait Composition and Assemblage Structure Analyses of Lacustrine Fishes: Synthesizing a Proposal for Better Fishing Practices. Water. 2024; 16(16):2333. https://doi.org/10.3390/w16162333

Chicago/Turabian Style

Díaz, Gustavo, Evelyn Habit, Roberto Urrutia, Aliro Manosalva, Ricardo O. Barra, and Ricardo Figueroa. 2024. "Trait Composition and Assemblage Structure Analyses of Lacustrine Fishes: Synthesizing a Proposal for Better Fishing Practices" Water 16, no. 16: 2333. https://doi.org/10.3390/w16162333

APA Style

Díaz, G., Habit, E., Urrutia, R., Manosalva, A., Barra, R. O., & Figueroa, R. (2024). Trait Composition and Assemblage Structure Analyses of Lacustrine Fishes: Synthesizing a Proposal for Better Fishing Practices. Water, 16(16), 2333. https://doi.org/10.3390/w16162333

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