Introduction
Parasites live in and/or on other organisms, obtaining the necessary resources for their survival, such as shelter, protection, or food (Goater et al. Reference Goater, Goater and Esch2014). Although parasites are often associated with diseases, zoonoses, and economic losses, they contribute significantly to biodiversity and biomass across ecosystems, as well as support essential ecosystem functions (Frainer et al. Reference Frainer, McKie, Amundsen, Knudsen and Lafferty2018; Hudson et al. Reference Hudson, Dobson and Lafferty2006). Despite this, parasite biodiversity is underestimated or unknown in many regions, for different reasons, including geographical biases and lack of knowledge of the host fauna itself (Poulin Reference Poulin2014; Poulin et al. Reference Poulin, Presswell, Bennett, de Angeli Dutra and Salloum2023). In this sense, studies surveying parasite assemblages in locations not yet studied are extremely important for the knowledge of local biodiversity, especially in regions that suffer from anthropogenic pressures, as is the case in many continental aquatic environments worldwide (Morley Reference Morley2007).
The Brazilian semi-arid zone, also known as the Caatinga domain, covers 70% of the Northeastern region of the country and is characterized by a high potential for evapotranspiration. As a result, the aquatic ecosystems in this region are either intermittent or artificial waterbodies, such as reservoirs (Silva et al. 2017). Such artificial waterbodies are widespread in the Caatinga domain due to water scarcity and long periods of meteorological drought, alongside the ever-increasing human need for water (Dantas et al. Reference Dantas, da Silva and Santos2020). These reservoirs exhibit low outflow and high residence time, leading to nutrient accumulation and increasing their vulnerability to eutrophication and pollution (Barbosa et al. Reference Barbosa, Medeiros, Brasil, Cordeiro, Crispim and Silva2012). Moreover, they have fragmented natural fluvial habitats, resulting in habitat loss and disconnection, which have driven species loss, invasions, and biotic homogenization (Yamada et al. Reference Yamada, Bongiovani, Yamada and Da Silva2017).
Changes on abiotic and biotic factors might lead to variation in parasite distribution, for example, due to the different life cycle complexities (Anderson et al. Reference Anderson and Sukhdeo2010). Hence, the most diverse host communities in theory will support a higher diversity of parasites and more stable trophic links (Hudson et al. Reference Hudson, Dobson and Lafferty2006). As drivers of biodiversity, native parasites can shape the dynamics of the community, and the introduction of host species can lead to different scenarios (Chalkowski et al. Reference Chalkowski, Lepczyk and Zohdy2018), including enemy release (Schatz and Park Reference Schatz and Park2023) and release from the enemy effect (Lacerda et al. Reference Lacerda, Takemoto, Poulin and Pavanelli2013a), or may lead to spillover and/or spillback of parasites, disrupting trophic interactions (Britton et al. Reference Britton2013; Kelly et al. Reference Kelly, Paterson, Townsend, Poulin and Tompkins2009; Llaberia-Robledillo et al. Reference Llaberia-Robledillo, Balbuena, Sarabeev and Llopis-Belenguer2022). Parasites might represent an early indicator to assess impacts on the community dynamics affected by anthropogenic events (Falkenberg et al. Reference Falkenberg, Golzio, Pessanha, Patrício, Vendel and Lacerda2019; Lacerda et al. Reference Lacerda, Roumbedakis, Bereta Junior, Nuñer, Petrucio and Martins2018; Sures et al. Reference Sures, Nachev, Schwelm, Grabner and Selbach2023), and monitoring these dynamics in vulnerable systems, such as Caatinga domain reservoirs, can provide fundamental and faster answers regarding to those impacts (Marcogliese et al. Reference Marcogliese2005; Palm Reference Palm and Mehlhorn2011).
Fishes are the most parasitized vertebrates (Poulin & Morand Reference Poulin and Morand2004), and among the known diversity of Brazil’s freshwater ichthyofauna, approximately 11% have been reported as hosts (Eiras et al. Reference Eiras, Takemoto, Pavanelli and Adriano2011). There are approximately 400 fish species in the Caatinga domain, distributed across 137 genera, 34 families, and 7 orders (Lima et al. Reference Lima, Ramos, Silva and Rosa2017). The Paraíba River Basin, which is predominantly situated in the Caatinga domain, is home to a diversity of 47 freshwater fish species, distributed across 38 genera, 20 families, and 6 orders (Ramos et al. Reference Ramos, Lima, Costa, da Silva, Avellar and Oliveira-Silva2018), and the Mamanguape River Basin is home to 32 freshwater fish species distributed across 26 genera, 16 families, and 6 orders (Oliveira-Silva et al. Reference Oliveira-Silva, Ramos, Carvalho-Rocha, Viana, Avellar and Ramos2018). Both basins are located in the Paraíba geographical state, one of the semi-arid areas most densely populated in the world and a region with significant socio-economic concerns as a consequence of water deficits caused by recurrent meteorological and hydrological droughts (Dantas et al. Reference Dantas, da Silva and Santos2020).
Despite the high diversity of potential hosts, a significant portion of the parasite fauna in Brazil remains undocumented (Luque and Poulin Reference Luque and Poulin2007). The main aim of this study was therefore to expand data on fish parasite diversity, providing the first inventory for fish parasite interactions in eight reservoirs in the Caatinga domain and widen our understanding of ichthyoparasites in these ecologically vulnerable and socio-economically crucial ecosystems. In addition, current and future changes plan to deal with water scarcity in the region, such as a water diversion project and its consequences for the aquatic biota (Sousa et al. Reference Sousa, Falkenberg, Lima, Winkeler, Ramos, Lustosa-Costa, Menezes and Lacerda2025), emphasizing the timeliness of the present inventory.
Material and methods
Study area
The Paraíba River Basin (37º 20’ to 36º 10’ W and 7º 20’ to 8º 20’ S), on the Borborema Plateau, represents 32% of the Paraíba state area and drains total or partially 85 municipalities (Xavier et al. Reference Xavier, Dornellas, Maciel and Bú2012). A total of eight reservoirs were selected: Poções Reservoir (Municipality of Monteiro), Camalaú Reservoir (Municipality of Camalaú), Epitácio Pessoa Reservoir (Municipality of Boqueirão), Acauã Reservoir (Municipality of Itatuba), Cordeiro Reservoir (Municipality of Congo), Sumé Reservoir (Municipality of Sumé), Taperoá Reservoir (Municipality of Taperoá), and Araçagi Reservoir in the Mamanguape River Basin (Municipality of Araçagi) (Figure 1).
Figure 1. Reservoirs of the Paraíba and Mamanguape River basins used in this study, and where they are geographically inserted.
Sample collection
Sampling was carried out as part of the Long-Term Ecological Project (Projeto Ecológico de Longa Duração Rio Paraíba Integrado; PELD-RIPA). Due to the hydrological seasonality of the reservoirs (Barbosa et al. Reference Barbosa, Medeiros, Brasil, Cordeiro, Crispim and Silva2012), samples were collected during two distinct periods: one corresponding to the rainy season (May to June 2022) and another to the dry season (December 2021 to January 2022). At each reservoir, sampling was performed using three types of fish-collecting gear: gillnets, trawl nets, and cast nets. Two sets of gillnets were placed, with measures of 40 meters in length and 1.5 meters in height, and consisting of four 10-meter sections with different mesh sizes (60 mm, 80 mm, 100 mm, and 120 mm). The gillnets were installed in the littoral zone of the reservoirs in the late afternoon and retrieved after 12 hours. Additionally, two trawl nets measuring 10 m in length and 1.5 m in height were used, along with 6 casts of cast nets (16 palms with a mesh size of 2.5 cm). This sampling effort was conducted separately for each reservoir during both the rainy and dry seasons. Samplings were conducted with authorization from the Brazilian Biodiversity Authorization and Information System (SISBIO) (License number: 83108-4) and were registered at the National Genetic Heritage Management System (SISGEN) (Access Number: AB202CE).
Fish were euthanized with eugenol solution and fixed in 10% formalin, which was injected mainly into the abdominal cavity to ensure the preservation of organs and any parasites (Lucena et al. Reference Lucena, Calegari, Pereira and Dallegrave2013; Malabarba and Reis Reference Malabarba and Reis1987). In the laboratory, fish were identified (Ramos et al. Reference Ramos, Lima, Costa, da Silva, Avellar and Oliveira-Silva2018) and necropsied, with all potential infection sites examined – external tegument, nostrils, fins, mesentery, eyes, gills, heart, intestine, stomach, liver, kidneys, gallbladder, swim bladder, and gonads. Selected structures were observed under a stereomicroscope, and the parasites found were stored in microtubes with 70% ethanol (Eiras et al. Reference Eiras, Takemoto and Pavanelli2006).
Before parasite identification, a series of processing methods was used to aid visualization: Nematoda specimens were clarified in lactic acid or Amman’s lactophenol; Copepoda and Monogenea were mounted in slides with Grey & Wess’s medium; and Digenea, Cestoda, and Acanthocephala were stained with Acetic Carmine (Eiras et al. Reference Eiras, Takemoto and Pavanelli2006). To identify parasites, literature sources including Boxshall and Montú (Reference Boxshall and Montú1997), Cohen et al. (Reference Cohen, Justo and Kohn2013), Gibson et al. (Reference Gibson, Bray and Jones2002), Moravec (Reference Moravec1998), and Yamaguti (Reference Yamaguti1963) were used. The specimens were identified to the lowest taxonomic level possible and grouped by the lower identification, sometimes with the use of morphospecies. All individuals identified at least to the family level were deposited at the Coleção de Invertebrados Paulo Young (CIPY), Universidade Federal da Paraíba, João Pessoa, except for those who only had damaged specimens (deposit numbers: UFPB.NEMA-170 to 175; UFPB.CRUST-7262 to 7266; UFPB.PLAT-97 to 124).
Data analysis
For the data analysis, only the non-encysted parasites were considered. Parasitological indices (prevalence, mean intensity, and mean abundance) were calculated for the host-parasite interaction data following Bush et al. (1997) using the software Quantitative Parasitology (QPweb, version 1.0.15). All subsequent data analysis was completed using R (version 4.4.2) (R Core Team, 2024), and the ‘vegan’ (Oksanen et al, Reference Oksanen, Simpson, Blanchet, Kindt, Legendre, Minchin, O’Hara, Solymos, Stevens, Szoecs, Wagner, Barbour, Bedward, Bolker, Borcard, Carvalho, Chirico, De Caceres, Durand, Evangelista, FitzJohn, Friendly, Furneaux, Hannigan, Hill, Lahti, McGlinn, Ouellette, Ribeiro Cunha, Smith, Stier, Ter Braak and Weedon2024), ‘dplyr’ (Wickham et al. Reference Wickham, François, Henry, Müller and Vaughan2023), ‘iNEXT’ (Hsieh et al. Reference Hsieh, K Ma and Chao2024), ‘ggplot2’ (Wickham Reference Wickham2016), and ‘bipartite’ (Dormann et al. Reference Dormann, Gruber and Frund2008) packages. Rarefaction was completed to understand whether the sampling effort was sufficient for each host species to represent the parasite diversity. For the rarefaction curve, the ‘q = 0’ method was used to rarefy and extrapolate based on parasite richness per host (Chao et al. Reference Chao, Gotelli, Hsieh, Sander, Ma, Colwell and Ellison2014). The overall host-parasite network for the region was visualised as a directed bipartite network, with nodes ordered by their similarity of interactions between hosts and parasites using correspondence analysis (CCA - Dormann et al. Reference Dormann, Gruber and Frund2008).
Data are available at the DATA-PB repository (Lacerda & de Lima, Reference Lacerda and de Lima2025), and a version-controlled repository is also available for the code and analysis (https://github.com/civlima/fish-parasite-diversity).
Results
A total of 1,170 fish were sampled and necropsied, belonging to 21 species: Astyanax bimaculatus (Linnaeus 1758) (n=282), Characidium bimaculatum Fowler 1941 (n=3), Cichla monoculus Spix & Agassiz 1831 (n=22), Cichlasoma orientale Kullander 1983 (n=107), Geophagus brasiliensis (Quoy & Gaimard 1824) (n=137), Hemigrammus marginatus Ellis 1911 (n=63), Hoplias malabaricus (Bloch 1794) (n=70), Hypostomus pusarum (Starks 1913) (n=27), Leporinus piau Fowler 1941 (n=29), Moenkhausia costae (Steindachner 1907) (n=3), Oreochromis niloticus (Linnaeus 1758) (n=239), Plagioscion squamosissimus (Heckel 1840) (n=8), Poecilia vivipara Bloch & Schneider 1801 (n=24), Prochilodus brevis Steindachner 1875 (n=72), Psalidodon fasciatus (Cuvier 1819) (n=1), Psectrogaster rhomboides Eigenmann & Eigenmann 1889 (n=7), Saxatilia brasiliensis (Bloch 1792) (syn. Crenicichla brasiliensis) (n=49), Serrapinnus piaba (Lütken 1875) (n=4), Steindachnerina notonota (Miranda Ribeiro 1937) (n=19), Synbranchus marmoratus Bloch 1795 (n=1), and Triportheus signatus (Garman 1890) (n=1).
Among the fishes collected, 42% of the individuals (n=491) were parasitized by at least one unencysted parasite, three species were parasitized only by cysts (C. bimaculatum, P. rhomboides, and S. piaba), and three species were not parasitized (P. fasciatus, S. marmoratus, and T. signatus). The rarefaction curve showed that A. bimaculatus is the host species most well-represented in parasite diversity, also highlighting O. niloticus as a host species with a high parasite diversity and greater abundance compared with A. bimaculatus. Cichla monoculus, P. brevis, and H. malabaricus supported high abundances of parasites, although parasite diversity was low (Figure 2).
Figure 2. Rarefaction curve of parasite diversity based on species richness across host species on Paraíba and Mamanguape basins reservoirs.
A total of 3,567 non-cyst parasite specimens were identified, belonging to 52 taxa across seven parasitological groups. The groups recorded here were Monogenea (23 taxa), Digenea (10), Cestoda (1), Nematoda (12), Acanthocephala (1), Copepoda (4), and Argulidae (1) associated with 15 parasitized fish species (Figure 3; Table 1).
Figure 3. Relative abundance of the major parasite groups identified across all host species.
Table 1. Parasite indices for each host-parasite interaction. P (%) = prevalence; MI = mean intensity; MA = mean abundance; IS = infection site; CI = confidence interval. *Only one host infected, standard deviation cannot be calculated. EY = eyes; FI = fins; GI = gills; GO = gonads; HE = heart; IC = intestinal caeca; IN = intestine; KI = kidney; LI = liver; ME = mesentery; SB = swim bladder; (M) = Metacercaria; (L) = Larva.
The most abundant taxa were Cichlidogyrus sp. 2 (Monogenea), parasitizing O. niloticus, and Proteocephalus microscopicus (Cestoda), parasitizing C. monoculus (Figure 4), which also presented the highest intensity value among the community. Other high values of mean intensities found were in the host H. malabaricus, for the digeneans Sphincterodiplostomum spp. and Dendrorchis sp. 1. High prevalences were present on Procamallanus (Spirocamallanus) inopinatus (Nematoda) in the host L. piau, Austrodiplostomum compactum (Digenea) in P. squamosissimus, and Miracetyma piraya (Copepoda) in P. brevis (Table 1).
Figure 4. (a) New geographical record of Cichlidogyrus sp. 2, parasitizing O. niloticus and new interaction with C. monoculus; (b) New geographical record of Unilatus aff. anoculus, parasitizing H. pusarum; (c) New geographical record of Dendrochis sp. 1, parasitizing H. malabaricus and a new interaction with P. squamosissimus.
There were 38 previously unrecorded parasite taxa for the Paraíba and Mamanguape River Basins. Additionally, 23 unknown host-parasite interactions were documented for the first time through this study, including the invasive host species C. monoculus, with the new record of interaction with the copepod Lamproglena monodi and the nematoda Procamallanus (Spirocamallanus) neocaballeroi (Figure 5).
Figure 5. Fish-parasite interactions of reservoirs in Paraíba state, highlighting new geographical records and new interactions.
Discussion
Expanding our understanding of host-parasite interactions is critical to fully understand the impacts of anthropogenic activities on biodiversity. Here, we developed the knowledge of ichthyoparasites in an understudied region of Brazil, identifying previously unobserved parasite taxa and interactions for both native and invasive fishes. The data produced here contributes significantly to the inventory of fish-parasite interactions, with only a few existing studies on fish parasites, mainly at a community level, in Brazilian semi-arid freshwater systems. Regarding the Brazilian semi-arid region, the parasite diversity is hardly documented; the first studies considering fish parasite communities were developed in the state of Ceará along the Jaguaribe River Basin (Falkenberg et al. Reference Falkenberg, de Lima, Yamada, Ramos and Lacerda2024a; Falkenberg et al. Reference Falkenberg, de Lima, Yamada, Ramos and Lacerda2024b). Although the region is highly dependent on reservoirs, there is no data on the fish parasite fauna in these freshwater systems, nor in the state of Paraíba at a community level. That is why this study represents the first steps towards unveiling the fish-parasite diversity of a unique freshwater system under anthropogenic impact, which may alter the structure and processes of an entire aquatic ecosystem and its fauna. This work builds on information we have in the proximal Jaguaribe River Basin (Falkenberg et al. Reference Falkenberg, de Lima, Yamada, Ramos and Lacerda2024a; Falkenberg et al. Reference Falkenberg, de Lima, Yamada, Ramos and Lacerda2024b), the Amazon region (Thatcher Reference Thatcher2006), and the Upper Paraná River floodplain (Takemoto et al. Reference Takemoto, Pavanelli, Lizama, Lacerda, Yamada, Moreira, Ceschini and Bellay2009), and wider information about Brazilian freshwater ichthyoparasitology (Eiras et al. Reference Eiras, Takemoto, Pavanelli and Adriano2011). Furthermore, it contributes to the continental knowledge, adding to data on South American and Neotropical fish parasites (Cohen et al. Reference Cohen, Justo and Kohn2013; Kohn et al. Reference Kohn, Fernandes and Cohen2007; Luque & Poulin Reference Luque and Poulin2007; Luque et al. Reference Luque, Pereira, Alves, Oliva and Timi2017; Moravec Reference Moravec1998).
High abundances of invasive fish species are found on the Brazilian semi-arid, such as the Nile tilapia (Oreochromis niloticus), which has been well-established in the Paraíba reservoirs since the introduction in the 1950s, when fisheries were started in the Brazilian Northeast (Ramos et al. Reference Ramos, Lima, Costa, da Silva, Avellar and Oliveira-Silva2018). An endemic cichlid from the Brazilian Northeast, Ciclhasoma orientale, popularly known as ‘cará’ or ‘cará preto’, occupies the same niche as O. niloticus (Berbel-Filho et al. Reference Berbel-Filho, Martinez, Ramos, Torres and Lima2016). Cichlasoma orientale presented a low parasite diversity in this study, even when compared to its invasive competitor O. niloticus. Furthermore, the Amazon species Cichla monoculus, popularly known as ‘tucunaré’, has been introduced in the semi-arid reservoirs (Attayde et al. Reference Attayde, Brasil and Menescal2011, Chellappa et al. Reference Chellappa, Câmara, Chellappa, Beveridge and Huntingford2003) and showed a high abundance of parasites per host individual, although the lower diversity compared to O. niloticus. The cestode species P. microscopicus was identified in C. monoculus, indicating the parasite was introduced with the Amazonian cichlid, as already documented for the Tocantins River with the invasive host Cichla piquiti (Lacerda et al. Reference Lacerda, Takemoto, Poulin and Pavanelli2013a). To better evaluate the host-parasite dynamics of biological invasions in the region, it is necessary to combine molecular studies at the invaded community level and biogeographical studies on the parasite fauna of hosts in their native range (Lacerda et al. Reference Lacerda, Takemoto, Poulin and Pavanelli2013a).
Native host species can be under-represented, due to difficulty of capturing or even the decrease of their populations due to competition with the invasive species (Reid et al. Reference Reid, Carlson, Creed, Eliason, Gell, Johnson, Kidd, MacCormack, Olden, Ormerod, Smol, Taylor, Tockner, Vermaire, Dudgeon and Cooke2019), reducing the sampling of parasites on those hosts. Hence, the absence of parasites in P. fasciatus, S. marmoratus, and T. signatus in the present study does not necessarily indicate that these species are not parasitized, but instead indicates the need to increase the efforts on these species. The most abundant species in this study, A. bimaculatus, was the native host species that showed the highest parasite diversity for the semi-arid reservoirs, reinforcing the correlation between the parasitic richness and the abundance of dominant fish species (Poulin and Morand Reference Poulin and Morand2004; Telfer and Bown Reference Telfer and Bown2012).
High prevalence values were observed for the following interactions: M. piraya (Copepoda) in P. brevis (44.4%), A. compactum (Digenea) in P. squamosissimus (62.5%), Anisakidae sp. 2 (Nematoda) in M. costae (33.3%), and P. (S.) inopinatus (Nematoda) in L. piau (79.3%). Despite the high prevalence of other groups, the most species-rich of freshwater parasites is the Class Monogenea (Luque & Poulin Reference Luque and Poulin2007), which presented 23 taxa recorded in this study. This is a group of ectoparasites known for their high host specificity (Poulin Reference Poulin1992), corroborated by our results, as only two monogenean taxa were found in more than one host species. The highest richness of this group was in the genus Cichlidogyrus, parasites of Nile tilapia (O. niloticus), of which we recorded 12 species. Although its host has well-established populations as an invasive species, its parasites are only starting to be identified in the Brazilian semi-arid. In the last years, a few studies approached new descriptions of Monogenean group (Diniz et al. Reference Diniz, Sousa, Yamada and Yamada2025; Silva et al. Reference Silva, Silva and Yamada2021; Silva et al. Reference Silva, Falkenberg and Yamada2025; Yamada et al. Reference Yamada, Diniz, Sousa, Yamada and Tavares-Dias2024), and we look forward to have the relations of parasites with native hosts, such as Cichlasoma orientale, that was found infected with one individual of Cichlidogyrus sp. 2, clarified in further studies with its implications to the native fish fauna.
Cestodes were found to be strongly abundant when present, besides showing the highest intensity among the parasite community, showing a highly aggregated distribution in the host population. In most fish, the cestodes were found in their encysted form, suggesting that the analysed fish serve as an important intermediate component in the life cycle of those parasites.
In 2017, the Paraíba River Basin received an inter-basin water transfer from the São Francisco River transposition (SF-IWT), to ensure water security in the semi-arid region of Paraíba (Silva et al. Reference Silva, Ramos, Carvalho, Brito, Ramos, Rosa, Sánchez-Botero, Novaes, Costa and Lima2020). Water quality and environmental parameters have been shown to be changing under the SF-IWT in the Paraíba reservoirs (Barbosa et al. Reference Barbosa, Severiano, Cavalcante, Lucena-Silva, Mendes, Barbosa, Silva, Oliveira and Molozzi2021). For aquatic organisms, such as fish, parasite transmission is altered by conditions in the aquatic environment, and initial results have shown that these might be impacting host-parasite relationships (Falkenberg et al. Reference Falkenberg, de Lima, Yamada, Ramos and Lacerda2024a). Water transfers can also result in species introductions, including fishes and their parasites (Dobson and May Reference Dobson and May1987), as recorded for the Paraíba Basin with the introduction of Moenkhausia costae after the SF-IWT (Ramos et al. Reference Ramos, Lustosa-Costa, Lima, Barbosa and Menezes2021; Sousa et al. Reference Sousa, Falkenberg, Lima, Winkeler, Ramos, Lustosa-Costa, Menezes and Lacerda2025). New interactions present unknown consequences for native fauna, potentially causing environmental imbalances or even extinction of endemic species (Lacerda et al. Reference Lacerda, Yamada, Antonucci, Dias, Pavanelli, Takemoto and Eiras2013b).
Although in this study we expanded our understanding of parasite communities, providing new geographical records (67–86% of taxa identified) for the region, it is important to recognize that several species could not be identified as species level, due to low abundance of the parasites/hosts, or due to complex taxonomic identification that requires integrative taxonomy efforts, as evidenced by recent studies that describe new species in Brazilian semi-arid aquatic ecosystems (Diniz et al. Reference Diniz, Sousa, Yamada and Yamada2025; Silva et al. Reference Silva, Silva and Yamada2021; Silva et al. Reference Silva, Falkenberg and Yamada2025; Yamada et al. Reference Yamada, Diniz, Sousa, Yamada and Tavares-Dias2024). Considering this, there is an urgent need to expand studies on fish parasite biodiversity, as the first steps to monitor and understand the ecological impacts of significant anthropogenic alterations in these critically important freshwater ecosystems.
Acknowledgements
The authors acknowledge the LAHMP/UFPB team Marcylenne Santana, Gilson do Nascimento Melo, and Igor Winkeler for their help with the lab procedures.
Financial support
This work is supported by Paraíba State Research Foundation, FAPESQ-PB, Brazil, as part of the doctoral thesis of V.M.M.L. (grant 16/2022), supervised by A.C.F.L. (grants FAPESQ PB Universal 3090/2021; CNPq/CONFAP-FAPs/PELD nº 23/2024, PELD RIPA 445968/2024-9; CNPq/MCTI/FNDCT/CT-Hidro nº 63/2022, 409348/2022-8, PRONEX/FAPESQ-PB 027-2023) and F.M.W. The sampling collection was supported by Universal/CNPq (R.F.M., Process Number: 421997/2018-4); This work was also supported by FUNCAP through postdoctoral fellowships (J.M.F., grant 0213-00077.01.01/23), (P.O.F.Y., grant FPD-0213-00301.01.01/23); by CNPq (F.H.Y., 304502/2022-7 and 174814/2023-2; T.P.A.R, 02654/2024-7); and by SNI-ANII and PEDECIBA (F.T.M.).
Competing interest
The authors declare none.
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