Open-access Preliminary assessment of the fish fauna of the Canandé Forest Reserve, Ecuadorian Chocó, with comments on its trophic role

Evaluación preliminar de la ictiofauna de la Reserva Forestal de Canandé, Chocó del Ecuador, con comentarios sobre su función trófica

Abstract

Ecuador’s coastal region has an exceptional freshwater biodiversity. Despite its biological richness, the region’s aquatic ecosystems face significant threats due to deforestation, pollution, unsustainable fishing and climate change. Ichthyological research becomes essential to understand current species diversity and to develop effective conservation strategies. Located within this region is the Canandé reserve, created to protect one of the last remnants of threatened tropical forests. This study aims to provide a first overview of the ichthyological fauna of the reserve and to highlight the interconnection between aquatic and terrestrial habitats. Two water bodies were sampled and 14 fish species from 11 different families were identified. During an initial sampling in 2023, 12 species were found, and two more were added in subsequent expeditions. Although all species encountered are classified as ‘Least Concern’ by the IUCN, one characin, Pseudochalceus bohlkei, is nationally ‘Endangered’. The ichthyological community of the reserve includes species that also feed on terrestrial resources like amphibians. At the same time, fish serve as prey for other terrestrial animals, suggesting a complex dynamic between aquatic and terrestrial ecosystems in the reserve. It highlights the importance of further research on the connectivity of aquatic and terrestrial ecosystems in the neotropics.

Keywords
checklist; ecology; food webs; freshwater; rivers

Resumen:

La región costera de Ecuador posee una excepcional biodiversidad de agua dulce. A pesar de su riqueza biológica, los ecosistemas acuáticos de la región se enfrentan a importantes amenazas debido a la deforestación, la contaminación, la pesca insostenible y el cambio climático. La investigación ictiológica se hace imprescindible para comprender la diversidad actual de especies y desarrollar estrategias de conservación eficaces. Dentro de esta región se encuentra la reserva de Canandé, creada para proteger uno de los últimos remanentes de bosques tropicales amenazados. Este estudio pretende ofrecer una primera visión general de la fauna ictiológica de la reserva y poner de relieve la interconexión entre los hábitats acuáticos y terrestres. Se muestrearon dos masas de agua y se identificaron 14 especies de peces de 11 familias diferentes. Durante un muestreo inicial en 2023, se encontraron 12 especies, a las que se añadieron otras dos en expediciones posteriores. Aunque todas las especies encontradas están clasificadas como de «Preocupación Menor» por la UICN, un characín, Pseudochalceus bohlkei, está «En Peligro» a escala nacional. La comunidad ictiológica de la reserva incluye especies que también se alimentan de recursos terrestres, como los anfibios. Al mismo tiempo, los peces sirven de presa a otros animales terrestres, lo que sugiere una compleja dinámica entre los ecosistemas acuáticos y terrestres de la reserva. Esto pone de relieve la importancia de seguir investigando la conectividad de los ecosistemas acuáticos y terrestres en los neotrópicos.

Introduction

The coastal northwest of Ecuador hosts an exceptional freshwater biodiversity that has long captured the attention of ichthyologists and experts in the study of aquatic ecosystems (Barriga 1994). However, human impact, such as unsustainable fishing, agricultural encroachment and pollution, as well as climate change threaten the health of aquatic ecosystems in the region (Aguirre et al. 2021). Thus, ichthyofaunistic research is crucial, not only to document the current species diversity, but also to help design effective conservation and restoration strategies.

Barriga (2012) identified eleven ichthyohydrographic zones in Ecuador. The main criterion for their delimitation was the composition of fish communities, which are the outcome of historical, geographic, as well as biotic and abiotic factors and processes. The study lists 948 freshwater and intertidal fish species in Ecuador, accounting for 7.3% of the world’s freshwater species, and 17% of the neotropical ichthyofauna. Of these, 824 species are strictly freshwater fish, and 36% are endemic to Ecuador (Barriga 2012). In northwestern Ecuador, especially in the Chocó, researchers such as Jiménez-Prado (2015) and Barriga (1989, 1991, 1994, 2012), as well as Aguirre (2017, 2019a, 2019b, 2021), have provided valuable contributions concerning the biological richness of the ichthyofauna in the past.

As a well-known biodiversity hotspot, the Chocó rainforest is a particularly important Ecuadorian ecoregion, comprising lowland to lower montane forest ecosystems on the Pacific side of the country. Although being under heavy pressure of habitat destruction and conversion (López et al. 2010, Luna et al. 2020), there are also officially and privately protected areas. A particularly important one is the Canandé Forest Reserve, owned and managed by the Ecuadorian NGO Jocotoco (www.jocotoco.org). The reserve is located in the province of Esmeraldas and was established with the intention to conserve one of the last remnants of Ecuador’s Chocó rainforests. Currently the reserve is the centre of a large-scale, international research project on the restoration and recovery of rainforests and the interactions of its inhabitants (www.reassembly.de, Blüthgen et al. 2020). This project investigates various organisms from plants and soil arthropods to frogs, birds, bats and monkeys. One important group of organisms that was not included so far is fish. Fish are not only a key taxonomic group in freshwater ecosystems but are also an important food source for humans, as well as other terrestrial animals in tropical regions. With this study, we thus aim to give a first overview about the fish fauna of a small forest stream close to the Canandé research station, as well as a mid-sized river close by. Furthermore, we want to give a summary of the known food web interactions of the recorded fish species.

Material and Methods

1. Study area

We sampled two streams, a larger one called “Río Colombiano” between the Canandé station and the village Hoja Blanca (Fig. 1A, B), and one small forest stream next to the Canandé Research Station (Fig. 1C, D). Both carry crystal clear water, except after heavy rains.

Figure 1
Photos of the sampled streams. A.-B. The large stream called “Río Colombiano”; and C.-D. a small forest stream, located close to the Canandé field station.

The larger river has a streambed with gravel and larger rocks. On average it is 10 m wide, and the water level is shallow (< 50 cm), exceeding 100 cm only in a very few deeper, almost stagnant pools. Those deeper pools often contain large assemblages of dead wood and leaf litter on sandy soil. This river flows through rainforest remnants, in some places reduced to very narrow (< 50 m) bands of riparian forest, in others, close to the road to Hoja Blanca, trees are lacking completely. Further upstream the river is close to cacao plantations. On 9th March 2023 at 8 pm, the water temperature measured 23.6°C and conductivity was 123 µS, pH and potential chemicals in the water could not be measured.

The small stream, draining old-growth and old-regenerating rainforest, showed a strongly fluctuating water level of 5–50 cm depending on rain, a sandy to gravely streambed, with constrictions by larger boulders (Fig. 1D), and deeper, flow-calmed troughs behind them. During mid-day of the 24th April 2024 the stream width varied between 91–215 cm along a 100 m transect, while the visible streambed extended between 3 and 7.5 m. Upstream the river comes close to a road, but not to plantations. The average water speed (mean of 10 measurements along 100 m) on 24th April 2024 was 0.21 m/s (max. 0.37 m/s, min 0.10 m/s). Water parameters were measured a year before during similar conditions at 8 pm on 4th March 2023. Water temperature was 23.2°C, conductivity was 177µS, pH was not measured. Both water bodies contained only very little to no visible trash or anthropogenic litter. In addition to the two sampled water bodies, we also report further random observations of other little streams in the reserve, very much resembling the small described one.

2. Field work

We have been searching for fish on several days in three time periods; during March 2023, November 2023, and in March-May 2024. Our main aim focused on amphibian communities; fish data were only sampled opportunistically. Fish were assessed with three different methods. Small hand nets of 30 × 20 cm and 1 mm mesh width were used mainly at night to catch resting fish in shallow water. Nylon funnel traps of 75 × 28 × 28 cm (5 mm mesh width) with two entrances of 6 cm diameter, were placed overnight in different areas of the streams and checked the next day. Finally, a self-made seine net of 6 × 1.5 m (5 mm mesh width) was used to close off whole sections of the stream and drive the fish to the bank where they were caught with hand nets. After identification and taking pictures, all caught individuals were released back to the streams, no specimens were collected. Species were identified based on morphology, mainly following Jiménez-Prado et al. (2015).

Results

In the two streams, we caught 14 species of fish (Table 1), belonging to eleven families, namely Trichomycteridae, Astroblepidae, Heptapteridae, Loricariidae, Poeciliidae, Lebiasinidae, Characidae (3 species), Erythrinidae, Cichlidae (2 species), Oxudercidae and Synbranchidae. The abundance in Table 1 represents the number of fish recorded per species.

Table 1
Taxonomic list of ichthyofauna we report for the Canandé Reserve, northwestern Ecuador. The abundance refers to the number of individuals observed in the Río Colombiano (R) and the small forest stream close to the station (FS) Further microhabitat information is given in the comments.

Additionally, we observed juveniles of Oreochromis aureus, a West African cichlid species commonly bred for food supply in small trenches along the road within the reserve. In the natural water bodies, we did not find any individuals. Therefore, we did not include them into the assessment of the Canandé fish fauna.

Discussion

During the fieldwork in early 2023, we were able to find and catch 12 fish species in the Canandé Reserve (Fig. 2). Two more visits to the “Río Colombiano” in November 2023 and March 2024 revealed two additional species (Fig. 3). Due to our limited sampling method, and by not collecting vouchers, it is likely that we missed some fish species in the larger river and potentially overlooked cryptic species among those we caught. Barriga (2012) documented 57 species for the ichthyohydrographic area of Esmeraldas; thus, we recorded approximately 25% of this diversity in two rivers within less than ten days of sampling.

Figure 2
Photos of the live fish, caught in March 2023. A. Trichomycterus banneaui; B. Astroblepus brachycephalus; C. Pimelodella modestus; D. Rineloricaria jubata; E. Pseudopoecilia fria, male and female; F. Lebiasina multimaculata; G. Pseudochalceus bohlkei; H. Bryconamericus dahli; I. Hyphessobrycon daguae; J. Hoplias malabaricus; K. Mesoheros festae; L. Andinoacara blombergi.
Figure 3
Photos of two additional fish species, found in November 2023 and March 2024 in the “Río Colombiano”. A. Awaous transandeanus; B.-C. Synbranchus marmoratus.

1. Environmental quality and anthropogenic impacts

Not surprisingly, we detected a much lower fish fauna in the small forest streams (three species versus thirteen species in “Río Colombiano”), this difference can be explained by the variation in microhabitats, water parameters, and environmental quality. Smaller streams offer limited habitat diversity regarding depth, current velocity, substrate type, and structural complexity, which directly influences fish diversity (Grossman & Freeman 1987, Casatti 2005, Vlach et al. 2005). Moreover, these smaller streams experience stronger seasonal fluctuations in flow and water temperature, becoming more vulnerable to heat stress, oxygen depletion, and organic pollution during dry seasons (Gutiérrez-Fonseca et al. 2020). This could be one reason why of the three species (Astroblepus brachycephalus, Trichomycterus banneaui, Pseudochalceus bohlkei), which we found in small streams, the two most abundant were catfish, being capable of enteral respiration.

Although our sampling sites are located within a protected reserve, some anthropogenic impacts still exist in the surrounding landscape, including agricultural expansion and selective logging. However, the detection of the gobiid Awaous transandeanus, classified as an indicator species for good water quality (CID-PUCESE & PRAS-MAE, 2014), suggests that, at least in parts of the Río Colombiano, environmental quality remains relatively intact. Nevertheless, continued environmental monitoring is recommended to detect subtle habitat changes that may affect sensitive species and the long-term conservation of the whole native ichthyofauna.

2. Species-specific microhabitat use

The species detected showed potential microhabitat specialization. For example, some characins like Bryconamericus dahli were abundant in the main channel, forming schools. In contrast, Pseudochalceus bohlkei was restricted to structurally complex microhabitats such as undercut banks and submerged woody debris. These microhabitat associations highlight the importance of preserving habitat heterogeneity for maintaining species diversity.

All caught cichlid specimens were small and probably juveniles or subadults, suggesting ontogenetic habitat segregation, a well-documented phenomenon in cichlids (Galis 1993, Kohda et al. 2008, Joyce et al. 2016). Further studies using techniques such as electrofishing could help clarify adult distribution patterns within the river system.

3. Taxonomic differentiation and distribution

The most abundant group in “Río Colombiano”, were characins. Bryconamericus dahli, showed some variation in colour. In some animals the dark eyespot on the base of the tail was almost absent, in others quite strong. Furthermore, some showed reflecting greenish colour, whereas others were more uniform silver. Román–Valencia et al. (2015) described a new Bryconamericus species from northwestern Ecuador which might occur in sympatry with B. dahli. As we caught the one H. daguae with a large school of B. dahli, a systematic investigation of a larger number of characin individuals as well as some dissections would be necessary to fully understand the species composition of characins swarms in the Rio Colombiano.

Pseudochalceus bohlkei was found to always swim in smaller groups. Within those, single individuals had longer fins and were more colourful than others, leading us to assume that this species may move in territorial harems instead of forming large swarms in the open water.

Citing Fowler (1911) and Laaz & Torres (2010), Jiménez Prado et al. (2015) state that Lebiasina aureoguttata mainly occurs in southwestern Ecuador (Rio Chimbo). Still, there have been observations of the species ~350 km further north in the Santiago-Cayapas River system (Jiménez Prado et al. 2015). While the Canandé area is located very close to Rio Esmeraldas it still belongs to the Santiago-Cayapas drainage. The sampled river “Rio Colombiano” flows into Rio Hoja Blanca which than flows eastwards and eventually into Rio Cayapas (https://www.openstreetmap.org/#map = 15/0.57905/−79.15838). Our finding therefore confirms that there are at least two separate distribution areas of L. aureoguttata in Ecuador.

4. Trophic ecology and ecosystem interactions

While fish communities in small to medium sized tropical streams have been shown to predominantly rely on autochthonous algae (Reis et al. 2020), terrestrial resources like leaf litter and terrestrial animals can be highly important food for fish as well (Brett et al. 2017). For example, terrestrial arthropods falling into the water serve as an important energy input into aquatic food webs, in lakes (Mehner et al. 2005) as well as in streams (Cloe & Garman 1996, Kawaguchi et al. 2003, Chan et al. 2008). Hence, we assume the large number and diversity of insects in the Ecuadorian Chocó to also play a crucial role in the food webs of the streams in Canandé.

While other terrestrial organisms probably play minor roles as food resource for fish, fish still can be important predators of them. This applies especially for amphibians, closely dependant on water bodies (e.g. Petranka et al. 1987, Martín-Torrijos et al. 2016, Remon et al. 2016). Fish do not only prey on adult frogs and toads, but especially on aquatic tadpoles and eggs, even if Gascon (1992) indicated fish to be only subordinate predators on tadpoles in neotropical systems. Many cases of fish-on-frog predation have been reported from the neotropics in the past, of which some do also refer to fish species found in Canandé. Individuals of Hoplias malabaricus have been observed preying on frogs of the genera Pristimantis (Delaix-Zaqueo et al. 2017) and Leptodactylus (Andrade et al. 2012). Both genera occur in the Canandé reserve with several species. Another erythrinid species, Hoplerythrinus unitaeniatus as well as a catfish of the genus Astroblepus were documented preying on poisonous Atelopus toads (Dias Lima et al. 2019, Cruz-García et al. 2023). Astroblepus catfish are found regularly in our study area and dwell in different kinds of streams where anurans, including Rhinella spp., Boana spp., Hyloscirtus palmeri and several centrolenid species have been found to reproduce. Therefore, we believe Astroblepus brachycephalus to be a potential predator of all these species, especially for their larvae. While Braga et al. (2008) states the eel Synbranchus marmoratus does not predate on tadpoles in the wild, other authors disagree (Junges et al. 2010, Maffei et al. 2011) and several experimental studies used the species as a model predator for Rhinella and Boana tadpoles (Junges et al. 2010 & 2012, Curi et al. 2021). We hence assume S. marmoratus to be at least an opportunistic predator of tadpoles in the Canandé area.

At the same time, fish are an important food source for terrestrial vertebrates (Suter 1991, Silva et al. 2014), especially birds, e.g. kingfishers and mammals like otters. In the Ecuadorian Chocó, there is the Neotropical Otter (Lontra longicaudis) and the Water Opossum (Chironectes minimus), of which the latter was also seen by the authors. Further taxa like some snakes are specialized or at least occasionally prey on fish. Tropidophis boulengeri is known to prey on Astroblepus in Canandé (Dwyer et al. 2018, Griesbaum et al. 2023). Pliocercus euryzonus, which is also present in the area, was recorded to feed on Trichomycterus in Colombia (Cardona et al. 2022).

Although we assume that fish consumption by terrestrial animals plays a small role in forming aquatic communities, compared to piscivorous fish, further research of terrestrial piscivores, especially birds, will help to better understand the food webs of the Canandé Reserve and the whole Chocó Rainforest. Furthermore, we want to encourage further research on the trophic relations between fish and terrestrial insects in the region.

5. Conservation considerations

Most of the recorded fish species are categorized as Least Concern by the IUCN (2024). However, Pseudochalceus bohlkei is considered “Endangered” in the national Red List of Ecuador (Aguirre et al. 2019a), even though in 2015 Jiménez Prado et al. classified it as only “rare”. On a national level, foremost a striking lack of data is visible: three of our recorded species are listed as “Data Deficient” (Trichomycterus banneaui, Astroblepus brachycephalus, Rineloricaria jubata) and five species have not even been assessed (Lebiasina aureoguttata, Hyphessobrycon daguae, Hoplias malabaricus, Awaous transandeanus, Synbranchus marmoratus). Aguirre et al. (2021) explicitly states that many more species of Ecuadorian fish might be threatened. Thus, to foster the conservation of Ecuadorian freshwater fish species, more systematic surveys and monitoring projects of water bodies including smaller forest streams are necessary. As freshwater ecosystems and their biodiversity are heavily impacted by human use, not only by fishery (Jiménez-Prado et al. 2015) but also by riparian land use, habitat alteration (Sánchez-Garcés 2017, Morabowen et al. 2019), and introduction of alien species, like Oreochromis aureus, a better understanding of trophic links and dependencies between the aquatic and terrestrial realm is also crucial for a sustainable forest management in the region.

Acknowledgments

We are grateful to ‘Fundación Jocotoco’ and its CEO Martin Schaefer, for permission to conduct research in the Canandé Reserve, and Katrin Krauth and Julio Carvajal for logistic support. We thank Nico Blüthgen, David Donoso, María-José Endara, Martin Schaefer, and Edith Villa for coordination and administration, and the Ministerio del Ambiente, Agua y Transición Ecológica for granting research permits through Contrato Marco MAE-DNBCM-2021-0187. This study was supported by the Deutsche Forschungsgemeinschaft (DFG) by funding the Research Unit REASSEMBLY (FOR 5207; sub-project SP2, grant RO 3064/5-1). Lastly, we want to thank Pablo Arguello, beknown expert of Ecuadorian fish, for his help and advice on the identity of the Lebiasina sp.

Data Availability

The original data only consists of photographs and geodata. Thus, we decided to publish it in a very convenient way, so it can be accessed and downloaded openly. The data can be found on iNaturalist under the following link: https://www.inaturalist.org/observations?nelat=0.8191313492076389&nelng=-78.71484976281675&swlat=-0.01580168281627335&swlng=-79.70361929406675&taxon_id=47178&user_id=frederic_griesbaum

References

  • AGUIRRE, W.E., NUGRA SALAZAR, F., SÁNCHEZ-GARCÉS, G., NAVARRETE-AMAYA, R., & VALDIVIEZO RIVERA, J. 2017. Range expansion of the genus sicydium (Teleostei: Gobiidae) to coastal mountain streams of southwestern Ecuador and possibly northwestern Peru. Check List, 13(1), 1–8. https://doi.org/10.15560/13.1.1
    » https://doi.org/10.15560/13.1.1
  • AGUIRRE, W., ANAGUANO-YANCHA, F., BURGOS-MORÁN, R., CARRILLO-MORENO, C., GUARDERAS, L., JÁCOME-NEGRETE, I., & VALDIVIEZO-RIVERA, J. (2019a). Lista roja de los peces dulceacuícolas de Ecuador Ministerio del Ambiente, DePaul University, Wildlife Conservation Society-Ecuador (WCS), Universidad Estatal Amazónica, Universidad Indoamérica, Instituto Quichua de Biotecnología Sacha Supai, Universidad Central del Ecuador, Pontificia Universidad Católica del Ecuador Sede en Esmeraldas, Instituto Nacional de Pesca, Universidad del Azuay, Instituto Nacional de Pesca, Universidad Central del Ecuador, Antonio Torres, Universidad de Guayaquil e Instituto Nacional de Biodiversidad. Quito, Ecuador.
  • AGUIRRE, W.E., YOUNG, A., NAVARRETE-AMAYA, R., VALDIVIEZO-RIVERA, J., JIMÉNEZ-PRADO, P., CUCALÓN, R.V., & SHERVETTE, V.R. (2019b). Vertebral number covaries with body form and elevation along the western slopes of the Ecuadorian Andes in the Neotropical fish genus Rhoadsia (Teleostei: Characidae). Biological Journal of the Linnean Society, 126(4), 706–720. https://doi.org/10.1093/biolinnean/biz023
    » https://doi.org/10.1093/biolinnean/biz023
  • AGUIRRE, W.E., ALVAREZ-MIELES, G., ANAGUANO-YANCHA, F., BURGOS MORÁN, R., CUCALÓN, R.V., ESCOBAR-CAMACHO, D., & ZÁRATE HUGO, E. (2021). Conservation threats and future prospects for the freshwater fishes of Ecuador: A hotspot of Neotropical fish diversity. Journal of Fish Biology, 99(4), 1158–1189. https://doi.org/10.1111/jfb.14881
    » https://doi.org/10.1111/jfb.14881
  • ANDRADE, E., JÚNIOR, T., JÚNIOR, J., & LEITE, J. (2012). Predation by native fish and feeding by crab species on Leptodactylus macrosternum Miranda-Ribeiro, 1926 (Anura: Leptodactylidae) in northeastern Brazil. Herpetology Notes, 5, 173–175.
  • BARRIGA, R. (1989). Peces de la Reserva Étnica y Forestal AWA, Ecuador Noroccidental Politécnica Nacional, 14(2)
  • BARRIGA, R. (1991). Peces de agua dulce Politécnica, 16(3), 7–88.
  • BARRIGA, R. (1994). Peces del Noroeste del Ecuador Politécnica, 19(2), 43–154.
  • BARRIGA, R. (2012). Lista de peces de agua dulce e intermareales del Ecuador Politécnica, 30(3), 83–119.
  • BLÜTHGEN, N., DONOSO, D. A., ENDARA, M.J., SCHAEFER, M., BREHM, G., DORMANN, C., et al. (2020). Reassembly of species interaction networks – Resistance, resilience and functional recovery of a rainforest ecosystem. Proposal for the 1st phase (2021–2025) of the DFG Research unit 5207. Privately published, Technical University Darmstadt.
  • BRAGA, A.L.C., DOS SANTOS POMPEU, P., CARVALHO, R.F., & FERREIRA, R.L. (2008). Dieta e crescimento de Synbranchus marmoratus (Bloch, 1975)(Pisces, Synbranchiformes) durante período de pré-estivação em uma lagoa marginal da bacia do São Francisco, Minas Gerais. Revista Brasileira de Zoociências, 10(2), 133–138.
  • BRETT, M.T., BUNN, S.E., CHANDRA, S., GALLOWAY, A.W.E., GUO, F., KAINZ, M.J., & Wehr, J.D. (2017). How important are terrestrial organic carbon inputs for secondary production in freshwater ecosystems? Freshwater Biology, 62, 833–853. https://doi.org/10.1111/fwb.12935
    » https://doi.org/10.1111/fwb.12935
  • CARDONA, J.P.D., FRANCO, E.G., MEJÍA, M.C.C., & SALGAR, J.C.A. (2022). Predation by a Cope’s False Coralsnake, Pliocercus euryzonus (Cope 1862) (Dipsadidae), on a catfish (Trichomycteridae: Trichomycterus) in the northern Andes of Colombia. Reptiles & Amphibians, 29(1), 95–97. https://doi.org/10.1163/156853222X16423687932912
    » https://doi.org/10.1163/156853222X16423687932912
  • CASATTI, L. (2005). Fish assemblage structure in a first-order stream, southeastern Brazil: Longitudinal distribution, seasonality, and microhabitat diversity. Biota Neotropica, 5, 75–83. https://doi.org/10.1590/S1676-06032005000100008
    » https://doi.org/10.1590/S1676-06032005000100008
  • CID-PUCESE, PRAS-MAE. (2014). Consultoría para la continuación de análisis de impactos de la minería aurífera en los cantones Eloy Alfaro y San Lorenzo de la Provincia de Esmeraldas. Producto 8.7 Informe Final de observación de calidad de agua en los cantones Eloy Alfaro y San Lorenzo, Esmeraldas
  • CHAN, E.K., ZHANG, Y., & DUDGEON, D. (2008). Arthropod ‘rain’ into tropical streams: the importance of intact riparian forest and influences on fish diets. Marine and Freshwater Research, 59(8), 653–660. https://doi.org/10.1071/MF07191
    » https://doi.org/10.1071/MF07191
  • CLOE, W., & GARMAN, G. (1996). The energetic importance of terrestrial arthropod inputs to three warm‐water streams. Freshwater Biology, 36(1), 104-114. https://doi.org/10.1046/j.1365-2427.1996.00080.x
    » https://doi.org/10.1046/j.1365-2427.1996.00080.x
  • CRUZ-GARCÍA, K., ZAPATA, N., & PEREZ-CORREA, J. (2023). First documented case of predation on the Río Pescado Jambato Toad (Atelopus balios Peters, 1973) by the Andean Catfish (Astroblepus sp.) on Cerro de Hayas, Naranjal, Ecuador. Herpetology Notes, 16, 423–425.
  • CURI, L.M., CUZZIOL BOCCIONI, A.P., PELTZER, P.M., ATTADEMO, A.M., BASSÓ, A., LEÓN, E.J., & LAJMANOVICH, R.C. (2022). Signals from predators, injured conspecifics, and pesticide modify the swimming behavior of the gregarious tadpole of the Dorbigny’s Toad, Rhinella dorbignyi (Anura: Bufonidae). Canadian Journal of Zoology, 100, 19–27. https://doi.org/10.1139/cjz-2021-0210
    » https://doi.org/10.1139/cjz-2021-0210
  • DELAIX-ZAQUEO, M., MELO-SAMPAIO, P.R., DE OLIVEIRA, I.L., MIRANDA, G.K.D., & DE CALDERON, L.A. (2017). The opportunistic behaviour of a common predator in aquatic systems in Amazonia: Predation on robber-frog Pristimantis cf. fenestratus by trahira Hoplias malabaricus (Bloch, 1794). Herpetology Notes, 10, 425–427.
  • DIAS LIMA, J., FERREIRA LIMA, J.R., & FERREIRA SOBRINHO, A. (2019). Predation of a harlequin toad, Atelopus hoogmoedi Lescure, 1974, by the gold wolf fish, Hoplerythrinus unitaeniatus (Spix & Agassiz, 1829), in a stream of the Cajari River Extractive Reserve, Amapá, Brazil. Herpetology Notes, 12, 587–589.
  • DWYER, Q., ARTEAGA, A., BARRIO-AMOROS, C., & FLAGE, A. (2018). Trachyboa boulengeri Diet. Herpetological Review, 49, 359–360.
  • FOWLER, H.W. (1911). New freshwater fishes from western Ecuador. Proceedings of the Academy of Natural Sciences of Philadelphia, 63, 493–520.
  • GALIS, F. (1993). Interactions between the pharyngeal jaw apparatus, feeding behaviour, and ontogeny in the cichlid fish, Haplochromis piceatus: A study of morphological constraints in evolutionary ecology. Journal of Experimental Zoology, 267(2), 137–154. https://doi.org/10.1002/jez.1402670204
    » https://doi.org/10.1002/jez.1402670204
  • GASCON, C. (1992). Aquatic predators and tadpole prey in central Amazonia: Field data and experimental manipulations. Ecology, 73(3), 971–980. https://doi.org/10.2307/1940657
    » https://doi.org/10.2307/1940657
  • GRIESBAUM, F., LINDNER, T., BOCK, S., ERNST, M., NEIRA-SALAMEA, K., MOREIRA, V., et al. (2023). Nine predation events by snakes from the Chocó rainforest of Ecuador. Herpetology Notes, 16, 749–756.
  • GROSSMAN, G.D., & FREEMAN, M.C. (1987). Microhabitat use in a stream fish assemblage. Journal of Zoology, 212(1), 151–176. https://doi.org/10.1111/j.1469-7998.1987.tb04086.x
    » https://doi.org/10.1111/j.1469-7998.1987.tb04086.x
  • GUTIÉRREZ-FONSECA, P.E., RAMÍREZ, A., PRINGLE, C.M., TORRES, P.J., MCDOWELL, W.H., COVICH, A., et al. (2020). When the rainforest dries: Drought effects on a montane tropical stream ecosystem in Puerto Rico. Freshwater Science, 39(2), 197–212. https://doi.org/10.1086/707073
    » https://doi.org/10.1086/707073
  • JIMÉNEZ-PRADO, P. (2015). Guía de peces para aguas continentales en la vertiente occidental del Ecuador Pontificia Universidad Católica del Ecuador Sede Esmeraldas.
  • JOYCE, B.J., DEMERS, E.E., CHIVERS, D.P., FERRARI, M.C., & BROWN, G.E. (2016). Risk-induced neophobia is constrained by ontogeny in juvenile convict cichlids. Animal Behaviour, 114, 37–43. https://doi.org/10.1016/j.anbehav.2016.02.005
    » https://doi.org/10.1016/j.anbehav.2016.02.005
  • JUNGES, C.M., LAJMANOVICH, R.C., PELTZER, P.M., ATTADEMO, A.M., BASSÓ, A. (2010). Predator–prey interactions between Synbranchus marmoratus (Teleostei: Synbranchidae) and Hypsiboas pulchellus tadpoles (Amphibia: Hylidae): Importance of lateral line in nocturnal predation and effects of fenitrothion exposure. Chemosphere, 81(10), 1233–1238.https://doi.org/10.1016/j.chemosphere.2010.01.052
    » https://doi.org/10.1016/j.chemosphere.2010.01.052
  • JUNGES, C.M., PELTZER, P.M., LAJMANOVICH, R.C., ATTADEMO, A.M., ZENKLUSEN, M.C., & BASSÓ, A. (2012). Toxicity of the fungicide trifloxystrobin on tadpoles and its effect on fish–tadpole interaction. Chemosphere, 87(11), 1348–1354. https://doi.org/10.1016/j.chemosphere.2012.02.061
    » https://doi.org/10.1016/j.chemosphere.2012.02.061
  • KAWAGUCHI, Y., TANIGUCHI, Y., & NAKANO, S. (2003). Terrestrial invertebrate inputs determine the local abundance of stream fishes in a forested stream. Ecology, 84(3), 701-708.
  • KOHDA, M., SHIBATA, J.Y., AWATA, S., GOMAGANO, D., TAKEYAMA, T., HORI, M., et al. (2008). Niche differentiation depends on body size in a cichlid fish: A model system of a community structured according to size regularities. Journal of Animal Ecology, 77(5), 859–868. https://doi.org/10.1111/j.1365-2656.2008.01408.x
    » https://doi.org/10.1111/j.1365-2656.2008.01408.x
  • LAAZ, E., & A. TORRES. 2010. Lista sistemática de peces de la Cuenca del río Guayas, disponible en: http://condor.depaul.edu/~waguirre/fishwestec/lista_peces_guayas.pdf
    » http://condor.depaul.edu/~waguirre/fishwestec/lista_peces_guayas.pdf
  • LÓPEZ, S., SIERRA, R., & TIRADO, M. (2010). Tropical deforestation in the Ecuadorian Chocó: Logging practices and socio-spatial relationships. Geographical Bulletin, 51(1), 3–11 https://doi.org/10.1163/001672610X12645838722227
    » https://doi.org/10.1163/001672610X12645838722227
  • LUNA, T.O., EGUIGUREN, P., GÜNTER, S., TORRES, B., & DIETER, M. (2020). What drives household deforestation decisions? Insights from the Ecuadorian lowland rainforests. Forests, 11, 1–12. https://doi.org/10.3390/f11010001
    » https://doi.org/10.3390/f11010001
  • MAFFEI, F., UBAID, F.K., & JIM, J. (2011). Anurofauna em área de cerrado aberto no município de Borebi, estado de São Paulo, Sudeste do Brasil: Uso do habitat, abundância e variação sazonal. Biota Neotropica, 11, 221–233. https://doi.org/10.1590/S1676-06032011000100019
    » https://doi.org/10.1590/S1676-06032011000100019
  • MARTÍN-TORRIJOS, L., SANDOVAL-SIERRA, J.V., MUÑOZ, J., DIÉGUEZ-URIBEONDO, J., BOSCH, J., & GUAYASAMIN, J.M. (2016). Rainbow trout (Oncorhynchus mykiss) threaten Andean amphibians. Neotropical Biodiversity, 2(1), 26–36. https://doi.org/10.1080/23766808.2016.1159075
    » https://doi.org/10.1080/23766808.2016.1159075
  • MEHNER, T., IHLAU, J., DÖRNER, H., & HÖLKER, F. (2005). Can feeding of fish on terrestrial insects subsidize the nutrient pool of lakes?. Limnology and Oceanography, 50(6), 2022–2031.
  • MORABOWEN, A., CRESPO-PÉREZ, V., & RÍOS-TOUMA, B. (2019). Effects of agricultural landscapes and land uses in highly biodiverse tropical streams of the Ecuadorian Choco. Inland Waters, 9(3), 289–300.
  • PETRANKA, J.W., KATS, L.B., & SIH, A. (1987). Predator-prey interactions among fish and larval amphibians: Use of chemical cues to detect predatory fish. Animal Behaviour, 35(2), 420–425. https://doi.org/10.1016/S0003-3472(87)80072-5
    » https://doi.org/10.1016/S0003-3472(87)80072-5
  • REIS, R.E., KULLANDER, S.O., & FERRARIS, C.J. (Eds.). (2003). Checklist of the freshwater fishes of South and Central America. Porto Alegre: EDIPUCRS.
  • REIS, A.S., ALBRECHT, M.P., & BUNN, S.E. (2020). Food web pathways for fish communities in small tropical streams. Freshwater Biology, 65(5), 893–907. https://doi.org/10.1111/fwb.13491
    » https://doi.org/10.1111/fwb.13491
  • REMON, J., BOWER, D.S., GASTON, T.F., CLULOW, J., & MAHONY, M.J. (2016). Stable isotope analyses reveal predation on amphibians by a globally invasive fish (Gambusia holbrooki). Aquatic Conservation: Marine and Freshwater Ecosystems, 26(4), 724–735. https://doi.org/10.1002/aqc.2673
    » https://doi.org/10.1002/aqc.2673
  • ROMÁN-VALENCIA, C., RUIZ-C.R.I., TAPHORN, B.D., JIMÉNEZ-PRADO, P., & GARCÍA-ALZATE, C.A. (2015). A new species of Bryconamericus (Characiformes, Stevardiinae, Characidae) from the Pacific coast of northwestern Ecuador, South America. Animal Biodiversity and Conservation, 38(2), 241–252.
  • ROMÁN-VALENCIA, C. (2000). Bryconamericus dahli sp. n. (Pisces: Characidae) de la vertiente Caribe de Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 24(93), 469–474. https://revistas.accefyn.org.co/index.php/revista/article/view/900
    » https://revistas.accefyn.org.co/index.php/revista/article/view/900
  • SÁNCHEZ-GARCÉS, G.C. (2017). A review of amphidromous freshwater fishes of the Chocó biogeographical region (Colombia and Ecuador): diversity, ecology, fisheries and conservation. Cybium, 41(2), 157–169.
  • SCHAEFER, S.A., & FERNÁNDEZ, L. (2009). Relationships of trichomycterid catfishes (Teleostei: Siluriformes): evidence from morphology and molecules. In L. R. Malabarba, R. E. Reis, R. P. Vari, Z. M. Lucena & C. A. Lucena (Eds.), Phylogeny and classification of Neotropical fishes (pp. 337–357). Porto Alegre: EDIPUCRS.
  • SCHAEFER, S.A. (2003). Astroblepidae (Andean climbing catfishes). In R.E. Reis, S.O. Kullander & C.J. Ferraris Jr. (Eds.), Checklist of the freshwater fishes of South and Central America (pp. 312–313). Porto Alegre: EDIPUCRS.
  • SILVA, E.R., ROSAS, F.C.W., & ZUANON, J. (2014). Feeding ecology of the giant otter (Pteronura brasiliensis) and the Neotropical otter (Lontra longicaudis) in Jaú National Park, Amazon, Brazil. Journal of Natural History, 48(7–8), 465–479. https://doi.org/10.1080/00222933.2013.865429
    » https://doi.org/10.1080/00222933.2013.865429
  • SUTER, W. (1991). Effects of piscivorous birds on freshwater fish populations—a review. Journal of Ornithology, 132, 29–45. https://doi.org/10.1007/BF01643022
    » https://doi.org/10.1007/BF01643022
  • VARI, R.P. (1991). Systematics of the neotropical characiform genus Pseudochalceus (Pisces: Characiformes: Characidae). Proceedings of the Biological Society of Washington, 104(4), 776–793. https://www.biodiversitylibrary.org/page/34603498
    » https://www.biodiversitylibrary.org/page/34603498
  • VLACH, P., DUSEK, J., SVÁTORA, M., & MORAVEC, P. (2005). Fish assemblage structure, habitat and microhabitat preference of five fish species in a small stream. Folia Zoologica Praha, 54(4), 421–428.
  • WEITZMAN, S.H., & WEITZMAN, M.J. (2003). Lebiasinidae (Pencil fishes). In R.E. Reis, S.O. Kullander & C.J. Ferraris Jr. (Eds.), Checklist of the freshwater fishes of South and Central America (pp. 241–251). Porto Alegre: EDIPUCRS.
  • WIJKMARK, N., KULLANDER, S.O., & SALAZAR, R.E.B. (2012). Andinoacara blombergi, a new species from the río Esmeraldas basin in Ecuador and a review of A. rivulatus (Teleostei: Cichlidae). Ichthyological Exploration of Freshwaters, 23(2), 117–137.

Edited by

  • Associate Editor
    Rosana Mazzoni

Publication Dates

  • Publication in this collection
    08 Sept 2025
  • Date of issue
    2025

History

  • Received
    10 Feb 2025
  • Accepted
    25 July 2025
location_on
Instituto Virtual da Biodiversidade | BIOTA - FAPESP a/c Dr. José Augusto Salim, Rua Charles Darwin s/n -- Bloco M 13083-862, Campinas, São Paulo, Brasil., +55 (19) 35216168 - Campinas - SP - Brazil
E-mail: contato@biotaneotropica.org.br
rss_feed Acompanhe os números deste periódico no seu leitor de RSS
Ir para o topo Reportar erro