Abstract
The Yarinacocha lake is an emblematic ecosystem of the Peruvian Amazon, representing the main point of fish landing and a tourist attraction in the city of Pucallpa. The wide fauna diversity in this area has made it the target of various studies, although for fish species most of them were focused on commercial species. In this work, we carried out the first ichthyofauna species inventory of the Yarinacocha lake, sampling throughout the entire lake during the rainy and dry seasons and considering also all previously recorded species deposited in the Ichthyological Collection of the MUSM with the same locality. A total of 164 fish species were recorded, representing 10 orders (plus Eupercaria), 34 families and 116 genera. Characiformes was the order with more species (68 spp., 41.5%) followed by Siluriformes (59 spp., 36%), Cichliformes (17 spp., 10.5%), and Gymnotiformes (8 spp., 4.9%). The most highly represented families, including almost 55.5% of the total diversity obtained, were Characidae (23 spp., 14%), followed by Cichlidae (17 spp., 10.4%), Loricariidae (14 spp., 8.5%), Pimelodidae (13 spp., 7.9%), Doradidae (13 spp., 7.4%) and Anostomidae (12 spp., 7.3%). From the total fish species recorded in this study, only 22 are considered protected species following the IUCN criteria and 109 species have commercial importance, including 90 ornamental species (54.8%). Our results contribute to the knowledge of the ichthyofauna of the Yarinacocha lake and can be used as a starting point for its conservation and sustainable management over time.
Keywords
Amazon; Ichthyofauna; Inventory; Neotropical; South America
Resumo
O lago Yarinacocha é um ecossistema emblemático da Amazônia peruana, representando o principal ponto de desembarque de peixes e também uma atração turística da cidade de Pucallpa. A grande diversidade faunística nesta área a tornou alvo de vários estudos, embora para as espécies de peixes a maioria deles tenha sido focada em espécies comerciais. Neste trabalho, realizamos o primeiro inventário da ictiofauna do lago Yarinacocha, amostrando todo o lago durante as estações chuvosa e seca, e também considerando todas as espécies depositadas na Coleção Ictiológica do MUSM dessa localidade. Um total de 164 espécies de peixes foram registradas, representando 10 ordens (mais Eupercaria), 34 famílias e 116 gêneros. Characiformes foi a ordem com mais espécies (68 espécies, 41,5%), seguida por Siluriformes (59 espécies, 36%), Cichliformes (17 espécies, 10,5%) e Gymnotiformes (8 espécies, 4,9%). As famílias mais representadas, incluindo quase 55,5% da diversidade total obtida, foram Characidae (23 spp., 14%), seguido por Cichlidae (17 spp., 10,4%), Loricariidae (14 spp., 8,5%), Pimelodidae (13 spp., 7,9%), Doradidae (13 spp., 7,4%) e Anostomidae (12 spp., 7,3%). Do total de espécies de peixes registradas neste estudo, apenas 22 são consideradas espécies protegidas seguindo os critérios da IUCN e 109 espécies têm importância comercial, incluindo 90 espécies ornamentais (54,8%). Nossos resultados contribuem para o conhecimento da ictiofauna do lago Yarinacocha e podem ser utilizados como ponto de partida para sua conservação e manejo sustentável ao longo do tempo.
Palavras-chave
Amazonia; América do Sul; Ictiofauna; Inventário; Neotropical
Introduction
The Peruvian Amazonian ichthyofauna has been the object of numerous and in-depth studies, which register more than 800 species (Ortega et al. 2012ORTEGA, H., HIDALGO, M. TREVEJO, G., CORREA, E., CORTIJO, A.M., MEZA, V. & ESPINO, J. 2012. Lista anotada de los peces de aguas continentales del Perú: Estado actual del conocimiento, distribución, usos y aspectos de conservación. Segunda Edición. Ministerio del Ambiente, Dirección General de Diversidad Biológica - Museo de Historia Natural, UNMSM. p. 56.). Among lotic water bodies, the most important assessments include the Amazon (Goulding et al. 2003GOULDING, M., BARTHEM, R., FORSBERG, B., CAÑAS, C. & ORTEGA, H. 2003. Las fuentes del Amazonas: Ríos, vida y conservación de la cuenca del Madre de Dios. Asociación para la Conservación de la Cuenca Amazónica (ACCA)/Amazon Conservation Association (ACA). Gráfica Biblos S.A., Lima, Perú. 198.), Huallaga (Ortega et al. 2007ORTEGA, H., GUERRA, H. & RAMÍREZ, R. 2007. The Introduction of Non-native Fishes into Freshwater Systems of Peru, in: BERT T.M. (Ed.), Ecological and Genetic Implications of Aquaculture Activities. Methods and Technologies in Fish Biology and Fisheries, vol 6. Springer, Dordrecht, pp.247–278. https://doi.org/10.1007/978-1-4020-6148-6_14
https://doi.org/10.1007/978-1-4020-6148-...
), Madre de Dios (Barthem et al. 2003BARTHEM, R., GOULDING, M., FORSBERG, B., CAÑAS, C. & ORTEGA, H. 2003. Aquatic Ecology of the Río Madre de Dios. Lima: Biblos. 117 p.), Aguaytía (Quezada et al. 2017QUEZADA-GARCIA, M., HIDALGO, M., TARAZONA, J. & ORTEGA, H. 2017. Ictiofauna de la cuenca del río Aguaytía, Ucayali, Perú. Revista peruana de biología, 24(4):331–342.) and Las Piedras (Carvalho et al. 2011CARVALHO, T.P., ESPINO, J., MÁXIME, E., QUISPE, R., RENGIFO, B., ORTEGA, H. & ALBERT, J.S. 2011. Fishes from the lower Urubamba river near Sepahua, Amazon Basin, Peru. Check List, 7(4):413–442. http://dx.doi.org/10.15560/7.4.413
https://doi.org/10.15560/7.4.413...
) River basins. Likewise, the lentic aquatic ecosystems of the Peruvian Amazon play an important ecological role and contribute to the support of economic activities of the local population (García-Vásquez et al. 2009GARCÍA-DÁVILA, C., SÁNCHEZ, H., FLORES, M., MEJIA, J., ANGULO, C., CASTRO-RUIZ, D., ESTIVALS, G., GARCÍA-VÁSQUEZ, A., ALONSO, J., CARVAJAL, F., MOREAU, J., NUÑEZ, J., RENNO, J.F., TELLO, S., MONTREUIL, V. & DUPONCHELLE, F. 2009. Life-history characteristics of the large Amazonian migratory catfish Brachyplatystoma rousseauxii in the Iquitos region, Peru. Journal of Fish Biology, 75(10):2527–2551.) due to its value as a source of hydrobiological resources. This is partially the consequence of oxbows migration, which promotes this enormous diversity forming sinuous rivers and lakes during the course changes (Nagel et al. 2022NAGEL, G.W., NOVO, E.M.L.M., MARTIN, V.S., CAMPOS-SILVA, J.V., BARBOSA, C.C.F. & BONNET, M.P. 2022. Impacts of meander migration on the Amazon riverine communities using Landsat time series and cloud computing. Sci. Total Environ, 806, 150449. https://doi.org/10.1016/j.scitotenv.2021.150449
https://doi.org/10.1016/j.scitotenv.2021...
), and the flood pulses that comprise annual oscillations of the average water level (Junk et al. 1989JUNK, W.J., BAYLEY, P.B. & SPARKS, R.E. 1989 The flood pulse concept in river– floodplain systems. Special Publication of the Canadian Journal of Fisheries and Aquatic Sciences, 106:10–127.), allowing the interconnection of lakes and small water bodies during the floods (Bartletta et al. 2010BARLETTA, M., JAUREGUIZAR, A.J., BAIGUN, C., FONTOURA, N.F., AGOSTINHO, A.A., ALMEIDA-VAL, V.M.F, VAL, A.L., TORRES, R.A., JIMENES-SEGURA, L.F., GIARRIZZO, T., FABRÉ, N.N., BATISTA, V.S., LASSO, C., TAPHORN, D.C., COSTA, M.F., CHAVES, P.T., VIEIRA, J.P. & CORRÊA, M.F.M. 2010. Fish and aquatic habitat conservation in South America: a continental overview with emphasis on neotropical systems. Journal of Fish Biology, 76:2118–2176. https://doi.org/10.1111/j.1095-8649.2010.02684.x
https://doi.org/10.1111/j.1095-8649.2010...
).
The Yarinacocha lake, located in the district of Yarinacocha, department of Ucayali, known regionally as “cocha” or “tipishca” by shipibo-conibo populations, is approximately 20 km long, with an area of 13.4 km2, a maximum depth of 19 m and average width of 650 m (Campbell et al. 2017CAMPBELL, E., ALFARO-SHIGUETO, J., GODLEY, B. & MANGEL, J. 2017. Abundance estimate of the Amazon River Dolphin (Inia geoffrensis) and the tucuxi (Sotalia fluviatilis) in southern Ucayali, Peru. Latin American Journal of Aquatic Research. 45:957–969. 10.3856/vol45-issue5-fulltext-11
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). Its meandering origin formed by the erosion of Ucayali River banks (Neuendorf et al. 2005NEUENDORF, K.K.E., MEHL, J.P. & JACKSON, J.A. 2005. Glossary of Geology. 5 ed. Berlin, Heidelberg, New York: Springer-Verlag.) has allowed the occurrence of an enormous diversity, not yet fully monitored, but which supports important fishing activities in the region, local consumption and tourist destination (García-Dávila et al. 2018GARCÍA, G.A., VARGAS, G., NOLORBE, C., NÚÑEZ, J., MARIAC, C., DUPONCHELLE, F. & RENNO, J.F. 2018. Peces De Consumo de la Amazonía Peruana. Instituto de Investigaciones de la Amazonía Peruana (IIAP). Iquitos, Peru. 218.). During the floods of the Ucayali River in January, February and March, the entire lake acts as a riverside highway for trade and transportation (Campbell et al. 2017CAMPBELL, E., ALFARO-SHIGUETO, J., GODLEY, B. & MANGEL, J. 2017. Abundance estimate of the Amazon River Dolphin (Inia geoffrensis) and the tucuxi (Sotalia fluviatilis) in southern Ucayali, Peru. Latin American Journal of Aquatic Research. 45:957–969. 10.3856/vol45-issue5-fulltext-11
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). The rest of the year it is supplied with water from small tributaries that can be streams or “caños”, which are small channels that connect the lake with other water bodies and represents a key habitat for many species of fish reproduction (MINAM 2021MINAM (MINISTERIO DEL AMBIENTE). 2021. Línea base de los peces ornamentales con fines de bioseguridad en el Peru. https://bioseguridad.minam.gob.pe/wp-content/uploads/2021/12/ldb_pecesornam_2021.pdf
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). However, as a result of the human populations growth on the riverbanks (hamlets, communities and the population of the Yarinacocha district), restaurants and tourism, the use of toxic substances in illegal fishing and by hospital wastewater that fail biodiversity and human health (personal comments by José Riofrío), this ecosystem is highly polluted by sewage and solid waste that could affect biodiversity (Rondon-Espinoza et al. 2022RONDÓN-ESPINOZA, J., GAVIDIA, C., GONZÁLEZ, R. & RAMOS, D. 2022. Water Quality and Microbiological Contamination across the Fish Marketing Chain: A Case Study in the Peruvian Amazon (Lagoon Yarinacocha). Water. 14. 1465. https://doi.org/10.3390/w14091465
https://doi.org/10.3390/w14091465...
). These characteristics and threats as a whole, give Yarinacocha lake an emblematic meaning covering cultural, economic and ecological aspects worthy of being preserved and studied.
Scientific knowledge of the fish species composition at Yarinacocha lake is limited. A recent study reports that the diversity of fish species that arrives at the Yarinacocha lake landing stage, including a large part of the Utuquinía and Callería sub-basins, and to a lesser extent Loreto and Aguaytía-San Alejandro (Salazar-Ramirez et al. 2021SALAZAR-RAMÍREZ, L.E., RIOFRÍO-QUIJANDRÍA, J.C., ZAVALETA-FLORES, J.O. & RUBIO-RODRÍGUEZ, J.A. 2021. Análisis de los desembarques de la pesca comercial en Yarinacocha (Ucayali, Perú) entre 2015-2019. Revista de Investigaciones Veterinarias del Perú, 32(4):e20931. http://dx.doi.org/10.15381/rivep.v32i4.20931
https://doi.org/10.15381/rivep.v32i4.209...
). In their results, 63 species between 2015 and 2019 were reported, being the most abundant “carachama” (Loricariidae), “piro” (Doradidae), “palometa” (Mylossoma spp.) and “sardina” (Triportheus spp.) all of them recorded only as common names that may include several species. Other studies include monitoring of fishing landings in the city of Pucallpa (Vela et al. 2016) and characterization of commercial fisheries in the Ucayali River basin (Zorrilla et al. 2016), without offering precise data on the ichthyofauna of the lake. Also, these studies focus on large fishes with commercial interest, while neglecting the smaller species that contain the highest biomass (Barletta et al. 2010BARLETTA, M., JAUREGUIZAR, A.J., BAIGUN, C., FONTOURA, N.F., AGOSTINHO, A.A., ALMEIDA-VAL, V.M.F, VAL, A.L., TORRES, R.A., JIMENES-SEGURA, L.F., GIARRIZZO, T., FABRÉ, N.N., BATISTA, V.S., LASSO, C., TAPHORN, D.C., COSTA, M.F., CHAVES, P.T., VIEIRA, J.P. & CORRÊA, M.F.M. 2010. Fish and aquatic habitat conservation in South America: a continental overview with emphasis on neotropical systems. Journal of Fish Biology, 76:2118–2176. https://doi.org/10.1111/j.1095-8649.2010.02684.x
https://doi.org/10.1111/j.1095-8649.2010...
) and to date, no other publication has reported the total diversity of the ichthyofauna of the Yarinacocha lake.
This lack of monitoring and rapid identification tools for the fish fauna of Yarinacocha lake encourages the existence of erroneous records in fishing statistics (García-Dávila et al. 2018) and could be covering up the depredation of the fish populations.
For the above stated, we have formulated as the main purpose of this work to present the first fish species checklist of Yarinacocha Lake, updating the information of continental catalogs (Reis et al. 2003REIS, R.E., KULLANDER, S.O. & FERRARIS, JR. C.J. 2003. Check List of the Freshwater Fishes of South and Central America. Edipucrs, Porto Alegre, 742.), species threatened status (IUCN 2022IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org (last access in 01 Aug 2022).
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) and economic importance of species reported.
Material and Methods
A total of 15 sampling sites were evaluated (Fig. 1, Table 1), covering variable environments inside Yarinacocha lake including streams, water channels and shores (Fig. 2).
Sampling efforts occurred during the start of the rainy season in October 2020 and in the dry season in July 2022, mostly during daylight. The collection of specimens involved bottom trawls of 10 and 20 m nets with 5 mm internodes, used on the beaches and areas with vegetation, trunks and leaves; cast nets of 15 m in open areas and channels; and gillnets of 30 m, blocking the passage of fish during the beginning of the day. Obtained specimens were anesthetized in a clove oil solution, muscle tissues were fixed in 96% ethanol (for molecular studies), vouchers and other specimens were fixed in 10% formalin by 48 hours and then preserved in 70% ethanol. Expeditions had a Fish Collection Permit for research purposes under PRODUCE license number 132/2021.
Species identifications to the lowest taxonomic level were conducted consulting the taxonomic literature (Galvis et al. 2006GALVIS, G., MOJICA, J.I., DUQUE, S.R., CASTELLANOS, C., SÁNCHEZ-DUARTE, P., ARCE, M., GUTIÉRREZ, A., JIMÉNEZ, L.F., SANTOS, M., VEJARANO-RIVADENEIRA, S., ARBELÁEZ, F., PRIETO, E. & LEIVA, M. 2006. Peces del medio Amazonas. Región de Leticia. Serie de Guías Tropicales de Campo Nº 5. Conservación Internacional. Editorial Panamericana, Formas e Impresos. Bogotá, Colombia. 548., Queiroz et al. 2013QUEIROZ, L.J.D., VILARA, G.T., OHARA, W.M., PIRES, T.H.D.S., ZUANON, J.A.S. & DÓRIA, C.R. 2013. Peixes do Rio Madeira y Cuyari Pirá Ketá, vol. 1, 2, 3. 1172., van der Sleen & Albert 2018VAN DER SLEEN, P. & ALBERT, J.S. 2017. Field guide to the fishes of the Amazon, Orinoco and Guianas. Princeton University Press, Princeton, USA.) and identification keys for Characiformes (Géry, 1977GÉRY, J. 1977. Characoids of the World. TFH Publications, Neptune City. 672 pp.; Vari, 1991; Malabarba, 2004MALABARBA, M.C.S.L. 2004. Revision of the Neotropical genus Triportheus Cope, 1872 (Characiformes: Characidae). Neotropical Ichthyology 2:167–204. https://doi.org/10.1590/S1679-6225200400040000
https://doi.org/10.1590/S1679-6225200400...
), Siluriformes (Burgess, 1989BURGESS, W.E. 1989. An atlas of freshwater and marine catfishes. A preliminary survey of the Siluriformes. TFH Publication, Neptune City, Canada, 28. p. 305–325.; Albert, 2001ALBERT, J.S. 2001. Species diversity and phylogenetic systematics of American knifefishes (Gymnotiformes, Teleostei). Misc. Publ. Mus. Zool. University of Michigan 190:1–129.; Littmann et al. 2021LITTMANN, M.W., LUNDBERG, J.G. & ROCHA, M.S. 2021. Revision of the South American catfish genus Hypophthalmus (Siluriformes, Pimelodidae) with descriptions of two new species from the Amazon and Orinoco Basins. Proceedings of the Academy of Natural Sciences of Philadelphia, 167(1), 171–223.), Cichliformes (Kullander, 1986KULLANDER, S.O. 1986. Cichlid fishes of the Amazon River drainage of Peru. Department of Vertebrate Zoology, Research Division, Swedish Museum of Natural History, Stockholm, Sweden. 394.; Kullander and Ferreira, 2006KULLANDER, S.O. & FERREIRA, E.J.G. 2006. A review of the South American cichlid genus Cichla, with descriptions of nine new species (Teleostei: Cichlidae). Ichthyological Exploration of Freshwaters, 17:289–398.; Arbour et al. 2014ARBOUR, J.H., SALAZAR, R.E.B. & LÓPEZ-FERNÁNDEZ, H. 2014. A new species of Bujurquina (Teleostei: Cichlidae) from the río Danta, Ecuador, with a key to the species in the genus. Copeia:79–86.) and Gymnotiformes (Mago-Leccia, 1994MAGO-LECCIA, F. 1994. Electric Fishes of the Continental Waters of America. Peces Eléctricos de las Aguas Continentales de América. Biblioteca de la Academia de Ciencias Físicas, Matemáticas y Naturales. Vol XXIX. Caracas, Venezuela. 226.; Crampton et al. 2016CRAMPTON, W.G.R., DE SANTANA, C.D., WADDELL, J.C. & LOVEJOY, N.R. 2016. A taxonomic revision of the Neotropical electric fish genus Brachyhypopomus (Ostariophysi: Gymnotiformes: Hypopomidae), with descriptions of 15 new species. Neotropical Ichthyology, v. 14, n. 04:e150146. https://doi.org/10.1590/1982-0224-20150146
https://doi.org/10.1590/1982-0224-201501...
; de Santana et al. 2019DE SANTANA, C.D., CRAMPTON, W.G.R., DILLMAN, C.B., FREDERICO, R.G., SABAJ, M.H., COVAIN, R., READY, J., ZUANON, J., DE OLIVEIRA, R.R., MENDES-JÚNIOR, R.N., BASTOS, D.A., TEIXEIRA, T.F., MOL, J., OHARA, W., CASTRO e CASTRO, N., PEIXOTO, L.A., NAGAMACHI, C., SOUSA, L., MONTAG, L.F.A, RIBERIRO F., WADDELL, J.C., PIORSKY, N.M., VARI, R.P. & WOSIAKI, W.B. 2019. Unexpected species diversity in electric eels with a description of the strongest living bioelectricity generator. Nat Commun 10, 4000. https://doi.org/10.1038/s41467-019-11690-z
https://doi.org/10.1038/s41467-019-11690...
); and valid names were confirmed following Fricke et al. (2022)FRICKE, R., ESCHMEYER, W.N. & VAN DER LAAN, R. 2022. Catalog of fishes: genera, species, references. Calif. Acad. Sci.. Classification follows the current phylogenetic arrangement of bony fishes sensu according to Betancur et al. (2017)BETANCUR-R, R., WILEY, E.O., ARRATIA, G., ACERO, A., BAILLY, N., MIYA, M., LECOINTRE, G. & ORTÍ, G. 2017. Phylogenetic classification of bony fishes. BMC Evol Biol17, 162. https://doi.org/10.1186/s12862-017-0958-3
https://doi.org/10.1186/s12862-017-0958-...
and for Characiformes the classification proposed by Oliveira et al. (2011)OLIVEIRA, C., AVELINO, G.S., ABE, K.T., MARIGUELA, T.C., BENINE, R.C., ORTÍ, G., VARI, R.P. & CORRÊA E CASTRO, R.M. 2011. Phylogenetic relationships within the speciose family Characidae (Teleostei: Ostariophysi: Characiformes) based on multilocus analysis and extensive ingroup sampling. BMC evolutionary biology, 11(1):1–25. https://doi.org/10.1186/1471-2148-11-275
https://doi.org/10.1186/1471-2148-11-275...
. To corroborate geographical distribution of species we used CLOFFSCA (Reis et al. 2003REIS, R.E., KULLANDER, S.O. & FERRARIS, JR. C.J. 2003. Check List of the Freshwater Fishes of South and Central America. Edipucrs, Porto Alegre, 742.), digital platforms like SpeciesLink (http://www.splink.org.br/) and FishNet2 (http://www.fishnet2.net/) and Nijssen and Isbrücker (1986) for additional taxonomic revisions and species descriptions.Vouchers were deposited in the Ichthyology Collection of Museo de Historia Natural of the Universidad Nacional Mayor de San Marcos, Lima, Peru (MUSM).
Specimens collected in the Yarinacocha lake by previous expeditions and deposited in the MUSM Fish Collection, were also recorded after their identifications were verified. In our results, only native species were included without considering invasive or exotic species because they lacked vouchers in the scientific collection; however, its presence was discussed due to previous reports from the Ministerio del Ambiente (MINAM).
The commercial species were classified into ornamental fish (IIAP 2011IIAP. 2011. Peces ornamentales amazónicos: catálogo 2011. Instituto de Investigaciones de la Amazonía Peruana, Iquitos, Peru. 218. https://hdl.handle.net/20.500.12921/138
https://hdl.handle.net/20.500.12921/138...
, García-Dávila et al. 2020GARCÍA-DÁVILA, C., ESTIVALS, G., MEJIA, J., FLORES, M., ANGULO, C., SÁNCHEZ, H., NOLORBE C., CHUQUIPIONDO, C., CASTRO-RUIZ, D., GARCÍA, A., ORTEGA, H., PINEDO, L., MARIAC, C., DUPONCHELLE, F., OLIVEIRA, C., RÖMER, U. & RENNO J.F. 2020. Peces ornamentales de la Amazonia Peruana. Instituto de Investigaciones de la Amazonía Peruana (IIAP). Iquitos, Peru. 503 pp.) and fish for consumption (García-Dávila et al. 2018). The threatened status of the species was derived from IUCN (2022IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org (last access in 01 Aug 2022).
https://www.iucnredlist.org...
) criteria available in https://www.iucnredlist.org/.
Results and Discussion
The total of previously reported species is 63 (Vela et al. 2016; Zorrilla et al. 2016; Salazar-Ramirez et al. 2021) reporting only Characiformes and Siluriformes. However, as indicated above, those studies did not include precise data on the location of capture, making impossible a comparison of the total richness previously reported in relation to the results presented here which only include fishes from Yarinacocha lake.
Our contribution carries a very high value due to the fact of representing the first ichthyofauna inventory of Yarinacocha lake in its entire extension, assessing a wide range of habitats, including almost all possible species, not only those with commercial value (Fig. 3). In this manner, a total 164 fish species were recorded (Table 2) representing 10 orders (plus Eupercaria), 34 families and 116 genera.
Some fish species collected in Yarinacocha lake, Ucayali basin, Peru. Scale bar 1 cm. A) Abramites hypselonotus, B) Schizodon fasciatus, C) Leporinus cf. parae, D) Aphyocharax pusillus, E) Ctenobrycon hauxwellianus, F) Tetragonopterus argenteus, G) Hydrolycus scomberoides, H) Prochilodus nigricans, I) Hoplias malabaricus, J) Pygocentrus nattereri, K) Adontosternarchus balaenops, L) Crenicichla proteus, M) Heros efasciatus, N) Cichla monoculus, O) Pterophyllum scalare, P) Mesonauta mirificus, Q) Satanoperca jurupari, R) Hypoclinemus mentalis, S) Corydoras multiradiatus, T) Oxydoras niger, U) Trachelyopterus galeatus, V) Hemiodontichthys acipenserinus, W) Sorubim lima, X) Pseudoplatystoma punctifer, Y) Synbranchus marmoratus.
Fish fauna recorded for the Yarinacocha lake. The list includes the Peruvian common name, economic importance of the species as ornamental fish (O) according to IIAP (2011IIAP. 2011. Peces ornamentales amazónicos: catálogo 2011. Instituto de Investigaciones de la Amazonía Peruana, Iquitos, Peru. 218. https://hdl.handle.net/20.500.12921/138
https://hdl.handle.net/20.500.12921/138... ) and García Dávila et al. (2021), or fish for consumption (C) according to García-Dávila et al. (2018), IUCN category (LC = Least Concern, DD = Data Deficient, NT = Near Threatened, CR = Critically Endangered, EN = Endangered, VU = Vulnerable). In addition, the catalog number of the voucher specimens deposited in the MUSM fish collections is also being considered.
The order with the most species richness was Characiformes with 68 species (41.5%) followed by Siluriformes with 59 species (36%), Cichliformes with 17 species (10.5%) and Gymnotiformes with eight species (4.9%). These orders represent 92.6% of the total species (Fig. 4). Clupeiformes, with four species, and remaining five orders (plus Eupercaria), with one or two species for each, represent 7.4% of total species. The most highly represented family was Characidae with 23 species (14%), followed by Cichlidae with 17 species (10.4%), Loricariidae with 14 species (8.5%), Pimelodidae with 13 species (7.9%), Doradidae with 13 species (7.4%) and Anostomidae with 12 species (7.3%); together represent 55.5% of the total species (Fig. 5). As in the Neotropical freshwater habitats, the ichthyofauna belongs to the Ostariophysi and in South America mostly represented by Characiformes, Siluriformes and Gymnotiformes (Reis et al. 2016REIS, R.E., ALBERT, J.S., DI DARIO, F., MINCARONE, M.M., PETRY, P. & ROCHA, L.A. 2016. Fish biodiversity and conservation in South America. J Fish Biol, 89:12–47. https://doi.org/10.1111/jfb.13016
https://doi.org/10.1111/jfb.13016...
). In the Peruvian Amazon, the families Characidae, Loricariidae y Cichlidae, concentrate the greatest diversity of species (Ortega et al. 2012ORTEGA, H., HIDALGO, M. TREVEJO, G., CORREA, E., CORTIJO, A.M., MEZA, V. & ESPINO, J. 2012. Lista anotada de los peces de aguas continentales del Perú: Estado actual del conocimiento, distribución, usos y aspectos de conservación. Segunda Edición. Ministerio del Ambiente, Dirección General de Diversidad Biológica - Museo de Historia Natural, UNMSM. p. 56.) and represent some of the dominant families in the composition of Amazonian species (Dagosta & de Pinna 2019DAGOSTA, F. & DE PINNA, M. 2019. The Fishes of the Amazon: Distribution and Biogeographical Patterns, with a Comprehensive List of Species. Bulletin of the American Museum of Natural History. 431. 1. https://doi.org/10.1206/0003-0090.431.1.1
https://doi.org/10.1206/0003-0090.431.1....
).
Another interesting novelty is Leporinus subniger, with distribution in the Upper Amazon Basin (Colombia and Ecuador) (Britski & Birindelli 2008BRITSKI, H.A. & BIRINDELLI, J.L.O. 2008. Description of a new species of the genus Leporinus Spix (Characiformes: Anostomidae) from the rio Araguaia, Brazil, with comments on the taxonomy and distribution of L. parae and L. lacustris. Neotropical Ichthyology, (1):45–51. https://doi.org/10.1590/S1679-62252008000100005
https://doi.org/10.1590/S1679-6225200800...
), species not previously reported by previous peruvian inventories (Ortega et al. 2012ORTEGA, H., HIDALGO, M. TREVEJO, G., CORREA, E., CORTIJO, A.M., MEZA, V. & ESPINO, J. 2012. Lista anotada de los peces de aguas continentales del Perú: Estado actual del conocimiento, distribución, usos y aspectos de conservación. Segunda Edición. Ministerio del Ambiente, Dirección General de Diversidad Biológica - Museo de Historia Natural, UNMSM. p. 56., Meza-Vargas et al. 2021MEZA-VARGAS, V., FAUSTINO-FUSTER, D.R., CHUCTAYA, J., HIDALGO, M. & ORTEGA TORRES H. 2021. Lista de especies de peces de agua dulce de Loreto, Perú. Revista Peruana de Biología 28 (especial): e21911. https://doi.org/10.15381/rpb.v28iespecial.21911
https://doi.org/10.15381/rpb.v28iespecia...
, Chuctaya et al. 2022CHUCTAYA, J., MEZA-VARGAS, V., FAUSTINO-FUSTER, D.R., HIDALGO, M. & ORTEGA, H. 2022. Lista De Especies De Peces De La Cuenca Del Río Ucayali, Perú. Revista Peruana De Biología 29 (4):e20049. https://doi.org/10.15381/rpb.v29i4.20049
https://doi.org/10.15381/rpb.v29i4.20049...
) representing a new record for the department of Ucayali.
The following species could not be identified to the species level Hemiodus aff. microlepis belongs to the Hemiodus microlepis species group, previously reported for the Ucayali River, being a species not formally described (Nogueira et al. 2021NOGUEIRA, A.F., OLIVEIRA, C., LANGEANI, F. & NETTO-FERREIRA, A.L. 2021. Overlooked biodiversity of mitochondrial lineages in Hemiodus (Ostariophysi, Characiformes). Zoologica Scripta, 337–351. https://doi.org/10.1111/zsc.12469
https://doi.org/10.1111/zsc.12469...
), Leporinus cf. parae, a member of the Leporinus friderici species complex (Silva-Santos et al. 2018SILVA-SANTOS, R., RAMIREZ, J.L., GALETTI, JR. P.M. & FREITAS, P.D. 2018. Molecular Evidences of a Hidden Complex Scenario in Leporinus cf. friderici. Frontiers in Genetics, 9:1–9. https://doi.org/10.3389/fgene.2018.00047
https://doi.org/10.3389/fgene.2018.00047...
), and Leporinus aff. amazonicus represents putative undescribed species. Leporinus is the most species-rich genus within Anostomidae and is considered one of the richest genera within Characiformes (Garavello and Britski 2003GARAVELLO, J.C. & BRITSKI, H.A. 2003. Family Anostomidae. In Check List of the Freshwater Fishes of South and Central America (R.E. Reis, S.O. Kullander, C.J. Ferraris Jr., eds). Porto Alegre: EDIPUCRS, p. 71–84., Ramirez et al. 2016RAMIREZ, J.L., CARVALHO-COSTA, L.F., VENERE, P.C., CARVALHO, D.C., TROY, W.P. & GALETTI, P.M. 2016. Testing monophyly of the freshwater fish Leporinus (Characiformes. Anostomidae) through molecular analysis. J. Fish Biol. 88:1204–1214. doi: https://doi.org/10.1111/jfb.12906
https://doi.org/10.1111/jfb.12906...
) and a thorough taxonomic review of this group is needed (Silva-Santos et al. 2018SILVA-SANTOS, R., RAMIREZ, J.L., GALETTI, JR. P.M. & FREITAS, P.D. 2018. Molecular Evidences of a Hidden Complex Scenario in Leporinus cf. friderici. Frontiers in Genetics, 9:1–9. https://doi.org/10.3389/fgene.2018.00047
https://doi.org/10.3389/fgene.2018.00047...
). Rineloricaria sp. could only be identified to the genus level without coinciding with R. morrowi or R. wolfei, the two species described for the region by Fowler (1940), further studies will be needed to classify these specimens. Moenkhausia aff. dichroura belongs to the M. dichroura species group, because it’s distributed only in the La Plata basin (Paraguay and lower Paraná rivers) and the morphotype found in the Amazon basin would be a possible new species (Britzke 2011BRITZKE, R. 2011. Revisão taxonômica do grupo Moenkhausia dichroura (Kner, 1858) (Characiformes: Characidae). Msc Thesis, Universidade Estadual Paulista, Botucatu, São Paulo, Brasil.). Several species have the same color pattern, such as M. dichroura, M. intermedia, M. barbouri, M. bonita and also the species Schultzites axelrodi, which in some cases may make it difficult to correctly identify this species group.
Regarding the species of economic importance, 109 species have commercial importance, of which 90 species (54.8%) are ornamentals, 26 species are used for both purposes and 20 species only for consumption, including Prochilodus nigricans (boquichico) which contributes the highest percentage of catches (Riofrío 1998, Riofrío 2002RIOFRÍO, J.C. 2002. Aspectos biométricos y reproductivos de boquichico Prochilodus nigricans Agassiz, 1829 (Pisces: Prochilodontidae) en Ucayali, Perú. Revista Peruana de Biología, 9(2):111–115. https://doi.org/10.15381/rpb.v9i2.2529.
https://doi.org/10.15381/rpb.v9i2.2529...
, Wasiw et al. 2012WASIW, J., RIOFRIO, J. & MUÑOZ, V. 2012. Monitoreo de la pesquería comercial en Pucallpa y Yarinacocha, 2010 (Ucayali – Perú). Informe IMARPE, 39(3-4):288–293. https://hdl.handle.net/20.500.12958/2235
https://hdl.handle.net/20.500.12958/2235...
, Salazar-Ramirez et al. 2021) and large migratory catfish, with high commercial value, such as Pseudoplatystoma punctifer (doncella) and Phractocephalus hemioliopterus (peje torre), whose presence indicates the importance of this ecosystem in its migratory route, since this type of whitewater ecosystems are the most important for commercial migratory species (Duponchelle et al. 2021DUPONCHELLE, F., ISAAC, V.J., RODRIGUES DA COSTA DORIA, C., VAN DAMME, P.A., HERRERA-R, G.A., ANDERSON, E.P., CRUZ, R.E.A., HAUSER, M., HERMANN, T., AGUDELO, E., BONILLA-CASTILLO, C., BARTHEM, R., FREITAS, C.E.C., GARCÍA-DÁVILA, C., GARCÍA-VASQUEZ, A., RENNO, J.F. & CASTELLO, L. 2021. Conservation of migratory fishes in the Amazon basin. Aquatic Conservation: Mar Freshw Ecosyst, 31: 1087–1105. https://doi.org/10.1002/aqc.3550
https://doi.org/10.1002/aqc.3550...
). Therefore, the majority of species recorded for Yarinacocha lake (66.7%) represent important economic resources. Nevertheless, the abundances of these resources can decline due to human impacts including habitat alteration, water pollution, overfishing, exotic species introduction and other factors (van der Sleen & Albert 2021).
The reduction in landings in Yarinacocha lake has been reported in recent years (Salazar-Ramirez et al. 2021). Species of consumption of high commercial demand such as Colossoma macropomum (gamitana) and Piaractus brachypomus (paco) have almost disappeared from landings, being replaced by small-sized species (Riofrío 1998, Wasiw et al. 2012WASIW, J., RIOFRIO, J. & MUÑOZ, V. 2012. Monitoreo de la pesquería comercial en Pucallpa y Yarinacocha, 2010 (Ucayali – Perú). Informe IMARPE, 39(3-4):288–293. https://hdl.handle.net/20.500.12958/2235
https://hdl.handle.net/20.500.12958/2235...
, Salazar-Ramirez et al. 2021). This evidences the decrease of fish populations for consumption due to overexploitation.
The global ornamental fish industry is a market expansion grid, and moves approximately US$ 15 billion/year, including equipment, accessories, supplies and publications (Cheong 1996CHEONG, L. 1996. Overview of the current international trade in ornamental fish, with special reference to Singapore. Revue scientifique et technique (International Office of Epizootics), 15(2):445–481.). There are 350 to 400 million ornamental fish sold, where 70% are produced in captivity and 63% are exported by developing countries generating about U.S. $ 202 million/year (Chao et al. 2001CHAO, N.L. 2001. Fisheries, diversity and conservation of ornamental fish of the Rio Negro River, Brazil- a review of Project Piaba (1989–99). 2001. In Conservation and Management of Ornamental Fish Resources of the Rio Negro Basin, Amazonia, Brazil- Project Piaba (L.N. Chao, P. Petry, G. Prang, L. Sonneschein, & M. Tlusty, eds). Manaus: University of Amazonas Press, p. 161–204.). This market is dominated by freshwater fishes and the Amazon basin is a key supplier of wild freshwater fishes to the ornamental trade (Moreau and Coomes 2007MOREAU, M. & COOMES, O. 2007. Aquarium fish exploitation in western Amazonia: Conservation issues in Perú. Environmental Conservation, 34(1):12–22. doi:10.1017/S0376892907003566
https://doi.org/10.1017/S037689290700356...
). In the Peruvian Amazon, most of these resources are the product of direct extraction from the natural environment (MINAM 2021) and Yarinacocha lake is one of the leading regions for ornamental fish extraction in the Ucayali department (PNIPA 2021PNIPA (PROGRAMA NACIONAL DE INNOVACIÓN EN PESCA Y ACUICULTURA). 2021. Estudio de prospectiva: cadena de valor de peces ornamentales. https://hdl.handle.net/20.500.12864/302
https://hdl.handle.net/20.500.12864/302...
). Although the collection of fish from tropical lentic environments can be carried out almost throughout the entire year, this activity can lead to direct depletion of wild populations (Andrews 1990ANDREWS, C. 1990. The ornamental fish trade and fish conservation. Journal of Fish Biology, 37:53–59. https://doi.org/10.1111/j.1095-8649.1990.tb05020.x
https://doi.org/10.1111/j.1095-8649.1990...
).
In recent years, the global trade for freshwater specimens has led to the overexploitation of native species, and the destruction of these habitats by invasive species (Chang et al. 2009CHANG, A.L., GROSSMAN, J.D., SPEZIO, T.S., WEISKEL, H.W., BLUM, J.C., BURT, J.W., MUIR, A.A., PIOVIA-SCOTT, J., VEBLEN, K.E. & GROSHOLZ, E.D. 2009. Tackling aquatic invasions: risks and opportunities for the aquarium fish industry. Biological Invasions, 11(4):773–785.). An example of an exotic species found in Peru would be Trichopodus trichopterus (Pallas 1770), order Anabantiformes, family Osphronemidae, that was reported in Yarinacocha in 2016, in the La Restinga fishing area (MINAM 2021MINAM (MINISTERIO DEL AMBIENTE). 2021. Línea base de los peces ornamentales con fines de bioseguridad en el Peru. https://bioseguridad.minam.gob.pe/wp-content/uploads/2021/12/ldb_pecesornam_2021.pdf
https://bioseguridad.minam.gob.pe/wp-con...
). This species is a tropical freshwater fish native to Southeast Asia, known as the three spot gourami, and has a specialized organ (organ labyrinth) that allows them to breathe oxygen from the air (Blank & Burggren 2014BLANK, T. & BURGGREN, W. 2014. Hypoxia-induced developmental plasticity of the gills and air-breathing organ of Trichopodus trichopterus. J Fish Biol, 84:808–826. https://doi.org/10.1111/jfb.12319
https://doi.org/10.1111/jfb.12319...
), helping them to adapt to an environment of low oxygen concentration in the water (Degani et al. 2021DEGANI, G., VEKSLER-LUBLINSKY, I. & MEERSON, A. 2021. Markers of Genetic Variation in Blue Gourami (Trichogaster trichopterus) as a Model for Labyrinth Fish. Biology. 10(3):228. https://doi.org/10.3390/biology10030228
https://doi.org/10.3390/biology10030228...
). Specimens of this group were introduced in 1970 for ornamentation (Ortega et al. 2007ORTEGA, H., GUERRA, H. & RAMÍREZ, R. 2007. The Introduction of Non-native Fishes into Freshwater Systems of Peru, in: BERT T.M. (Ed.), Ecological and Genetic Implications of Aquaculture Activities. Methods and Technologies in Fish Biology and Fisheries, vol 6. Springer, Dordrecht, pp.247–278. https://doi.org/10.1007/978-1-4020-6148-6_14
https://doi.org/10.1007/978-1-4020-6148-...
) and have been reported for the city of Iquitos (department of Loreto, Peru) (Meza-Vargas et al. 2021). The effects of exotic species on lentic water bodies can be complex (Ortega and Hidalgo 2008ORTEGA, H. & HIDALGO, M. 2008. Health & Management. Freshwater fishes and aquatic habitats in Peru: Current knowledge and conservation. Aquatic Ecosystem doi: https://doi.org/10.1080/14634980802319135
https://doi.org/10.1080/1463498080231913...
), although the impact of T. trichopterus on native species is uncertain, the introduction of exotic species may imply a risk of co-introduction of parasites, especially with phylogenetically similar native fauna (Trujillo-González et al. 2018TRUJILLO-GONZÁLEZ, A., BECKER, J., VAUGHAN, D.B. & HUTSON, K.S. 2018. Monogenean parasites infect ornamental fish imported to Australia. Parasitology Research. 117:995–1011. https://doi.org/10.1007/s00436-018-5776-z
https://doi.org/10.1007/s00436-018-5776-...
), and generate problems of predation, competition for food and occupation of niches (Meza-Vargas et al. 2021). Therefore, T. trichopterus can be detrimental to native fish and further studies are required to assess the real impact on native species.
Freshwater fish may currently be one of the most threatened vertebrate groups, mainly based on the more than 5,000 species assessed by the IUCN, where the main threats include habitat modification, fragmentation and destruction; the introduction of invasive species; fisheries overexploitation; environment pollution; and climate change (Reid 2013REID, G.M. 2013. Introduction to freshwater fishes and their conservation. Int. Zoo. Yb. 47:1–5.). Thus, in the long term, it is estimated that 20% of the world’s freshwater fish should be in the vulnerable, threatened or extinct category in recent decades (Revenga et al. 1998REVENGA, C., MURRAY, S., ABRAMOVITZ, J. & HAMMOND, A. 1998. Watersheds of the World: Ecological Value and Vulnerability. Washington (DC) World Resources Institute, Washington, DC.).
In this study, 22 species are considered protected species (13.4%) following the IUCN criteria. One species, Arapaima gigas, was categorized as “Data deficient” (DD), therefore the knowledge about the biology and ecology of this species is limited, in particular the structure of its natural populations (Vitorino et al. 2017VITORINO, C.A., NOGUEIRA, F., SOUZA, I.L, ARARIPE, J. & VENERE, P.C. 2017. Low Genetic Diversity and Structuring of the Arapaima (Osteoglossiformes, Arapaimidae) Population of the Araguaia-Tocantins Basin.Front. Genet. 8:159. https://doi.org/10.3389/fgene.2017.00159
https://doi.org/10.3389/fgene.2017.00159...
) and further information on the species is required. The natural populations of this species have historically been reduced or even eradicated near the main cities (Castello et al. 2011CASTELLO, L., STEWART, D.J. & ARANTES, C.C. 2011. Modeling population dynamics and conservation of Arapaima in the Amazon. Reviews in Fish Biology and Fisheries, 21(3):623–640 https://doi.org/10.1007/s11160-010-9197-z
https://doi.org/10.1007/s11160-010-9197-...
). Currently, its landing in Yarinacocha lake is almost nil (Salazar-Ramírez et al. 2021SALAZAR-RAMÍREZ, L.E., RIOFRÍO-QUIJANDRÍA, J.C., ZAVALETA-FLORES, J.O. & RUBIO-RODRÍGUEZ, J.A. 2021. Análisis de los desembarques de la pesca comercial en Yarinacocha (Ucayali, Perú) entre 2015-2019. Revista de Investigaciones Veterinarias del Perú, 32(4):e20931. http://dx.doi.org/10.15381/rivep.v32i4.20931
https://doi.org/10.15381/rivep.v32i4.209...
). The other 21 species are considered “Least concern” (LC), which means that after being evaluated, it doesn’t meet any of the criteria that define the other categories. Most of the registered species lack information that allows them to be classified in some category, therefore further studies are necessary for an adequate evaluation of these species and of the Peruvian ichthyofauna in general.
Overall, Yarinacocha lake is an ecosystem with a high diversity of fish species that are valuable resources for the local people. Nonetheless, these resources are vulnerable to pollution, overexploitation and the presence of exotic species that affect native populations. Our study contributes to the knowledge of the ichthyofauna of the Yarinacocha lake and can be used as a starting point for its conservation and sustainable management over time.
Acknowledgments
We thank Roberto Quispe, Gian Pier Valenzuela, Ricardo López, Julio Ramirez, Flavio Lima for the help in the identification of some species and help in the field. Authors were supported by Concytec- Banco Mundial “Mejoramiento y Ampliación de los Servicios del Sistema Nacional de Ciencia Tecnología e Innovación Tecnológica” 8682-PE, through the Programa Nacional de Investigación Científica y Estudios Avanzados (ProCiencia) grants number 022-2019- FONDECYT-BM and 363-2019-FONDECYT.
Data Availability
Supporting data are available at: Britzke, Ricardo, 2022, “Fishes from Yarinacocha lake: an emblematic Amazonian ecosystem in the lower Ucayali River basin, Pucallpa, Peru”, https://doi.org/10.48331/scielodata.4FNJMY, SciELO Data, DRAFT VERSION.
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Publication Dates
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Publication in this collection
12 May 2023 -
Date of issue
2023
History
-
Received
03 Oct 2022 -
Accepted
28 Mar 2023