Accelerated Diversification Explains the Exceptional Species Richness of Tropical Characoid Fishes.

The Neotropics harbor the most species-rich freshwater fish fauna on the planet, but the timing of that exceptional diversification remains unclear. Did the Neotropics accumulate species steadily throughout their long history, or attain their remarkable diversity recently? Biologists have long debated the relative support for these museum and cradle hypotheses, but few phylogenies of megadiverse tropical clades have included sufficient taxa to distinguish between them. We used 1288 ultraconserved element loci spanning 293 species, 211 genera, and 21 families of characoid fishes to reconstruct a new, fossil-calibrated phylogeny and infer the most likely diversification scenario for a clade that includes a third of Neotropical fish diversity. This phylogeny implies paraphyly of the traditional delimitation of Characiformes because it resolves the largely Neotropical Characoidei as the sister lineage of Siluriformes (catfishes), rather than the African Citharinodei. Time-calibrated phylogenies indicate an ancient origin of major characoid lineages and reveal a much more recent emergence of most characoid species. Diversification rate analyses infer increased speciation and decreased extinction rates during the Oligocene at around 30 Ma during a period of mega-wetland formation in the proto-Orinoco-Amazonas. Three species-rich and ecomorphologically diverse lineages (Anostomidae, Serrasalmidae, and Characidae) that originated more than 60 Ma in the Paleocene experienced particularly notable bursts of Oligocene diversification and now account collectively for 68% of the approximately 2150 species of Characoidei. In addition to paleogeographic changes, we discuss potential accelerants of diversification in these three lineages. While the Neotropics accumulated a museum of ecomorphologically diverse characoid lineages long ago, this geologically dynamic region also cradled a much more recent birth of remarkable species-level diversity. [Biodiversity; Characiformes; macroevolution; Neotropics; phylogenomics; ultraconserved elements.].

The combined effects of climate change and river fragmentation on the distribution of Andean Amazon fishes.

Upstream range shifts of freshwater fishes have been documented in recent years due to ongoing climate change. River fragmentation by dams, presenting physical barriers, can limit the climatically induced spatial redistribution of fishes. Andean freshwater ecosystems in the Neotropical region are expected to be highly affected by these future disturbances. However, proper evaluations are still missing. Combining species distribution models and functional traits of Andean Amazon fishes, coupled with dam locations and climatic projections (2070s), we (a) evaluated the potential impacts of future climate on species ranges, (b) investigated the combined impact of river fragmentation and climate change and (c) tested the relationships between these impacts and species functional traits. Results show that climate change will induce range contraction for most of the Andean Amazon fish species, particularly those inhabiting highlands. Dams are not predicted to greatly limit future range shifts for most species (i.e., the Barrier effect). However, some of these barriers should prevent upstream shifts for a considerable number of species, reducing future potential diversity in some basins. River fragmentation is predicted to act jointly with climate change in promoting a considerable decrease in the probability of species to persist in the long-term because of splitting species ranges in smaller fragments (i.e., the Isolation effect). Benthic and fast-flowing water adapted species with hydrodynamic bodies are significantly associated with severe range contractions from climate change.

To see or not to see: molecular evolution of the rhodopsin visual pigment in neotropical electric fishes.

Functional variation in rhodopsin, the dim-light-specialized visual pigment, frequently occurs in species inhabiting light-limited environments. Variation in visual function can arise through two processes: relaxation of selection or adaptive evolution improving photon detection in a given environment. Here, we investigate the molecular evolution of rhodopsin in Gymnotiformes, an order of mostly nocturnal South American fishes that evolved sophisticated electrosensory capabilities. Our initial sequencing revealed a mutation associated with visual disease in humans. As these fishes are thought to have poor vision, this would be consistent with a possible sensory trade-off between the visual system and a novel electrosensory system. To investigate this, we surveyed rhodopsin from 147 gymnotiform species, spanning the order, and analysed patterns of molecular evolution. In contrast with our expectation, we detected strong selective constraint in gymnotiform rhodopsin, with rates of non-synonymous to synonymous substitutions lower in gymnotiforms than in other vertebrate lineages. In addition, we found evidence for positive selection on the branch leading to gymnotiforms and on a branch leading to a clade of deep-channel specialized gymnotiform species. We also found evidence that deleterious effects of a human disease-associated substitution are likely to be masked by epistatic substitutions at nearby sites. Our results suggest that rhodopsin remains an important component of the gymnotiform sensory system alongside electrolocation, and that photosensitivity of rhodopsin is well adapted for vision in dim-light environments.

Comparative phylogeography of trans-Andean freshwater fishes based on genome-wide nuclear and mitochondrial markers.

The Neotropical region represents one of the greatest biodiversity hot spots on earth. Despite its unparalleled biodiversity, regional comparative phylogeographic studies are still scarce, with most focusing on model clades (e.g. birds) and typically examining a handful of loci. Here, we apply a genome-wide comparative phylogeographic approach to test hypotheses of codiversification of freshwater fishes in the trans-Andean region. Using target capture methods, we examined exon data for over 1,000 loci combined with complete mitochondrial genomes to study the phylogeographic history of five primary fish species (>150 individuals) collected from eight major river basins in Northwestern South America and Lower Central America. To assess their patterns of genetic structure, we inferred genealogical concordance taking into account all major aspects of phylogeography (within loci, across multiple genes, across species and among biogeographic provinces). Based on phylogeographic concordance factors, we tested four a priori biogeographic hypotheses, finding support for three of them and uncovering a novel, unexpected pattern of codiversification. The four emerging inter-riverine patterns are as follows: (a) Tuira + Atrato, (b) Ranchería + Catatumbo, (c) Magdalena system and (d) Sinú + Atrato. These patterns are interpreted as shared responses to the complex uplifting and orogenic processes that modified or sundered watersheds, allowing codiversification and speciation over geological time. We also find evidence of cryptic speciation in one of the species examined and instances of mitochondrial introgression in others. These results help further our knowledge of the historical geographic factors shaping the outstanding biodiversity of the Neotropics.

Fishes (Actinopterygii) of the rapids and associated environments in the lower Vaupés River Basin: an undiscovered Colombian Amazon diversity.

The Vaupés River stands out as one of the few within the Amazon basin due to its numerous rapids. These riverine fast-flowing sections not only provide habitat to highly specialized fishes but also function as natural barriers hindering the movement of fish along its course. During a fish-collecting expedition in the lower Vaupés River basin in Colombia, 95 species were registered belonging to 30 families and seven orders. Despite recent inventories in the region, our comprehensive sampling efforts particularly focused on the rapids and associated rheophilic fauna, allowing us to contribute the first records of four fish species in Colombia (Mylopluslucienae Andrade, Ota, Bastos & Jégu, 2016, Tometesmakue Jégu, Santos & Jégu, 2002, also first record of the genus, Leptodoraspraelongus (Myers & Weitzman, 1956), and Eigenmanniamatintapereira Peixoto, Dutra & Wosiacki, 2015) and six presumably undescribed species (i.e., Jupiaba sp., Moenkhausia sp., Phenacogaster sp., Bunocephalus sp., Hemiancistrus sp., and Archolaemus sp.). In this study, a commented list of the ichthyofauna of these environments is presented, as well as a photographic catalog of fish species integrated into the CaVFish Project - Colombia.

A regional perspective on the diversity and conservation of tropical Andean fishes.

The tropical Andes harbor an extraordinarily varied concentration of species in a landscape under increasing pressure from human activities. Conservation of the region's native plants and animals has received considerable international attention, but the focus has been on terrestrial biota. The conservation of freshwater fauna, particularly the conservation of fishes, has not been emphasized. Tropical Andean fishes are among the most understudied vertebrates in the world. We estimate that between 400 and 600 fish species inhabit the diverse aquatic environments in the region. Nearly 40% of these species are endemic. Tropical Andean fishes are vulnerable to ongoing environmental changes related to deforestation, water withdrawals, water pollution, species introductions, and hydropower development. Additionally, their distributions and population dynamics may be affected by hydrologic alterations and warmer water temperatures associated with projected climate change. Presently, at least three species are considered extinct, some populations are endangered, and some species are likely to decline or disappear. The long-term persistence of tropical Andean fishes will depend on greater consideration of freshwater systems in regional conservation initiatives.

Applied science facilitates the large-scale expansion of protected areas in an Amazonian hot spot.

Meeting international commitments to protect 17% of terrestrial ecosystems worldwide will require >3 million square kilometers of new protected areas and strategies to create those areas in a way that respects local communities and land use. In 2000-2016, biological and social scientists worked to increase the protected proportion of Peru's largest department via 14 interdisciplinary inventories covering >9 million hectares of this megadiverse corner of the Amazon basin. In each landscape, the strategy was the same: convene diverse partners, identify biological and sociocultural assets, document residents' use of natural resources, and tailor the findings to the needs of decision-makers. Nine of the 14 landscapes have since been protected (5.7 million hectares of new protected areas), contributing to a quadrupling of conservation coverage in Loreto (from 6 to 23%). We outline the methods and enabling conditions most crucial for successfully applying similar campaigns elsewhere on Earth.

A database of freshwater fish species of the Amazon Basin.

The Amazon Basin is an unquestionable biodiversity hotspot, containing the highest freshwater biodiversity on earth and facing off a recent increase in anthropogenic threats. The current knowledge on the spatial distribution of the freshwater fish species is greatly deficient in this basin, preventing a comprehensive understanding of this hyper-diverse ecosystem as a whole. Filling this gap was the priority of a transnational collaborative project, i.e. the AmazonFish project - https://www.amazon-fish.com/. Relying on the outputs of this project, we provide the most complete fish species distribution records covering the whole Amazon drainage. The database, including 2,406 validated freshwater native fish species, 232,936 georeferenced records, results from an extensive survey of species distribution including 590 different sources (e.g. published articles, grey literature, online biodiversity databases and scientific collections from museums and universities worldwide) and field expeditions conducted during the project. This database, delivered at both georeferenced localities (21,500 localities) and sub-drainages grains (144 units), represents a highly valuable source of information for further studies on freshwater fish biodiversity, biogeography and conservation.

Photafish system: An affordable device for fish photography in the wild.

This paper describes the Photafish System, an innovative technique for capturing high-quality digital images of live freshwater fishes and other aquatic organisms in the field. This system may improve and facilitate the visual documentation of such organisms to illustrate guidebooks, populate online databases and support scientific papers, particularly taxonomic ones. The system was designed and tested to produce Taxonomically Informative Photos (TIPs) of living specimens and employs an Aquarium Assembly for Photography (AAP) distinguished by its portability and low cost. Using Neotropical freshwater fishes as a model, we propose an optimized protocol for documenting the morphology of live specimens to support taxonomy, as well as awareness and conservation of freshwater biodiversity. This paper addresses the technical aspects and challenges of field studio photography in remote locations. Tricks gleaned from personal experiences with nature photography are also shared. Finally, we briefly discuss the advantages and weaknesses of our approach.

Species delimitation of neotropical Characins (Stevardiinae): Implications for taxonomy of complex groups.

Accurate species delimitation is crucial for studies of phylogeny, phylogeography, ecology, conservation and biogeography. The limits of species and genera in the Characidae family are controversial due to its uncertain phylogenetic relationships, high level of morphological homoplasy and the use of ambiguous morphological characters for descriptions. Here we establish species boundaries for Bryconamericus, Hemibrycon, Knodus and Eretmobrycon (Stevardiinae: Characidae), previously diagnosed with morphology, using three different barcoding approaches (GMYC, PTP, ABGD). Results revealed that species delimitation was successful by the use of a single-gene approach and by following a workflow in the context of integrative taxonomy, making evident problems and mistakes in the cataloging of Characidae species. Hence, it was possible to infer boundaries at genus level for clusters in the trees (GMYC and PTP) and automatic partitions (ABGD) which were consistent with some of recent taxonomic changes proposed in Characidae. We found that discordance cases between methods were linked to limitations of the methods and associated to putative species cluster closely related, some historically problematic in their diagnosis and identification. Furthermore, we suggested taxonomic changes and possibly new species, revealing a high degree of hidden diversity. Finally, we propose a workflow as a fast, accurate and objective way to delimit species from mitochondrial DNA sequences and to help clarify the classification of this group.

Mitochondrial genomes of the South American electric knifefishes Eigenmannia humboldtii (Steindachner 1878), Eigenmannia limbata (Schreiner and Miranda Ribeiro 1903), Sternopygus aequilabiatus (Humboldt 1805) and Sternopygus macrurus (Bloch and Schneider 1801), (Gymnotiformes, Sternopygidae).

We report four mitochondrial genomes of South American electric knifefishes, derived from target capture and Illumina sequencing (HiSeq 2500 PE100). Two trans-Andean species Eigenmannia humboldtii (mitochondrial consensus genome of 25 individuals) and Sternopygus aequilabiatus (mitochondrial consensus genome of 30 individuals) from Colombia and two cis-Andean species Eigenmannia limbata from Suriname and Sternopygus macrurus from Argentina. Regarding Eigenmannia humboldtii, Eigenmannia limbata, and Sternopygus macrurus mitochondrial genomes have 13 protein-coding genes, 1 D-loop, 2 ribosomal RNAs, 22 transfer RNAs, and are 13,394 bp, 10,921 bp, and 13,013 bp in length respectively, for Sternopygus aequilabiatus mitochondrial genomes have 13 protein-coding genes, 2 ribosomal RNAs, 22 transfer RNAs, and is 14,270 bp in length.

Mitochondrial genomes of the headstanders Megaleporinus muyscorum (Steindachner 1900) and Megaleporinus obtusidens (Valenciennes 1837), (Characiformes, Anostomidae).

We report two mitochondrial genomes of headstanders, derived from target capture and Illumina sequencing (HiSeq 2500 PE100). One trans-Andean species Megaleporinus muyscorum (mitochondrial consensus genome of 25 individuals) from Colombia and one cis-Andean species M. obtusidens from Argentina. Regarding M. muyscorum, mitochondrial genome has 13 protein-coding genes, 1 D-loop, 2 ribosomal RNAs, 21 transfer RNAs, and is 14,434 bp in length, for M. obtusidens mitochondrial genome has 13 protein-coding genes, 2 ribosomal RNAs, 22 transfer RNAs, and is 15,546 bp in length.

Fishes of the Cusiana River (Meta River basin, Colombia), with an identification key to its species.

The Cusiana River sub-basin has been identified as a priority conservation area in the Orinoco region in Colombia due to its high species diversity. This study presents an updated checklist and identification key for fishes of the Cusiana River sub-basin. The checklist was assembled through direct examination of specimens deposited in the main Colombian ichthyological collections. A total of 2020 lots from 167 different localities from the Cusiana River sub-basin were examined and ranged from 153 to 2970 m in elevation. The highest number of records were from the piedmont region (1091, 54.0 %), followed by the Llanos (878, 43.5 %) and Andean (51, 2.5 %). 241 species distributed in 9 orders, 40 families, and 158 genera were found. The fish species richness observed (241), represents 77.7 % of the 314 estimated species (95 % CI=276.1-394.8). The use of databases to develop lists of fish species is not entirely reliable; therefore taxonomic verification of specimens in collections is essential. The results will facilitate comparisons with other sub-basins of the Orinoquia, which are not categorized as areas of importance for conservation in Colombia.

ResumenLa sub-cuenca del rio Cusiana ha sido designada como una de las áreas prioritarias para la conservación en la región del Orinoco en Colombia debido a su alta diversidad de especies. Este estudio presenta una lista actualizada y una clave de identificación para los peces del área. Para ello se revisaron los especímenes depositados en las principales colecciones ictiológicas colombianas. Se examinaron un total de 2020 lotes de 167 localidades diferentes de la sub-cuenca del río Cusiana, que oscilaron entre 153 y 2970 m en altitud. El mayor número de registros corresponde a la región del piedemonte (1091, 54.0%), seguida por los Llanos (878, 43.5%) y Andina (51, 2.5%). Encontramos 241 especies distribuidas en 9 órdenes, 40 familias y 158 géneros. La riqueza de especies de peces observada (241), representan el 77.7% de las 314 especies estimadas (IC 95% = 276.1-394.8). El uso de bases de datos para elaborar listas de especies de peces no es del todo fiable, por lo tanto, es esencial la verificación taxonómica de los especímenes en las colecciones. Estos resultados permitirán evaluar otras sub-cuencas de la Orinoquia, las cuales no están categorizadas como áreas de importancia para la conservación en Colombia.

Fragmentation of Andes-to-Amazon connectivity by hydropower dams.

Andes-to-Amazon river connectivity controls numerous natural and human systems in the greater Amazon. However, it is being rapidly altered by a wave of new hydropower development, the impacts of which have been previously underestimated. We document 142 dams existing or under construction and 160 proposed dams for rivers draining the Andean headwaters of the Amazon. Existing dams have fragmented the tributary networks of six of eight major Andean Amazon river basins. Proposed dams could result in significant losses in river connectivity in river mainstems of five of eight major systems-the Napo, Marañón, Ucayali, Beni, and Mamoré. With a newly reported 671 freshwater fish species inhabiting the Andean headwaters of the Amazon (>500 m), dams threaten previously unrecognized biodiversity, particularly among endemic and migratory species. Because Andean rivers contribute most of the sediment in the mainstem Amazon, losses in river connectivity translate to drastic alteration of river channel and floodplain geomorphology and associated ecosystem services.

Checklist of the freshwater fishes of Colombia: a Darwin Core alternative to the updating problem.

The present work is part of a process to create a Catalogue of the Freshwater Fishes of Colombia and consisted in the depuration and updating of the taxonomic and geographic components of the checklist of the freshwater fishes of Colombia. An exhaustive revision of the 1435 species recorded in 2008 was necessary to: 1. Add new species described since 2009 and species originally described from Colombia but inadvertently omitted in 2008; 2. Add new records of already described species; 3. Delete species whose presence in Colombia was not supported by voucher specimens in ichthyological collections; and 4. Revise the geographic distribution of the species listed in 2008. This process resulted in the following numbers: 1. Total number of freshwater fish species in Colombia: 1494; 2. Number of species recorded by hydrographic region - Amazon: 706, Orinoco: 663, Caribbean: 223, Magdalena-Cauca: 220, Pacific: 130; and 3. Number of endemic species: 374 (76% from the trans-Andean region). Updating the current checklist is a fundamental requirement to ensure its incorporation in the decision-making process with regard to the conservation of Colombian aquatic species and ecosystems, which are facing transformation processes as a result of activities such as mining, construction of hydroelectric plants, expansion of the agricultural frontier and subsequent deforestation, industrial and domestic pollution, development of waterways, introduction of exotic species, and climate change.

ResumenEl presente trabajo es parte de un proceso para crear un Catálogo de Peces de Agua Dulce de Colombia y consistió en la depuración y actualización de los componentes taxonómico y geográfico del listado de peces de agua dulce de Colombia. Una revisión exhaustiva de las 1435 especies registradas en 2008 fue necesaria para: 1. Adicionar las especies nuevas descritas desde 2009 y especies descritas originalmente para Colombia pero omitidas inadvertidamente en 2008; 2. Adicionar nuevos registros de especies ya descritas; 3. Eliminar especies cuya presencia en Colombia no estaba soportada por especímenes en colecciones ictiológicas; y 4. Revisar la distribución geográfica de las especies listadas en 2008. Este proceso resultó en las siguientes cifras: 1. Número total de especies de peces de agua dulce en Colombia: 1494; 2. Número de especies registradas por región hidrográfica - Amazonas: 706, Orinoco: 663, Caribe: 223, Magdalena-Cauca: 220, Pacífico: 130; y 3. Número de especies endémicas 374 (76% distribuidas en la región transandina). La actualización del presente listado es un requerimiento fundamental para asegurar su incorporación en el proceso de toma de decisiones en lo concerniente a la conservación de las especies y ecosistemas acuáticos colombianos, los cuales están enfrentando procesos de transformación, como resultado de actividades como la minería, construcción de plantas hidroeléctricas, expansión de la frontera agropecuaria y subsecuente desforestación, polución industrial y doméstica, desarrollo de hidrovías, introducción de especies exóticas y cambio climático.

Description of a new species of the genus Chaetostoma from the Orinoco River drainage with comments on Chaetostoma milesi Fowler, 1941 (Siluriformes: Loricariidae).

Chaetostoma joropo n. sp. is described from the piedmont of the Orinoco River drainage in Colombia. The new species has been long confused with Chaetostoma milesi, a species with similar overall morphology and color pattern that is restricted to the Magdalena-Cauca River Basin. We diagnose the new species on the basis of morphology as well as a precise description of the color pattern. Chaetostoma joropo n. sp. is also easily distinguished from C. formosae the most similar species and other species inhabiting the Orinoco River drainage in Colombia. Data on ontogenetic variation and sexual dimorphism are provided, as well as natural history notes and remarks on the usage of the name Chaetostoma milesi for specimens from both the Orinoco and Magdalena-Cauca drainages. A discussion on the usage of the name Chaetostoma platyrhynchus is also provided given its current instability in the literature.

Complete mitochondrial genome sequence of Grundulus bogotensis (Humboldt, 1821).

The Grundulus bogotensis is an Endangered fish in Colombia. In this study, we report the complete mitochondrial DNA sequences of G. bogotensis. The entire genome comprised 17.123 bases and a GC content of 39.84%. The mitogenome sequence of G. bogotensis would contribute to better understand population genetics, and evolution of this lineage. Molecule was deposited at the GenBank database under the accession number KM677190.

Evolution of electric communication signals in the South American ghost knifefishes (Gymnotiformes: Apteronotidae): A phylogenetic comparative study using a sequence-based phylogeny.

The electric communication signals of weakly electric ghost knifefishes (Gymnotiformes: Apteronotidae) provide a valuable model system for understanding the evolution and physiology of behavior. Apteronotids produce continuous wave-type electric organ discharges (EODs) that are used for electrolocation and communication. The frequency and waveform of EODs, as well as the structure of transient EOD modulations (chirps), vary substantially across species. Understanding how these signals have evolved, however, has been hampered by the lack of a well-supported phylogeny for this family. We constructed a molecular phylogeny for the Apteronotidae by using sequence data from three genes (cytochrome c oxidase subunit 1, recombination activating gene 2, and cytochrome oxidase B) in 32 species representing 13 apteronotid genera. This phylogeny and an extensive database of apteronotid signals allowed us to examine signal evolution by using ancestral state reconstruction (ASR) and phylogenetic generalized least squares (PGLS) models. Our molecular phylogeny largely agrees with another recent sequence-based phylogeny and identified five robust apteronotid clades: (i) Sternarchorhamphus+Orthosternarchus, (ii) Adontosternarchus, (iii) Apteronotus+Parapteronotus, (iv) Sternarchorhynchus, and (v) a large clade including Porotergus, 'Apteronotus', Compsaraia, Sternarchogiton, Sternarchella, and Magosternarchus. We analyzed novel chirp recordings from two apteronotid species (Orthosternarchus tamandua and Sternarchorhynchus mormyrus), and combined data from these species with that from previously recorded species in our phylogenetic analyses. Some signal parameters in O. tamandua were plesiomorphic (e.g., low frequency EODs and chirps with little frequency modulation that nevertheless interrupt the EOD), suggesting that ultra-high frequency EODs and "big" chirps evolved after apteronotids diverged from other gymnotiforms. In contrast to previous studies, our PGLS analyses using the new phylogeny indicated the presence of phylogenetic signals in the relationships between some EOD and chirp parameters. The ASR demonstrated that most EOD and chirp parameters are evolutionarily labile and have often diversified even among closely related species.

Variation in freshwater fish assemblages along a regional elevation gradient in the northern Andes, Colombia.

Studies on elevation diversity gradients have covered a large number of taxa and regions throughout the world; however, studies of freshwater fish are scarce and restricted to examining their changes along a specific gradient. These studies have reported a monotonic decrease in species richness with increasing elevation, but ignore the high taxonomic differentiation of each headwater assemblage that may generate high ß-diversity among them. Here, we analyzed how fish assemblages vary with elevation among regional elevation bands, and how these changes are related to four environmental clines and to changes in the distribution, habitat use, and the morphology of fish species. Using a standardized field sampling technique, we assessed three different diversity and two structural assemblage measures across six regional elevation bands located in the northern Andes (Colombia). Each species was assigned to a functional group based on its body shape, habitat use, morphological, and/or behavioral adaptations. Additionally, at each sampling site, we measured four environmental variables. Our analyses showed: (1) After a monotonic decrease in species richness, we detected an increase in richness in the upper part of the gradient; (2) diversity patterns vary depending on the diversity measure used; (3) diversity patterns can be attributed to changes in species distribution and in the richness and proportions of functional groups along the regional elevation gradient; and (4) diversity patterns and changes in functional groups are highly correlated with variations in environmental variables, which also vary with elevation. These results suggest a novel pattern of variation in species richness with elevation: Species richness increases at the headwaters of the northern Andes owing to the cumulative number of endemic species there. This highlights the need for large-scale studies and has important implications for the aquatic conservation of the region.