New records of native and introduced fish species in a river basin of Western Ecuador, the Chocó-Darien Ecoregion, using DNA barcoding.

DNA barcoding, based on mitochondrial markers, is widely applied in species identification and biodiversity studies. The aim of this study was to establish a barcoding reference database of fishes inhabiting the Cube River from Western Ecuador in the Chocó-Darien Global Ecoregion (CGE), a threatened ecoregion with high diversity and endemism, and evaluate the applicability of using barcoding for the identification of fish species. Barcode sequences were obtained from seven orders, 17 families, 23 genera and 26 species, which were validated through phylogenetic analysis, morphological measurements, and literature review. Our results showed that 43% of fish species in this region are endemic, confirmed the presence of known species in the area, and included the addition of three new records of native (Hoplias microlepis, Rhamdia guatemalensis and Sicydium salvini) and an introduced species (Xiphophorus maculatus) to Ecuador. In addition, eight species were barcoded for the first time. Species identification based on barcoding and morphology showed discrepancy with species lists from previous studies in the CGE, suggesting that the current baseline of western fishes of Ecuador is still incomplete. Because this study analyzed fishes from a relatively small basin (165 km2), more molecular-based studies focusing on fish are needed to achieve a robust sequence reference library of species inhabiting Western Ecuador. The new sequences of this study will be useful for future comparisons and biodiversity monitoring, supporting the application of barcoding tools for studying fish diversity in genetically unexplored regions and to develop well-informed conservation programs.

Rediscovery of Rhyacoglanis pulcher (Boulenger, 1887) (Siluriformes: Pseudopimelodidae), a rare rheophilic bumblebee catfish from Ecuadorian Amazon.

Rhyacoglanis pulcher is a rare Neotropical rheophilic bumblebee catfish known only from the type locality in the Cis-Andean Amazon region, Ecuador, and the type-species of the genus. So far, the three syntypes collected in 1880 were the only specimens unambiguously associated to the name R. pulcher available in scientific collections. Recently, a specimen was discovered in a fast-flowing stretch of the Villano river, a tributary of the Curaray river, Napo river basin, Ecuador, representing a new record after nearly 140 years. Here, we present this new record, identified by morphology, provide the DNA barcode sequence of the specimen, and propose why the species of Rhyacoglanis are scarce in zoological collections. Additionally, we discuss the intraspecific variation in the color pattern observed in R. pulcher.

Validating the occurrence of the giant mottled eel (Anguilla marmorata) in the Galápagos Islands.

The giant mottled eel (Anguilla marmorata) is distributed mostly in the Indo-West Pacific. However, a few records indicate the presence of this eel in the Tropical Central and East Pacific. In April 2019, an eel specimen was caught in a small stream in San Cristobal Island, Galápagos. Morphological and molecular characters (16S and Cytb mtDNA sequences) confirmed the species as A. marmorata Quoy & Gaimard, 1824. The re-discovery of A. marmorata in Galápagos supports the hypothesis of an eastward range expansion from the west, probably through the North Equatorial Counter-Current.

Human impacts outpace natural processes in the Amazon.

Amazonian environments are being degraded by modern industrial and agricultural activities at a pace far above anything previously known, imperiling its vast biodiversity reserves and globally important ecosystem services. The most substantial threats come from regional deforestation, because of export market demands, and global climate change. The Amazon is currently perched to transition rapidly from a largely forested to a nonforested landscape. These changes are happening much too rapidly for Amazonian species, peoples, and ecosystems to respond adaptively. Policies to prevent the worst outcomes are known and must be enacted immediately. We now need political will and leadership to act on this information. To fail the Amazon is to fail the biosphere, and we fail to act at our peril.

Effects of experimental warming on two tropical Andean aquatic insects.

Temperatures have increased around the globe, affecting many ecosystems, including high-elevation Andean streams where important aquatic insect species coexist. Depending on the magnitude of change, warming could lead to the mortality of sensitive species, and those tolerant to rising water temperatures may exhibit differences in growth rates and development. Taxon-specific optimal temperature ranges for growth determine how high or low temperatures alter an organism's body size. In this study, we observed the effects of different climate change scenarios (following three scenarios of the 2021 IPCC predictions) in two aquatic insect species distributed in high-elevation streams in Ecuador: the mayfly Andesiops peruvianus (Ephemeroptera: Baetidae) and the caddisfly Anomalocosmoecus illiesi (Trichoptera: Limnephilidae). We assessed how increased water temperatures affect larval growth rates and mortality during a 10-day microcosm experiment. Our results showed that Andesiops peruvianus was more thermally sensitive than Anomalocosmoecus illiesi. Mortality was higher (more than 50% of the individuals) in mayflies than in caddisflies, which presented mortality below 12% at +2.5°C and +5°C. Mortality in mayflies was related to lower dissolved oxygen levels in increased temperature chambers. Higher temperatures affected body size and dry mass with a faster growth rate of Andesiops peruvianus larvae at experimentally higher temperatures, suggesting an important response of this hemimetabolous species to stream temperatures. For Anomalocosmoecus illiesi, we did not find significant changes in mortality, body size or growth rate in response to temperature changes during our experiment. In situ outcomes of species survival and growth in Andean streams are difficult to predict. Nevertheless, our results suggest that at only +2.5°C, a water temperature increase affected the two insect taxa differentially, leading to a drastic outcome for one species' larvae while selecting for a more tolerant species. Our study suggests that climate change might produce significant mortality and growth rate effects on ectotherm tropical aquatic insects, especially Andean mayflies, which showed higher sensitivity to increased water temperature scenarios.

Reducing adverse impacts of Amazon hydropower expansion.

Proposed hydropower dams at more than 350 sites throughout the Amazon require strategic evaluation of trade-offs between the numerous ecosystem services provided by Earth's largest and most biodiverse river basin. These services are spatially variable, hence collective impacts of newly built dams depend strongly on their configuration. We use multiobjective optimization to identify portfolios of sites that simultaneously minimize impacts on river flow, river connectivity, sediment transport, fish diversity, and greenhouse gas emissions while achieving energy production goals. We find that uncoordinated, dam-by-dam hydropower expansion has resulted in forgone ecosystem service benefits. Minimizing further damage from hydropower development requires considering diverse environmental impacts across the entire basin, as well as cooperation among Amazonian nations. Our findings offer a transferable model for the evaluation of hydropower expansion in transboundary basins.

Life history and secondary production of Anomalocosmoecusilliesi Marlier, 1962 (Trichoptera, Limnephilidae) in a small stream in the northern Ecuadorian Paramo.

Life history of benthic faunas of tropical high-altitude cold environments are poorly studied. Here, monthly larval and adult data are presented for Anomalocosmoecusilliesi at Saltana Stream in Ecuador. In cold conditions throughout the year (6 °C), this species showed an asynchronous and continuous production. Larval density showed two peaks in August and April. All five larval instars were present in most months. Using the size-frequency method an annual rate of secondary production per biomass of 4.8 was calculated. The measured biomass was 785 mg/m2.

Impacts of detritivore diversity loss on instream decomposition are greatest in the tropics.

The relationship between detritivore diversity and decomposition can provide information on how biogeochemical cycles are affected by ongoing rates of extinction, but such evidence has come mostly from local studies and microcosm experiments. We conducted a globally distributed experiment (38 streams across 23 countries in 6 continents) using standardised methods to test the hypothesis that detritivore diversity enhances litter decomposition in streams, to establish the role of other characteristics of detritivore assemblages (abundance, biomass and body size), and to determine how patterns vary across realms, biomes and climates. We observed a positive relationship between diversity and decomposition, strongest in tropical areas, and a key role of abundance and biomass at higher latitudes. Our results suggest that litter decomposition might be altered by detritivore extinctions, particularly in tropical areas, where detritivore diversity is already relatively low and some environmental stressors particularly prevalent.

Latitude dictates plant diversity effects on instream decomposition.

Running waters contribute substantially to global carbon fluxes through decomposition of terrestrial plant litter by aquatic microorganisms and detritivores. Diversity of this litter may influence instream decomposition globally in ways that are not yet understood. We investigated latitudinal differences in decomposition of litter mixtures of low and high functional diversity in 40 streams on 6 continents and spanning 113° of latitude. Despite important variability in our dataset, we found latitudinal differences in the effect of litter functional diversity on decomposition, which we explained as evolutionary adaptations of litter-consuming detritivores to resource availability. Specifically, a balanced diet effect appears to operate at lower latitudes versus a resource concentration effect at higher latitudes. The latitudinal pattern indicates that loss of plant functional diversity will have different consequences on carbon fluxes across the globe, with greater repercussions likely at low latitudes.

Temperature dependence of metabolic rate in tropical and temperate aquatic insects: Support for the Climate Variability Hypothesis in mayflies but not stoneflies.

A fundamental gap in climate change vulnerability research is an understanding of the relative thermal sensitivity of ectotherms. Aquatic insects are vital to stream ecosystem function and biodiversity but insufficiently studied with respect to their thermal physiology. With global temperatures rising at an unprecedented rate, it is imperative that we know how aquatic insects respond to increasing temperature and whether these responses vary among taxa, latitudes, and elevations. We evaluated the thermal sensitivity of standard metabolic rate in stream-dwelling baetid mayflies and perlid stoneflies across a ~2,000 m elevation gradient in the temperate Rocky Mountains in Colorado, USA, and the tropical Andes in Napo, Ecuador. We used temperature-controlled water baths and microrespirometry to estimate changes in oxygen consumption. Tropical mayflies generally exhibited greater thermal sensitivity in metabolism compared to temperate mayflies; tropical mayfly metabolic rates increased more rapidly with temperature and the insects more frequently exhibited behavioral signs of thermal stress. By contrast, temperate and tropical stoneflies did not clearly differ. Varied responses to temperature among baetid mayflies and perlid stoneflies may reflect differences in evolutionary history or ecological roles as herbivores and predators, respectively. Our results show that there is physiological variation across elevations and species and that low-elevation tropical mayflies may be especially imperiled by climate warming. Given such variation among species, broad generalizations about the vulnerability of tropical ectotherms should be made more cautiously.

New Ecuadorian records of the eyeless banjo catfish Micromyzon akamai (Siluriformes: Aspredinidae) expand the species range and reveal intraspecific morphological variation.

Two specimens of Micromyzon akamai, an eyeless and miniaturized species previously known only from the deep channels of the eastern Amazon basin in Brazil, are reported from the Curaray River, a tributary of the Napo River in Ecuador. The new specimens are the first records of Micromyzon in the headwaters of the Amazon River and the first records of M. akamai outside Brazil. External morphological characters and a phylogenetic analysis of cytochrome c oxidase I (coI) gene support the identification of the new specimens as M. akamai. Nevertheless, the new specimens also indicate that some features previously hypothesized to be apomorphic for M. akamai are intraspecifically variable.

Darwin Returns to the Galapagos: Genetic and Morphological Analyses Confirm the Presence of Tramea darwini at the Archipelago (Odonata, Libellulidae).

The status of the Tramea species present in the Galapagos Islands (Odonata, Libellulidae) has been the subject of a long-standing debate among odonatologists. Here, we use molecular and morphological data to analyze a series of specimens from this genus collected in 2018 from the Islands of San Cristobal, Isabela, and Santa Cruz, with the aim of determining their relationship with Tramea calverti Muttkowski and with their currently considered senior synonym T. cophysa Hagen. We combined sequencing of mitochondrial and nuclear DNA with morphological examination of several specimens of Tramea, including representatives of continental T. cophysa and T. calverti. Our molecular analyses place the Tramea from Galapagos in the same clade as T. calverti, with T. cophysa as a closely related species. The morphological analyses found only one consistent difference between T. cophysa and T. calverti: the presence of an accessory lobe in the male vesica spermalis of T. cophysa that is absent in T. calverti and in the Tramea from Galapagos. In agreement with our genetic results, the overall morphological differences documented by us indicate that the Galapagos material examined is conspecific with T. calverti. In light of this, and following the principle of priority in taxonomic nomenclature, Tramea calverti Muttkowski, 1910 should hereafter be considered a junior synonym of Tramea darwini Kirby, 1889.

Philogenia gaiae sp. nov. (Zygoptera: Philogeniidae) and description of the female of P. macuma Dunkle, 1986, two species from the Ecuadorean lowland rainforest.

Philogenia gaiae sp. nov. (Holotype ♂, Ecuador, Orellana, Tiputini Biodiversity Station, -0.6349, -76.1501, 241 m, 13 xii 2012, A. Cordero-Rivera M. Sánchez-Herrera leg., in MUAE) from the helena group is described, illustrated, diagnosed and compared with morphologically close species of the genus. Philogenia gaiae can be distinguished from its most similar congener P. minteri Dunkle, 1986 by the enlarged cerci and the club-like paraprocts. We also describe the female of P. macuma Dunkle, 1986, from a pair collected at Jatún Sacha Biological Reserve, which is also similar to P. gaiae and can be distinguished by the intersternite morphology, ovipositor length and vulvar lamina shape.

Validating anthropogenic threat maps as a tool for assessing river ecological integrity in Andean-Amazon basins.

Anthropogenic threat maps are commonly used as a surrogate for the ecological integrity of rivers in freshwater conservation, but a clearer understanding of their relationships is required to develop proper management plans at large scales. Here, we developed and validated empirical models that link the ecological integrity of rivers to threat maps in a large, heterogeneous and biodiverse Andean-Amazon watershed. Through fieldwork, we recorded data on aquatic invertebrate community composition, habitat quality, and physical-chemical parameters to calculate the ecological integrity of 140 streams/rivers across the basin. Simultaneously, we generated maps that describe the location, extent, and magnitude of impact of nine anthropogenic threats to freshwater systems in the basin. Through seven-fold cross-validation procedure, we found that regression models based on anthropogenic threats alone have limited power for predicting the ecological integrity of rivers. However, the prediction accuracy improved when environmental predictors (slope and elevation) were included, and more so when the predictions were carried out at a coarser scale, such as microbasins. Moreover, anthropogenic threats that amplify the incidence of other pressures (roads, human settlements and oil activities) are the most relevant predictors of ecological integrity. We concluded that threat maps can offer an overall picture of the ecological integrity pattern of the basin, becoming a useful tool for broad-scale conservation planning for freshwater ecosystems. While it is always advisable to have finer scale in situ measurements of ecological integrity, our study shows that threat maps provide fast and cost-effective results, which so often are needed for pressing management and conservation actions.

A global perspective on tropical montane rivers.

Tropical montane rivers (TMR) are born in tropical mountains, descend through montane forests, and feed major rivers, floodplains, and oceans. They are characterized by rapid temperature clines and varied flow disturbance regimes, both of which promote habitat heterogeneity, high biological diversity and endemism, and distinct organisms' life-history adaptations. Production, transport, and processing of sediments, nutrients, and carbon are key ecosystem processes connecting high-elevation streams with lowland floodplains, in turn influencing soil fertility and biotic productivity downstream. TMR provide key ecosystem services to hundreds of millions of people in tropical nations. In light of existing human-induced disturbances, including climate change, TMR can be used as natural model systems to examine the effects of rapid changes in abiotic drivers and their influence on biodiversity and ecosystem function.

Global patterns and drivers of ecosystem functioning in rivers and riparian zones.

River ecosystems receive and process vast quantities of terrestrial organic carbon, the fate of which depends strongly on microbial activity. Variation in and controls of processing rates, however, are poorly characterized at the global scale. In response, we used a peer-sourced research network and a highly standardized carbon processing assay to conduct a global-scale field experiment in greater than 1000 river and riparian sites. We found that Earth's biomes have distinct carbon processing signatures. Slow processing is evident across latitudes, whereas rapid rates are restricted to lower latitudes. Both the mean rate and variability decline with latitude, suggesting temperature constraints toward the poles and greater roles for other environmental drivers (e.g., nutrient loading) toward the equator. These results and data set the stage for unprecedented "next-generation biomonitoring" by establishing baselines to help quantify environmental impacts to the functioning of ecosystems at a global scale.

Biodiversity of leaf litter fungi in streams along a latitudinal gradient.

Global patterns of biodiversity have emerged for soil microorganisms, plants and animals, and the extraordinary significance of microbial functions in ecosystems is also well established. Virtually unknown, however, are large-scale patterns of microbial diversity in freshwaters, although these aquatic ecosystems are hotspots of biodiversity and biogeochemical processes. Here we report on the first large-scale study of biodiversity of leaf-litter fungi in streams along a latitudinal gradient unravelled by Illumina sequencing. The study is based on fungal communities colonizing standardized plant litter in 19 globally distributed stream locations between 69°N and 44°S. Fungal richness suggests a hump-shaped distribution along the latitudinal gradient. Strikingly, community composition of fungi was more clearly related to thermal preferences than to biogeography. Our results suggest that identifying differences in key environmental drivers, such as temperature, among taxa and ecosystem types is critical to unravel the global patterns of aquatic fungal diversity.

Narrow thermal tolerance and low dispersal drive higher speciation in tropical mountains.

Species richness is greatest in the tropics, and much of this diversity is concentrated in mountains. Janzen proposed that reduced seasonal temperature variation selects for narrower thermal tolerances and limited dispersal along tropical elevation gradients [Janzen DH (1967) Am Nat 101:233-249]. These locally adapted traits should, in turn, promote reproductive isolation and higher speciation rates in tropical mountains compared with temperate ones. Here, we show that tropical and temperate montane stream insects have diverged in thermal tolerance and dispersal capacity, two key traits that are drivers of isolation in montane populations. Tropical species in each of three insect clades have markedly narrower thermal tolerances and lower dispersal than temperate species, resulting in significantly greater population divergence, higher cryptic diversity, higher tropical speciation rates, and greater accumulation of species over time. Our study also indicates that tropical montane species, with narrower thermal tolerance and reduced dispersal ability, will be especially vulnerable to rapid climate change.

Determinants of food resource assimilation by stream insects along a tropical elevation gradient.

Food resource availability varies along gradients of elevation where riparian vegetative cover exerts control on the relative availability of allochthonous and autochthonous resources in streams. Still, little is known about how elevation gradients can alter the availability and quality of resources and how stream food webs respond. We sampled habitat characteristics, stable isotope signatures (δ13C, δ15N, δ2Η) and the carbon, nitrogen and phosphorus composition of basal food resources and insects in 11 streams of similar size along an elevation gradient from 1260 to 4045 m on the northeastern slope of the Ecuadorian Andean-Amazon region. Algal-based (autochthonous) food resources primarily supported insects occurring at higher elevations, but at low elevations there was a shift to greater allochthony, corresponding with lower light availability and reduced epilithon resource abundance. Additionally, percent phosphorus (%P) of both autochthonous and allochthonous food resources and of body tissue for some abundant insect taxa (stonefly Anacroneuria and mayfly Andesiops) declined with increasing elevation, despite the greater autochthony at high elevation. Allochthonous food resources were always a lower quality food resource, as indicated by higher C:N, N:P, and lower %P, across elevation in comparison to autochthonous resources, but autochthonous resources had higher %P than allochthonous resources across all elevations and comprised a greater portion of high-elevation insect resource assimilation. Aquatic insects may be able to compensate for the lower quality of both resource types at high elevations through altered body stoichiometry, even though higher quality autochthonous-based foods are in high abundance at high elevations.

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.