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Integrating marine historical ecology into management of Alaska’s Pacific cod fishery for climate readiness

The Pacific cod ( Gadus macrocephalus ) fishery was closed in 2020 after a rapid decline in biomass caused by the marine heat waves of 2014–2019. Pacific cod are exceptionally thermally sensitive and management of this fishery is now challenged by increasingly unpredictable climate conditions. Fisheries monitoring is critical for climate readiness, but short-term monitoring data may be inadequate for recognizing and anticipating change under rapid climate changes. We propose an interdisciplinary, marine historical ecology framework that looks to long-term records (local and traditional knowledge, history, archaeology, and paleoclimatology) to capture a long range of ecological variability and provide historical context for management. In order to connect to contemporary fisheries management, this framework must be built on a common vocabulary and an understanding of the key metrics used in fisheries stock assessments. Here, we propose metrics derived from Pacific cod stock assessment and synthesize information relevant to understanding the effects of past warming periods on cod populations across the Gulf of Alaska and Bering Sea. This case study provides a framework for thinking about how to use these historical records in the context of fisheries management under rapidly changing climate conditions.

Alaska

Manipulation of the Symbiodiniaceae microbiome confers multigenerational impacts on symbioses and reproductive ecology of its Exaiptasia diaphana host

Symbiodiniaceae-associated microbiota strongly influence cnidarian symbioses. We systematically reduced the bacterial and fungal communities associated with Symbiodiniaceae to study potential effects on the cnidarian holobiont Exaiptasia diaphana (Aiptasia). Clonal anemones were inoculated with xenic Breviolum minutum (SSB01) and microbiome-manipulated cultures after antibacterial or antifungal treatment. The asexual reproduction of pedal laceration allowed for three generations of clonal aposymbiotic Aiptasia to be utilised in this study, from the initial adult generation (G0), to the first (G1), and second (G2) generation. We inoculated small and large G1 Aiptasia with SSB01 algae and monitored onset of symbiosis, rate of algal proliferation, and holobiont characteristics. Sequencing the 16S and 18S rRNA gene regions identified significant differences in the bacterial and fungal communities of the G0 and G1 generations, alongside differences between the size classes of small and large G1 anemones. The microbiome of larger G1 individuals was distinct to the smaller G1 anemones, suggesting a microbiome maturation process. Control Breviolum minutum cultures exhibited a significantly greater proliferation rate in large G1 anemones when compared to antibacterial or antifungal treated cultures, whereas the opposite trend was documented in the small G1 anemones. Although no differences were observed between algal photochemical parameters, or the growth and polyp activity of G1 juveniles, we observed a significant influence in the production of G2 clones between treatments. Overall, we provide strong ecological implications of manipulating Symbiodiniaceae microbiome, not for the algae themselves, but for the maturation of the host Aiptasia, as well as for the cnidarian holobiont over multiple generations.

The ISME Journal

Disturbance is the primary determinant of food chain length when the top predator is constant

Food chain length (FCL) is a primary determinant of food web structure and is hypothesized to be influenced by habitat size, productivity, and disturbance. Understanding the environmental characteristics that determine food chain length can assist in understanding how food webs may be impacted due to changes in habitats and environmental characteristics. This study examines the impact of hydrologic disturbance on stream food webs when the top predator is constant. We analyzed FCL in less disturbed groundwater flashy streams and more disturbed runoff flashy streams using stable isotopes. Despite no difference in species richness or fish density, food chains in more disturbed streams had a lower FCL compared to food chains in more stable streams. Assemblage analysis showed that flow regime and drainage area significantly impacted individual species abundances. The more disturbed runoff flashy streams had higher proportions of primary consumer fish, such as the algivorous Campostoma sp. (Stonerollers), which likely drives the reduced FCL. Drainage area and land cover had non-significant relationships with FCL. Shifting community structure due to hydrologic variability likely leads to differences in diet of Micropterus dolomieu (Smallmouth Bass), and thus a difference in FCL.

Arkansas, Missouri, Oklahoma

Assessing environmental drivers of denitrification in restored riverine floodplains

Restoration of impaired floodplains is an increasingly prevalent strategy for alleviating water quality concerns and reducing downstream flooding at watershed scales. Floodplains temporarily store water and slow flow velocity to promote sedimentation during overbank flooding and remove inorganic nitrogen from floodwater and groundwater via denitrification. Evaluating the impacts of different restoration strategies on denitrification can inform more strategic investments into floodplain modifications that improve water quality outcomes. Our research investigates how denitrification rates in floodplains respond to environmental factors that are actionable from an engineering perspective through design and water resources management. We seasonally measured soil denitrification enzyme activity and various environmental characteristics in 4 floodplains with different restoration design and management approaches at the confluence of the Wabash and Tippecanoe Rivers in Indiana, United States. Our results showed that denitrification rates in an agricultural floodplain were significantly lower than in restored floodplains with native vegetation. Certain soil conditions characteristic of floodplain wetlands were associated with higher denitrification, particularly elevated total nitrogen, moisture, silt, and organic matter contents. Vegetation species composition was correlated with denitrification rates. This link may reflect the direct effects of vegetation on soil conditions, such as supplying labile organic carbon, or indirect effects, such as vegetation acting as an indicator of hydrologic regime and land use. Denitrification seasonally varied, peaking in winter when nitrate supply from rivers draining agricultural watersheds in the region is also high. Substrate limitation of soil denitrification enzyme activity was most significant during the summer when overbank flooding, which replenishes soil nitrogen stocks, rarely occurs. Our findings indicate that denitrification capacity will likely be maximized in riverine floodplains that are restored as wetlands with diverse native vegetation and enhanced hydrologic connectivity. Such restoration activities promote higher denitrification rates via elevated moisture, fine sediment deposition, and soil organic matter.

Tippecanoe River, Wabash River

Overstorey mortality promotes juvenile piñon pine growth during favourable weather at cooler, wetter sites

Hotter droughts have resulted in widespread tree die-off events globally, frequently leading to regeneration failure. Dry forest recovery often depends on the growth and survival of extant juvenile trees. However, it is unclear how microenvironmental changes following overstorey tree die-off affect juvenile trees, particularly in dryland systems where tree recruitment is typically limited by water availability and heat stress. We simulated an overstorey tree die-off event by girdling trees in piñon-juniper woodlands across the south-western United States. We sampled juvenile piñon pine growth from live and dead overstorey treatments across six study sites spanning a regional latitudinal gradient and local elevational gradients. We examined how juvenile branch and needle growth differed between live and dead overstorey treatments, and whether responses varied with weather conditions and juvenile tree size following overstorey mortality. We found greater juvenile branch and needle growth under dead compared with live overstorey trees for 2 years following overstorey mortality at mid- and high-elevation sites which are typically cooler and wetter than the other sites. These observed growth releases were contingent on favourable post-mortality weather conditions. Higher growth under dead overstorey occurred at sites experiencing near-average climatic water deficits compared with sites experiencing above-average climatic water deficits. Growth at all sites increased from the first to second year after overstorey mortality. Across sites, growth was unrelated to juvenile tree size. Synthesis . Our results underscore differentiation in juvenile responses to overstorey tree die-off driven by local site conditions and weather across the range of Pinus edulis . Overstorey mortality resulted in consistently higher juvenile growth only at climatically favourable sites and during favourable weather, while unmeasured microsite differences could help account for variation observed at the hottest and driest site. Results from less climatically favourable sites suggest that overstorey trees neither directly limit nor facilitate juvenile growth, though further study over longer timeframes is needed to resolve the pace and magnitude of potential recovery or decline. Overall, juvenile vigour may be promoted following overstorey mortality only in a narrow spatial (site) and temporal (weather) environmental context, suggesting additional vulnerabilities for piñon populations under more arid conditions.

Arizona, Colorado

Nonstationary demographic state-space models using unreplicated counts for species undergoing environmental stressors

A fundamental task in ecological statistics is to estimate abundance and growth rate distributions from wildlife monitoring data to inform conservation management. Modeling time series of wildlife populations presents a number of challenges from both statistical and ecological perspectives, including discreteness; lack of replication; nonstationarity; and observation, demographic, and other phenomenological processes. Nonstationary dynamics are often exhibited by populations undergoing environmental stressors. Models must account for these characteristics to produce reliable estimates of abundance and trends, yet estimation can be challenging with unreplicated data. We propose nonstationary demographic state-space models using unreplicated counts for populations undergoing environmental stressors. A reduced growth rate model matches the complexity of the unreplicated count data, and a fecundity bound on growth rate distributions allows the separation of processes affecting growth rates like environmental stressors from those affecting abundance external to growth rates like migration. NDSSMs allow for the embedding of nonstationary model components, and we explore the use of changepoints, volatility clustering, and migration processes. We apply the proposed nonstationary models in case studies of herons affected by predator/competitor reestablishment and three bat species affected by a fungal pathogen causing white-nose syndrome. Nonstationary models outperform stationary models and generalized linear mixed effects models according to model scoring and visual inspection of predictions, and provide estimates more consistent with published values. Incorporating migration improves model fit universally, even with approximate one-way immigration, most likely because populations are extirpated, recolonized, and increase multiple-fold over the upper bound set by species fecundity. In addition, estimates of the timing and severity of the environmental stressor differed for models with migration. Including nonstationary and demographic components in a fecundity-bounded growth rate model improves inference and benefits interpretability of hyperparameters. In turn, this adjusts uncertainties in predictions of abundance and growth rates over time, providing the ingredients needed for informed conservation analysis and for directing future monitoring of at-risk species.

Journal of Agricultural, Biological and Environmen

Keeping pace: Migrating sea lampreys adjust swim speed in response to shifting water velocity to maintain a constant ground speed

Upstream migrating fish must balance energetic costs with environmental and ecological constraints. Theoretical models predict that fish can conserve energy by adjusting swim speed in response to changing water velocity, but detailed field tests of these predictions remain rare. We used fine-scale acoustic telemetry and hydrodynamic modeling to track adult sea lamprey (Petromyzon marinus) migrating through a river and evaluated how movement speeds varied with environmental conditions and individual traits. Swim speed increased with faster water velocities, while ground speed remained constant, supporting the hypothesis that lampreys regulate swim effort to maintain a fixed migration pace. The average ground speed aligned with a predicted optimum of ∼1 body length per second. Both swim and ground speeds declined slightly in deeper water, suggesting a trade-off between movement efficiency and predator avoidance. These findings provide field-based support for a theoretical model of energy optimization and underscore how migratory fish adaptively adjust behavior to local hydrodynamic conditions. Our results highlight the value of using high-resolution tracking and environmental modeling to reveal behavioral tactics that underlie efficient migration strategies.

Michigan

Hydrologic variability and groundwater age of springs in eastern Oregon and northern Nevada, USA

The ecological importance of springs in semiarid regions is far greater than their small size and sparse distribution, yet little is known about the hydrologic functioning of these systems. During 2016–22, 261 springs were visited in the volcanic terrane of eastern Oregon and northern Nevada. When conditions were suitable, measurements of discharge, water temperature, and specific conductance were made, and samples for the analysis of carbon-14, tritium, and water stable isotopes (WSI) were collected. A subset of 60 springs was revisited during different seasons in the same year and during the dry season in multiple years to evaluate variability in discharge, chemistry, and groundwater age. Specific conductance and WSI varied considerably among springs across the study area but were unexpectedly stable across seasons and years at individual springs. Seasonal and interannual variability in spring discharge was related to the residence time of the discharging groundwater. Springs discharging older groundwater (10 3 –10 4 years) had significantly less variability in their discharge compared to springs discharging younger groundwater (10 0 –10 1 years). Variability among springs discharging younger groundwater included cessation of late-summer discharge at 18 % of the repeat-visit springs. A logistic regression model predicted the age of discharging spring water with 89 % accuracy using only the spring latitude, longitude, elevation, and δ 2 H value. This study framework provides a simple, inexpensive, and robust method to provisionally assess the hydrologic behavior of springs having little or no prior information in understudied, semiarid regions across the globe.

Nevada, Oregon

Forecasting water levels using the ConvLSTM algorithm in the Everglades, USA

Forecasting water levels in complex ecosystems like wetlands can support effective water resource management, ecological conservation, and understanding surface and groundwater hydrology. Predictive models can be used to simulate the complex interactions among natural processes, hydrometeorological factors, and human activities. The Greater Everglades in the USA is a well-known example of an ecosystem where complexity has motivated adoption of machine learning algorithms in water level prediction studies. This paper aims to contribute to extending existing machine learning algorithms by integrating spatiotemporal data with deep-learning algorithms in the forecasting process. In this study, a deep-learning model is developed to predict water levels on a regional scale, covering a large area of approximately 9,138 square kilometers in the Everglades ecosystem. This model has the architecture of Convolutional Long Short-Term Memory which can deal with spatiotemporal data by capturing both spatial and temporal dependencies in the training data. The forecasting capabilities of this model (referred to as the global model) are assessed by comparing the global model to two Artificial Neural Networks developed at two different gaging stations, referred to here as local models. One local model is developed at a gaging station directly influenced by nearby water control structures, whereas the other is developed at a gaging station located farther away from these structures. By leveraging data from the Everglades Depth Estimation Network spanning from January 2002 to May 2023, the global and local models were trained to forecast water levels with a two-day lead time. Our findings suggest that both the global and local models perform with approximately the same level of accuracy, with Mean Absolute Relative Error values ranging from 0.38% to 1.4% at the selected stations. The developed global model has demonstrated strong potential as a standalone forecasting tool for the entire study area in the Everglades and could eliminate the need for developing multiple local models. This finding also highlights how machine learning can capture complex spatial and temporal relationships to generate accurate water level predictions on a regional scale.

Florida

Perspectives and insights gained from applying landscape ecological approaches to the Upper Mississippi riverscape

Advances in remote sensing, spatial data collection, hydraulic and geospatial models, and the maturation of landscape ecology during the early 2000s spurred the concept of rivers as landscapes or ‘riverscapes’. Since that time, the Upper Mississippi River Restoration (UMRR) Program - one of the world’s longest running large-river ecosystem restoration and monitoring programs - has applied landscape ecological principles to the Upper Mississippi River System (UMRS). I examine spatial patterns of aquatic nutrient concentrations, submersed aquatic vegetation, and fish and mussel communities, finding that within-river patterns are often patchy in nature, likely resulting from how nutrient processes and community traits interact with spatial variability in hydraulic exchange patterns. Local-scale patchy spatial patterns in the UMRS are embedded in system-scale longitudinal gradients and a hierarchical view of the river has provided a foundation for strategic river restoration planning. Key Words: Aquatic Habitat, hydraulic connectivity, patch, restoration, riverscape

BioScience

Birds, breakpoints, and baselines: How citizen science data can reveal ecological boundaries in Kenya’s Upper Tana watershed

Tropical watersheds are increasingly threatened by climate change, land-use conversion, and resource extraction, yet conventional biodiversity monitoring in these systems is often spatially and temporally limited. Citizen science offers a complementary approach, enabling biodiversity data collection over large areas that can supplement professional scientific surveys. We analyzed 10 years (2012–2022) of Upper Tana Watershed bird data from the Kenya Bird Map project, covering 114 pentads (9 × 9 km) within a 17,000 km 2 watershed to assess patterns of bird community composition and distribution across this watershed and to also evaluate the effects of environmental variables, seasonality, and sampling effort to help inform improvements in future citizen science projects. Citizen (or community) scientists recorded 575 species (>50% of Kenya’s total avifauna) in 74 families. Asymptotic species accumulation indicates that most probable species present in the watershed were detected. Threshold indicator taxa analysis revealed distinct ecological boundaries along elevation (∼1,500 m), precipitation (∼1,100 mm), and mean temperature (∼19°C) gradients, corresponding to a turnover from xeric savanna to mesic montane forest assemblages. Notably, bird communities showed little seasonal differentiation between wet and dry periods, consistent with dominance by resident year-round species. Data limitations including uneven survey distribution and frequency, absence of abundance metrics, and coarse representation of local environmental conditions that likely reduced our ability to detect fine-scale species–habitat relationships. Addressing these gaps through spatially balanced sampling at greater resolution, greater survey frequency in underrepresented areas, and improved capture of habitat metrics could strengthen the use of citizen-science bird data for watershed bioassessment. Our findings demonstrate that structured citizen-science initiatives can identify ecological boundaries and inform adaptive management of tropical socio-ecological systems under rapid environmental change.

Upper Tana watershed

Editorial: From cold seeps to hydrothermal vents: Geology, chemistry, microbiology, and ecology in marine and coastal environments

This Research Topic compiles contemporary studies on cold seeps, hydrothermal vents, mud volcanoes, and related seafloor features that are associated with focused fluid emissions and the transfer of carbon, other chemical species, and sometimes heat from the geosphere to the ocean. Because these features sometimes tap fluids and gas originating kilometers below the seafloor, they provide an important window into deep processes that are otherwise inaccessible to scientists. At the shallow portion of their journey, migrating fluids nearing the seafloor contribute to a range of unique biological, physical, and chemical processes within the sediments themselves and at the sediment-water interface. Seafloor fluid emissions play a critical role in global biogeochemical cycles, ocean chemistry, and possibly even climate change. Seafloor leakage points often emit hydrocarbon gases (especially methane and CO 2 ) and are sometimes the loci for deposition of seafloor minerals that have economic value. A burgeoning area of research focuses on natural products generated at these features, seeking compounds with potential pharmaceutical or other applications. Multidisciplinary studies have become routine for characterization of seafloor fluid emission sites, attesting to the inseparability of geologic, physical, chemical, and biological processes in these settings. It is increasingly common for researchers to combine in a single research cruise: subbottom imaging and seafloor mapping; porewater and water column geochemistry and gas sampling; sediment retrieval for lithologic, biostratigraphic, and solid phase analyses; and studies of benthic and subseafloor communities at the microbial to macrofaunal scales. This multidisciplinary approach has the advantage of ensuring the spatial and temporal coincidence of surveys and samples, an important factor at highly dynamic seafloor fluid emission sites. In addition, researchers often use remotely operated vehicles (ROVs), autonomous underwater vehicles (AUVs), or human-occupied vehicles (HOVs) to record video of the seafloor, compile photomosaics, collect targeted samples, and survey with high-resolution geophysical near-seafloor systems, providing a degree of detail about seafloor fluid emission sites that is unprecedented compared to most areas of the deep ocean. While rarer, long-term cabled observatories or shorter-term deployments of portable observatories are also used at some loci for seafloor fluid flux and are particularly helpful for capturing temporal variations at these dynamic features. Here we summarize the Research Topic’s contribution to multidisciplinary seafloor emission studies in the categories of cold seeps, mud volcanoes, and hydrothermal vents. Figure 1 shows the geographic distribution of the studies in this Research Topic and key features referred to in this Introduction.

Frontiers in Earth Science

King Rails in southeastern Arkansas are majority resident

King Rails ( Rallus elegans ), a declining species of secretive marsh bird, have a mixed migratory strategy, with some individuals migrating and some being year-round residents. Determining the distribution of migratory versus resident King Rails is critical to guiding habitat management and assessing risks to populations. We examined the migratory status of adult King Rails at the Choctaw West Wildlife Management Area in southeastern Arkansas, the only known site in Arkansas with multiple breeding pairs. Of the 23 adult King Rails we tagged in 2022 and 2023, two birds migrated away from the local area, one in each year. One migrated 145 km, and the other 450 km. Our findings suggest that the line between the migratory and resident range of Arkansas King Rails could be north of our site.

Arkansas

Estimation of parental abundance using hierarchical Bayesian modeling with data augmentation

Pedigree-based estimation methods leverage the fact that each offspring in a cohort is genotypically “marked” by its parents and represent a recent and promising toolset for estimating population dynamics. This includes pedigree accumulation estimators that model the “accumulation” of inferred unique parents within a given cohort to estimate parental abundance. Unlike close-kin mark-recapture approaches, which rely on intercohort comparisons, pedigree accumulation modeling can be completed solely using intracohort samples. This is particularly advantageous for semelparous species, where intercohort pairs are impossible and adult life stages can be difficult to sample without affecting their likelihood of successfully reproducing. Previous work has evaluated a range of estimators for such datasets, concluding that the non-parametric Chao estimator provides the most accurate and precise estimates for feasible levels of sampling effort. We used simulated data to evaluate an alternative estimator based on hierarchical modeling and data augmentation in a Bayesian framework. Results indicate that estimates from the hierarchical Bayesian estimator had comparable accuracy and better precision than both the previously tested Chao1 estimator and the improved iChao formulation across a range of sample sizes and sex ratios. Furthermore, the Bayesian estimator was far more robust to simulated errors in pedigree reconstruction, especially the presence of false negatives. Hierarchical Bayesian pedigree accumulation models can also provide additional insight into underlying reproductive ecology through their use of an explicit observation process, allowing for the incorporation or estimation of species- and population-specific reproductive dynamics. More broadly, the parametric nature of these models offers opportunities to efficiently pool information among datasets as well as to propagate uncertainty within more complex models.

Ecology and Evolution

Family life is critical for migration and survival in a long-lived social bird

Extended parental care is common among birds and particularly well known among long-lived migratory species. Yet, factors influencing the duration of parental care are poorly understood, and empirical evidence for the functional importance of family life for offspring development is scarce. Using unique tracking and genetic data of 55 adult and juvenile migratory Bewick’s swans, among which 18 parent-offspring pairs and 13 sibling pairs, we quantify variation in family bond duration and identify the cause and consequences of their dissolution. Families generally broke up during the second half of the first northward migration of the offspring, when offspring were nearly one year old. Earlier separation occurred on the wintering grounds in response to disturbance ( n = 10 juveniles) and led to delayed migration and, critically, lower survival (40 vs. 86%). Our results provide rare empirical evidence that family bonds, as a pathway for social learning, are not just important, but vital to the migration and survival of young family-living birds.

Behavioral Ecology and Sociobiology

The effect of temperature, flow, density and disease on smallmouth bass (Micropterus dolomieu) early life growth rate

Individual growth rates of fish are influenced by gradually changing environmental conditions and rapid-onset events. However, there is a paucity of information on drivers of spatiotemporal variability in growth rates of fishes across large spatial extents. Using a 36-year dataset of smallmouth bass ( Micropterus dolomieu ) length-at-age data across 10 river reaches in Pennsylvania, USA ( n = 54,068 individuals), we estimated annual early life growth rates (mm year −1 ) and assessed the effects of relative abundance, summer temperature, summer streamflow rate and a disease outbreak on growth rate. We found that temperature had a positive effect on growth rate, varied spatially, and had a stronger effect in diseased reaches. There was also evidence for an impact of disease on early life growth through density-dependent mechanisms, with growth rates increasing from ∼85 to 101 mm·year −1 following disease-related decreases in abundance. This study adds to our understanding of the factors that shape early life growth, including rapid-onset events like disease which can have immediate and lasting effects on growth rate trajectory.

Pennsylvania

Satellite time series analysis to quantify changing climax ciénegas using a state and transition model approach

Ciénegas are rare wetlands in arid landscapes of the North American Southwest, historically providing critical ecological and hydrological functions but increasingly threatened by changing climate and land use pressures. This study quantifies changes in ciénega condition and floodplain dynamics using a state-and-transition model (STM) informed by expert knowledge and remote sensing. Key factors include woody plant encroachment, water availability, and soil aggradation. We mapped 31 ciénegas with high-resolution imagery and analyzed Landsat data (1985–2023) to assess vegetation health and moisture using the Normalized Difference Vegetation Index (NDVI) and Normalized Difference Infrared Index (NDII). Results show substantial interannual variability in phenology, water stress, and soil moisture, with regional drying and elevation strongly influencing ciénega resilience. We classified ciénegas into three functional states—healthy, desiccated, and dormant—and mapped their 2023 condition. Trend analyses indicate most ciénegas exhibit greening despite drought, though localized variability underscores the need for site-specific management. None are in a stable climax (reference) state; rather, they transition among states in response to external drivers. Increasing woody plant cover and surface drying, likely linked to declining regional water tables, favor deep-rooted species over wetland grasses—a pattern mirrored in adjacent control plots. Spatially explicit analysis revealed intra-ciénega variability often masked by aggregated data, highlighting the importance of high-resolution monitoring. Seasonal and long-term trends provide context for understanding ciénega dynamics, including degradation and restoration pathways. This study emphasizes the importance of groundwater conservation and demonstrates how remote sensing supports long-term monitoring. The STM framework offers a practical tool for adaptive management to sustain freshwater resources in arid environments.

Arizona, New Mexico

Riverscape heterogeneity shapes population diversity for a migratory fish

Habitat patch dynamics can scale up to influence population demography and diversity with implications for resilience to environmental stochasticity. But how the spatial arrangement and size of habitat patches interact with other components of habitat heterogeneity to shape population diversity at larger spatial scales is not well understood. For riverine fishes, there is increasing evidence that tributary streams provide critical demographic support to main stem rivers. However, the extent to which main stem rivers rely on demographic contributions from tributaries, and the factors underlying this dependence, have not been assessed. Here, we used genetic stock identification to evaluate the effect of tributaries on population diversity of Yellowstone cutthroat trout ( Oncorhynchus virginalis bouvieri ) occupying the main stem Snake River, Wyoming, USA. We found that the main stem relied almost entirely on tributaries for demographic support, but main stem composition varied spatially among river sections. Distance between habitat patches, catchment area, and groundwater availability acted in concert to determine the contribution of specific tributaries to the main stem, but contributions were ultimately modulated by habitat connectivity. We also found evidence for multi-scale spatial structure in tributary contributions, providing insight into untested drivers of main stem river population diversity. Our results demonstrate how spatially discrete and distributed riverscape attributes influence population diversity at broader spatial scales, illustrating how ecosystem resilience emerges from the dynamic, two-way exchange of individuals and energy across habitat networks. Management plans for large rivers that address the ecological contributions of tributaries may be needed to achieve optimal outcomes. Similarly, conservation strategies that exclusively focus on headwater streams may fail to capture the broader habitat requirements necessary to maintain robust cold-water fish populations and associated recreational fisheries, particularly under global environmental change.

Wyoming