Macroscale controls determine the recovery of river ecosystem productivity following flood disturbances.

River ecosystem function depends on flow regimes that are increasingly modified by changes in climate, land use, water extraction, and flow regulation. Given the wide range of variation in flow regime modifications and autotrophic communities in rivers, it has been challenging to predict which rivers will be more resilient to flow disturbances. To better understand how river productivity is disturbed by and recovers from high-flow disturbance events, we used a continental-scale dataset of daily gross primary production time series from 143 rivers to estimate growth of autotrophic biomass and ecologically relevant flow disturbance thresholds using a modified population model. We compared biomass recovery rates across hydroclimatic gradients and catchment characteristics to evaluate macroscale controls on ecosystem recovery. Estimated biomass accrual (i.e., recovery) was fastest in wider rivers with less regulated flow regimes and more frequent instances of biomass removal during high flows. Although disturbance flow thresholds routinely fell below the estimated bankfull flood (i.e., the 2-y flood), a direct comparison of disturbance flows estimated by our biomass model and a geomorphic model revealed that biomass disturbance thresholds were usually greater than bed disturbance thresholds. We suggest that primary producers in rivers vary widely in their capacity to recover following flow disturbances, and multiple, interacting macroscale factors control productivity recovery rates, although river width had the strongest overall effect. Biomass disturbance flow thresholds varied as a function of geomorphology, highlighting the need for data such as bed slope and grain size to predict how river ecosystems will respond to changing flow regimes.

Light and flow regimes regulate the metabolism of rivers.

Mean annual temperature and mean annual precipitation drive much of the variation in productivity across Earth's terrestrial ecosystems but do not explain variation in gross primary productivity (GPP) or ecosystem respiration (ER) in flowing waters. We document substantial variation in the magnitude and seasonality of GPP and ER across 222 US rivers. In contrast to their terrestrial counterparts, most river ecosystems respire far more carbon than they fix and have less pronounced and consistent seasonality in their metabolic rates. We find that variation in annual solar energy inputs and stability of flows are the primary drivers of GPP and ER across rivers. A classification schema based on these drivers advances river science and informs management.

Genetic erosion in an endangered desert fish during a megadrought despite long-term supportive breeding.

Human water use combined with a recent megadrought have reduced river and stream flow through the southwest United States and led to periodic drying of formerly perennial river segments. Reductions in snowmelt runoff and increased extent of drying collectively threaten short-lived, obligate aquatic species, including the endangered Rio Grande silvery minnow (Hybognathus amarus). This species is subject to boom-and-bust population dynamics, under which large fluctuations in abundance are expected to lower estimates of effective population size and erode genetic diversity over time. Rates of diversity loss are also affected by additions of hatchery-origin fish used to supplement the wild population. We used demographic and genetic data from wild and hatchery individuals to examine the relationship of genetic diversity and effective population size to abundance over the last two decades. Genetic diversity was low during the early 2000s, but diversity and demographic metrics stabilized after the hatchery program was initiated and environmental conditions improved. Yet, from 2017 onward, allelic diversity declined (Cohen's d = 1.34) and remained low despite hatchery stocking and brief wild population recovery. Across the time series, single-sample estimates of effective population size based on linkage disequilibrium (LD Ne ) were positively associated (r = 0.53) with wild abundance and total abundance, but as the proportion of hatchery-origin spawners increased, LD Ne declined (r = -0.55). Megadrought limited wild spawner abundance and precluded refreshment of hatchery brood stocks with wild fish; hence, we predict a riverine population increasingly dominated by hatchery-origin individuals and accelerated loss of genetic diversity despite supplementation. We recommend an adaptive and accelerated management plan that integrates river flow management and hatchery operations to slow the pace of genetic diversity loss exacerbated by megadrought.

El uso humano del agua, combinado con una megasequía reciente, ha reducido el flujo de los ríos y arroyos en el suroeste de los Estados Unidos y ha provocado la seca periódica de segmentos de ríos que antes eran perennes. Las reducciones en la escorrentía del deshielo y el aumento de la sequía amenazan colectivamente a especies obligatoriamente acuáticas de vida corta, incluyendo la amenazada carpa chamizal (Hybognathus amarus). Esta especie está sujeta a una dinámica poblacional de explosión y colapso, bajo la cual se espera que grandes fluctuaciones en la abundancia reduzcan las estimaciones del tamaño efectivo de la población y erosionen la diversidad genética con el tiempo. Las tasas de pérdida de la diversidad también se ven afectadas por la adición de peces procedentes de criaderos usados para suplementar la población silvestre. Utilizamos datos demográficos y genéticos de individuos silvestres y de criaderos para examinar la relación entre la diversidad genética y el tamaño efectivo de la población con la abundancia durante las últimas dos décadas. La diversidad genética fue baja a principios de los 2000, pero las métricas de diversidad y demografía estabilizaron después de que se inició el programa de criadero y mejoraron las condiciones ambientales. Sin embargo, a partir de 2017, la diversidad alélica disminuyó (d de Cohen = 1,34) y se mantuvo baja a pesar de la suplementación con individuos de criaderos y la breve recuperación de la población silvestre. A lo largo del tiempo, las estimativas de muestras individuales del tamaño efectivo de la población basados en el desequilibrio de ligamiento (LD Ne) estaban asociadas positivamente (r = 0,53) con la abundancia silvestre y la abundancia total, pero a medida que la proporción de desovadores originados en criaderos aumentó, el LD Ne disminuyó (r = -0,55). La megasequía limitó la abundancia de desovadores silvestres e impidió el reabastecimiento de las poblaciones en cautiverio con peces silvestres; por lo tanto, predecimos una población ribereña cada vez más dominada por individuos procedentes de criaderos y una pérdida acelerada de diversidad genética a pesar de la suplementación. Recomendamos un plan de gestión adaptativo y acelerado que integre la gestión del caudal del río y las operaciones de criaderos para frenar el ritmo de pérdida de diversidad genética exacerbada por la megasequía.

Invader removal triggers competitive release in a threatened avian predator.

Changes in the distribution and abundance of invasive species can have far-reaching ecological consequences. Programs to control invaders are common but gauging the effectiveness of such programs using carefully controlled, large-scale field experiments is rare, especially at higher trophic levels. Experimental manipulations coupled with long-term demographic monitoring can reveal the mechanistic underpinnings of interspecific competition among apex predators and suggest mitigation options for invasive species. We used a large-scale before-after control-impact removal experiment to investigate the effects of an invasive competitor, the barred owl (Strix varia), on the population dynamics of an iconic old-forest native species, the northern spotted owl (Strix occidentalis caurina). Removal of barred owls had a strong, positive effect on survival of sympatric spotted owls and a weaker but positive effect on spotted owl dispersal and recruitment. After removals, the estimated mean annual rate of population change for spotted owls stabilized in areas with removals (0.2% decline per year), but continued to decline sharply in areas without removals (12.1% decline per year). The results demonstrated that the most substantial changes in population dynamics of northern spotted owls over the past two decades were associated with the invasion, population expansion, and subsequent removal of barred owls. Our study provides experimental evidence of the demographic consequences of competitive release, where a threatened avian predator was freed from restrictions imposed on its population dynamics with the removal of a competitively dominant invasive species.

Models of underlying autotrophic biomass dynamics fit to daily river ecosystem productivity estimates improve understanding of ecosystem disturbance and resilience.

Directly observing autotrophic biomass at ecologically relevant frequencies is difficult in many ecosystems, hampering our ability to predict productivity through time. Since disturbances can impart distinct reductions in river productivity through time by modifying underlying standing stocks of biomass, mechanistic models fit to productivity time series can infer underlying biomass dynamics. We incorporated biomass dynamics into a river ecosystem productivity model for six rivers to identify disturbance flow thresholds and understand the resilience of primary producers. The magnitude of flood necessary to disturb biomass and thereby reduce ecosystem productivity was consistently lower than the more commonly used disturbance flow threshold of the flood magnitude necessary to mobilize river bed sediment. The estimated daily maximum percent increase in biomass (a proxy for resilience) ranged from 5% to 42% across rivers. Our latent biomass model improves understanding of disturbance thresholds and recovery patterns of autotrophic biomass within river ecosystems.

Range-wide sources of variation in reproductive rates of northern spotted owls.

We conducted a range-wide investigation of the dynamics of site-level reproductive rate of northern spotted owls using survey data from 11 study areas across the subspecies geographic range collected during 1993-2018. Our analytical approach accounted for imperfect detection of owl pairs and misclassification of successful reproduction (i.e., at least one young fledged) and contributed further insights into northern spotted owl population ecology and dynamics. Both nondetection and state misclassification were important, especially because factors affecting these sources of error also affected focal ecological parameters. Annual probabilities of site occupancy were greatest at sites with successful reproduction in the previous year and lowest for sites not occupied by a pair in the previous year. Site-specific occupancy transition probabilities declined over time and were negatively affected by barred owl presence. Overall, the site-specific probability of successful reproduction showed substantial year-to-year fluctuations and was similar for occupied sites that did or did not experience successful reproduction the previous year. Site-specific probabilities for successful reproduction were very small for sites that were unoccupied the previous year. Barred owl presence negatively affected the probability of successful reproduction by northern spotted owls in Washington and California, as predicted, but the effect in Oregon was mixed. The proportions of sites occupied by northern spotted owl pairs showed steep, near-monotonic declines over the study period, with all study areas showing the lowest observed levels of occupancy to date. If trends continue it is likely that northern spotted owls will become extirpated throughout large portions of their range in the coming decades.

Exploring metapopulation-scale suppression alternatives for a global invader in a river network experiencing climate change.

Invasive species can dramatically alter ecosystems, but eradication is difficult, and suppression is expensive once they are established. Uncertainties in the potential for expansion and impacts by an invader can lead to delayed and inadequate suppression, allowing for establishment. Metapopulation viability models can aid in planning strategies to improve responses to invaders and lessen invasive species' impacts, which may be particularly important under climate change. We used a spatially explicit metapopulation viability model to explore suppression strategies for ecologically damaging invasive brown trout (Salmo trutta), established in the Colorado River and a tributary in Grand Canyon National Park. Our goals were to estimate the effectiveness of strategies targeting different life stages and subpopulations within a metapopulation; quantify the effectiveness of a rapid response to a new invasion relative to delaying action until establishment; and estimate whether future hydrology and temperature regimes related to climate change and reservoir management affect metapopulation viability and alter the optimal management response. Our models included scenarios targeting different life stages with spatially varying intensities of electrofishing, redd destruction, incentivized angler harvest, piscicides, and a weir. Quasi-extinction (QE) was obtainable only with metapopulation-wide suppression targeting multiple life stages. Brown trout population growth rates were most sensitive to changes in age 0 and large adult mortality. The duration of suppression needed to reach QE for a large established subpopulation was 12 years compared with 4 with a rapid response to a new invasion. Isolated subpopulations were vulnerable to suppression; however, connected tributary subpopulations enhanced metapopulation persistence by serving as climate refuges. Water shortages driving changes in reservoir storage and subsequent warming would cause brown trout declines, but metapopulation QE was achieved only through refocusing and increasing suppression. Our modeling approach improves understanding of invasive brown trout metapopulation dynamics, which could lead to more focused and effective invasive species suppression strategies and, ultimately, maintenance of populations of endemic fishes.

Las especies invasoras pueden alterar dramáticamente un ecosistema, pero erradicarlas es complicado y suprimirlas es costoso una vez que están establecidas. Las incertidumbres en el potencial de expansión y el impacto de un invasor pueden derivar en una supresión retardada e inadecuada que permite el establecimiento. Los modelos de viabilidad meta poblacional pueden auxiliar en la planeación de estrategias para mejorar las respuestas ante especies invasoras y disminuir su impacto, lo cual puede ser particularmente importante ante el cambio climático. Usamos un modelo meta poblacional espacialmente explícito para explorar las estrategias de supresión usadas con la trucha café (Salmo trutta), una especie invasora y dañina establecida en el Río Colorado en el Parque Nacional del Gran Cañón. Nuestros objetivos fueron estimar la efectividad de las estrategias enfocadas en diferentes etapas de vida y subpoblaciones dentro de una meta población; cuantificar la efectividad de una respuesta rápida ante una nueva invasión en relación a retardar la acción hasta que ocurra el establecimiento; y estimar si los sistemas térmicos e hidrológicos relacionados con el cambio climático y la gestión de cuencas afectarán la viabilidad meta poblacional y alterarán la respuesta óptima de gestión en el futuro. Nuestros modelos incluyeron escenarios enfocados en diferentes etapas de vida con intensidades espacialmente variables de pesca eléctrica, destrucción de redes, cultivo incentivado de pescadores, piscicidas y un dique. La cuasi extinción (CE) sólo se obtuvo con una supresión a nivel meta poblacional enfocada en múltiples etapas de vida. Las tasas de crecimiento poblacional de la trucha fueron más sensibles a los cambios en edad cero y una gran mortalidad adulta. La duración de la supresión requerida para llegar a la CE para una subpoblación grande establecida fue de doce años en comparación con los cuatro de una respuesta rápida a una nueva invasión. Las subpoblaciones aisladas fueron vulnerables a la supresión; sin embargo, las subpoblaciones conectadas por medio de tributarios incrementaron la persistencia meta poblacional al fungir como refugios climáticos. La escasez de agua, cambios impulsores en el almacenamiento de la cuenca y el calentamiento subsecuente causarían declinaciones de la trucha, pero la CE meta poblacional sólo se logró con el reenfoque e incremento de la supresión. Nuestra estrategia de modelado mejora el entendimiento de las dinámicas meta poblacionales de la trucha café invasora, lo cual podría llevar a estrategias de supresión más enfocadas y efectivas y, finalmente, al mantenimiento de las poblaciones de peces endémicos. Exploración de alternativas a la supresión a escala meta poblacional de un invasor mundial en una red de ríos que experimenta el cambio climático.

Migration timing and tributary use of spawning flannelmouth sucker (Catostomus latipinnis).

Spawning phenology and associated migrations of fishes are often regulated by factors such as temperature and stream discharge, but flow regulation of mainstem rivers coupled with climate change might disrupt these cues and affect fitness. Flannelmouth sucker (Catostomus latipinnis) persisting in heavily modified river networks are known to spawn in tributaries that might provide better spawning habitat than neighboring mainstem rivers subject to habitat degradation (e.g., embedded sediments, altered thermal regimes, and disconnected floodplains). PIT tag data and radio telemetry were used to quantify the timing and duration of flannelmouth sucker tributary spawning migrations in relation to environmental cues in McElmo Creek, a tributary of the San Juan River in the American Southwest. We also tested the extent of the tributary migration and assessed mainstem movements prior to and after tributary migrations. Additionally, multiyear data sets of PIT detections from other tributaries in the Colorado River basin were used to quantify interannual and cross-site variation in the timing of flannelmouth sucker spawning migrations in relation to environmental cues. The arrival and residence times of fish spawning in McElmo Creek varied among years, with earlier migration and a 3-week increase in residence time in relatively wet years compared to drier years. Classification tree analysis suggested a combination of discharge- and temperature-determined arrival timing. Of fish PIT tagged in the fall, 56% tagged within 10 km of McElmo Creek spawned in the tributary the following spring, as did 60% of radio-tagged fish, with a decline in its use corresponding to increased distance of tagging location. A broader analysis of four tributaries in the Colorado River basin, including McElmo Creek, found photoperiod and temperature of tributary and mainstem rivers were the most important variables in determining migration timing, but tributary and mainstem discharge also aided in classification success. The largest tributary, the Little Colorado River, had more residential fish or fish that stayed for longer periods (median = 30 days), whereas McElmo Creek fish stayed an average of just 10 days in 2022. Our results generally suggest that higher discharge, across years or across sites, results in extended use of tributaries by flannelmouth suckers. Conservation actions that limit water extraction and maintain natural flow regimes in tributaries, while maintaining open connection with mainstem rivers, may benefit migratory species, including flannelmouth suckers.

Quantifying the conservation status and abundance trends of wildlife communities with detection-nondetection data.

Effective conservation requires understanding species' abundance patterns and demographic rates across space and time. Ideally, such knowledge should be available for whole communities because variation in species' dynamics can elucidate factors leading to biodiversity losses. However, collecting data to simultaneously estimate abundance and demographic rates of communities of species is often prohibitively time intensive and expensive. We developed a multispecies dynamic N-occupancy model to estimate unbiased, community-wide relative abundance and demographic rates. In this model, detection-nondetection data (e.g., repeated presence-absence surveys) are used to estimate species- and community-level parameters and the effects of environmental factors. To validate our model, we conducted a simulation study to determine how and when such an approach can be valuable and found that our multispecies model outperformed comparable single-species models in estimating abundance and demographic rates in many cases. Using data from a network of camera traps across tropical equatorial Africa, we then used our model to evaluate the statuses and trends of a forest-dwelling antelope community. We estimated relative abundance, rates of recruitment (i.e., reproduction and immigration), and apparent survival probabilities for each species' local population. The antelope community was fairly stable (although 17% of populations [species-park combinations] declined over the study period). Variation in apparent survival was linked more closely to differences among national parks than to individual species' life histories. The multispecies dynamic N-occupancy model requires only detection-nondetection data to evaluate the population dynamics of multiple sympatric species and can thus be a valuable tool for examining the reasons behind recent biodiversity loss.

La conservación efectiva requiere del entendimiento de los patrones de abundancia de las especies a lo largo del tiempo y el espacio. Sería ideal que dicho conocimiento estuviera disponible para todas las comunidades ya que la variación en la dinámica de las especies puede esclarecer los factores que llevan a la pérdida de la biodiversidad. Sin embargo, la recolección de información para estimar simultáneamente las tasas demográficas y de abundancia de las comunidades de especies con frecuencia es cara y consume tiempo. Desarrollamos un modelo multiespecies dinámico de ocupación-N para estimar la tasa demográfica y de abundancia relativas sin sesgos y en toda la comunidad. En este modelo usamos información de detección-no detección (p. ej.: censos repetidos de presencia-ausencia) para estimar los parámetros a nivel comunitario y de especie y los efectos de los factores ambientales. Para validar nuestro modelo, realizamos un estudio de simulación para determinar cómo y cuándo dicha estrategia puede ser valiosa y descubrimos que nuestro modelo multiespecies superó a los modelos comparables de una sola especie en la estimación de las tasas demográficas y de abundancia en muchos casos. Usamos nuestro modelo con datos de una red de cámaras trampa ubicadas a lo largo de África ecuatorial para evaluar los estados y tendencias de una comunidad forestal de antílopes. Estimamos la abundancia relativa, tasa de reclutamiento (es decir, reproducción e inmigración) y las probabilidades de supervivencia aparente para la población local de cada especie. La comunidad de antílopes fue bastante estable (aunque el 17% de las poblaciones [combinaciones especie-parque] declinaron durante el periodo de estudio). La variación en la supervivencia aparente estuvo vinculada con mayor cercanía a las diferencias entre los parques nacionales que a la historia de vida de cada especie individual. El modelo multiespecies dinámico de ocupación-N requiere solamente información de detección-no detección para evaluar las dinámicas poblacionales de muchas especies simpátricas y por lo tanto puede ser una herramienta valiosa para examinar las razones detrás de la pérdida reciente de la biodiversidad.

Water storage decisions will determine the distribution and persistence of imperiled river fishes.

Managing the world's freshwater supply to meet societal and environmental needs in a changing climate is one of the biggest challenges for the 21st century. Dams provide water security; however, the allocation of dwindling water supply among reservoirs could exacerbate or ameliorate the effects of climate change on aquatic communities. Here, we show that the relative sensitivity of river thermal regimes to direct impacts of climate change and societal decisions concerning water storage vary substantially throughout a river basin. In the absence of interspecific interactions, future Colorado River temperatures would appear to benefit both endemic and nonnative fish species. However, endemic species are already declining or extirpated in locations where their ranges overlap with warmwater nonnatives and changes in water storage may lead to warming in some of the coolest portions of the river basin, facilitating further nonnative expansion. Integrating environmental considerations into ongoing water storage negotiations may lead to better resource outcomes than mitigating nonnative species impacts after the fact.

A need for speed in Bayesian population models: a practical guide to marginalizing and recovering discrete latent states.

Bayesian population models can be exceedingly slow due, in part, to the choice to simulate discrete latent states. Here, we discuss an alternative approach to discrete latent states, marginalization, that forms the basis of maximum likelihood population models and is much faster. Our manuscript has two goals: (1) to introduce readers unfamiliar with marginalization to the concept and provide worked examples and (2) to address topics associated with marginalization that have not been previously synthesized and are relevant to both Bayesian and maximum likelihood models. We begin by explaining marginalization using a Cormack-Jolly-Seber model. Next, we apply marginalization to multistate capture-recapture, community occupancy, and integrated population models and briefly discuss random effects, priors, and pseudo-R2 . Then, we focus on recovery of discrete latent states, defining different types of conditional probabilities and showing how quantities such as population abundance or species richness can be estimated in marginalized code. Last, we show that occupancy and site-abundance models with auto-covariates can be fit with marginalized code with minimal impact on parameter estimates. Marginalized code was anywhere from five to >1,000 times faster than discrete code and differences in inferences were minimal. Discrete latent states and fully conditional approaches provide the best estimates of conditional probabilities for a given site or individual. However, estimates for parameters and derived quantities such as species richness and abundance are minimally affected by marginalization. In the case of abundance, marginalized code is both quicker and has lower bias than an N-augmentation approach. Understanding how marginalization works shrinks the divide between Bayesian and maximum likelihood approaches to population models. Some models that have only been presented in a Bayesian framework can easily be fit in maximum likelihood. On the other hand, factors such as informative priors, random effects, or pseudo-R2 values may motivate a Bayesian approach in some applications. An understanding of marginalization allows users to minimize the speed that is sacrificed when switching from a maximum likelihood approach. Widespread application of marginalization in Bayesian population models will facilitate more thorough simulation studies, comparisons of alternative model structures, and faster learning.

Brackish Tidal Marsh Management and the Ecology of a Declining Freshwater Turtle.

Water management practices in tidal marshes of the San Francisco Bay Estuary, California are often aimed at increasing suitable habitat for threatened fish species and sport fishes. However, little is known about how best to manage habitat for other sensitive status species like the semiaquatic freshwater Western Pond Turtle (Actinemys marmorata) that is declining throughout much of its range. Here, we examined the basking activity, abundance, survival, and growth of Western Pond Turtles at two brackish water study sites in Suisun Marsh, California that differed in how they were managed, with one having passive management (i.e., no active water regulation) and another having active management (i.e., water regulated for seasonal hunting). Our results revealed that basking activity was greatest when salinity, water stage, and air temperatures were low, shortwave radiation was high, and wind levels were intermediate. These preferred habitat characteristics often reflected conditions that were naturally maintained at the passively managed, muted tidal site. We also found that turtles were more abundant and had higher survival rates in the passively managed habitat compared to the actively managed habitat (201-323 turtles/km2 and 96% survival versus 11-135 turtles/km2 and 77% survival, respectively). Finally, characteristic growth constants from von Bertalanffy models showed that turtles grew more quickly in passively managed habitat compared to the actively managed habitat. Our results suggest that management strategies for this sensitive status species may be more effective if they protect passively managed muted tidal systems that limit or delay extreme cycles of salinity and water levels and conserve elevated terrestrial buffer zones adjacent to muted and full tidal systems.

The past and future roles of competition and habitat in the range-wide occupancy dynamics of Northern Spotted Owls.

Slow ecological processes challenge conservation. Short-term variability can obscure the importance of slower processes that may ultimately determine the state of a system. Furthermore, management actions with slow responses can be hard to justify. One response to slow processes is to explicitly concentrate analysis on state dynamics. Here, we focus on identifying drivers of Northern Spotted Owl (Strix occidentalis caurina) territorial occupancy dynamics across 11 study areas spanning their geographic range and forecasting response to potential management actions. Competition with Barred Owls (Strix varia) has increased Spotted Owl territory extinction probabilities across all study areas and driven recent declines in Spotted Owl populations. Without management intervention, the Northern Spotted Owl subspecies will be extirpated from parts of its current range within decades. In the short term, Barred Owl removal can be effective. Over longer time spans, however, maintaining or improving habitat conditions can help promote the persistence of northern spotted owl populations. In most study areas, habitat effects on expected Northern Spotted Owl territorial occupancy are actually greater than the effects of competition from Barred Owls. This study suggests how intensive management actions (removal of a competitor) with rapid results can complement a slower management action (i.e., promoting forest succession).

Inferring species interactions through joint mark-recapture analysis.

Introduced species are frequently implicated in declines of native species. In many cases, however, evidence linking introduced species to native declines is weak. Failure to make strong inferences regarding the role of introduced species can hamper attempts to predict population viability and delay effective management responses. For many species, mark-recapture analysis is the more rigorous form of demographic analysis. However, to our knowledge, there are no mark-recapture models that allow for joint modeling of interacting species. Here, we introduce a two-species mark-recapture population model in which the vital rates (and capture probabilities) of one species are allowed to vary in response to the abundance of the other species. We use a simulation study to explore bias and choose an approach to model selection. We then use the model to investigate species interactions between endangered humpback chub (Gila cypha) and introduced rainbow trout (Oncorhynchus mykiss) in the Colorado River between 2009 and 2016. In particular, we test hypotheses about how two environmental factors (turbidity and temperature), intraspecific density dependence, and rainbow trout abundance are related to survival, growth, and capture of juvenile humpback chub. We also project the long-term effects of different rainbow trout abundances on adult humpback chub abundances. Our simulation study suggests this approach has minimal bias under potentially challenging circumstances (i.e., low capture probabilities) that characterized our application and that model selection using indicator variables could reliably identify the true generating model even when process error was high. When the model was applied to rainbow trout and humpback chub, we identified negative relationships between rainbow trout abundance and the survival, growth, and capture probability of juvenile humpback chub. Effects on interspecific interactions on survival and capture probability were strongly supported, whereas support for the growth effect was weaker. Environmental factors were also identified to be important and in many cases stronger than interspecific interactions, and there was still substantial unexplained variation in growth and survival rates. The general approach presented here for combining mark-recapture data for two species is applicable in many other systems and could be modified to model abundance of the invader via other modeling approaches.

Integrating count and detection-nondetection data to model population dynamics.

There is increasing need for methods that integrate multiple data types into a single analytical framework as the spatial and temporal scale of ecological research expands. Current work on this topic primarily focuses on combining capture-recapture data from marked individuals with other data types into integrated population models. Yet, studies of species distributions and trends often rely on data from unmarked individuals across broad scales where local abundance and environmental variables may vary. We present a modeling framework for integrating detection-nondetection and count data into a single analysis to estimate population dynamics, abundance, and individual detection probabilities during sampling. Our dynamic population model assumes that site-specific abundance can change over time according to survival of individuals and gains through reproduction and immigration. The observation process for each data type is modeled by assuming that every individual present at a site has an equal probability of being detected during sampling processes. We examine our modeling approach through a series of simulations illustrating the relative value of count vs. detection-nondetection data under a variety of parameter values and survey configurations. We also provide an empirical example of the model by combining long-term detection-nondetection data (1995-2014) with newly collected count data (2015-2016) from a growing population of Barred Owl (Strix varia) in the Pacific Northwest to examine the factors influencing population abundance over time. Our model provides a foundation for incorporating unmarked data within a single framework, even in cases where sampling processes yield different detection probabilities. This approach will be useful for survey design and to researchers interested in incorporating historical or citizen science data into analyses focused on understanding how demographic rates drive population abundance.

Benefits of the destinations, not costs of the journeys, shape partial migration patterns.

The reasons that lead some animals to seasonally migrate, and others to remain in the same area year-round, are poorly understood. Associations between traits, such as body size, and migration provide clues. For example, larger species and individuals are more likely to migrate. One explanation for this size bias in migration is that larger animals are capable of moving faster (movement hypothesis). However, body size is linked to many other biological processes. For instance, the energetic balances of larger animals are generally more sensitive to variation in food density because of body size effects on foraging and metabolism and this sensitivity could drive migratory decisions (forage hypothesis). Identifying the primary selective forces that drive migration ultimately requires quantifying fitness impacts over the full annual migratory cycle. Here, we develop a full annual migratory cycle model from metabolic and foraging theory to compare the importance of the forage and movement hypotheses. We parameterize the model for Galapagos tortoises, which were recently discovered to be size-dependent altitudinal migrants. The model predicts phenomena not included in model development including maximum body sizes, the body size at which individuals begin to migrate, and the seasonal timing of migration and these predictions generally agree with available data. Scenarios strongly support the forage hypothesis over the movement hypothesis. Furthermore, male Galapagos tortoises on Santa Cruz Island would be unable to grow to their enormous sizes without access to both highlands and lowlands. Whereas recent research has focused on links between traits and the migratory phases of the migratory cycle, we find that effects of body size on the non-migratory phases are far more important determinants of the propensity to migrate. Larger animals are more sensitive to changing forage conditions than smaller animals with implications for maintenance of migration and body size in the face of environmental change.

Dynamic N-occupancy models: estimating demographic rates and local abundance from detection-nondetection data.

Occupancy modeling is a widely used analytical technique for assessing species distributions and range dynamics. However, occupancy analyses frequently ignore variation in abundance of occupied sites, even though site abundances affect many of the parameters being estimated (e.g., extinction, colonization, detection probability). We introduce a new model ("dynamic N-occupancy") capable of providing accurate estimates of local abundance, population gains (reproduction/immigration), and apparent survival probabilities while accounting for imperfect detection using only detection/nondetection data. Our model utilizes heterogeneity in detection based on variations in site abundances to estimate latent demographic rates via a dynamic N-mixture modeling framework. We validate our model using simulations across a wide range of values and examine the data requirements, including the number of years and survey sites needed, for unbiased and precise estimation of parameters. We apply our model to estimate spatiotemporal heterogeneity in abundances of barred owls (Strix varia) within a recently invaded region in Oregon (USA). Estimates of apparent survival and population gains are consistent with those from a nearby radio-tracking study and elucidate how barred owl abundances have increased dramatically over time. The dynamic N-occupancy model greatly improves inferences on individual-level population processes from occupancy data by explicitly modeling the latent population structure.

Allometric and temporal scaling of movement characteristics in Galapagos tortoises.

Understanding how individual movement scales with body size is of fundamental importance in predicting ecological relationships for diverse species. One-dimensional movement metrics scale consistently with body size yet vary over different temporal scales. Knowing how temporal scale influences the relationship between animal body size and movement would better inform hypotheses about the efficiency of foraging behaviour, the ontogeny of energy budgets, and numerous life-history trade-offs. We investigated how the temporal scaling of allometric patterns in movement varies over the course of a year, specifically during periods of motivated (directional and fast movement) and unmotivated (stationary and tortuous movement) behaviour. We focused on a recently diverged group of species that displays wide variation in movement behaviour - giant Galapagos tortoises (Chelonoidis spp.) - to test how movement metrics estimated on a monthly basis scaled with body size. We used state-space modelling to estimate seven different movement metrics of Galapagos tortoises. We used log-log regression of the power law to evaluate allometric scaling for these movement metrics and contrasted relationships by species and sex. Allometric scaling of movement was more apparent during motivated periods of movement. During this period, allometry was revealed at multiple temporal intervals (hourly, daily and monthly), with values observed at daily and monthly intervals corresponding most closely to the expected one-fourth scaling coefficient, albeit with wide credible intervals. We further detected differences in the magnitude of scaling among taxa uncoupled from observed differences in the temporal structuring of their movement rates. Our results indicate that the definition of temporal scales is fundamental to the detection of allometry of movement and should be given more attention in movement studies. Our approach not only provides new conceptual insights into temporal attributes in one-dimensional scaling of movement, but also generates valuable insights into the movement ecology of iconic yet poorly understood Galapagos giant tortoises.

Flow management and fish density regulate salmonid recruitment and adult size in tailwaters across western North America.

Rainbow and brown trout have been intentionally introduced into tailwaters downriver of dams globally and provide billions of dollars in economic benefits. At the same time, recruitment and maximum length of trout populations in tailwaters often fluctuate erratically, which negatively affects the value of fisheries. Large recruitment events may increase dispersal downriver where other fish species may be a priority (e.g., endangered species). There is an urgent need to understand the drivers of trout population dynamics in tailwaters, in particular the role of flow management. Here, we evaluate how flow, fish density, and other physical factors of the river influence recruitment and mean adult length in tailwaters across western North America, using data from 29 dams spanning 1-19 years. Rainbow trout recruitment was negatively correlated with high annual, summer, and spring flow and dam latitude, and positively correlated with high winter flow, subadult brown trout catch, and reservoir storage capacity. Brown trout recruitment was negatively correlated with high water velocity and daily fluctuations in flow (i.e., hydropeaking) and positively correlated with adult rainbow trout catch. Among these many drivers, rainbow trout recruitment was primarily correlated with high winter flow combined with low spring flow, whereas brown trout recruitment was most related to high water velocity. The mean lengths of adult rainbow and brown trout were influenced by similar flow and catch metrics. Length in both species was positively correlated with high annual flow but declined in tailwaters with high daily fluctuations in flow, high catch rates of conspecifics, and when large cohorts recruited to adult size. Whereas brown trout did not respond to the proportion of water allocated between seasons, rainbow trout length increased in rivers that released more water during winter than in spring. Rainbow trout length was primarily related to high catch rates of conspecifics, whereas brown trout length was mainly related to large cohorts recruiting to the adult size class. Species-specific responses to flow management are likely attributable to differences in seasonal timing of key life history events such as spawning, egg hatching, and fry emergence.

The roles of competition and habitat in the dynamics of populations and species distributions.

The role of competition in structuring biotic communities at fine spatial scales is well known from detailed process-based studies. Our understanding of competition's importance at broader scales is less resolved and mainly based on static species distribution maps. Here, we bridge this gap by examining the joint occupancy dynamics of an invading species (Barred Owl, Strix varia) and a resident species (Northern Spotted Owl, Strix occidentalis caurina) in a 1000-km study area over a 22-year period. Past studies of these competitors have focused on the dynamics of one species at a time, hindering efforts to parse out the roles of habitat and competition and to forecast the future of the resident species. In addition, while these studies accounted for the imperfect detection of the focal species, no multi-season analysis of these species has accounted for the imperfect detection of the secondary species, potentially biasing inference. We analyzed survey data using models that combine the general multistate-multi-season occupancy modeling framework with autologistic modeling, allowing us to account for important aspects of our study system. We found that local extinction probability increases for each species when the other is present; however, the effect of the invader on the resident is greater. Although the species prefer different habitats, these habitats are highly correlated at the patch scale, and the impacts of invader on the resident are greatest in patches that would otherwise be optimal. As a consequence, competition leads to a weaker relationship between habitat and Northern Spotted Owl occupancy. Colonization and extinction rates of the invader are closely related to neighborhood occupancy, and over the first half of the study the availability of colonists limited the rate of population growth. Competition is likely to exclude the resident species, both through its immediate effects on local extinction and by indirectly lowering colonization rates as Northern Spotted Owl occupancy declines. Our analysis suggests that dispersal limitation affects both the invasion dynamics and the scale at which the effects of competition are observed. We also provide predictions regarding the potential costs and benefits of managing Barred Owl populations at different target levels.