Ecological success of no-take marine protected areas: Using population dynamics theory to inform a global meta-analysis.

Adaptively managing marine protected areas (MPAs) requires accurately assessing whether established MPAs are achieving their goals of protecting and conserving biomass, especially for harvested populations. Ecological MPA assessments commonly compare inside of the MPA to a reference point outside of and/or before implementation (i.e., calculating "response ratios"). Yet, MPAs are not simple ecological experiments; by design, protected populations interact with those outside, and population dynamic responses can be nonlinear. This complicates assessment interpretations. Here, we used a two-patch population model to explore how MPA response ratios (outside-inside, before-after, and before-after-control-impact [BACI]) for fished populations behave under different conditions, like whether the population is receiving a sustainable larval supply or if it is declining despite protection from harvest. We then conducted a Bayesian evaluation of MPA effects on fish and invertebrate populations based on data collected from 82 published studies on 264 no-take MPAs worldwide, using the results of an earlier global meta-analysis as priors. We considered the effects of calculating different summary metrics on these results, drawing on the theoretical insights from our population model as a comparative framework. We demonstrate that not all response ratio comparison types provide the same information: For example, outside-inside and BACI comparisons can fail to detect population decline within MPAs, whereas before-after comparisons likely detect that pattern. Considering these limitations, we nonetheless found that MPAs globally are producing positive outcomes, with on average greater biomass, density, and organism size within their boundaries than reference sites. However, only a small portion of studies (18 of 82) provided the temporal data necessary to determine that protection, on average, has led to increased abundance of populations within MPAs over time. These findings demonstrate the importance of considering the underlying system dynamics when assessing MPA effects. Assuming that large outside-inside or BACI response ratios always reflect large and net positive conservation effects may lead to misleading conclusions, we recommend that: (1) when assessing specific MPA effects, empirical findings be considered alongside theoretical knowledge relevant to that MPA system, and (2) management should respond to the local conditions and outcomes, rather than a blanket expectation for positive MPA effects.

Pulse disturbances in age-structured populations: Life history predicts initial impact and recovery time.

Understanding population responses to discrete 'pulsed' environmental disturbances is essential to conservation and adaptive management. Populations of concern can be driven to low levels by disturbance, and understanding interspecific differences in recovery trajectories is necessary for evaluating management options. We analysed single-species models to investigate the demographic and management factors determining the two components of population 'resilience': the magnitude of initial impact on population abundance, and duration of the recovery time. We simulated age-structured populations with density-dependent recruitment, subjected to a pulse disturbance consisting of a period of increased mortality of either the juvenile age class or all age classes, and calculated both impact and return time. For illustration, we used demographic parameters from a suite of 16 fish species. We formulated the model as a renewal equation, allowing us to describe disturbance impacts mathematically as a convolution. We also included nonlinear dynamics, representing populations that recover to a steady state; this is more realistic (in most cases) than prior analyses of resilience in linear models without density-dependence. When the disturbance affected only one or a few young age-classes, longevity was the major life-history determinant of impact and recovery time. Shorter-lived species endured greater impacts when disturbed because each age class is a greater proportion of the population. However, shorter-lived species also had faster recovery times, for the same reason. When disturbance affected adult age-classes, the impact was more immediate and no longer affected by species' longevity, though the effect of longevity on recovery time remained. These results improve our understanding of interspecific differences in resilience and increase our ability to make predictions for adaptive management. Additionally, formulating the problem as a renewal equation and using mathematical convolutions allows us to quantify how disturbances with different time courses (not just an immediate, constant level of disturbance but gradually increasing or decreasing levels of disturbance) would have different effects on population resilience: delayed responses for species in which biomass is concentrated in older age classes, and for disturbances that become progressively more severe.

Entender las respuestas de la población a perturbaciones ambientales, específicamente a pulsadas individuales, es esencial para la conservación y la gestión adaptativa. Las poblaciones de interés pueden reducirse a niveles bajas debido a la perturbación, y es necesario entender las diferencias interespecíficas en las trayectorias de recuperación para evaluar las opciones de gestión. Analizamos modelos para especies individuales para investigar los factores demográficos y de gestión que determinan los dos componentes de la 'resiliencia' de la población: la magnitud del impacto inicial sobre la abundancia de la población y la duración del tiempo de recuperación. Simulamos poblaciones estructuradas por edad con reclutamiento que depende de la densidad, las sometimos a una perturbación pulsada que consiste en un período de mayor mortalidad del grupo etário juvenil o de todos los grupos etários, y calculamos tanto el impacto como el tiempo de retorno. A modo de ilustración, utilizamos parámetros demográficos de un conjunto de 16 especies de peces. Formulamos el modelo como una ecuación de renovación, lo que nos permite describir matemáticamente los impactos de las perturbaciones como una convolución. También incluimos dinámicas no lineales que representan poblaciones que se recuperan hacia un estado estable; esto es más realista (en la mayoría de los casos) que los análisis previos de resiliencia en modelos lineales sin la dependencia de la densidad. Cuando la perturbación ha afectado a uno o a algunos pocos grupos etários jóvenes, la longevidad fue el principal determinante de la historia de vida del impacto y el tiempo de recuperación. Las especies de vida más corta sufrieron mayores impactos cuando fueron perturbadas porque cada grupo etáreo representa una mayor proporción de la población. Sin embargo, las especies con vidas más cortas también tuvieron tiempos de recuperación más rápidos, por la misma razón. Cuando la perturbación afectó a los grupos etários adultos, el impacto fue más inmediato y ya no se vio afectado por la longevidad de las especies, aunque se mantuvo el efecto de la longevidad sobre el tiempo de recuperación. Estos resultados mejoran nuestra comprensión de las diferencias interespecíficas de la resiliencia y aumentan nuestra capacidad para hacer predicciones con fin a la gestión adaptativa. Además, formular el problema como una ecuación de renovación y usar convoluciones matemáticas nos permite cuantificar cómo las perturbaciones con distintos lapsos de tiempo (no solo un nivel de perturbación constante e inmediato, sino niveles de perturbación que aumentan o disminuyen gradualmente) tendrían diferentes efectos sobre la resiliencia de la población: respuestas tardías para especies en las que la biomasa se concentra en grupos etários de mayor edad y para perturbaciones que se vuelven progresivamente más severas.

Pelagic marine refugia and climatically sensitive areas in an eastern boundary current upwelling system.

Refugia are areas relatively buffered from contemporary climate change that enable the persistence of valued physical, ecological, or sociocultural resources. Spatially identifying refugia is important for conservation and applied management. Yet the concept of refugia has not been broadly extended to marine ecosystems. Here, we analyze data from a unique and long-term (1999-2015) standardized survey of pelagic marine and anadromous species off Oregon and Washington in the northern California Current to identify such refugia. We use quantitative approaches to assess locations with high species richness and community persistence relative to local and basin-scale environmental fluctuations. We have identified a potential climate change refugial zone along the continental shelf of Washington State in the Northeastern Pacific Ocean, characterized by a species-rich community with low interannual temporal community change. This region contrasts with adjacent areas to the south and offshore that have lower species richness, and higher temporal species community change. Also, using spatially variant generalized additive mixed models, we identify areas with species compositions that are more influenced by basin-scale climatic fluctuations than others. We propose that upwelling regions with retentive topographic features, such as wide continental shelves, can function as marine refugia for pelagic fauna, whereas offshore locations are potentially more climatically sensitive and experience high temporal change in species composition. Further identification of these marine refugia using in situ data for pelagic biodiversity and climatically sensitive areas can help guide management in the face of inevitable climatically driven change.

Eight decades of sampling reveal a contemporary novel fish assemblage in coastal nursery habitats.

In order to adequately monitor biodiversity trends through time and their responses to natural or anthropogenic impacts, researchers require long time series that are often unavailable. This general lack of datasets that are several decades or longer makes establishing a background or baseline of diversity metrics difficult - especially when attempting to understand species composition changes against a backdrop of climate and ecological variability. Here, we present an analysis of a community of juvenile nearshore fishes based on nearly 8 decades of highly standardized Norwegian survey records. Using multivariate statistical techniques, we: (i) characterize the change in taxonomic community composition through time, (ii) determine whether there has been an increase in warm-water affinity species relative to their cold water affinity counterparts, and (iii) characterize the temporal change in the species' functional trait assemblage. Our results strongly indicate a shift toward a novel fish assemblage between the late 1990s and 2000s. The context of changes within the most recent two decades is in stark contrast to those during the 1960s and 1970s, but similar to those during the previous warm period during the 1930s and 1940s. This novel assemblage is tightly linked to the warming temperatures in the region portrayed by the increased presence of warm-water species and a higher incidence of pelagic, planktivorous species. The results indicate a clear influence of ocean temperature on the region's juvenile fish community that points to climate-mediated effects on the species assemblages of an important fish nursery area.

The use of mesoscale eddies by juvenile loggerhead sea turtles (Caretta caretta) in the southwestern Atlantic.

Marine animals, such as turtles, seabirds and pelagic fishes, are observed to travel and congregate around eddies in the open ocean. Mesoscale eddies, large swirling ocean vortices with radius scales of approximately 50-100 km, provide environmental variability that can structure these populations. In this study, we investigate the use of mesoscale eddies by 24 individual juvenile loggerhead sea turtles (Caretta caretta) in the Brazil-Malvinas Confluence region. The influence of eddies on turtles is assessed by collocating the turtle trajectories to the tracks of mesoscale eddies identified in maps of sea level anomaly. Juvenile loggerhead sea turtles are significantly more likely to be located in the interiors of anticyclones in this region. The distribution of surface drifters in eddy interiors reveals no significant association with the interiors of cyclones or anticyclones, suggesting higher prevalence of turtles in anticyclones is a result of their behavior. In the southern portion of the Brazil-Malvinas Confluence region, turtle swimming speed is significantly slower in the interiors of anticyclones, when compared to the periphery, suggesting that these turtles are possibly feeding on prey items associated with anomalously low near-surface chlorophyll concentrations observed in those features.