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Climate change is affecting marine ecosystems in many ways including rising temperatures and ocean acidification. From 2014-2016, an extensive marine heatwave extended along the west coast of North America and had devastating effects on numerous species during this period, including bull kelp (Nereocystis luetkeana). Bull kelp is an important foundation species in coastal ecosystems that can be affected by marine heat waves and ocean acidification, however these impacts have not been investigated on sensitive early life stages. To determine the effects of changing temperatures and carbonate levels on Northern California’s bull kelp populations, we collected sporophylls from mature bull kelp individuals in Point Arena, CA. At the Bodega Marine Laboratory, we released spores from field-collected bull kelp, and cultured microscopic gametophytes in a common garden experiment with a fully factorial design crossing modern conditions (11.63±0.54°C and pH 7.93±0.26) with observed extreme climate conditions (15.56±0.83°C and 7.64±0.32pH). Our results found that both increased temperature and decreased pH influenced growth and egg production of bull kelp microscopic stages. Increased temperature generally resulted in decreased gametophyte survival and offspring production. In contrast, decreased pH had less of an effect, but generally resulted in increased gametophyte survival and offspring production. Additionally, we found that increased temperature significantly impacted reproductive timing by causing female gametophytes to produce offspring earlier than under ambient temperature conditions. Our findings inform better predictions of the impacts of climate change on coastal ecosystems as well as provide key insight into environmental dynamics regulating the bull kelp lifecycle.

Adaptation across environmental gradients has been demonstrated in numerous systems with extensive dispersal, despite high gene flow and consequently low genetic structure. The speed and mechanisms by which such adaptation occurs remain poorly resolved, but are critical to understanding species spread and persistence in a changing world. Here, we investigate these mechanisms in the European green crab Carcinus maenas, a globally distributed invader. We focus on a northwestern Pacific population that spread across12 degrees of latitude in 10 years from a single source, following its introduction35 years ago. Using six locations spanning1500 km, we examine genetic structure using 9376 single nucleotide polymorphisms (SNPs). We find high connectivity among five locations, with significant structure between these locations and an enclosed lagoon with limited connectivity to the coast. Among the five highly connected locations, the only structure observed was a cline driven by a handful of SNPs strongly associated with latitude and winter temperature. These SNPs are almost exclusively found in a large cluster of genes in strong linkage disequilibrium that was previously identified as a candidate for cold tolerance adaptation in this species. This region may represent a balanced polymorphism that evolved to promote rapid adaptation in variable environments despite high gene flow, and which now contributes to successful invasion and spread in a novel environment. This research suggests an answer to the paradox of genetically depauperate yet successful invaders: populations may be able to adapt via a few variants of large effect despite low overall diversity.

Using species distribution models (SDMs), we predicted the distribution of 170 plant species under different climatic scenarios (current and future climatic conditions) and used this information to create invasion risk maps to identify potential invasion hot spots in California. The risk of invasion by individual species was also assessed using species’ predicted area in combination with some biological traits associated with invasiveness (growth form, reproduction mechanisms, and age of maturity). A higher number of species would find suitable climatic conditions along the coast; the Central Western (CW) and South Western (SW) were ecoregions where a higher number of species were predicted. Overall, hot spots of species distribution were similar under current and future climatic conditions; however, individual species’ predicted area (increase or decrease) was variable depending on the climate change scenario and the greenhouse gas emission. Out of the 170 species assessed, 22% ranked as high-risk species, with herbs, grasses, and vines accounting for 78% within this risk class, and a high proportion (67%) of Asteraceae species ranked as high risk. This study suggests that current climatic conditions of the central and south coastal regions of California would be considered as hot spots of new invasions, and for some species this risk might increase with hotter and drier future climatic conditions.

Marine foundation species are critical to the structure and resilience of coastal ecosystems and provide key ecosystem services. Since many have suffered severe population declines, restoration of foundation species has been undertaken worldwide. The Olympia oyster (Ostrea lurida) is a foundation species, and the restoration of depleted populations is a priority for maintaining ecosystem function of estuaries along the west coast of North America. Here, we synthesize all native oyster restoration projects conducted from California, USA, to British Columbia, Canada, and analyze project goals, methods, and outcomes. Currently, restoration projects are spread unevenly across the species’ range, driven by locally varying goals and implemented with contrasting approaches. We highlight the value of regional strategic planning and decision support tools to evaluate project design and methods for restoration, including the selection of substrates and the targeted use of aquaculture. We recommend future investment in larger projects, which our analysis found were more cost-effective, but which have been relatively rare for this species. We also recommend that funders support monitoring over broader temporal and spatial scales than in the past to better characterize long-term effects of restoration on oyster populations and the services they provide beyond the project footprint. We found that most projects successfully supported native oysters and engaged local communities, and recommend similar efforts to continue to enhance understanding of Olympia oysters, which remain unfamiliar to many coastal residents. We believe that the results of this synthesis are broadly applicable to marine foundation species generally, and can inform restoration and conservation efforts worldwide.

The introduction of nonnative invasive pests is among the many threats facing coastal ecosystems worldwide. Managing these pests often requires considerable effort and resources, and community scientists can be essential for providing the capacity needed for management and monitoring activities. In response to the invasion of a Northern California estuary by the predatory European green crab, a collaborative team of academic researchers and community scientists initiated a local eradication program. The green crab is listed among the world's 100 worst invaders, and threatened both native species and commercial shellfisheries. The program dramatically reduced the green crab population over a 5-year period, but it rebounded, which necessitated a switch in project goals from eradication to population suppression. Community scientists were essential for facilitating this switch by providing the necessary capacity to quantify population characteristics and maintain reduced crab populations. The result was a sustainable program that successfully maintained low green crab densities, which will likely improve habitat for native species.

Strong competitive ability of introduced plant species has been frequently stated as a key factor promoting successful invasion. The dynamics of invasive species may depend on their abilities to compete for resources and exploit disturbances relative to native species. This study compares the growth performance of the invasive blackberry (Rubus niveusThunb.) with four of the most common woody native species of the Scalesia forest in the Galapagos Islands. Using a series of greenhouse and field studies, the growth rate and biomass production of native species alone and in competition withR. niveuswas compared under different water and light stress conditions.Rubus niveusshowed a faster growth rate and biomass production than the native species as well as a broad tolerance to light and water stress conditions. Competitive ability was also assessed by looking at the seedbank and regeneration processes after herbicide control in the field. Although the number ofR. niveusseedlings that germinated from the soil samples was considerably larger than that of native species, recruitment of the invasive on the experimental plots was limited. Overall,R. niveusshowed superior competitive ability over native species with comparable growth forms, suggesting a possible mechanism that enablesR. niveusto successfully invade a wide range of habitats in the Galapagos Islands. Implementing an integrated management strategy including biological control, seedbank reduction, and active revegetation, should improve the efficiency ofR. niveusmanagement, enabling the restoration of degraded vegetation in the Galapagos.

Estuaries represent steep stress gradients for aquatic organisms, with abiotic stress due to temperature and salinity typically increasing with distance into estuary. Invertebrate communities and their predators are strongly influenced by these stress gradients. The environmental stress model predicts that the importance of predation in structuring communities decreases with increasing environmental stress. Estuaries contain a stress gradient for marine organisms this includes salinity, temperature, and other abiotic properties. Additionally, estuaries are hotspots for biological invasions; increased stress tolerance among non-native species could change the predictions of the environmental stress model. In this study, we investigate how introduced species alter the predictions of the environmental stress model by examining the effects of predators on sessile invertebrates across an estuarine gradient. To do this, we deployed recruitment plates across the estuarine gradient of Tomales Bay, California, USA using various caging treatments over the summer of 2019. We found that the effect of predation changed across sites, with the mid-estuary site experiencing the greatest reductions in prey abundance and prey species richness when exposed to predators. This was likely to be due to higher proportions of non-native prey and predator taxa mid-estuary, including solitary ascidians, which are highly susceptible to predation. Overall, predation did not follow the predictions of the environmental stress model, but rather followed the abundance of functional groups with non-native species, whose distribution could be mediated by environmental stress gradients. We suggest that this may be a general result and that communities subject to large numbers of stress-tolerant invaders may have high rates of consumption in high stress areas, contrasting predictions by previous models.

Invasive species, both plants and animals, are a long-standing threat to the National Parks of the United States. For nearly two decades the National Park Service has implemented a service-wide invasive plant management program without a commensurate program focusing on invasive animals. While individual park units are struggling to sufficiently address the threat of invasive terrestrial and aquatic animal species, a system-wide effort could bring the resources and capacity needed to address a challenge of this magnitude. We present our key findings from a detailed review about invasive animal species and their management by the National Park Service. We assert that the global threat of invasive animals substantially undermines the National Park Service mission. Coordinated action could improve the ability for the National Park Service to meet the challenge, and partnering with neighboring agencies and invasive species networks outside of the National Park Service is essential for success. Public engagement, cooperation and support is also critical and can be accomplished through strategic engagement efforts. Finally, the National Park Service would benefit from the development of an invasive animal program that includes structured decision support, adaptive management and monitoring, the organizational structure to meet the highest needs, and capitalizing on the significant opportunities that exist through the appropriate use of emerging technologies.

Antipredator behavior is known to have a strong effect on prey population dynamics. While there have been many studies of antipredator behavior in population dynamic models, none have examined how antipredator behavior interacts with predator foraging mode. To examine this process, we incorporated predator and prey velocities into a simple tritrophic food chain. In this model, antipredator behavior allows prey to respond to predators by slowing their velocity in response to predator density. Prey can slow their velocity to hide from predators, but this in turn reduces their ability to consume resources, creating a trade-off between hiding and foraging. We examined the effects of both fast-moving “mobile” predators and slow-moving “sit-and-wait” predators on equilibrium prey density and amplitude of predator-prey cycles. We found that antipredator behavior was ineffective against mobile predators, but it was very effective against sit-and-wait predators. Antipredator responses to sit-and-wait predators reduced top-down control and allowed prey density to increase with increased carrying capacity. Furthermore, antipredator responses to sit-and-wait predators eliminated population cycles within the community, whereas antipredator behavior had no effect on population cycles within mobile predator communities. Therefore, our model demonstrates predator foraging mode must be taken into account when examining predator-prey cycles. We discuss the potential implications of this model for invasive species and for trophic cascades.

As biological invasions continue to increase globally, eradication programs have been undertaken at significant cost, often without consideration of relevant ecological theory. Theoretical fisheries models have shown that harvest can actually increase the equilibrium size of a population, and uncontrolled studies and anecdotal reports have documented population increases in response to invasive species removal (akin to fisheries harvest). Both findings may be driven by high levels of juvenile survival associated with low adult abundance, often referred to as overcompensation. Here we show that in a coastal marine ecosystem, an eradication program resulted in stage-specific overcompensation and a 30-fold, single-year increase in the population of an introduced predator. Data collected concurrently from four adjacent regional bays without eradication efforts showed no similar population increase, indicating a local and not a regional increase. Specifically, the eradication program had inadvertently reduced the control of recruitment by adults via cannibalism, thereby facilitating the population explosion. Mesocosm experiments confirmed that adult cannibalism of recruits was size-dependent and could control recruitment. Genomic data show substantial isolation of this population and implicate internal population dynamics for the increase, rather than recruitment from other locations. More broadly, this controlled experimental demonstration of stage-specific overcompensation in an aquatic system provides an important cautionary message for eradication efforts of species with limited connectivity and similar life histories.

Understanding how the biological invasion is driven by environmental factors will improve model prediction and advance early detection, especially in the context of accelerating anthropogenic ecological changes. Although a large body of studies has examined how favorable environments promote biological invasions, a more comprehensive and mechanistic understanding of invasive species response to unfavorable/stressful conditions is still developing. Grass invasion has been problematic across the globe; in particular, C4 grass invaders, with high drought tolerance, adaptations to high temperatures, and high water use efficiency, could become more severe. Here, we conducted a rigorous microcosm experiment, with one of the most damaging invasive C4 grass - cogongrass (Imperata cylindrica), to explore how cogongrass responds to soil water and nutrient stress. We further integrated the results of the microcosm study with a species distribution model to (1) corroborate greenhouse results with field observations and (2) validate the robustness of our findings at sub-continental scales. Both the microcosm experiments and species distribution model agreed that soil water stress had a stronger impact on cogongrass than the nutrient one. New vegetative growth of cogongrass continued to be inhibited by the prior water stress. The significant water effect on cogongrass total biomass was supported by the finding that both allometric and biochemical traits of cogongrass did not show significant responses to the changes in water treatment. Different to the conventional wisdom that nutrient enrichment plays a bigger role in facilitating biological invasions, this study highlighted the possibility that water conditions may have a more substantial effect on some aggressive invaders. Therefore, an important implication of this study on biological conservation is that field managers might take advantage of the negative effect of global drought on some invasive species to increase the efficiency of their controlling efforts because invasive species may become more vulnerable under drought effect.

Conservation aquaculture is becoming an important tool to support the recovery of declining marine species and meet human needs. However, this tool comes with risks as well as rewards, which must be assessed to guide aquaculture activities and recovery efforts. Olympia oysters (Ostrea lurida) provide key ecosystem functions and services along the west coast of North America, but populations have declined to the point of local extinction in some estuaries. Here, we present a species-level, range-wide approach to strategically planning the use of aquaculture to promote recovery of Olympia oysters. We identified 12 benefits of culturing Olympia oysters, including identifying climate-resilient phenotypes that add diversity to growers’ portfolios. We also identified 11 key risks, including potential negative ecological and genetic consequences associated with the transfer of hatchery-raised oysters into wild populations. Informed by these trade-offs, we identified ten priority estuaries where aquaculture is most likely to benefit Olympia oyster recovery. The two highest scoring estuaries have isolated populations with extreme recruitment limitation—issues that can be addressed via aquaculture if hatchery capacity is expanded in priority areas. By integrating social criteria, we evaluated which project types would likely meet the goals of local stakeholders in each estuary. Community restoration was most broadly suited to the priority areas, with limited commercial aquaculture and no current community harvest of the species, although this is a future stakeholder goal. The framework we developed to evaluate aquaculture as a tool to support species recovery is transferable to other systems and species globally; we provide a guide to prioritizing local knowledge and developing recommendations for implementation by using transparent criteria. Our collaborative process engaging diverse stakeholders including managers, scientists, Indigenous Tribal representatives, and shellfish growers can be used elsewhere to seek win-win opportunities to expand conservation aquaculture where benefits are maximized for both people and imperiled species.

Burrowing animals can profoundly influence the structure of surrounding communities, as well as the performance of individual species. Changes in the community structure of burrowing animals or plants together with changing abiotic parameters could shift the influence of burrowers on surrounding habitats. For example, prior studies in salt marshes suggest that fiddler crabs stimulate cordgrass production, but leaf-grazing crabs suppress cordgrass production. Unfortunately, testing this prediction and others are impeded because few studies have examined crab impacts on the plant community and across multiple sites, multiple years, or both. This challenges our ability to predict how burrowing animals will influence plant community structure, and when and where these impacts will occur. We manipulated the densities of the dominant burrowing crabs in plant assemblages dominated by Pacific cordgrass (Spartina foliosa) and perennial pickleweed (Sarcocornia pacifica) at three sites in southern California for three years (2016, 2017, 2018). Crab impacts on plant community structure differed among each of our three sites. In contrast to our predictions, (1) leaf-grazing crabs (Pachygrapsus crassipes) had positive effects on cordgrass cover at one site and no effect on cordgrass production at a nearby site in the same marsh and (2) fiddler crabs (Uca crenulata) did not stimulate cordgrass production at another marsh. Because crabs affected traits of cordgrass, but not pickleweed, in the direction consistent with changes in cordgrass cover, we propose that marsh-specific crab effects on community structure were largely mediated through changes in cordgrass, as opposed to pickleweed. Importantly, crabs facilitated cordgrass during marsh-wide cordgrass loss, suggesting that crabs may mitigate environmental stress for this ecologically important plant. Because cordgrass abundance can be a critical measure of marsh functioning and is often a restoration target, we suggest that managing cordgrass populations would benefit from additional information about crab populations and their impacts among years, and among and within marshes.

Extreme climate events, such as drought, are becoming increasingly important drivers of plant community change, yet little is known about their impacts on invasive plants. Further, drought impacts may be altered by other anthropogenic stressors, such as eutrophication. We found drought dramatically reduced density of invasive Lepidium latifolium in salt marshes, and this die-back was mitigated by nutrient addition. In a 3-yr field experiment (2014-2016) conducted during an unprecedented drought (2012-2015), we tracked the effects of drought and nutrient addition on the plant community. We conducted this research at four salt marshes across a salinity gradient in the San Francisco Bay, California, USA. We manipulated paired native and invaded plots, one-half of which were treated monthly with N and P for 1.5 yr during the most intense period of the drought and one subsequent wet winter. In addition, we monitored unmanipulated L. latifolium-invaded transects within our freshest and most saline sites throughout the three years of our manipulative experiment and one additional wet winter. We documented a dramatic die-back of invasive L. latifolium during extreme drought, with reductions in stem density (52-100%) and height (17-47%) that were more severe at low salinity sites than high salinity sites. We found nutrient application lessened the effect of drought on L. latifolium stem density, but not height. In native plots, extreme drought reduced native plant cover (4-24%), but nutrient addition mitigated this impact. Interestingly, native plants in invaded plots did not suffer reductions in cover due to drought, perhaps because they were simultaneously benefiting from the die-back of the invader. Our results show drought negatively impacted both native and invasive plants and this impact was stronger on the invader, which experienced persistent declines two years after the end of the drought. However, by mitigating the effect of drought on invasive plants, nutrient addition potentially erased the advantage drought provided native plants over invasive plants under ambient nutrient conditions.

Recruitment of new propagules into a population can be a critical determinant of adult density. We examined recruitment dynamics in the Olympia oyster (Ostrea lurida), a species occurring almost entirely in estuaries. We investigated spatial scales of interannual synchrony across 37 sites in eight estuaries along 2500 km of Pacific North American coastline, predicting that high vs. low recruitment years would coincide among neighboring estuaries due to shared exposure to regional oceanographic factors. Such synchrony in recruitment has been found for many marine species and some migratory estuarine species, but has never been examined across estuaries in a species that can complete its entire life cycle within the same estuary. To inform ongoing restoration efforts for Olympia oysters, which have declined in abundance in many estuaries, we also investigated predictors of recruitment failure. We found striking contrasts in absolute recruitment rate and frequency of recruitment failure among sites, estuaries, and years. Although we found a positive relationship between upwelling and recruitment, there was little evidence of synchrony in recruitment among estuaries along the coast, and only limited synchrony of sites within estuaries, suggesting recruitment rates are affected more strongly by local dynamics within estuaries than by regional oceanographic factors operating at scales encompassing multiple estuaries. This highlights the importance of local wetland and watershed management for the demography of oysters, and perhaps other species that can complete their entire life cycle within estuaries. Estuaries with more homogeneous environmental conditions had greater synchrony among sites, and this led to the potential for estuary-wide failure when all sites had no recruitment in the same year. Environmental heterogeneity within estuaries may thus buffer against estuary-wide recruitment failure, analogous to the portfolio effect for diversity. Recruitment failure was correlated with lower summer water temperature, higher winter salinity, and shorter residence time, all indicators of stronger marine influence on estuaries. Recruitment failure was also more common in estuaries with limited networks of nearby adult oysters. Large existing oyster networks are thus of high conservation value, while estuaries that lack them would benefit from restoration efforts to increase the extent and connectivity of sites supporting oysters. This article is protected by copyright. All rights reserved.

Summary Climate change is predicted to intensify the impacts of invasive species by enhancing their performance relative to their native counterparts. However, few studies have compared the performance of invasive predators and native prey, despite the fact that non-native predators are well known to disrupt native communities. The ‘trophic sensitivity hypothesis’ suggests that predators are less tolerant of increasing environmental stress than their prey, whereas the ‘tolerant invaders hypothesis’ suggests that invaders are more tolerant than native species due to selection during the introduction process. It is therefore unclear how invasive predators will respond to increasing climate stressors. We coupled physiological measurements (thermal tolerance, thermal optima, salinity tolerance, predation rate) with environmental time-series data to assess the effects of warming and extreme low salinity events on non-native predators (gastropods) and native prey (oysters) from a coastal ecosystem. In general support of the trophic sensitivity hypothesis, we found that both non-native predators exhibited lower thermal optima relative to native prey, lower salinity tolerance and one predator was less tolerant of warming. However, because warming tolerance was extremely high (i.e. habitat temperature is 7·9–21 °C below thermal tolerance), near-term warming may first increase predator performance (consumption and growth rates), with negative effects on prey. Low salinity will likely produce heterogeneous effects on predator–prey interactions due to varying watershed sizes among estuaries that control the duration of low salinity events. The trophic sensitivity hypothesis may be a useful framework for understanding community responses to extreme climate change, which portends a decoupling of predator–prey interactions. However, we conclude that this hypothesis must be evaluated in environmental context and that coupling physiological metrics with in situ environmental data offers the best predictive power of near-term climate change impacts on invaded communities. Within our study system, warming is likely to intensify the impacts of both invasive predators, which may greatly reduce the abundance of the native oyster, a species of conservation and restoration focus.

Variability in the effectiveness of biological control programs can be due to a myriad of abiotic and biotic interactions. Here, we explore these interactions in relation to the suboptimal biological control on the invasive water hyacinth, Pontederia (=Eichhornia) crassipes, by two weevils, Neochetina bruchi and N. eichhorniae, in the Sacramento-San Joaquin River Delta in California, USA (Delta). We conducted monthly surveys from 2015 to 2017 throughout 1667 km2 to analyze the impacts of temperature, nutrients, river flow, pathogens and genetic variation in relation to the performance of these biological control agents. Weevils (predominantly N. bruchi) demonstrated a positive numerical response to increased nitrogen content in water hyacinth crown and roots. Weevil densities and water hyacinth biomass declined significantly during periods of high river flow. We found negative impacts of cooler temperatures on weevil performance, with a significant lagged-response of weevil densities in the field to water temperature from three months prior and a lack of population growth under laboratory conditions simulating winter air temperatures in the Delta (3.9–13.9 °C). We confirmed the presence of a microsporidian pathogen, with 10.7% average prevalence of infection across four seasons and low infection loads. In addition, we found low genetic diversity of N. bruchi in the Delta compared to native populations, low population genetic structure and a lack of genetic isolation by distance among metapopulations. Overall we conclude that cooler temperatures and periods of high river flow are likely the main factors limiting the current biological control program in the Delta.

The intentional introduction of exotic species through classical biological control programs provides unique opportunities to examine the consequences of population movement and ecological processes for the genetic diversity and population structure of introduced species. The weevils Neochetina bruchi and N.eichhorniae (Coleoptera: Curculionidae) have been introduced globally to control the invasive floating aquatic weed, Eichhornia crassipes, with variable outcomes. Here, we use the importation history and data from polymorphic microsatellite markers to examine the effects of introduction processes on population genetic diversity and structure. We report the first confirmation of hybridization between these species, which could have important consequences for the biological control program. For both species, there were more rare alleles in weevils from the native range than in weevils from the introduced range. N.eichhorniae also had higher allelic richness in the native range than in the introduced range. Neither the number of individuals initially introduced nor the number of introduction steps appeared to consistently affect genetic diversity. We found evidence of genetic drift, inbreeding, and admixture in several populations as well as significant population structure. Analyses estimated two populations and 11 sub-clusters for N.bruchi and four populations and 23 sub-clusters for N.eichhorniae, indicating divergence of populations during and after introduction. Genetic differentiation and allocation of introduced populations to source populations generally supported the documented importation history and clarified pathways in cases where multiple introductions occurred. In populations with multiple introductions, genetic admixture may have buffered against the negative effects of serial bottlenecks on genetic diversity. The genetic data combined with the introduction history from this biological control study system provide insight on the accuracy of predicting introduction pathways from genetic data and the consequences of these pathways for the genetic variation and structure of introduced species.

Some efforts to reduce invasive populations have paradoxically led to population increases. This phenomenon, referred to as overcompensation, occurs when reduced intraspecific pressures increase juvenile survival or maturation rates, leading to increased population size. Overcompensation in response to eradication efforts could derail management efforts, so it would be beneficial to evaluate the likelihood of overcompensation prior to removal. We conducted a series of experiments to examine the potential for overcompensation of a non-native population of the European green crab, Carcinus maenas, which was being removed in Bodega Harbor, California. First, we examined the impact of adults on juvenile survival by measuring adult cannibalism on juveniles in the presence and absence of alternative prey, and the survival of tethered juveniles at varying adult densities. Second, we examined how adult presence affected juvenile short-term foraging and growth rates. Although adult presence reduced juvenile short-term foraging, we detected only minimal cannibalism and found no evidence that adults greatly reduce juvenile growth or survivorship. These results suggest that overcompensation is not likely to occur in this population in response to removal. We assessed this prediction using pre- and post-removal surveys of juvenile recruitment in Bodega Harbor compared to nearby populations, testing for evidence of overcompensation. Relative juvenile abundance did not statistically increase in removal compared to reference populations, consistent with our conclusion from the experiments. This experimental approach which focuses on an organism’s population biology provides a tool to assess capacity for assessing the capacity for overcompensation in management strategies for invasive species.

Restoration of degraded ecosystems is an important societal goal, yet inadequate monitoring and the absence of clear performance metrics are common criticisms of many habitat restoration projects. Funding limitations can prevent adequate monitoring, but we suggest that the lack of accepted metrics to address the diversity of restoration objectives also presents a serious challenge to the monitoring of restoration projects. A working group with experience in designing and monitoring oyster reef projects was used to develop standardized monitoring metrics, units, and performance criteria that would allow for comparison among restoration sites and projects of various construction types. A set of four universal metrics (reef areal dimensions, reef height, oyster density, and oyster size–frequency distribution) and a set of three universal environmental variables (water temperature, salinity, and dissolved oxygen) are recommended to be monitored for all oyster habitat restoration projects regardless of their goal(s). In addition, restoration goal-based metrics specific to four commonly cited ecosystem service-based restoration goals are recommended, along with an optional set of seven supplemental ancillary metrics that could provide information useful to the interpretation of prerestoration and postrestoration monitoring data. Widespread adoption of a common set of metrics with standardized techniques and units to assess well-defined goals not only allows practitioners to gauge the performance of their own projects but also allows for comparison among projects, which is both essential to the advancement of the field of oyster restoration and can provide new knowledge about the structure and ecological function of oyster reef ecosystems.

Populations that span a large geographic range often experience a thermal gradient that can differentially affect the phenotypic response of individuals across the population. Variation in temperature has been shown to affect the final adult size of ectotherms, which is referred to as the temperature–size rule for ectotherms. Body size is a fundamentally important trait, as it can impact physiological performance, fecundity, longevity, and macroecological patterns. Hence, temperature may affect body size across a range, which can in turn influence maintenance of populations and ecological interactions. Here, we test whether biogeographic differences in size (carapace width) exist for a recent invasion of the non-native European green crab, Carcinus maenas, along the west coast of North America. We assembled trapping and temperature data collected from 10 sites along the western North American coast over a 5-year period. We also conducted a literature review of C. maenas size across their native range. Our results indicate that adult body size shows negative correlation with environmental temperature in both the native and invaded ranges, conforming to the temperature–size rule for ectotherms. Given the short time since colonization and lack of evident genetic structure across the invasive range, it may be that phenotypic plasticity in response to environmental temperature is driving this relationship. Forces that shape the phenotypic trajectory of species may play an important role in both invasion dynamics and subsequent ecological impacts.

The trade in marine and estuarine ornamental species has resulted in the introductions of some of the world’s worst invasive species, including the seaweed Caulerpa taxifolia and the lionfish Pterois volitans. We conducted an analysis of the historical introductions and establishments of marine and estuarine ornamental species in California using a database (‘NEMESIS’) and the contemporary fluxes (quantities, taxa) based on government records, direct observations of aquarium-bound shipments, and internet commerce. California is the major port of entry of marine ornamental species in the United States, which is the major global importing country. The vector was considered possibly responsible for twelve species introduced between 1853–2011, nine of which successfully established, including Caulerpa taxifolia (Mediterranean invasive strain). The flux of imported ornamental species was over 11 million individual animals in 2009 and included 37 taxa from at least six temperate countries, although the majority originated from the Indo-Pacific region. Almost 4,000 individuals representing at least 149 species were imported into San Francisco on a single day in 2012. Estimates of imported quantities were probably accurate within an order of magnitude but could be improved with data on interstate shipments and internet commerce. Importations of high concern included P. volitans, live rock, Chromis viridis (green chromis), and Cromileptes altivelis (panther or humpback grouper). The low historical establishment rate for ornamental species hypothetically could be explained in part by a low release rate or temperature mismatch, but the flux remains high and is a growing concern that could be addressed by heightened public education.

Although ballast water and hull fouling are widely recognized as important vectors for marine invasions, the risk posed by commercial aquaculture remains poorly quantified. To understand the importance of aquaculture as an invasion vector in California, we conducted an analysis of both current and historical introductions of marine and estuarine species associated with aquaculture using a comprehensive database (‘NEMESIS’) and permitting records for species imported into California. Our results showed that 126 non-native species associated with commercial aquaculture have been reported from California waters and 106 of these have become established. The vast majority are unintentional introductions linked to historical importation practices of the aquaculture industry. To understand the consequences of these invasions, we reviewed the literature on the impacts of mollusk and algal species introduced into California via aquaculture. Of the few studies we found, the majority demonstrated negative impacts on native species. Finally and significantly, we found that changes in aquaculture importation practices over the past decade have resulted in most shellfish currently being imported as larvae or juveniles. Consequently, rates of unintentional introductions have been reduced. We cautiously conclude that current aquaculture importation in California represents a minor risk as a vector for introductions of NIS.

Understanding how climate change may influence ecosystems depends substantially on its effects on foundation species, such as the ecologically important giant kelp (Macrocystis pyrifera). Despite its broad distribution along strong temperature and pH gradients and strong barriers to dispersal, the potential for local adaptation to climate change variables among kelp populations remains poorly understood. We assessed this potential by exposing giant kelp early life stages from genetically disparate populations in Chile and California to current and projected temperature and pH levels in common garden experiments. We observed high resistance at the haploid life stage to elevated temperatures with developmental failure appearing at the egg and sporophyte production stages among Chilean and high-latitude California populations, suggesting a greater vulnerability to climate- or ENSO-driven warming events. Additionally, populations that experience low pH events via strong upwelling, internal waves, or estuarine processes, produced more eggs per female under experimental low-pH conditions, which could increase fertilization success. These results enhance our ability to predict population extinctions and ecosystem range shifts under projected declines in ocean pH and increases in ocean temperature.

Conservation vs. eradication What's an ecologist to do when an endangered bird lives in an invasive grass? Ecosystems are complicated networks, with one species relying on another, and managing one species in isolation may damage other members of a community. Lampert et al. (see the Perspective by Buckley and Han) looked at the conflict between eradicating a damaging invasive grass species and protecting an endangered bird species that uses the grass as its home. The most effective management and restoration approach focused not on eradicating the invasive grass as quickly as possible but on making changes slowly enough that the birds could adapt. This approach may prove useful in other situations in which active restoration conflicts with other conservation goals. Science , this issue p. 1028 ; see also p. 975

Invasive species remain a major environmental problem in the world's oceans. Managing the vectors of introduction is the most effective means of mitigating this problem, but current risk assessments and management strategies are largely focused on species, not on vectors and certainly not on multiple simultaneous vectors. To highlight the issue that multiple vectors contribute to invasions, we analyzed the historical and contemporary contributions of eight maritime vectors to the establishment of nonindigenous species in California, where most species were associated with two to six vectors. Vessel biofouling looms larger than ballast water as a major vector and a management opportunity, but aquaculture risk appears reduced from historic levels. Standardized data on species abundances in each vector are lacking for a robust cross-vector assessment, which could be obtained in a proof-of-concept “vector blitz.” Management must shift away from one or two target vectors to coordination across multiple vectors.

The objective of this study was to evaluate the efficacy of integrating a traditional sediment quality triad approach with selected sublethal chronic indicators in resident species in assessing sediment quality in four salt marshes in northern California, USA. These included the highly contaminated (Stege Marsh) and relatively clean (China Camp) marshes in San Francisco Bay and two reference marshes in Tomales Bay. Toxicity potential of contaminants and benthic macroinvertebrate survey showed significant differences between contaminated and reference marshes. Sublethal responses (e.g., apoptotic DNA fragmentation, lipid accumulation, and glycogen depletion) in livers of longjaw mudsucker (Gillichthys mirabilis) and embryo abnormality in lined shore crab (Pachygrapsus crassipes) also clearly distinguished contaminated and reference marshes, while other responses (e.g., cytochrome P450, metallothionein) did not. This study demonstrates that additional chronic sublethal responses in resident species under field exposure conditions can be readily combined with sediment quality triads for an expanded multiple lines of evidence approach. This confirmatory step may be warranted in environments like salt marshes in which natural variables may affect interpretation of toxicity test data. Qualitative and quantitative integration of the portfolio of responses in resident species and traditional approach can support a more comprehensive and informative sediment quality assessment in salt marshes and possibly other habitat types as well. © 2013 .

By changing ecosystem processes and altering the physical landscape, invasive ecosystem engineers can have substantial impacts on ecosystem functions and human economies and may facilitate other non-native species. Eradication programs in terrestrial and aquatic systems aim to reverse the impacts of invasive species and return the system to its pre-invasion conditions. Despite an extensive focus on the impacts of both native and non-native ecosystem engineers, the consequences of removing invasive ecosystem engineers, particularly in coastal ecosystems, are largely unknown. In this study, we quantified changes in a benthic community following the eradication of the invasive ecosystem engineer, hybrid cordgrass Spartina, in San Francisco Bay, California. We used field experimental manipulations to test for persistent effects of both aboveground and belowground structural modifications of the invasive plant on the benthic community. We found significant effects of the invasive plant more than four years following eradication. Experimental modification of the above- vs. belowground structure of this ecosystem engineer revealed taxonomic specific effects resulting in hysteresis in the recovery of the benthic food webs. We found that these “legacy effects” resulted from two specific mechanisms: (1) delayed breakdown of belowground structures (stems, roots) and (2) persistence of other invasive species whose invasion was facilitated by the ecosystem engineer. Both of these mechanisms are likely to occur in similar systems where belowground structures breakdown more slowly or where other associated long-lived invaders persist. Our work is among the first to quantify the slow rate of change in food web and community processes and the persistent legacy effects of an invasive ecosystem engineer in a coastal ecosystem. We suggest that this delayed transition to pre-invasion conditions could resemble an alternate state that would be misidentified without a sufficient monitoring interval or recovery duration, with consequences for future management and restoration activity planning.

Our ability to predict the identity of future invasive alien species is largely based upon knowledge of prior invasion history. Emerging alien species-those never encountered as aliens before-therefore pose a significant challenge to biosecurity interventions worldwide. Understanding their temporal trends, origins, and the drivers of their spread is pivotal to improving prevention and risk assessment tools. Here, we use a database of 45,984 first records of 16,019 established alien species to investigate the temporal dynamics of occurrences of emerging alien species worldwide. Even after many centuries of invasions the rate of emergence of new alien species is still high: One-quarter of first records during 2000-2005 were of species that had not been previously recorded anywhere as alien, though with large variation across taxa. Model results show that the high proportion of emerging alien species cannot be solely explained by increases in well-known drivers such as the amount of imported commodities from historically important source regions. Instead, these dynamics reflect the incorporation of new regions into the pool of potential alien species, likely as a consequence of expanding trade networks and environmental change. This process compensates for the depletion of the historically important source species pool through successive invasions. We estimate that 1-16% of all species on Earth, depending on the taxonomic group, qualify as potential alien species. These results suggest that there remains a high proportion of emerging alien species we have yet to encounter, with future impacts that are difficult to predict.

Climate change and biological invasions are primary threats to global biodiversity that may interact in the future. To date, the hypothesis that climate change will favour non-native species has been examined exclusively through local comparisons of single or few species. Here, we take a meta-analytical approach to broadly evaluate whether non-native species are poised to respond more positively than native species to future climatic conditions. We compiled a database of studies in aquatic and terrestrial ecosystems that reported performance measures of non-native (157 species) and co-occurring native species (204 species) under different temperature, CO(2) and precipitation conditions. Our analyses revealed that in terrestrial (primarily plant) systems, native and non-native species responded similarly to environmental changes. By contrast, in aquatic (primarily animal) systems, increases in temperature and CO(2) largely inhibited native species. There was a general trend towards stronger responses among non-native species, including enhanced positive responses to more favourable conditions and stronger negative responses to less favourable conditions. As climate change proceeds, aquatic systems may be particularly vulnerable to invasion. Across systems, there could be a higher risk of invasion at sites becoming more climatically hospitable, whereas sites shifting towards harsher conditions may become more resistant to invasions.

Extreme climatic events (ECEs) – such as unusual heat waves, hurricanes, floods, and droughts – can dramatically affect ecological and evolutionary processes, and these events are projected to become more frequent and more intense with ongoing climate change. However, the implications of ECEs for biological invasions remain poorly understood. Using concepts and empirical evidence from invasion ecology, we identify mechanisms by which ECEs may influence the invasion process, from initial introduction through establishment and spread. We summarize how ECEs can enhance invasions by promoting the transport of propagules into new regions, by decreasing the resistance of native communities to establishment, and also sometimes by putting existing non-native species at a competitive disadvantage. Finally, we outline priority research areas and management approaches for anticipating future risks of unwanted invasions following ECEs. Given predicted increases in both ECE occurrence and rates of species introductions around the globe during the coming decades, there is an urgent need to understand how these two processes interact to affect ecosystem composition and functioning.

The abundance of nearly one-quarter of the world's shorebird species is declining. At the same time, the number of non-native species in coastal ecosystems is increasing rapidly. In some cases, non-native species may affect negatively the abundance and diversity of shorebird prey species. We conducted an experimental study of the effects of the introduced European green crab (Carcinus maenas) on prey consumption by wintering Dunlin (Calidris alpina) in a central California estuary. We placed green crabs and Dunlin sequentially in field enclosures and measured changes in density of benthic invertebrate prey (e.g. polychaetes and small clams), Dunlin biomass, and gut contents of both Dunlin and crabs and observed foraging behavior of Dunlin. Green crabs significantly affected Dunlin foraging success through both direct and indirect multitrophic linkages. In enclosures with high densities of green crabs, crab foraging reduced the availability of polychaetes, and Dunlin consumed significantly fewer polychaetes compared with Dunlin in enclosures without crabs. High densities of green crabs were also associated with increased availability of small clams. Dunlin consumed significantly more small clams compared with Dunlin in enclosures without crabs. In our literature survey of studies of effects of non-native invasive species on shorebirds, we found three prior experiments that addressed the effect of non-native invasive species on shorebirds. Results of two of these studies showed positive direct effects of non-native invertebrates on shorebirds, 1 showed negative direct effects of a non-native plant on shorebirds through habitat conversion, and none showed indirect effects of non-native invertebrates. We suggest future management of shorebirds explicitly examine how non-native marine species, particularly invertebrates, directly and indirectly affect shorebirds. Resumen: La abundancia de aproximadamente la cuarta parte de las especies de aves playeras del mundo esta declinando. Al mismo tiempo, el numero de especies no esta incrementando rapidamente en los ecosistemas costero. En alguno casos, la especies no nativas pueden afectar negativamente a la abundancia y diversidad de especies que son presa de aves playeras. Realizamos un estudio experimental de los efectos del cangrejo introducido Carcinus maenas sobre el consumo de presas por Calidris alpina en un estuario en el centro de California. Secuencialmente colocamos C. maenas y C. alpina en encierros y medimos cambios en la densidad de invertebrados bentonicos (poliquetos y almejas), biomasa de C. alpina y del contenido estomacal de aves y cangrejos y observamos la conducta de forrajeo de C. alpina. Los cangrejos afectaron significativamente el exito de forrajeo tanto en relaciones multitroficas directas como indirectas. En los encierros con densidades altas de cangrejos, el forrajeo de cangrejos redujo la disponibilidad de poliquetos y C. alpina consumio significativamente menos poliquetos en comparacion con encierros sin cangrejos. Las altas densidades de cangrejos tambien se asociaron con incremento en la disponibilidad de almejas pequenas. C. alpina consumio significativamente mas almejas pequenas en comparacion con encierros sin cangrejos. En nuestra revision de literatura de estudios sobre efectos de especies invasoras no nativas sobre aves playeras, encontramos 3 experimentos previos que abordaron el efecto de especies invasoras no nativas sobre aves playeras. Los resultados de 2 de esos estudios mostraron efectos positivos directos de invertebrados no nativos sobre las aves playeras, 1 mostro efectos negativos directos de una planta no nativa sobre aves playeras mediante el cambio de habitat, y ninguno mostro efectos indirectos de los invertebrados no nativos. Sugerimos que para el manejo futuro de aves playeras se examine explicitamente el efecto directo o indirecto de especies marinas no nativas, particularmente invertebrados, sobre las aves playeras.

Many non-native plants in the US have become problematic invaders of native and managed ecosystems, but a new generation of invasive species may be at our doorstep. Here, we review trends in the horticultural trade and invasion patterns of previously introduced species and show that novel species introductions from emerging horticultural trade partners are likely to rapidly increase invasion risk. At the same time, climate change and water restrictions are increasing demand for new types of species adapted to warm and dry environments. This confluence of forces could expose the US to a range of new invasive species, including many from tropical and semiarid Africa as well as the Middle East. Risk assessment strategies have proven successful elsewhere at identifying and preventing invasions, although some modifications are needed to address emerging threats. Now is the time to implement horticulture import screening measures to prevent this new wave of plant invasions.