Biogeographic and seasonal differences in consumer pressure underlie strong predation in the tropics.

Biotic interactions play a critical role in shaping patterns of global biodiversity. While several macroecological studies provide evidence for stronger predation in tropical regions compared with higher latitudes, results are variable even within the tropics, and the drivers of this variability are not well understood. We conducted two complementary standardized experiments on communities of sessile marine invertebrate prey and their associated predators to test for spatial and seasonal differences in predation across the tropical Atlantic and Pacific coastlines of Panama. We further tested the prediction that higher predator diversity contributes to stronger impacts of predation, using both direct observations of predators and data from extensive reef surveys. Our results revealed substantially higher predation rates and stronger effects of predators on prey in the Pacific than in the Atlantic, demonstrating striking variation within tropical regions. While regional predator diversity was high in the Atlantic, functional diversity at local scales was markedly low. Peak predation strength in the Pacific occurred during the wet, non-upwelling season when ocean temperatures were warmer and predator communities were more functionally diverse. Our results highlight the importance of regional biotic and abiotic drivers that shape interaction strength and the maintenance of tropical communities, which are experiencing rapid environmental change.

Stage-specific overcompensation, the hydra effect, and the failure to eradicate an invasive predator.

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.

Balanced polymorphism fuels rapid selection in an invasive crab despite high gene flow and low genetic diversity.

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 across >12 degrees of latitude in 10 years from a single source, following its introduction <35 years ago. Using six locations spanning >1500 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.

Aquatic invasion patterns across the North Atlantic.

Biological invasions are a major driver of biodiversity loss and socioeconomic burden globally. As invasion rates accelerate worldwide, understanding past invasion dynamics is essential to inform predictions of future invaders and impacts. Owing to a high diversity of pathways and current biosecurity gaps, aquatic systems near urban centres are especially susceptible to alien species establishments. Here, we compiled and compared alien species lists for three different aquatic recipient regions spanning the North Atlantic: Chesapeake Bay, Great Lakes-St. Lawrence River and North and Baltic Seas. Each system is a major trade centre, with a history of invasions, and characterized by a strong natural salinity gradient. Our goal was to compare the alien species across systems, to test for similarities in the taxonomic composition and geographic origin as well as species overlap among the three regions. We selected specific macroinvertebrate, algal and fish taxa for analysis, to control for uneven taxonomic and biogeographic resolution across regions. Cumulatively, we identified 326 individual alien species established in these aquatic systems, with the North and Baltic Seas most invaded overall (163), followed by Great Lakes-St. Lawrence River (84) and Chesapeake Bay (79). Most invasions were from Ponto-Caspian, Eurasian, Northwest Pacific, Northwest Atlantic and North American origins, and mostly comprised Arthropoda, Chordata, Mollusca and Annelida. However, origins and taxonomies differed significantly among destinations, with Ponto-Caspian species particularly successful invaders to the North and Baltic Seas then Great Lakes-St. Lawrence River, but less so to Chesapeake Bay. Nevertheless, approximately eight-tenths of invaders established in only one region, indicating disparate invasion patterns and a high potential for future aliens to accrue from increasingly diverse source pools and pathways. These results support biosecurity strategies that consider a broad range of geographic origins and taxonomic groups to limit the translocation, arrival and spread of alien species worldwide.

Invasion of the body snatchers: the role of parasite introduction in host distribution and response to salinity in invaded estuaries.

In dynamic systems, organisms are faced with variable selective forces that may impose trade-offs. In estuaries, salinity is a strong driver of organismal diversity, while parasites shape species distributions and demography. We tested for trade-offs between low-salinity stress and parasitism in an invasive castrating parasite and its mud crab host along salinity gradients of two North Carolina rivers. We performed field surveys every six to eight weeks over 3 years to determine factors influencing parasite prevalence, host abundance, and associated taxa diversity. We also looked for signatures of low-salinity stress in the host by examining its response (time-to-right and gene expression) to salinity. We found salinity and temperature significantly affected parasite prevalence, with low-salinity sites (less than 10 practical salinity units (PSU)) lacking infection, and populations in moderate salinities at warmer temperatures reaching prevalence as high as 60%. Host abundance was negatively associated with parasite prevalence. Host gene expression was plastic to acclimation salinity, but several osmoregulatory and immune-related genes demonstrated source-dependent salinity response. We identified a genetic marker that was strongly associated with salinity against a backdrop of no neutral genetic structure, suggesting possible selection on standing variation. Our study illuminates how selective trade-offs in naturally dynamic systems may shape host evolutionary ecology.

Invasion history shapes host transcriptomic response to a body-snatching parasite.

By shuffling biogeographical distributions, biological invasions can both disrupt long-standing associations between hosts and parasites and establish new ones. This creates natural experiments with which to study the ecology and evolution of host-parasite interactions. In estuaries of the Gulf of Mexico, the white-fingered mud crab (Rhithropanopeus harrisii) is infected by a native parasitic barnacle, Loxothylacus panopaei (Rhizocephala), which manipulates host physiology and behaviour. In the 1960s, L. panopaei was introduced to the Chesapeake Bay and has since expanded along the southeastern Atlantic coast, while host populations in the northeast have so far been spared. We use this system to test the host's transcriptomic response to parasitic infection and investigate how this response varies with the parasite's invasion history, comparing populations representing (i) long-term sympatry between host and parasite, (ii) new associations where the parasite has invaded during the last 60 years and (iii) naïve hosts without prior exposure. A comparison of parasitized and control crabs revealed a core response, with widespread downregulation of transcripts involved in immunity and moulting. The transcriptional response differed between hosts from the parasite's native range and where it is absent, consistent with previous observations of increased susceptibility in populations lacking exposure to the parasite. Crabs from the parasite's introduced range, where prevalence is highest, displayed the most dissimilar response, possibly reflecting immune priming. These results provide molecular evidence for parasitic manipulation of host phenotype and the role of gene regulation in mediating host-parasite interactions.

Environmental factors drive the release of Perkinsus marinus from infected oysters.

Since the discovery of Perkinsus marinus as the cause of dermo disease in Crassostrea virginica, salinity and temperature have been identified as the main environmental drivers of parasite prevalence. However, little is known about how these variables affect the movement of the parasite from host to water column. In order to elucidate how environmental factors can influence the abundance of this parasite in the water column, we conducted a series of experiments testing the effects of time of day, temperature and salinity on the release of P. marinus cells from infected oysters. We found that P. marinus cells were released on a diurnal cycle, with most cells released during the hottest and brightest period of the day (12:00-18:00). Temperature also had a strong and immediate effect on the number of cells released, but salinity did not, only influencing the intensity of infection over the course of several months. Taken together, our results demonstrate that (1) the number of parasites in the water column fluctuates according to a diurnal cycle, (2) temperature and salinity act on different timescales to influence parasite abundance, and (3) live infected oysters may substantially contribute to the abundance of transmissive parasites in the water column under particular environmental conditions.

Drivers of future alien species impacts: An expert-based assessment.

Understanding the likely future impacts of biological invasions is crucial yet highly challenging given the multiple relevant environmental, socio-economic and societal contexts and drivers. In the absence of quantitative models, methods based on expert knowledge are the best option for assessing future invasion trajectories. Here, we present an expert assessment of the drivers of potential alien species impacts under contrasting scenarios and socioecological contexts through the mid-21st century. Based on responses from 36 experts in biological invasions, moderate (20%-30%) increases in invasions, compared to the current conditions, are expected to cause major impacts on biodiversity in most socioecological contexts. Three main drivers of biological invasions-transport, climate change and socio-economic change-were predicted to significantly affect future impacts of alien species on biodiversity even under a best-case scenario. Other drivers (e.g. human demography and migration in tropical and subtropical regions) were also of high importance in specific global contexts (e.g. for individual taxonomic groups or biomes). We show that some best-case scenarios can substantially reduce potential future impacts of biological invasions. However, rapid and comprehensive actions are necessary to use this potential and achieve the goals of the Post-2020 Framework of the Convention on Biological Diversity.

Predation shapes invertebrate diversity in tropical but not temperate seagrass communities.

The hypothesis that biotic interactions are stronger at lower relative to higher latitudes has a rich history, drawing from ecological and evolutionary theory. While this hypothesis suggests that stronger interactions at lower latitudes may contribute to the maintenance of contemporary patterns of diversity, there remain few standardized biogeographic comparisons of community effects of species interactions. Using marine seagrasses as a focal ecosystem of conservation importance and sessile marine invertebrates as model prey, we tested the hypothesis that predation is stronger at lower latitudes and can shape contemporary patterns of prey diversity. To further advance understanding beyond prior studies, we also explored mechanisms that likely underlie a change in interaction outcomes with latitude. Multiple observational and experimental approaches were employed to test for effects of predators, and the mechanisms that may underlie these effects, in seagrass ecosystems of the western Atlantic Ocean spanning 30° of latitude from the temperate zone to the tropics. In predator exclusion experiments conducted in a temperate and a tropical region, predation decreased sessile invertebrate abundance, richness and diversity on both natural and standardized artificial seagrass at tropical but not temperate sites. Further, predation reduced invertebrate richness at both local and regional scales in the tropics. Additional experiments demonstrated that predation reduced invertebrate recruitment in the tropics but not the temperate zone. Finally, direct observations of predators showed higher but variable consumption rates on invertebrates at tropical relative to temperate latitudes. Together, these results demonstrate that strong predation in the tropics can have consequential impacts on prey communities through discrete effects on early life stages as well as longer-term cumulative effects on community structure and diversity. Our detailed experiments also provide some of the first data linking large-scale biogeographic patterns, community-scale interaction outcomes and direct observation of predators in the temperate zone and tropics. Therefore, our results support the hypothesis that predation is stronger in the tropics, but also elucidate some of the causes and consequences of this variation in shaping contemporary patterns of diversity.

An experimental test of stationary lay-up periods and simulated transit on biofouling accumulation and transfer on ships.

Biofouling accumulation on ships' submerged surfaces typically occurs during stationary periods that render surfaces more susceptible to colonization than when underway. As a result, stationary periods longer than typical port residence times (hours to days), often referred to as lay-ups, can have deleterious effects on hull maintenance strategies, which aim to minimize biofouling impacts on ship operations and the likelihood of invasive species transfers. This experimental study tested the effects of different lay-up durations on the magnitude of biofouling, before and after exposure to flow, using fouling panels with three coating treatments (antifouling, foul-release, and controls), at two sites, and a portable field flume to simulate voyage sheer forces. Control panels subjected to extended stationary durations (28-, 45- and 60-days) had significantly higher biofouling cover and there was a 13- to 25-fold difference in biofouling accumulation between 10-days and 28-days of static immersion. Prior to flume exposure, the antifouling coating prevented biofouling accumulation almost entirely at one site and kept it below 20% at the other. Foul-release coatings also proved effective, especially after flume exposure, which reduced biofouling at one site from >52% to <6% cover (on average). The experimental approach was beneficial for co-locating panel deployments and flume processing using a consistent (standardized) flow regime on large panels across sites of differing conditions and biofouling assemblages. While lay-ups of commercial vessels are relatively common, inevitable, and unavoidable, it is important to develop a better understanding of the magnitude of their effects on biofouling of ships' submerged surfaces and to develop workable post-lay-up approaches to manage and respond to elevated biofouling accumulation that may result.

Diversity and microhabitat associations of Labyrinthula spp. in the Indian River Lagoon System.

Seagrasses create foundational habitats in coastal ecosystems. One contributing factor to their global decline is disease, primarily caused by parasites in the genus Labyrinthula. To explore the relationship between seagrass and Labyrinthula spp. diversity in coastal waters, we examined the diversity and microhabitat association of Labyrinthula spp. in 2 inlets on Florida's Atlantic Coast, the Indian River Lagoon (IRL) and Banana River. We used amplicon-based high throughput sequencing with 2 newly designed primers to amplify Labyrinthula spp. from 5 seagrass species, water, and sediments to determine their spatial distribution and microhabitat associations. The SSU primer set identified 12 Labyrinthula zero-radius operational taxonomic units (ZOTUs), corresponding to at least 8 putative species. The ITS1 primer set identified 2 ZOTUs, corresponding to at least 2 putative species. Based on our phylogenetic analyses, which include sequences from previous studies that assigned seagrass-related pathogenicity to Labyrinthula clades, all but one of the ZOTUs that we recovered with the SSU primers were from non-pathogenic species, while the 2 ZOTUs recovered with the ITS1 primers were from pathogenic species. Some of the ZOTUs were widespread across the sampling sites and microhabitats (e.g. SSU ZOTU_10), and most were present in more than one site. Our results demonstrate that targeted metabarcoding is a useful tool for examining the relationships between seagrass and Labyrinthula diversity in coastal waters.

Environmental DNA Metabarcoding: A Promising Tool for Ballast Water Monitoring.

Nonindigenous species are introduced worldwide with ballast water (BW). To prevent further introductions, oceanic BW exchange and BW treatment systems are utilized, but their performance needs to be evaluated. To that aim, characterizing BW communities is essential but usually relies on exhaustive sampling and morphological taxonomic identification, which does not always allow fine-scale taxonomic resolution. Through the analysis of BW samples from 11 vessels arriving to the Chesapeake Bay (USA), we evaluated the potential of environmental DNA (eDNA) metabarcoding for BW monitoring by assessing whether the impact of BW management type could be identified, analyzing the influence of BW sampling access locations on communities, and comparing the accuracy of eDNA for taxonomic assignment and identification of nonindigenous taxa. We found that (1) different sampling access locations of the same tank resulted in different communities, (2) communities from treated and exchanged BW differ, (3) signals of source port and of ocean exchange are observed, (4) eDNA metabarcoding results in more diversity than morphological taxonomy, and (5) the nonindigenous copepod Oithona davisae, not reported before in the Chesapeake Bay, is detected. Overall, this study highlights the potential of eDNA metabarcoding for BW monitoring, but more comprehensive sampling will be needed to optimize the approach.

Bioerosion in a changing world: a conceptual framework.

Bioerosion, the breakdown of hard substrata by organisms, is a fundamental and widespread ecological process that can alter habitat structure, biodiversity and biogeochemical cycling. Bioerosion occurs in all biomes of the world from the ocean floor to arid deserts, and involves a wide diversity of taxa and mechanisms with varying ecological effects. Many abiotic and biotic factors affect bioerosion by acting on the bioeroder, substratum, or both. Bioerosion also has socio-economic impacts when objects of economic or cultural value such as coastal defences or monuments are damaged. We present a unifying definition and advance a conceptual framework for (a) examining the effects of bioerosion on natural systems and human infrastructure and (b) identifying and predicting the impacts of anthropogenic factors (e.g. climate change, eutrophication) on bioerosion. Bioerosion is responding to anthropogenic changes in multiple, complex ways with significant and wide-ranging effects across systems. Emerging data further underscore the importance of bioerosion, and need for mitigating its impacts, especially at the dynamic land-sea boundary. Generalised predictions remain challenging, due to context-dependent effects and nonlinear relationships that are poorly resolved. An integrative and interdisciplinary approach is needed to understand how future changes will alter bioerosion dynamics across biomes and taxa.

Dry and wet periods drive rapid shifts in community assembly in an estuarine ecosystem.

The impacts of changing climate regimes on emergent processes controlling the assembly of ecological communities remain poorly understood. Human alterations to the water cycle in the western United States have resulted in greater interannual variability and more frequent and severe extremes in freshwater flow. The specific mechanisms through which such extremes and climate regime shifts may alter ecological communities have rarely been demonstrated, and baseline information on current impacts of environmental variation is widely lacking for many habitats and communities. Here, we used observations and experiments to show that interannual variation in winter salinity levels in San Francisco Bay controls the mechanisms determining sessile invertebrate community composition during the following summer. We found consistent community changes in response to decadal-scale dry and wet extremes during a 13-year period, producing strikingly different communities. Our results match theoretical predictions of major shifts in species composition in response to environmental forcing up to a threshold, beyond which we observed mass mortality and wholesale replacement of the former community. These results provide a window into potential future community changes, with environmental forcing altering communities by shifting the relative influences of the mechanisms controlling species distributions and abundances. We place these results in the context of historical and projected future environmental variation in the San Francisco Bay Estuary.

Classification of non-indigenous species based on their impacts: considerations for application in marine management.

Assessment of the ecological and economic/societal impacts of the introduction of non-indigenous species (NIS) is one of the primary focus areas of bioinvasion science in terrestrial and aquatic environments, and is considered essential to management. A classification system of NIS, based on the magnitude of their environmental impacts, was recently proposed to assist management. Here, we consider the potential application of this classification scheme to the marine environment, and offer a complementary framework focussing on value sets in order to explicitly address marine management concerns. Since existing data on marine NIS impacts are scarce and successful marine removals are rare, we propose that management of marine NIS adopt a precautionary approach, which not only would emphasise preventing new incursions through pre-border and at-border controls but also should influence the categorisation of impacts. The study of marine invasion impacts requires urgent attention and significant investment, since we lack the luxury of waiting for the knowledge base to be acquired before the window of opportunity closes for feasible management.

Ballast Water Exchange and Invasion Risk Posed by Intracoastal Vessel Traffic: An Evaluation Using High Throughput Sequencing.

Ballast water remains a potent vector of non-native aquatic species introductions, despite increased global efforts to reduce risk of ballast water mediated invasions. This is particularly true of intracoastal vessel traffic, whose characteristics may limit the feasibility and efficacy of management through ballast water exchange (BWE). Here we utilize high throughput sequencing (HTS) to assess biological communities associated with ballast water being delivered to Valdez, Alaska from multiple source ports along the Pacific Coast of the United States. Our analyses indicate that BWE has a significant but modest effect on ballast water assemblages. Although overall richness was not reduced with exchange, we detected losses of some common benthic coastal taxa (e.g., decapods, mollusks, bryozoans, cnidaria) and gains of open ocean taxa (e.g., certain copepods, diatoms, and dinoflagellates), including some potentially toxic species. HTS-based metabarcoding identified significantly differentiated biodiversity signatures from individual source ports; this signal persisted, though weakened, in vessels undergoing BWE, indicating incomplete faunal turnover associated with management. Our analysis also enabled identification of taxa that may be of particular concern if established in Alaskan waters. While these results reveal a clear effect of BWE on diversity in intracoastal transit, they also indicate continued introduction risk of non-native and harmful taxa.

Richness and distribution of tropical oyster parasites in two oceans.

Parasites can exert strong effects on population to ecosystem level processes, but data on parasites are limited for many global regions, especially tropical marine systems. Characterizing parasite diversity and distributions are the first steps towards understanding the potential impacts of parasites. The Panama Canal serves as an interesting location to examine tropical parasite diversity and distribution, as it is a conduit between two oceans and a hub for international trade. We examined metazoan and protistan parasites associated with ten oyster species collected from both Panamanian coasts, including the Panama Canal and Bocas del Toro. We found multiple metazoan taxa (pea crabs, Stylochus spp., Urastoma cyrinae). Our molecular screening for protistan parasites detected four species of Perkinsus (Perkinsus marinus, Perkinsus chesapeaki, Perkinsus olseni, Perkinsus beihaiensis) and several haplosporidians, including two genera (Minchinia, Haplosporidium). Species richness was higher for the protistan parasites than for the metazoans, with haplosporidian richness being higher than Perkinsus richness. Perkinsus species were the most frequently detected and most geographically widespread among parasite groups. Parasite richness and overlap differed between regions, locations and oyster hosts. These results have important implications for tropical parasite richness and the dispersal of parasites due to shipping associated with the Panama Canal.

Mini-review: Assessing the drivers of ship biofouling management--aligning industry and biosecurity goals.

Biofouling exerts a frictional and cost penalty on ships and is a direct cause of invasion by marine species. These negative consequences provide a unifying purpose for the maritime industry and biosecurity managers to prevent biofouling accumulation and transfer, but important gaps exist between these sectors. This mini-review examines the approach to assessments of ship biofouling among sectors (industry, biosecurity and marine science) and the implications for existing and emerging management of biofouling. The primary distinctions between industry and biosecurity in assessment of vessels biofouling revolve around the resolution of biological information collected and the specific wetted surface areas of primary concern to each sector. The morphological characteristics of biofouling and their effects on propulsion dynamics are of primary concern to industry, with an almost exclusive focus on the vertical sides and flat bottom of hulls and an emphasis on antifouling and operational performance. In contrast, the identity, biogeography, and ecology of translocated organisms is of highest concern to invasion researchers and biosecurity managers and policymakers, especially as it relates to species with known histories of invasion elsewhere. Current management practices often provide adequate, although not complete, provision for hull surfaces, but niche areas are well known to enhance biosecurity risk. As regulations to prevent invasions emerge in this arena, there is a growing opportunity for industry, biosecurity and academic stakeholders to collaborate and harmonize efforts to assess and manage biofouling of ships that should lead to more comprehensive biofouling solutions that promote industry goals while reducing biosecurity risk and greenhouse gas emissions.

Biodiversity differentially impacts disease dynamics across marine and terrestrial habitats.

The relationship between biodiversity and infectious disease, where increased biodiversity leads to decreased disease risk, originated from research in terrestrial disease systems and remains relatively underexplored in marine systems. Understanding the impacts of biodiversity on disease in marine versus terrestrial systems is key to continued marine ecosystem functioning, sustainable aquaculture, and restoration projects. We compare the biodiversity-disease relationship across terrestrial and marine systems, considering biodiversity at six levels: intraspecific host diversity, host microbiomes, interspecific host diversity, biotic vectors and reservoirs, parasite consumers, and parasites. We highlight gaps in knowledge regarding how these six levels of biodiversity impact diseases in marine systems and propose two model systems, the Perkinsus-oyster and Labyrinthula-seagrass systems, to address these gaps.

Geographic variation in marine invasions among large estuaries: effects of ships and time.

Coastal regions exhibit strong geographic patterns of nonnative species richness. Most invasions in marine ecosystems are known from bays and estuaries, where ship-mediated transfers (on hulls or in ballasted materials) have been a dominant vector of species introductions. Conspicuous spatial differences in nonnative species richness exist among bays, but the quantitative relationship between invasion magnitude and shipping activity across sites is largely unexplored. Using data on marine invasions (for invertebrates and algae) and commercial shipping across 16 large bays in the United States, we estimated (1) geographic variation in nonnative species richness attributed to ships, controlling for effects of salinity and other vectors, (2) changes through time in geographic variation of these ship-mediated invasions, and (3) effects of commercial ship traffic and ballast water discharge magnitude on nonnative species richness. For all nonnative species together (regardless of vector, salinity, or time period), species richness differed among U.S. coasts, being significantly greater for Pacific Coast bays than Atlantic or Gulf Coast bays. This difference also existed when considering only species attributed to shipping (or ballast water), controlling for time and salinity. Variation in nonnative species richness among Pacific Coast bays was strongly affected by these same criteria. San Francisco Bay, California, had over 200 documented nonnative species, more than twice that reported for other bays, but many species were associated with other (non-shipping) vectors or the extensive low-salinity habitats (unavailable in some bays). When considering only ship- or ballast-mediated introductions in high-salinity waters, the rate of newly detected invasions in San Francisco Bay has converged increasingly through time on that for other Pacific Coast bays, appearing no different since 1982. Considering all 16 bays together, there was no relationship between either (1) number of ship arrivals (from foreign ports) and number of introductions attributed to ships since 1982 or (2) volume of foreign ballast water discharge and number of species attributed to ballast water since 1982. These shipping measures are likely poor proxies for propagule supply, although they are sometimes used as such, highlighting a fundamental gap in data needed to evaluate invasion dynamics and management strategies.