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The human-mediated introduction of marine non-indigenous species is a centuries- if not millennia-old phenomenon, but was only recently acknowledged as a potent driver of change in the sea. We provide a synopsis of key historical milestones for marine bioinvasions, including timelines of (a) discovery and understanding of the invasion process, focusing on transfer mechanisms and outcomes, (b) methodologies used for detection and monitoring, (c) approaches to ecological impacts research, and (d) management and policy responses. Early (until the mid-1900s) marine bioinvasions were given little attention, and in a number of cases actively and routinely facilitated. Beginning in the second half of the 20th century, several conspicuous non-indigenous species outbreaks with strong environmental, economic, and public health impacts raised widespread concerns and initiated shifts in public and scientific perceptions. These high-profile invasions led to policy documents and strategies to reduce the introduction and spread of non-indigenous species, although with significant time lags and limited success and focused on only a subset of transfer mechanisms. Integrated, multi-vector management within an ecosystem-based marine management context is urgently needed to address the complex interactions of natural and human pressures that drive invasions in marine ecosystems.

The 2011 East Japan earthquake generated a massive tsunami that launched an extraordinary transoceanic biological rafting event with no known historical precedent. We document 289 living Japanese coastal marine species from 16 phyla transported over 6 years on objects that traveled thousands of kilometers across the Pacific Ocean to the shores of North America and Hawai‘i. Most of this dispersal occurred on nonbiodegradable objects, resulting in the longest documented transoceanic survival and dispersal of coastal species by rafting. Expanding shoreline infrastructure has increased global sources of plastic materials available for biotic colonization and also interacts with climate change–induced storms of increasing severity to eject debris into the oceans. In turn, increased ocean rafting may intensify species invasions.

Aim The introduction of aquatic non‐indigenous species (ANS) has become a major driver for global changes in species biogeography. We examined spatial patterns and temporal trends of ANS detections since 1965 to inform conservation policy and management. Location Global. Methods We assembled an extensive dataset of first records of detection of ANS (1965–2015) across 49 aquatic ecosystems, including the (a) year of first collection, (b) population status and (c) potential pathway(s) of introduction. Data were analysed at global and regional levels to assess patterns of detection rate, richness and transport pathways. Results An annual mean of 43 (±16 SD) primary detections of ANS occurred—one new detection every 8.4 days for 50 years. The global rate of detections was relatively stable during 1965–1995, but increased rapidly after this time, peaking at roughly 66 primary detections per year during 2005–2010 and then declining marginally. Detection rates were variable within and across regions through time. Arthropods, molluscs and fishes were the most frequently reported ANS. Most ANS were likely introduced as stowaways in ships’ ballast water or biofouling, although direct evidence is typically absent. Main conclusions This synthesis highlights the magnitude of recent ANS detections, yet almost certainly represents an underestimate as many ANS go unreported due to limited search effort and diminishing taxonomic expertise. Temporal rates of detection are also confounded by reporting lags, likely contributing to the lower detection rate observed in recent years. There is a critical need to implement standardized, repeated methods across regions and taxa to improve the quality of global‐scale comparisons and sustain core measures over longer time‐scales. It will be fundamental to fill in knowledge gaps given that invasion data representing broad regions of the world's oceans are not yet readily available and to maintain knowledge pipelines for adaptive management.

The emergence of new Arctic trade routes will probably change the global dynamics of invasive species, potentially affecting marine habitats and ecosystem functions, especially in coastal regions.

Latitudinal patterns of nonnative species richness suggest fewer successful invasions in the tropics, relative to temperate regions. One main hypothesis for this pattern is that biotic resistance to invasion is stronger in the tropics than at higher latitudes. Biotic resistance can limit the distribution and abundance of nonnative species and, in extreme cases, can prevent establishment. We provide the first experimental test of this hypothesis, comparing the strength of biotic resistance in a tropical and a temperate marine ecosystem. Predation is one mechanism of biotic resistance, and since predation can be stronger at lower latitudes, we predicted that predation will serve to increase biotic resistance more in the tropics than at higher latitude. We conducted predator-exclusion experiments on marine epifaunal communities, a heavily invaded system, focusing on nonnative tunicates as a model fauna. The effect of predation on species richness of nonnative tunicates was more than three times greater at sites in tropical Panama than in temperate Connecticut, consistent with the prediction of stronger biotic resistance in the tropics. In Connecticut, predation reduced the abundance of one nonnative tunicate but did not affect the abundances of any other nonnative tunicate species, and no species were excluded from communities. In contrast, predation resulted in striking reductions in abundance and often exclusion of nonnative tunicates from experimental communities in Panama. If proved to be general, latitudinal differences in the biotic resistance of communities to nonnative species establishment may help explain emerging patterns of global invasions.

Discoveries of persistent coastal species in the open ocean shift our understanding of biogeographic barriers. Floating plastic debris from pollution now supports a novel sea surface community composed of coastal and oceanic species at sea that might portend significant ecological shifts in the marine environment.

Global trade by merchant ships is a leading mechanism for the unintentional transfer of marine organisms, including non-indigenous species, to bays and estuaries worldwide. To reduce the likelihood of new invasions, ships are increasingly being required to manage their ballast water (BW) prior to discharge in coastal waters. In the United States, most overseas arrivals have been required to manage BW discharge since 2004, primarily through ballast water exchange (BWE), which flushes out ballast tanks in the open ocean (>200 miles from shore). Studies have found BWE to generally reduce the abundance of organisms, and the amount of water exchanged has been estimated at 96-100%. Despite its widespread use, the overall effect of this management strategy on net propagule supply through time has not been explored. Here, temporal changes in zooplankton concentrations and the volume of BW discharged in Chesapeake Bay, U.S. were evaluated, comparing pre-management era and post-management era time periods. Chesapeake Bay is a large port system that receives extensive BW discharge, especially from bulk cargo vessels (bulkers) that export coal overseas. For bulkers arriving from overseas, mean zooplankton concentrations of total and coastal indicator taxa in BW did not decline between pre- (1993-2000) and post management (2012-2013) eras, when controlling for season and sampling method. Moreover, bulkers discharged 21 million tonnes (82% of total for Chesapeake Bay) of overseas BW in 2013, representing a 374% increase in volume when compared to 2005. The combination of BW discharge volume and zooplankton concentration data indicates that (a) net propagule supply by bulkers has increased since BWE began in Chesapeake Bay; and (b) changes in vessel behaviour and trade have contributed strongly to this outcome. Specifically, the coal-driven increase in BW discharge volume from 2005-2013, concurrent with the onset of BWE regulations, worked to counteract intended results from BW management. A long-term analysis of bulker arrivals (1994-2013) reveals a 20-year minimum in arrival numbers in 2000, just when the implementation of BWE began. This study underscores the need to consider shifts in trade patterns, in order to advance and evaluate effective management strategies for biological invasions.

1 Propagule pressure plays a key role in the successful establishment of introduced species. Explaining invasion patterns, predicting future invasions and reducing invasion rates are priority areas of research and management, especially in marine systems, which need more detailed correlates and invasion predictors. 2 The commercial maritime shipping fleet is the most prolific long distance anthropogenic transfer mechanism (vector) of marine non-indigenous species on a global scale, causing invasions of coasts by a wide diversity of organisms. Although most vessel arrivals provide an opportunity for organism introductions, there are often substantial differences among ship types—in both their \"morphological traits\" (structural design) and \"behavioural ecology\" (cargo delivery model and operational tempo)-that influence propagule delivery by ballast water and biofouling, the two dominant sources or sub-vectors for ship-mediated species transfers. 3 We reviewed ship specialization and its implications for marine invasion and vector management. First, we identified factors that affect ship-mediated propagule delivery characteristics (number, identity, diversity and quality/condition), classifying these as ship type independent or dependent factors. Second, we compared the relevance of these factors for both ballast water and biofouling. Third, we estimated and compared the magnitude of several key factors affecting propagule delivery among seven major ship types. 4 Typical voyage speed varies by 74% and port residence time varies sixfold among ship types. Similarly, typical ballast water discharge varies by an order of magnitude among ship types. These and other ship type dependent factors affect propagule delivery characteristics, resulting in uneven magnitude of species transfer among ship types. 5 Policy implications. Variation among commercial ship types is rarely integrated into analyses of marine bioinvasions and proxy measures of propagule delivery. Their inclusion may lead to more robust explanation, prediction and management of marine invasions. Risk analyses that account for differences among ship types and prevailing traffic directionality will likely offer greater insight than null models, which treat ships equally. Furthermore, ballast treatment technologies and hull husbandry may advance to reduce species transfers more effectively when tailored for different ship types, recognizing the variation and operational constraints (that affect propagule delivery) among the diverse range of ship types.

Recreational boats and their marinas are important components of the hub-and-spoke model of invasion in coastal marine systems. Like most vectors, however, species transfers by boats are under-sampled and the extent of invasion among coastal bays is patchily sampled, with a bias towards larger urbanized bays with commercial port systems. We assessed both the hubs (bays with marinas) and the spokes (recreational boat movements and their vector biota) that form the basis of a non-shipping hub-and-spoke system for the coast of California. Non-native invertebrates and algae have been reported from most bays in California (26 of 27), with only five of these having commercial shipping ports. Recreational boats offer frequent opportunities for transfers of species among bays; 292,000 recreational boats were registered in coastal California, and more than half (54%) of surveyed recreational boat owners made voyages outside their home bays. Destinations for recreational boaters often included bays without commercial ports and areas of conservation value inside National Marine Sanctuaries. We recorded 158 taxa from the hulls and niche areas of 49 transient recreational boats upon arrival at Californian marinas. These included several non-native species that are already well-established at arrival locations and four that are not known from the state or sampling location. Biofouling abundance on vessels ranged from zero to remarkably high levels (estimated at 106 organisms) for vessels that traverse open coastal systems. Management opportunities and models exist for the recreational boat vector but, absent operational incentives, these are challenging to implement broadly. An initial management focus on areas recognised for their distinct marine habitats and biodiversity may be an effective implementation approach.

Species interactions are widely assumed to be stronger at lower latitudes, but surprisingly few experimental studies test this hypothesis, and none ties these processes to observed patterns of species richness across latitude. We report here the first experimental field test that predation is both stronger and has a disproportionate effect on species richness in the tropics relative to the temperate zone. We conducted predator-exclusion experiments on communities of sessile marine invertebrates in four regions, which span 32°° latitude, in the western Atlantic Ocean and Caribbean Sea. Over a three-month timescale, predation had no effect on species richness in the temperate zone. In the tropics, however, communities were from two to over ten times more species-rich in the absence of predators than when predators were present. While micro-and macro-predators likely compete for the limited prey resource in the tropics, micropredators alone were able to exert as much pressure on the invertebrate communities as the full predator community. This result highlights the extent to which exposure to even a subset of the predator guild can significantly impact species richness in the tropics. Patterns were consistent in analyses that included relative and total species abundances. Higher species richness in the absence of predators in the tropics was also observed when species occurrences were pooled across two larger spatial scales, site and region, demonstrating a consistent scaling relationship. These experimental results show that predation can both limit local species abundances and shape patterns of regional coexistence in the tropics. When preestablished diverse tropical communities were then exposed to predation for different durations, ranging from one to several days, species richness was always reduced. These findings confirmed that impacts of predation in the tropics are strong and consistent, even in more established communities. Our results offer empirical support for the long-held prediction that predation pressure is stronger at lower latitudes. Furthermore, we demonstrate the magnitude to which variation in predation pressure can contribute to the maintenance of tropical species diversity.