Searching for a Home Port in a Polyvectic World: Molecular Analysis and Global Biogeography of the Marine Worm Polydora hoplura (Annelida: Spionidae).

The spionid polychaete Polydora hoplura Claparède, 1868 is a shell borer widely occurring across the world and considered introduced in many areas. It was originally described in the Gulf of Naples, Italy. Adult diagnostic features are the palps with black bands, prostomium weakly incised anteriorly, caruncle extending to the end of chaetiger 3, short occipital antenna, and heavy sickle-shaped spines in the posterior notopodia. The Bayesian inference analysis of sequence data of four gene fragments (2369 bp in total) of the mitochondrial 16S rDNA, nuclear 18S, 28S rDNA and Histone 3 has shown that worms with these morphological features from the Mediterranean, northern Europe, Brazil, South Africa, Australia, Republic of Korea, Japan and California are genetically identical, form a well-supported clade, and can be considered conspecific. The genetic analysis of a 16S dataset detected 15 haplotypes of this species, 10 of which occur only in South Africa. Despite the high genetic diversity of P. hoplura in South Africa, we tentatively propose the Northwest Pacific, or at the most the Indo-West Pacific, as its home region, not the Atlantic Ocean or the Eastern Pacific Ocean. The history of the discovery of P. hoplura around the world appears to be intimately linked to global shipping commencing in the mid-19th century, followed by the advent of the global movement of commercial shellfish (especially the Pacific oyster Magallana gigas) in the 20th century, interlaced with continued, complex dispersal by vessels and aquaculture. Given that P. hoplura has been detected in only a few of the 17 countries where Pacific oysters have been established, we predict that it may already be present in many more regions. As global connectivity through world trade continues to increase, it is likely that novel populations of P. hoplura will continue to emerge.

Extent and reproduction of coastal species on plastic debris in the North Pacific Subtropical Gyre.

We show that the high seas are colonized by a diverse array of coastal species, which survive and reproduce in the open ocean, contributing strongly to its floating community composition. Analysis of rafting plastic debris in the eastern North Pacific Subtropical Gyre revealed 37 coastal invertebrate taxa, largely of Western Pacific origin, exceeding pelagic taxa richness by threefold. Coastal taxa, including diverse taxonomic groups and life history traits, occurred on 70.5% of debris items. Most coastal taxa possessed either direct development or asexual reproduction, possibly facilitating long-term persistence on rafts. Our results suggest that the historical lack of available substrate limited the colonization of the open ocean by coastal species, rather than physiological or ecological constraints as previously assumed. It appears that coastal species persist now in the open ocean as a substantial component of a neopelagic community sustained by the vast and expanding sea of plastic debris.

First report of marine debris as a species dispersal vector in the temperate Northwest Atlantic Ocean.

We provide the first report of the role of marine debris in transporting native and introduced species in the temperate Northwest Atlantic Ocean. Plastic was the most frequent biofouled material. Thirty-three attached species (five non-native) were found on rafted debris, 16 of which have not been previously reported as rafters. Forty-six percent of the attached invertebrate rafters (including three of the introduced species, the bryozoans Fenestrulina delicia and Tricellaria inopinata and the spirorbid Janua heterostropha) detected in this study reproduce by either direct development or produce larvae of short-term planktonic existence, suggesting that rafting on long-term, non-biodegradable debris may enhance their dispersal potential. We suggest that a prominent non-native species, the green alga Codium fragile fragile, may play a previously undetected role in the transport of marine debris and associated biofouling. Marine debris may further be a potentially significant source of biodiversity records; we detected two bryozoan species in our study region that were either previously unknown or had not been found for >75 years.

Global marine biosecurity and ship lay-ups: intensifying effects of trade disruptions.

Recent global trade disruptions, due to blockage of the Suez Canal and cascading effects of COVID-19, have altered the movement patterns of commercial ships and may increase worldwide invasions of marine non-indigenous species. Organisms settle on the hulls and underwater surfaces of vessels and can accumulate rapidly, especially when vessels remain stationary during lay-ups and delays. Once present, organisms can persist on vessels for long-periods (months to years), with the potential to release propagules and seed invasions as ships visit ports across the global transportation network. Shipborne propagules also may be released in increasing numbers during extended vessel residence times at port or anchor. Thus, the large scale of shipping disruptions, impacting thousands of vessels and geographic locations and still on-going for over two years, may elevate invasion rates in coastal ecosystems in the absence of policy and management efforts to prevent this outcome. Concerted international and national biosecurity actions, mobilizing existing frameworks and tools with due diligence, are urgently needed to address a critical gap and abate the associated invasion risks.

Out of taxonomic crypsis: A new trans-arctic cryptic species pair corroborated by phylogenetics and molecular evidence.

Cryptic species are a common phenomenon in cosmopolitan marine species. The use of molecular tools has often uncovered cryptic species occupying a fraction of the geographic range of the original morphospecies. Shipworms (Teredinidae) are marine bivalves, living in drift and fixed wood, many of which have a conserved morphology across cosmopolitan distributions. Herein novel and GenBank mitochondrial (cytochrome c oxidase subunit I) and nuclear (18S rRNA) DNA sequences are employed to produce a phylogeny of the Teredinidae and delimit a cryptic species pair in the Psiloteredo megotara complex. The anatomy, biogeography, and ecology of P. megotara, Psiloteredo sp. and Nototeredo edax are compared based on private and historic museum collections and a thorough literature review. Morphological and anatomical characters of P. megotara from the North Atlantic and Psiloteredo sp. from Japan were morphologically indistinguishable, and differ in pallet architecture and soft tissue anatomy from N. edax. The two Psiloteredo species were then delimited as genetically distinct species using four molecular-based methods. Consequently, the Northwest Pacific species, Psiloteredo pentagonalis, first synonymized with N. edax and then with P. megotara, is resurrected. Nototeredo edax, P. megotara and P. pentagonalis are redescribed based upon morphological and molecular characters. Phylogenetic analysis further revealed cryptic species complexes within the cosmopolitan species Bankia carinata and possibly additional cryptic lineages within the cosmopolitan Lyrodus pedicellatus.

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.

Mediators of invasions in the sea: life history strategies and dispersal vectors facilitating global sea anemone introductions.

Widespread non-native species tend to demonstrate an apparent lack of selectivity in habitat requirements, feeding regimes, and reproductive needs, while displaying a tendency to thrive in human-modified habitats. The high phenotypic plasticity typical of sessile, substrate-attached marine species may enhance their chances of survival and spread in a new region. Anthropogenic activities have changed marine habitats over a wide range of phenomena, including water temperature, community species composition, and the types of available substrates, creating new physical and biotic regimes that may contribute to the potential for successful species introduction. Here we examine ten species of sea anemones that have been introduced outside of their native range, and elucidate specific characteristics that are common among globally introduced sea anemones. Various life history strategies enable these species to survive and flourish through transport, introduction, establishment and spread, leading to the successful colonization of a new geographic area. Considering life history strategies and weighing of vector potential, we suggest conditions that facilitate introduction of these species, and identify species of sea anemones that may be introduced in the future in the face of changing climate and increased anthropogenic activities.

Scientists' warning on invasive alien species.

Biological invasions are a global consequence of an increasingly connected world and the rise in human population size. The numbers of invasive alien species - the subset of alien species that spread widely in areas where they are not native, affecting the environment or human livelihoods - are increasing. Synergies with other global changes are exacerbating current invasions and facilitating new ones, thereby escalating the extent and impacts of invaders. Invasions have complex and often immense long-term direct and indirect impacts. In many cases, such impacts become apparent or problematic only when invaders are well established and have large ranges. Invasive alien species break down biogeographic realms, affect native species richness and abundance, increase the risk of native species extinction, affect the genetic composition of native populations, change native animal behaviour, alter phylogenetic diversity across communities, and modify trophic networks. Many invasive alien species also change ecosystem functioning and the delivery of ecosystem services by altering nutrient and contaminant cycling, hydrology, habitat structure, and disturbance regimes. These biodiversity and ecosystem impacts are accelerating and will increase further in the future. Scientific evidence has identified policy strategies to reduce future invasions, but these strategies are often insufficiently implemented. For some nations, notably Australia and New Zealand, biosecurity has become a national priority. There have been long-term successes, such as eradication of rats and cats on increasingly large islands and biological control of weeds across continental areas. However, in many countries, invasions receive little attention. Improved international cooperation is crucial to reduce the impacts of invasive alien species on biodiversity, ecosystem services, and human livelihoods. Countries can strengthen their biosecurity regulations to implement and enforce more effective management strategies that should also address other global changes that interact with invasions.

First mussel settlement observed in Antarctica reveals the potential for future invasions.

Global biodiversity is both declining and being redistributed in response to multiple drivers characterizing the Anthropocene, including synergies between biological invasions and climate change. The Antarctic marine benthos may constitute the last biogeographic realm where barriers (oceanographic currents, climatic gradients) have not yet been broken. Here we report the successful settlement of a cohort of Mytilus cf. platensis in a shallow subtidal habitat of the South Shetland Islands in 2019, which demonstrates the ability of this species to complete its early life stages in this extreme environment. Genetic analyses and shipping records show that this observation is consistent with the dominant vectors and pathways linking southern Patagonia with the Antarctic Peninsula and demonstrates the potential for impending invasions of Antarctic ecosystems.

Trends in the detection of aquatic non-indigenous species across global marine, estuarine and freshwater ecosystems: A 50-year perspective.

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.

A Plasticene Lexicon.

As plastic pollution in the environment has increased rapidly in the last half century, so too has the study of the effects of plastic on marine, aquatic and terrestrial ecosystems. From this research, a series of new terms has emerged to describe the phenomena unique to the presence of plastic-based materials in nature. In this short note, we bring together disparate neologisms into a single lexicon with the aim to encourage use of a unified vocabulary to describe the new reality of ecological, chemical, and geological systems in the age of plastics.

Supporting Spartina: Interdisciplinary perspective shows Spartina as a distinct solid genus.

In 2014, a DNA-based phylogenetic study confirming the paraphyly of the grass subtribe Sporobolinae proposed the creation of a large monophyletic genus Sporobolus, including (among others) species previously included in the genera Spartina, Calamovilfa, and Sporobolus. Spartina species have contributed substantially (and continue contributing) to our knowledge in multiple disciplines, including ecology, evolutionary biology, molecular biology, biogeography, experimental ecology, biological invasions, environmental management, restoration ecology, history, economics, and sociology. There is no rationale so compelling to subsume the name Spartina as a subgenus that could rival the striking, global iconic history and use of the name Spartina for over 200 yr. We do not agree with the subjective arguments underlying the proposal to change Spartina to Sporobolus. We understand the importance of both the objective phylogenetic insights and of the subjective formalized nomenclature and hope that by opening this debate we will encourage positive feedback that will strengthen taxonomic decisions with an interdisciplinary perspective. We consider that the strongly distinct, monophyletic clade Spartina should simply and efficiently be treated as the genus Spartina.

Historical baselines in marine bioinvasions: Implications for policy and management.

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 invasion risk of species associated with Japanese Tsunami Marine Debris in Pacific North America and Hawaii.

Marine debris from the Great Tsunami of 2011 represents a unique transport vector for Japanese species to reach Pacific North America and Hawaii. Here we characterize the invasion risk of invertebrate species associated with tsunami debris using a screening-level risk assessment tool - the Canadian Marine Invasive Screening Tool (CMIST). Higher-risk invertebrate invaders were identified for each of five different ecoregions. Some of these are well-known global invaders, such as the mussel Mytilus galloprovincialis and the ascidian Didemnum vexillum which already have invasion histories in some of the assessed ecoregions, while others like the sea star Asterias amurensis and the shore crab Hemigrapsus sanguineus have yet to invade large portions of the assessed ecoregions but also are recognized global invaders. In general, the probability of invasion was lower for the Gulf of Alaska and Hawaii, in part due to lower climate matches and the availability of other invasion vectors.

Trait-based characterization of species transported on Japanese tsunami marine debris: Effect of prior invasion history on trait distribution.

Nearly 300 coastal marine species collected from >630 debris items from the 2011 Great East Japan earthquake and tsunami have landed alive along the North American Pacific coast and the Hawaiian Archipelago. We synthesized life history, environmental, and distributional traits for 103 of these species and compared species with (n=30) and without (n=62) known invasion histories. The species represent 12 phyla, and Mollusca, Crustacea, and Bryozoa accounted for 71 of the 103 species. The majority are native to the Northwest Pacific and the Central Indo-Pacific. Species with known invasion history were more common on artificial and hardpan substrates, in temperate reef, fouling, and flotsam habitats, at subtropical and tropical temperatures, and exhibited greater salinity tolerance than species with no prior invasion history. Thirty-five Japanese tsunami marine species without prior invasion history overlapped in ordination trait space with known invaders, indicating a subset of species in this novel assemblage that possess traits similar to species with known invasion history.

A Framework for Understanding Marine Cosmopolitanism in the Anthropocene.

Recent years have witnessed growing appreciation for the ways in which human-mediated species introductions have reshaped marine biogeography. Despite this we have yet to grapple fully with the scale and impact of anthropogenic dispersal in both creating and determining contemporary distributions of marine taxa. In particular, the past several decades of research on marine biological invasions have revealed that broad geographic distributions of coastal marine organisms-historically referred to simply as "cosmopolitanism"-may belie complex interplay of both natural and anthropogenic processes. Here we describe a framework for understanding contemporary cosmopolitanism, informed by a synthesis of the marine bioinvasion literature. Our framework defines several novel categories in an attempt to provide a unified terminology for discussing cosmopolitan distributions in the world's oceans. We reserve the term eucosmopolitan to refer to those species for which data exist to support a true, natural, and prehistorically global (or extremely broad) distribution. While in the past this has been the default assumption for species observed to exhibit contemporary cosmopolitan distributions, we argue that given recent advances in marine invasion science this assignment should require positive evidence. In contrast, neocosmopolitan describes those species that have demonstrably achieved extensive geographic ranges only through historical anthropogenic dispersal, often facilitated over centuries of human maritime traffic. We discuss the history and human geography underpinning these neocosmopolitan distributions, and illustrate the extent to which these factors may have altered natural biogeographic patterns. We define the category pseudocosmopolitan to encompass taxa for which a broad distribution is determined (typically after molecular investigation) to reflect multiple, sometimes regionally endemic, lineages with uncertain taxonomic status; such species may remain cosmopolitan only so long as taxonomic uncertainty persists, after which they may splinter into multiple geographically restricted species. We discuss the methods employed to identify such species and to resolve both their taxonomic status and their biogeographic histories. We argue that recognizing these different types of cosmopolitanism, and the important role that invasion science has played in understanding them, is critically important for the future study of both historical and modern marine biogeography, ecology, and biodiversity.

Transoceanic dispersal of the mussel Mytilus galloprovincialis on Japanese tsunami marine debris: An approach for evaluating rafting of a coastal species at sea.

Biofouled debris from the 2011 Great East Japan earthquake and tsunami has landed in the Northeast Pacific and along the Hawaiian Islands since 2012. As of 2017, >630 biofouled debris items with >320 living species of algae, invertebrates, and fish have been examined. The invasive mussel Mytilus galloprovincialis was present on >50% of those items. Size, reproduction, and growth of this filter-feeding species were examined to better understand long-distance rafting of a coastal species. The majority of mussels (79%) had developing or mature gametes, and growth rates averaged 0.075±0.018 SE mm/day. Structural and elemental (barium/calcium) analysis of mussel shells generated estimates of growth in coastal waters (mean=1.3 to 25mm total length), which provides an indication of residence times in waters along North America and the Hawaiian Islands prior to landing. Detailed studies of individual species contribute to our understanding of debris as a transport vector and aid efforts to evaluate potential risks associated with marine debris.