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.
Despite the Mediterranean being both a hotspot for recreational boating and for non‐indigenous species (NIS), no data currently exists on the recreational boating sector's contribution to the spread ...of NIS in this Sea.
To improve the basis for management decisions, a wide‐scale sampling study on the biofouling communities of recreational vessels and marinas was undertaken. Specifically, we surveyed over 600 boat owners and sampled the same boat hulls for NIS in 25 marinas across the Mediterranean, from France to Cyprus, to determine which factors (i.e. boat characteristics, travel behaviour, home marina) are associated with higher NIS richness on boat hulls.
Among the surveyed boats, we found recreational vessels to travel considerably, averaging 67 travel days and 7.5 visited marinas per annum. This results in a high potential for spreading NIS, especially as 71% of sampled vessels host at least one (and up to 11) NIS. Boats with high NIS richness strongly correlate with home marinas with high NIS richness. Over half of the vessels were carriers of NIS which were not yet present in the marinas they were visiting. The presence of biofouling in niche areas of the hull (i.e. in the cavities and metallic parts) emerges as the best predictor for NIS richness on boats, along with longer times since their last cleaning and antifouling applications. Interestingly, colonization of NIS occurred rapidly, even on boats that had recently had their hulls cleaned professionally.
Synthesis and applications. We demonstrate that recreational boating has a very high capacity for the spread of non‐indigenous species (NIS) in the Mediterranean, due to both high NIS richness on boats and extensive travel. To counteract the spread of NIS, routine monitoring for new NIS needs to be established for both marinas and vessels, along with frequent pontoon cleaning. Additionally, policy should require preliminary screenings for incoming vessels from new countries, especially those emanating from high‐risk marinas. The niche areas of the boat hulls should be checked first for biofouling, which was the best predictor for NIS richness since they often go overlooked with in‐water cleanings are rarely have antifouling applied to them.
We demonstrate that recreational boating has a very high capacity for the spread of non‐indigenous species (NIS) in the Mediterranean, due to both high NIS richness on boats and extensive travel. To counteract the spread of NIS, routine monitoring for new NIS needs to be established for both marinas and vessels, along with frequent pontoon cleaning. Additionally, policy should require preliminary screenings for incoming vessels from new countries, especially those emanating from high‐risk marinas. The niche areas of the boat hulls should be checked first for biofouling, which was the best predictor for NIS richness since they often go overlooked with in‐water cleanings are rarely have antifouling applied to them.
Ecology Letters (2010) 13: 235-245 A major aim in ecology is identifying determinants of invasiveness. We performed a meta-analysis of 117 field or experimental-garden studies that measured pair-wise ...trait differences of a total of 125 invasive and 196 non-invasive plant species in the invasive range of the invasive species. We tested whether invasiveness is associated with performance-related traits (physiology, leaf-area allocation, shoot allocation, growth rate, size and fitness), and whether such associations depend on type of study and on biogeographical or biological factors. Overall, invasive species had significantly higher values than non-invasive species for all six trait categories. More trait differences were significant for invasive vs. native comparisons than for invasive vs. non-invasive alien comparisons. Moreover, for comparisons between invasive species and native species that themselves are invasive elsewhere, no trait differences were significant. Differences in physiology and growth rate were larger in tropical regions than in temperate regions. Trait differences did not depend on whether the invasive alien species originates from Europe, nor did they depend on the test environment. We conclude that invasive alien species had higher values for those traits related to performance than non-invasive species. This suggests that it might become possible to predict future plant invasions from species traits.
AIM: The global sprawl of marine hard infrastructure (e.g. breakwaters, sea walls and jetties) can extensively modify coastal seascapes, but the knowledge of such impacts remains limited to local ...scales. We examined the regional‐scale effects of marine artificial habitats on the distribution and abundance of assemblages of ascidians, a key group of ecosystem engineer species in benthic fouling systems. LOCATION: Five hundred kilometers of coastline in the North Adriatic Sea. METHODS: We sampled a variety of natural reefs, marine infrastructures and marinas, and tested hypotheses about the role of habitat type and location in influencing the relative distribution and abundance of both native and non‐indigenous species. RESULTS: Assemblages differed significantly between natural and artificial habitats and among different types of artificial habitats. Non‐indigenous species were 2–3 times more abundant on infrastructures built along sedimentary coastlines than on natural rocky reefs or infrastructures built close to rocky coastlines. Conversely, native species were twice as abundant on natural reefs than on nearby infrastructures and were scarce to virtually absent on infrastructures built along sedimentary coasts. The species composition of assemblages in artificial habitats was more similar to that of marinas than of natural reefs, independently of their location. MAIN CONCLUSIONS: Our results show that marine infrastructures along sandy shores disproportionally favour non‐indigenous over native hard bottom species, affecting their spread at regional scales. This is particularly concerning for coastal areas that have low natural densities of rocky reef habitats. We discuss design and management options to improve the quality as habitat of marine infrastructures and to favour their preferential use by native species over non‐indigenous ones.
Summary
Concerns regarding the rapid loss of endemic biodiversity, and introduction and spread of non‐indigenous species, have focused attention on the need and ability to detect species present in ...communities at low abundance. However, detection of rare species poses immense technical challenges, especially for morphologically cryptic species, microscopic taxa and those beneath the water surface in aquatic ecosystems.
Next‐generation sequencing technology provides a robust tool to assess biodiversity, especially for detection of rare species. Here, we assess the sensitivity of 454 pyrosequencing for detection of rare species using known indicator species spiked into existing complex plankton samples. In addition, we develop universal small subunit ribosomal DNA primers for amplification of a wide range of taxa for detailed description of biodiversity in complex communities.
A universality test of newly designed primers for the hypervariable V4 region of the nuclear small subunit ribosomal DNA (V4‐nSSU) using a plankton sample collected from Hamilton Harbor showed that 454 pyrosequencing based on this universal primer pair can recover a wide range of taxa, including animals, plants (algae), fungi, blue‐green algae and protists.
A sensitivity test showed that 454 pyrosequencing based on newly designed universal V4‐nSSU primers was extremely sensitive for detection of very rare species. Pyrosequencing was able to recover spiked indicator species with biomass percentage as low as approximately 2·3 × 10−5% when 24 artificially assembled samples were tagged and sequenced in one PicoTiter plate (i.e. sequencing depth of an equivalent of 1/24 PicoTiter plate). In addition, spiked rare species were sometimes recovered as singletons (i.e. Operational Taxonomic Units represented by a single sequence), suggesting that at least some singletons are informative for recovering unique lineages in ‘rare biospheres’.
The method established here allows biologists to better investigate the composition of aquatic communities, especially for detection of rare taxa. Despite a small‐scale pyrosequencing effort, we demonstrate the extreme sensitivity of pyrosequencing using rare species spiked into plankton samples. We propose that the method is a powerful tool for detection of rare native and/or alien species.
Ecology Letters (2010) 13: 947-958 What determines invasiveness of alien organisms is among the most interesting and urgent questions in ecology. In attempts to answer this question, researchers ...compare invasive alien species either to native species or to non-invasive alien species, and this is done in either the introduced or native ranges. However, inferences that can be drawn from these comparisons differ considerably, and failure to recognize this could hamper the search for determinants of invasiveness. To increase awareness about this issue, we present a framework of the various comparisons that can be used to test for determinants of invasiveness, and the specific questions each comparison can address. Moreover, we discuss how different comparisons complement each other, and therefore should be used in concert. For progress in invasion biology, it is crucial to realize that different comparisons address different biological questions and that some questions can only be answered unambiguously by combining them.
Aim
Environmental DNA metabarcoding has recently emerged as a non‐invasive tool for aquatic biodiversity inventories, frequently surpassing traditional methods for detecting a wide range of taxa in ...most habitats. The major limitation currently impairing the large‐scale application of eDNA‐based inventories is the lack of species sequences available in public genetic databases. Unfortunately, these gaps are still unknown spatially and taxonomically, hindering targeted future sequencing efforts.
Innovation
We propose GAPeDNA, a user‐friendly web interface that provides a global overview of genetic database completeness for a given taxon across space and conservation status. As an application, we synthetized data from regional checklists for marine and freshwater fishes along with their IUCN conservation status to provide global maps of species coverage using the European Nucleotide Archive public reference database for 19 metabarcoding primers. This tool automatizes the scanning of gaps in these databases to guide future sequencing efforts and support the deployment of eDNA inventories at larger scale. This tool is flexible and can be expanded to other taxa and primers upon data availability.
Main conclusions
Using our global fish case study, we show that gaps increase towards the tropics where species diversity and the number of threatened species are the highest. It highlights priority areas for fish sequencing like the Congo, the Mekong and the Mississippi freshwater basins which host more than 60 non‐sequenced threatened fish species. For marine fishes, the Caribbean and East Africa host up to 42 non‐sequenced threatened species. By presenting the global genetic database completeness for several primers on any taxa and building an open‐access, updatable and flexible tool, GAPeDNA appears as a valuable contribution to support any kind of eDNA metabarcoding study.
With the entry into force of the International Maritime Organization's International Ballast Water Management Convention in September 2017, ships have begun to install and operate onboard ballast ...water management systems (BWMS) to reduce the number of live organisms in ballast water. Scientific methods were developed to assess the effectiveness of BWMS at reducing the number of live organisms in ballast water. However, detecting low organism concentrations in treated ballast water is challenging when considering the small sample volume (6 mL) analyzed for organisms in the 10–50 μm size class. The volume analyzed can be increased by concentrating the sample prior to analysis, but it is important to assess the effects of the sample concentration method due to potential cell loss experienced during the concentration step. Therefore, laboratory experiments were conducted to assess the effects of a gravity filtration method to concentrate samples to a factor of 40:1. Experiments were conducted for both low and high organism abundances. For unpreserved samples at low organism abundances (∼10 cells mL−1), concentrated samples had on average 31% fewer live cells mL−1 than unconcentrated samples for four out of five experiments. At high organism abundances (≥ 120 cells mL−1), unpreserved concentrated samples had on average 55% fewer live cells than unconcentrated samples. Alternatively, with preserved samples at low organism abundances, concentrated samples had on average 4.5× more cells than unconcentrated samples. At high organism abundances, concentrated samples had on average 6.4× more cells than unconcentrated samples. Differences were also observed between preserved and unpreserved samples. These findings can help to improve ballast water monitoring procedures and BWMS assessments, addressing a critical challenge to maritime environmental protection.
•Concentration by gravity filtration led to cell loss in live phytoplankton samples.•Alternate concentration methods should be explored to mitigate cell loss.•Concentrating had different effects on live and preserved samples.•Concentrating samples is important to improve precision at low abundances.•Counting six subsamples improved precision in low abundance samples.
At 726 the number of recorded multicellular non indigenous species (NIS) in the Mediterranean Sea is far higher than in other European Seas. Of these, 614 have established populations in the sea. 384 ...are considered Erythraean NIS, the balance are mostly ship and culture-introductions. In order to effectively implement EU Regulation on the prevention and management of the introduction and spread of invasive NIS and the Marine Strategy Framework Directive in the Mediterranean Sea it is crucial that this priority pathway is appropriately managed. Three potential impediments – incomplete and inaccurate data; unknown impacts; policy mismatch – hinder implementation. Current geographical, taxonomical and impact data gaps will be reduced only by instituting harmonized standards and methodologies for monitoring NIS populations in all countries bordering the Mediterranean Sea, prioritizing bridgehead sites and dispersal hubs. The option of implementing European environmental policies concerning marine NIS in member states alone may seem expedient, but piecemeal protection is futile. Since only 9 of the 23 states bordering the Mediterranean are EU member states, the crucial element for an effective strategy for slowing the influx of NIS is policy coordination with the Regional Sea Convention (Barcelona Convention) to ensure consistency in legal rules, standards and institutional structures to address all major vectors/pathways.
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•384 of the 614 NIS established in the Mediterranean were introduced via Suez Canal.•Erythraean NIS expanded westwards and northward in the past two decades.•To comply with regulations priority pathways should be appropriately managed.•Regulation enforcement depends on policy coordination with non EU Member States.•Data gaps reduced by standardizing methodologies in all Mediterranean countries.
A recent trend in invasion ecology relates the success of non‐indigenous species (NIS) to reduced control by enemies such as pathogens, parasites and predators (i.e. the enemy release hypothesis, ...ERH). Despite the demonstrated importance of enemies to host population dynamics, studies of the ERH are split – biogeographical analyses primarily show a reduction in the diversity of enemies in the introduced range compared with the native range, while community studies imply that NIS are no less affected by enemies than native species in the invaded community. A broad review of the invasion literature implies at least eight non‐exclusive explanations for this enigma. In addition, we argue that the ERH has often been accepted uncritically wherever (i) NIS often appear larger, more fecund, or somehow ‘better’ than either congeners in the introduced region, or conspecifics in the native range; and (ii) known enemies are conspicuously absent from the introduced range. However, all NIS, regardless of their abundance or impact, will lose natural enemies at a biogeographical scale. Given the complexity of processes that underlie biological invasions, we argue against a simple relationship between enemy ‘release’ and the vigour, abundance or impact of NIS.
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