Patterns of invasion in the Laurentian Great Lakes in relation to changes in vector activity

The Laurentian Great Lakes basin has been invaded by at least 182 non-indigenous species. A new invader is discovered every 28 weeks, which is the highest rate recorded for a freshwater ecosystem. Over the past century, invasions have occurred in phases linked to changes in the dominant vectors. The number of ship-vectored invaders recorded per decade is correlated with the intensity of vessel traffic within the basin. Ballast water release from ocean vessels is the putative vector for 65% of all invasions recorded since the opening of the St. Lawrence Seaway in 1959. As a preventive measure, ocean vessels have been required since 1993 to exchange their freshwater or estuarine ballast with highly saline ocean water prior to entering the Great Lakes. However, this procedure has not prevented ship-vectored species introductions. Most ships visiting the Great Lakes declare 'no ballast on board' (NOBOB) and are exempt from the regulation, even though they carry residual water that is discharged into the Great Lakes during their activities of off-loading inbound cargo and loading outbound cargo. Recently introduced species consist predominantly of benthic invertebrates with broad salinity tolerance. Such species are most likely to survive in a ballast tank following ballast water exchange, as well as transport in the residual water and tank sediments of NOBOB ships. Thus, the Great Lakes remain at risk of being invaded by dozens of euryhaline invertebrates that have spread into Eurasian ports from whence originates the bulk of foreign ships visiting the basin.     

Modeling the secondary spread of viral hemorrhagic septicemia virus (VHSV) by commercial shipping in the Laurentian Great Lakes

Researchers have only begun to study the role of shipping in the spread of invasive species in the Laurentian Great Lakes despite a well-documented history of introductions in these lakes due to ballast water release. Here, we determine whether ballast water discharge was a likely vector of spread of the fish disease, viral hemorrhagic septicemia virus genotype IVb (VHSV-IVb), throughout the Great Lakes and St. Lawrence Seaway. Three models were developed to assess whether the spread of VHSV was due to (1) chance (random model), or (2) ballast water discharge (location model), and whether (3) increased propagule pressure, as measured by the number of visitations by ships carrying ballast water from VHSV infected areas, increased the likelihood of a discharge location becoming infected with VHSV (propagule pressure model). The third model was also used to assess the probable point of initial introduction of VHSV. Presence and absence accuracies and weighted Cohen’s kappa were calculated to determine which models best predicted observed presences and absences of VHSV. Location models explain the patterns of VHSV detections better than random models, and inclusion of “propagule pressure” often improved model fit; however, the relationship is weak likely because of a long lag time between introduction and detection, a high rate of false negatives in reporting, and the possible contribution of other vectors of spread. Montreal was also identified as the more likely introduction site of VHSV, rather than Lake St. Clair, the site where the virus was first detected.     

The Importance of Ship Hull Fouling as a Vector of Species Introductions into the North Sea

Ships have long been recognized as a major vector for the introduction of non-native and harmful organisms. From 1992 to 1996 a shipping study was undertaken in Germany, focusing on the fauna transported by ships, to assess the importance of species introductions by international shipping traffic. Ballast water, tank sediment or hull fouling of 186 vessels was sampled. A total of 257 species were identified, ranging from Foraminifera to Teleostei, and 57% of the species sampled were considered to be non-native to the North Sea region, originating from elsewhere in the world including the north eastern Atlantic (west of the English Channel). Non-native species were recorded in 38% of all ballast water samples, 57% of all sediment samples and 96% of all hull samples, indicating that hull fouling is an important vector of introduction. Four species (1.6%) of unknown origin (cryptogenic species) were identified. The potential for establishment in the North Sea region of all non-native species found was classified into three categories based on the degree of similarity of climatic conditions in the North Sea and the donor region. Based on this criterion 19 of the species found in the fouling communities on ships' hulls were deemed to have a high potential for establishment in the North Sea.     

Domestic ships as a potential pathway of nonindigenous species from the Saint Lawrence River to the Great Lakes

Ballast water moved by transoceanic vessels has been recognized globally as a predominant vector for the introduction of aquatic nonindigenous species (NIS). In contrast, domestic ships operating within confined geographic areas have been viewed as low risk for invasions, and are exempt from regulation in consequence. We examined if the St. Lawrence River could serve as a source of NIS for the Laurentian Great Lakes by surveying ballast water carried by domestic vessels and comparing biological composition in predominant St. Lawrence River—Great Lakes port-pairs in order to determine the likelihood that NIS could be transported to, and survive in, the Great Lakes. Thirteen potential invaders were sampled from ballast water, while 26 taxa sampled from St. Lawrence River ports are not reported from the Great Lakes. The majority of NIS recorded in samples are marine species with low potential for survival in the Great Lakes, however two euryhaline species (copepod Oithona similis, and amphipod Gammarus palustris) and two taxa reported from brackish waters (copepod Microsetella norvegica and decapod Cancer irroratus) may pose a risk for invasion. In addition, four marine NIS were collected in freshwater samples indicating that at least a subset of marine species have potential as new invaders to the Great Lakes. Based on results from this study, the ports of Montreal, Sorel, Tracy and Trois Rivières appear to pose the highest risk for new ballast-mediated NIS from the St. Lawrence River to the Great Lakes.     

Molecular detection of invasive species in heterogeneous mixtures using a microfluidic carbon nanotube platform

Screening methods to prevent introductions of invasive species are critical for the protection of environmental and economic benefits provided by native species and uninvaded ecosystems. Coastal ecosystems worldwide remain vulnerable to damage from aquatic species introductions, particularly via ballast water discharge from ships. Because current ballast management practices are not completely effective, rapid and sensitive screening methods are needed for on-site testing of ships in transit. Here, we describe a detection technology based on a microfluidic chip containing DNA oligonucleotide functionalized carbon nanotubes. We demonstrate the efficacy of the chip using three ballast-transported species either established (Dreissena bugensis) or of potential threat (Eriocheir sinensis and Limnoperna fortuneii) to the Laurentian Great Lakes. With further refinement for on-board application, the technology could lead to real-time ballast water screening to improve ship-specific management and control decisions.     

Anguilla rostrata glass eel migration and recruitment in the estuary and Gulf of St Lawrence

This study describes catches of Anguilla rostrata glass eels and associated oceanographic conditions in the St Lawrence Estuary and Gulf. Ichthyoplankton survey data suggest that they enter the Gulf primarily in May, migrate at the surface at night, and disperse broadly once they have passed Cabot Strait. They arrive in estuaries beginning at about mid-June and through the month of July. Migration extends west up to Québec City, in the freshwater zone of the St Lawrence Estuary, 1000 km west of Cabot Strait. Anguilla rostrata glass eels travel between Cabot Strait and receiving estuaries at a straight-line ground speed of c. 10–15 km day⁻¹. Catches of fish per unit effort in estuaries in the St Lawrence system are much lower than those reported for the Atlantic coast of Canada. Low abundance of A. rostrata glass eels in the St Lawrence system may be due to cold surface temperatures during the migration period which decrease swimming capacity, long distances from the spawning ground to Cabot Strait and from Cabot Strait to the destination waters (especially the St Lawrence River), complex circulation patterns, and hypoxic conditions in bottom waters of the Laurentian Channel and the St Lawrence Estuary.     

Linking environmental conditions and ship movements to estimate invasive species transport across the global shipping network

Aim Some nations, and the International Maritime Organization, are moving towards requirements for managing ballast water to reduce the number of alien species transported and released. These and other measures will be most efficient when targeted at ships posing the greatest risks. Here, we analyse world‐wide ship movements and port environmental conditions to explore how these risk components differ across arriving ships. Location Global, with a case study of the Laurentian Great Lakes. Methods We gathered salinity and temperature data for all global shipping ports, and data for all global ship movements during a 12 ‐month period. We applied these data to the Laurentian Great Lakes to determine which global ports may donate new species to the Great Lakes via ship traffic, and which are most environmentally similar to the Great Lakes. Results We show that ships regularly travel to the Great Lakes from all major coastal, and many inland, regions of the world. Most global ports, and thus the species in them, are separated from the Great Lakes by no more than two ship voyages. Combined with a measure of environmental similarity among global ports, we identify ship routes likely to transport species adapted for survival in the Great Lakes and global regions that may be the source of increasing future invasions. Main conclusions The Great Lakes account for a small fraction of global shipping yet are closely connected to all other ports, and the species in them, by the shipping network. Our methods and data allow risks from individual ships to be ranked so that management activities can be targeted at ships most likely to introduce new invaders. Because our data sets are global, they could be applied to ship arrivals at any global port.     

Viability of invertebrate diapausing eggs exposed to saltwater: implications for Great Lakes’ ship ballast management

International shipping has been the dominant vector of nonindigenous species introductions to the Laurentian Great Lakes over the past century. Apparent ballast-mediated invasions have been recorded in recent years, despite the implementation of voluntary ballast water exchange regulations in 1989. Since unregulated no-ballast-on-board vessels currently dominate inbound traffic to the Great Lakes, it has been proposed that live or dormant organisms contained in residual ballast of these vessels may be partially responsible for recent invasions. Alternatively, euryhaline species may pose a significant invasion threat because they can potentially survive ballast exchange. In this study, we explored whether exposure to open-ocean water (32) reduced the viability of invertebrate diapausing eggs in ballast sediments. Sediments collected from three transoceanic ships and from three freshwater habitats were exposed to open-ocean seawater. Egg viability, assessed as the abundance of taxa hatched between exposed and unexposed sediments, was not affected by saltwater exposure in any experiment. Species richness of hatched diapausing eggs was reduced by saltwater exposure in only one of seven trials. Our results indicate that oligostenohaline zooplankton may pose an invasion risk because their diapausing eggs are largely resistant to exposure to open-ocean saltwater.     

Recent mass invasion of the North American Great Lakes by Ponto-Caspian species

The North American Great Lakes have been invaded and dramatically altered by more than 145 alien species. Many invasions have occurred during the past few decades because of the release of Eurasian ballast water from transoceanic ships. Current regulations require ships to exchange foreign ballast with highly saline water before entering the Great Lakes; this procedure should prevent colonization by strictly freshwater species, but species with broad salinity tolerance might survive transport in exchanged water. A recent series of invasions by euryhaline organisms from the Black and Caspian Seas region signals a new phase in the transformation of the Great Lakes - one that supports the concept of an 'invasional meltdown'.     

Stratigraphic and sedimentological evidence for late Wisconsinan sub-glacial outburst floods to Laurentian Fan

Sub-glacial meltwater produces a distinctive stratigraphic and sedimentological response on the continental margin. Seismo-stratigraphy of Laurentian Channel reveals thick till deposits at its seaward end that pass laterally into stratified sediment in deeper basins, that may record periods of water build up beneath the ice. Two scales of meltwater discharge are recognised: large scale that caused catastrophic erosion and transported large volumes of coarse sediment to the abyssal plain and smaller scale, yielding principally muddy sediment. Sub-glacial outburst floods from the Laurentian Channel ice stream delivered distinctive red sediment derived from Permian-Carboniferous strata of the Gulf of St. Lawrence directly to Laurentian Fan between ca. 17 and 14 ¹⁴C ka, separate from North Atlantic Heinrich events. On levees of Laurentian Fan, three major pulses of meltwater plume muds are separated by intervals dominated by hemipelagic sediments. These meltwater intervals are recognised distally as periods of plume sedimentation on the Scotian Slope and ice-rafting of hematite-stained quartz to the North Atlantic Ocean. In channels of Laurentian Fan, at least one major sediment transport event is recognised that eroded the upper slope and the major fan valleys, depositing a bed of gravel at least 3 m thick in the characteristically wide fan valleys and thick sand on the Sohm Abyssal Plain. The same event was probably responsible for giant flute-like scours. The age of the gravel bed is directly constrained only by the presence of local overlying Holocene sediment. Much of the surface of the gravel bed was re-worked by the 1929 “Grand Banks” turbidity current. An erosional event on the upper slope, likely correlative with the flood-generated gravel bed, has been dated at 16.5 ¹⁴C ka. Such large scale erosional flood events can be recognised back through several glacial cycles and have played an important role in the architectural evolution of Laurentian Fan.     

Efficacy of ‘saltwater flushing’ in protecting the Great Lakes from biological invasions by invertebrate eggs in ships’ ballast sediment

1. Mid‐ocean exchange and saltwater flushing were implemented as management practices to reduce the likelihood of new biological invasions in the Laurentian Great Lakes associated with ships’ ballast water and sediments. Despite this, there has been no formal assessment of the efficacy of these procedures. Here, we conduct a comparative analysis of community composition of dormant taxa transported by ballast sediment before and after regulations came into effect in 2006. 2. Ballast sediment samples were collected from 17 ships during the post‐regulation interval of 2007 and 2008. Invertebrate eggs were counted, hatched and species identified in the laboratory. Results were compared to similar samples collected from 39 ships between 2000 and 2002, prior to implementation of saltwater flushing regulations. 3. The estimated amount of residual ballast sediment transported by vessels was significantly lower during the post‐regulation period, ranging from <1 to 45 tonnes per ship, with an average of 5 tonnes. Mean density and number of dormant viable eggs per ship declined 91 and 81%, respectively. 4. Community composition also changed through time, with Rotifera accounting for 78% of taxa transported prior to regulation, whereas Cladocera and Copepoda each accounted for 38% of abundance post‐regulation. Although the number of non‐indigenous species (NIS) declined 73% per ship after 2006, the reduction was not statistically significant; however, the number of freshwater NIS – which pose the greatest risk of invasion for the Great Lakes – was significantly lowered. 5. Our comparative analysis suggests that ballast management regulations enacted in 2006 markedly reduced the probability of introduction of NIS via dormant eggs carried in ballast sediments.     

Introduced marine species of the North Sea coasts

About 80 non-indigenous species are assumed to have been introduced into the North Sea by transoceanic shipping and aquaculture. The number is certainly underestimated as most small organisms received insufficient attention at the species level. Also, the seafaring tradition of the North Sea countries is much longer than our biological surveys are. Most exotic invertebrates originate from the western Atlantic and were introduced by shipping, while most algae stem from the Pacific and came with the introduced oysters. A peak of newcomers was observed in the 1970s. Most of the arrivals became established in brackish environments, at harbor sites and in the vicinity of oyster farms, fouling on hard substrates or living as epibionts. A few live in sediments, are holoplanktonic or are parasites. At the open coast, approximately 6% of the macrobenthic species are exotics, while in estuaries their share is up to 20%. Most exotics have been encountered in the southern North Sea first, and many did not spread further north. About 25% of the established non-natives are widespread and attain locally high abundances. As a consequence, some inshore habitats are entirely dominated by exotics. The overall effect on the ecosystem seems to be more additive than one of displacement. This suggests that the coastal biota of the North Sea are quite capable of accommodating newcomers. However, this is no guarantee that the next introduced species may not cause severe ecological change or economic harm. There is a need to minimize the risk of unintentional introductions by ballast water treatment and by adhering to quarantine procedures in aquaculture. Current research on exotics in the North Sea is regarded as inadequate for proper evaluation and management requirements.     

The role of constructed reefs in non-indigenous species introductions and range expansions

Constructed reefs can contribute to non-indigenous species (NIS) introductions or range expansions in several ways. Reef materials that retain developed fouling communities or ballast, such as decommissioned petroleum platforms, inactive or derelict ships, and bridge rubble are potential NIS vectors. Habitat provided by reefs placed in areas devoid of natural hard bottom or structure may be colonized by NIS propagules dispersed from natural or anthropogenic sources. A network of reef structures may also create NIS corridors for linking previously unconnected areas. Due to its level of offshore habitat alteration, changing environmental conditions, volume of shipping and boating traffic, and subtropical location, the Gulf of Mexico may be particularly vulnerable to NIS introductions and has a number of documented NIS. Non-indigenous or invasive species concerns have delayed and increased costs for some recent reef construction projects. The linkages between NIS and constructed reefs are reviewed, and approaches for anticipating, assessing, and controlling introductions are recommended. Using basic information about NIS risks, reef planners can begin to evaluate unintended consequences and incorporate risk management measures to reduce future introductions. Prevention is the most effective risk reduction approach because controlling marine NIS after introduction is expensive and offers limited probability for success.     

Episodic global dispersal in shallow water marine organisms: the case history of the European shore crabs Carcinus maenas and C. aestuarii

Aim This paper evaluates global collection records, evidence of anthropogenic transport methods, and experimental and distributional data relative to temperature requirements to understand the historical and potential dispersal of a well‐known genus of estuarine crab. Location The records analysed are from temperate and tropical coastal ocean areas. Methods The study is based primarily on literature analysis and examination of museum specimens. Results The human‐mediated successful global dispersal of the European shore crabs Carcinus maenas (Linnaeus, 1758) and C. aestuarii (Nardo, 1847) occurred in three major episodes: around 1800, in the 1850s–70s, and in the 1980s–90s. The nineteenth century introductions occurred through transport by ships (probably in hull fouling or in solid ballast), while the introductions in the 1980s could have occurred through a greater variety of dispersal mechanisms (ships’ hull fouling and seawater system fouling; fouling on semisubmersible drilling platforms; ballast water; transport with fisheries products intended for food or bait; scientific research; releases from aquaria maintained for educational or scientific purposes; or intentional non‐governmental releases for human food production). These introductions have resulted in Carcinus’ establishment in five temperate regions outside of its native Europe in Atlantic North America, Australia, South Africa, Japan and Pacific North America, while releases into tropical regions have not established populations. C. maenas’ range in both its native and introduced regions appears to be regulated by similar temperature parameters, enabling an assessment of its potential distribution. Main conclusions The second episode of Carcinus’ global dispersal, the period from the 1850s to 1870s, may be part of a broader surge of world‐wide invasions caused by an increase in shipping.     

Relative Invasion Risk for Plankton across Marine and Freshwater Systems: Examining Efficacy of Proposed International Ballast Water Discharge Standards

Understanding the implications of different management strategies is necessary to identify best conservation trajectories for ecosystems exposed to anthropogenic stressors. For example, science-based risk assessments at large scales are needed to understand efficacy of different vector management approaches aimed at preventing biological invasions associated with commercial shipping. We conducted a landscape-scale analysis to examine the relative invasion risk of ballast water discharges among different shipping pathways (e.g., Transoceanic, Coastal or Domestic), ecosystems (e.g., freshwater, brackish and marine), and timescales (annual and per discharge event) under current and future management regimes. The arrival and survival potential of nonindigenous species (NIS) was estimated based on directional shipping networks and their associated propagule pressure, environmental similarity between donor-recipient ecosystems (based on salinity and temperature), and effects of current and future management strategies (i.e., ballast water exchange and treatment to meet proposed international biological discharge standards). Our findings show that current requirements for ballast water exchange effectively reduce invasion risk to freshwater ecosystems but are less protective of marine ecosystems because of greater environmental mismatch between source (oceanic) and recipient (freshwater) ecoregions. Future requirements for ballast water treatment are expected to reduce risk of zooplankton NIS introductions across ecosystem types but are expected to be less effective in reducing risk of phytoplankton NIS. This large-scale risk assessment across heterogeneous ecosystems represents a major step towards understanding the likelihood of invasion in relation to shipping networks, the relative efficacy of different invasion management regimes and seizing opportunities to reduce the ecological and economic implications of biological invasions.     

Importance of geographic origin for invasion success: A case study of the North and Baltic Seas versus the Great Lakes–St. Lawrence River region

Recently, several studies indicated that species from the Ponto‐Caspian region may be evolutionarily predisposed to become nonindigenous species (NIS); however, origin of NIS established in different regions has rarely been compared to confirm these statements. More importantly, if species from certain area/s are proven to be better colonizers, management strategies to control transport vectors coming from those areas must be more stringent, as prevention of new introductions is a cheaper and more effective strategy than eradication or control of established NIS populations. To determine whether species evolved in certain areas have inherent advantages over other species in colonizing new habitats, we explored NIS established in the North and Baltic Seas and Great Lakes–St. Lawrence River regions—two areas intensively studied in concern to NIS, highly invaded by Ponto‐Caspian species and with different salinity patterns (marine vs. freshwater). We compared observed numbers of NIS in these two regions to expected numbers of NIS from major donor regions. The expected numbers were calculated based on the available species pool from donor regions, frequency of shipping transit, and an environmental match between donor and recipient regions. A total of 281 NIS established in the North and Baltic Seas and 188 in the Great Lakes–St. Lawrence River. Ponto‐Caspian taxa colonized both types of habitats, saltwater areas of the North and Baltic Seas and freshwater of the Great Lakes–St. Lawrence River, in much higher numbers than expected. Propagule pressure (i.e., number of introduced individuals or introduction effort) is of great importance for establishment success of NIS; however in our study, either shipping vector or environmental match between regions did not clarify the high numbers of Ponto‐Caspian taxa in our study areas. Although we cannot exclude the influence of other transport vectors, our findings suggest that the origin of the species plays an important role for the predisposition of successful invaders.     

International dissemination of epidemic Vibrio cholerae by cargo ship ballast and other nonpotable waters.

In 1991 and 1992, toxigenic Vibrio cholerae O1, serotype Inaba, biotype El Tor, was recovered from nonpotable (ballast, bilge, and sewage) water from five cargo ships docked in ports of the U.S. Gulf of Mexico. Four of these ships had taken on ballast water in cholera-infected countries; the fifth took on ballast in a noninfected country. Isolates examined by pulsed-field gel electrophoresis were indistinguishable from the Latin American epidemic strain, C6707; however, they differed significantly from the endemic Gulf Coast strain (VRL 1984), the sixth-pandemic strain (569-B), and a V. cholerae non-O1 strain isolated from a ship arriving from a foreign port. On the basis of our findings, the Food and Drug Administration recommended that the U.S. Coast Guard issue an advisory to shipping agents and captains requesting that ballast waters be exchanged on the high seas before entry of ships into U.S. ports.     

船舶压载水携带外来物种及其对水域生态环境的影响

随着世界航运业的不断发展 ,船舶频繁地来往于世界各地 ,船舶压载水作为一种外来物种传播的有效媒介物 ,使得不同水域的物种得以传播到其他水域。这些物种种类繁多 ,普遍具有适应性较强 ,危害性较大等特点。本文就一些世界范围内广泛传播的典型外来物种进行了列举 ,并对它们的分类、发源地、传入媒介、习性、分布以及由于它们的入侵给当地的生态环境、人类健康、经济带来的不利影响以及当地所采取的一些治理措施等进行了较为详细地阐述。提出控制船舶压载水的排放以及研究较为有效的压载水治理方法是解决压载水携带外来物种入侵性传播的有效途径     

Assessing invasion risk across taxa and habitats: life stage as a determinant of invasion success

Aim Many aquatic invertebrates produce dormant life‐history stages as a means to endure inhospitable environments and to facilitate natural long‐distance dispersal, yet we have little understanding of the role of dormant stages as a mechanism for human‐mediated introductions of non‐indigenous species. We explore the survival of invertebrate dormant eggs in collected ships’ ballast sediment over a 1‐year period to determine relative invasion potential across taxa (i.e. rotifers, copepods, cladocerans and bryozoans) and different habitats (freshwater, marine). Location Canadian Atlantic and Pacific coasts and Laurentian Great Lakes. Methods During 2007 and 2008, 19 ballast samples were collected as a part of a larger study. The degradation rate of dormant eggs was assessed by enumerating dormant eggs and by conducting viability hatching experiments. Results Taxa examined included rotifers, copepods, anomopods, onychopods and bryozoans. Dormant eggs of rotifers degraded at the highest rate of all taxa examined, with no viable eggs remaining within 10 months. Copepods showed a less rapid degradation rate than rotifers. The degradation rate of anomopod dormant eggs was significantly slower than that of both rotifers and copepods. Onychopods and bryozoans did not visibly degrade at all over 12 months. Viability hatching experiments were successful for rotifers, copepods, and anomopods. Onychopods and bryozoans did not hatch during any of the three hatching trials. Main conclusions Dormancy is not equally beneficial to all invertebrate taxa. Our results indicate that dormant eggs of rotifers and copepods degrade at a rapid rate and may not pose high invasion risk. In contrast, the slow degradation rate of anomopod dormant eggs and the lack of degradation of onychopod and bryozoan dormant eggs could result in high invasion risk because of their accumulation in ballast tanks. Species having resistant dormant eggs mostly belong to freshwater taxa making freshwater habitats at higher invasion risk by dormant invertebrates than marine habitats.