Antarctica: The final frontier for marine biological invasions

Antarctica is experiencing significant ecological and environmental change, which may facilitate the establishment of non‐native marine species. Non‐native marine species will interact with other anthropogenic stressors affecting Antarctic ecosystems, such as climate change (warming, ocean acidification) and pollution, with irreversible ramifications for biodiversity and ecosystem services. We review current knowledge of non‐native marine species in the Antarctic region, the physical and physiological factors that resist establishment of non‐native marine species, changes to resistance under climate change, the role of legislation in limiting marine introductions, and the effect of increasing human activity on vectors and pathways of introduction. Evidence of non‐native marine species is limited: just four marine non‐native and one cryptogenic species that were likely introduced anthropogenically have been reported freely living in Antarctic or sub‐Antarctic waters, but no established populations have been reported; an additional six species have been observed in pathways to Antarctica that are potentially at risk of becoming invasive. We present estimates of the intensity of ship activity across fishing, tourism and research sectors: there may be approximately 180 vessels and 500+ voyages in Antarctic waters annually. However, these estimates are necessarily speculative because relevant data are scarce. To facilitate well‐informed policy and management, we make recommendations for future research into the likelihood of marine biological invasions in the Antarctic region.     

Evolutionary game theory as a framework for studying biological invasions

Although biological invasions pose serious threats to biodiversity, they also provide the opportunity to better understand interactions between the ecological and evolutionary processes structuring populations and communities. However, ecoevolutionary frameworks for studying species invasions are lacking. We propose using game theory and the concept of an evolutionarily stable strategy (ESS) as a conceptual framework for integrating the ecological and evolutionary dynamics of invasions. We suggest that the pathways by which a recipient community may have no ESS provide mechanistic hypotheses for how such communities may be vulnerable to invasion and how invaders can exploit these vulnerabilities. We distinguish among these pathways by formalizing the evolutionary contexts of the invader relative to the recipient community. We model both the ecological and the adaptive dynamics of the interacting species. We show how the ESS concept provides new mechanistic hypotheses for when invasions result in long- or short-term increases in biodiversity, species replacement, and subsequent evolutionary changes.     

Trends in marine biological invasions at local and regional scales: the Northeast Pacific Ocean as a model system

Introduced species are an increasing agent of global change. Biogeographic comparisons of introduced biotas at regional and global scales can clarify trends in source regions, invasion pathways, sink regions, and survey effort. We identify the Northeast Pacific Ocean (NEP; northern California to British Columbia) as a model system for analyzing patterns of marine invasion success in cool temperate waters. We review literature and field surveys, documenting 123 introduced invertebrate, algal, fish, and vascular plant species in the NEP. Major invasion pathways were shipping (hull fouling, solid and water ballast; 1500s-present) and shellfish (particularly oysters) and finfish imports (commonest from the 1870s to mid-1900s). The cumulative number of successful invasions over time increased at linear, quadratic, and exponential rates for different taxa, pathways, and regions within the NEP. Regional analysis of four major NEP estuaries showed that Puget Sound and the contiguous Straits had the most introduced species, followed by Humboldt Bay, Coos Bay and Willapa Bay. Data on cumulative shipping volumes predicted smaller-scale, but not larger-scale spatial patterns in the number of shipping-mediated invasions. We identify the major challenges in scaling up from regional to global invasion analysis in cool temperate regions. Retrospective analyses for distinct biogeographic regions such as the NEP provide insight into vector dynamics and regional invasibility, and are a necessary foundation for monitoring and managing global change caused by biotic invasions.     

海洋外来物种入侵生态学研究

海洋外来物种入侵已成为最为严重的全球性环境问题之一。海洋生态系统类型多样、环境复杂,其生物入侵的监测、控制与管理难度相对较大。我国对陆地外来生物的入侵已开展了较为深入的研究,但对于海洋外来生物的入侵研究仍处于起步阶段,对其入侵监测、入侵机制、入侵危害的程度以及防治等问题缺乏基础数据。本文在分析国内外海洋外来生物入侵现状的基础上,概述其入侵生态学研究形势及相关成果,包括海洋外来物种的入侵途径、入侵过程、入侵生态效应以及全球变化对入侵的影响等。海洋外来生物的入侵可能对海洋生态系统造成直接或间接的影响,如种间竞争破坏生态环境、与土著种杂交造成遗传污染、病原生物及有毒藻类导致海洋生态灾害加剧等。此外,从政策和法规、入侵风险评估、监测和公共宣传教育、生物信息系统和有效管理机制等方面提出对我国海洋外来物种入侵的防治策略。本研究为我国海洋外来物种的进一步研究提供了参考。     

The influence of antifouling practices on marine invasions

Vessel hull-fouling is increasingly recognised as one of the major vectors for the transfer of marine non-indigenous species. For hundreds of years, copper (Cu) has been used as a primary biocide to prevent the establishment of fouling assemblages on ships' hulls. Some non-indigenous fouling taxa continue to be transferred via hull-fouling despite the presence of Cu antifouling biocides. In addition, several of these species appear to enjoy a competitive advantage over similar native taxa within metal-polluted environments. This metal tolerance may further assist their establishment and spread in new habitats. This review synthesises existing research on the links between Cu and the invasion of fouling species, and shows that, with respect to the vector of hull-fouling, tolerance to Cu has the potential to play a role in the transfer of non-indigenous fouling organisms. Also highlighted are the future directions for research into this important nexus between industry, ecology and environmental management.     

Diluting the founder effect: cryptic invasions expand a marine invader's range

Most invasion histories include an estimated arrival time, followed by range expansion. Yet, such linear progression may not tell the entire story. The European green crab (Carcinus maenas) was first recorded in the US in 1817, followed by an episodic expansion of range to the north. Its population has recently exploded in the Canadian Maritimes. Although it has been suggested that this northern expansion is the result of warming sea temperatures or cold-water adaptation, Canadian populations have higher genetic diversity than southern populations, indicating that multiple introductions have occurred in the Maritimes since the 1980s. These new genetic lineages, probably from the northern end of the green crab's native range in Europe, persist in areas that were once thought to be too cold for the original southern invasion front. It is well established that ballast water can contain a wide array of nonindigenous species. Ballast discharge can also deliver genetic variation on a level comparable to that of native populations. Such gene flow not only increases the likelihood of persistence of invasive species, but it can also rapidly expand the range of long-established nonindigenous species.     

Biological invasions drive size increases in marine and estuarine invertebrates

Ecologists have long been fascinated by the morphological changes that species frequently undergo when introduced into new regions. In this study an unusual pattern of size change associated with the invasion of 19 species of marine and estuarine invertebrates is reported. The results show that the majority of species are significantly larger in the introduced range compared with the native range with little evidence for any decrease in size following invasion. This invasion‐driven increase in body size sharply contrasts with the pattern observed in many other taxa including plants, mammals and lizards, where invaders frequently exhibit post‐invasion decreases and increases in size. These size changes were not influenced by differences in latitude, sample size or length of time since invasion. Although several mechanisms, may explain the results, none have been demonstrated.     

Establishment failure in biological invasions: a case history of Littorina littorea in California, USA

Background

The early stages of biological invasions are rarely observed, but can provide significant insight into the invasion process as well as the influence vectors have on invasion success or failure.

Methodology/Principal Findings

We characterized three newly discovered populations of an introduced gastropod, Littorina littorea (Linné, 1758), in California, USA, comparing them to potential source populations in native Europe and the North American East Coast, where the snail is also introduced. Demographic surveys were used to assess spatial distribution and sizes of the snail in San Francisco and Anaheim Bays, California. Mitochondrial DNA was sequenced and compared among these nascent populations, and various populations from the North American East Coast and Europe, to characterize the California populations and ascertain their likely source. Demographic and genetic data were considered together to deduce likely vectors for the California populations. We found that the three large California L. littorea populations contained only adult snails and had unexpectedly high genetic diversity rather than showing an extreme bottleneck as typically expected in recent introductions. Haplotype diversity in Californian populations was significantly reduced compared to European populations, but not compared to East Coast populations. Genetic analyses clearly suggested the East Coast as the source region for the California introductions.

Conclusions and Significance

The California L. littorea populations were at an early, non-established phase of invasion with no evidence of recruitment. The live seafood trade is the most likely invasion vector for these populations, as it preferentially transports large numbers of adult L. littorea, matching the demographic structure of the introduced California L. littorea populations. Our results highlight continued operation of live seafood trade vectors and the influence of vectors on the demographic and genetic structure of the resulting populations, especially early stages of the invasion process.     

The genetics and genomics of marine fish invasions: a global review

Reviews in Fish Biology and Fisheries - We present a comprehensive review of the available global literature on the genetics and genomics of marine fish invasions. Overall this review provides data...     

Seed predation as a barrier to alien conifer invasions

Interactions between exotic plants and animals can play a major role in determining success or failure of plant introductions. Seed predation has been seen as important in explaining biotic resistance to plant invasion, but this hypothesis has rarely been tested. We studied seed predation on exotic forest plants on an island in Patagonia, Argentina where 43 pine species, including 60% of the world’s known invasive Pinaceae, were introduced ca. 80 years ago, but where exotics attain relatively high densities only near the original plantings. To test if seed predation limits exotic conifer establishment in this area, we compared seed predation in areas close to plantations (colonized by exotics) and far from them (not invaded). Seeds of exotics were preferred over seeds of native species, possibly because exotic seeds are bigger. Predation was more intense in areas far from plantations than in areas close to them, substantially reducing the chances of exotic seed establishment. Using automatic cameras, we found that both rodents and birds preyed on exotic seeds. This study suggests that native seed predators can be an important component of biological resistance to plant invasion.     

Aquaculture, non-native salmonid invasions and associated declines of native fishes in Northern Patagonian lakes

1. Even though intensive aquaculture production of salmonids in lakes occurs in many locations around the world published studies on the survival and reproductive success of escaped cultured salmonids in freshwater ecosystems are not common. A recent expansion of aquaculture in Chile has led it to become the world's second largest producer of cultured salmonids.
2. We document the recent history of escaped and self-sustaining salmonid populations over a wide spatial scale and a long temporal scale in Chilean Patagonian lakes. Our hypotheses are that salmonid density in lakes will be higher where there is intensive aquaculture, due to greater numbers of potential escapees. Secondly, if non-native salmonids have adverse impacts on native fishes, increases in the abundance of non-native species should be associated with decreases in relative abundance of native species. Finally, if the first two hypotheses are correct we anticipate that diets of salmonids may show evidence of predation on native fishes, diet overlap with native species, and evidence of the influence of feed from aquaculture operations in the diets of salmonids and native fishes.
3. We sampled six lakes with gill nets from 1992 to 2001. Our results show that the relative abundance of free-living salmonids is closely related to the level of fish farming production. Salmonids are the top predators and in lakes with fish farming the main prey item is native fishes. The relative abundance of native fishes has decreased, most likely due to predation by salmonids.
4. Our study contributes to the understanding of the effects of non-native salmonids in oligotrophic lakes, and it provides a starting point to judge the establishment of new fish farming sites in lakes around the world.     

Bilge water as a vector for the spread of marine pests: a morphological,metabarcoding and experimental assessment

Vessel movements are considered the primary anthropogenic pathway for the secondary spread of marine non-indigenous species. In comparison to the well-studied mechanisms of hull fouling and ballast water, the importance of bilge water for domestic and cross-regional spread of non-indigenous species is largely unknown and has the potential to compromise the overall effectiveness of biosecurity management actions. In this study, the diversity and abundance of biological material contained in bilge water from 30 small vessels (<20 m) was assessed using traditional and molecular identification tools (metabarcoding of the 18S rRNA gene). Laboratory-based studies were also used to investigate the relationship between voyage duration and propagule success. A large taxonomic diversity in organisms was detected, with 118 and 45 distinct taxa identified through molecular and morphological analyses, respectively. Molecular techniques identified five species recognised as non-indigenous to the study region in 23 of the 30 bilge water samples analysed. Larvae and fragments passed through an experimental bilge pump system relatively unharmed. Time spent in the bilge sump was found to affect discharge success, particularly of short-lived and sensitive larvae, but survival for 3 days was observed. Our findings show that bilge water discharges are likely to pose a non-negligible biosecurity threat and that further research to identify high-risk vessel operating profiles and potential mitigation measures are warranted.     

Evaluating dominance as a component of non-native species invasions

Many studies have quantified plant invasions by determining patterns of non‐native species establishment (i.e. richness and absolute cover). Until recently, dominance has been largely overlooked as a significant component of invasion. Therefore, we re‐examined a 6‐year data set of 323 0.1 ha plots within 18 vegetation types collected in the Grand Staircase‐Escalante National Monument from 1998 to 2003, including dominance (i.e. relative cover) in our analyses. We specifically focused on the non‐native species Bromus tectorum, a notable dominant annual grass in this system. We found that non‐native species establishment and dominance are both occurring in species‐rich, mesic vegetation types. Therefore, non‐native species dominance may result despite many equally abundant native species rather than a dominant few, and competitive exclusion does not seem to be a primary control on either non‐native species establishment or dominance in this study. Unlike patterns observed for non‐native species establishment, relative non‐native species cover could not be predicted by native species richness across vegetation types (R² < 0.001; P = 0.45). However, non‐native species richness was found to be positively correlated with relative non‐native species cover and relative B. tectorum cover (R² = 0.46, P < 0.01; R² = 0.17, P < 0.01). Analyses within vegetation types revealed predominantly positive relationships among these variables for the correlations that were significant. Regression tree analyses across vegetation types that included additional biotic and abiotic variables were a little better at predicting non‐native species dominance (PRE = 0.49) and B. tectorum dominance (PRE = 0.39) than at predicting establishment. Land managers will need to set priorities for control efforts on the more productive, species‐rich vegetation types that appear to be susceptible to both components of invasion.     

Effect of Argentine ant invasions on ground-dwelling arthropods in northern California riparian woodlands

Although the Argentine ant (Linepithema humile) is a widespread invasive species that displaces native ants throughout its introduced range, the effects of these invasions on arthropods other than ants remain poorly known. This study documents the consequences of Argentine ant invasions on ants and other ground-dwelling arthropods in northern California riparian woodlands. Baits and unbaited pitfall traps were used to sample different components of the arthropod communities at five pairs of uninvaded and invaded sites. Sites occupied by Argentine ants supported almost no native epigeic ants except for the winter-active Prenolepis imparis. Sites with Argentine ants averaged four to ten times more ant workers than did sites with native ants, but ant worker biomass did not differ between uninvaded and invaded sites. Argentine ants recruited to baits in invaded areas, on average, in less than half the time of native ants in uninvaded areas. Despite the loss of epigeic native ants, higher Argentine ant worker abundance, and faster recruitment by Argentine ants at invaded sites, pitfall trap samples from uninvaded and invaded areas contained similar abundances and diversities of non-ant arthropods. These findings suggest that Argentine ants and the native ants they displace interact with the ground-dwelling arthropods of these habitats in a similar manner. Received: 24 February 1997 / Accepted: 9 November 1997