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.     

Boom‐bust dynamics in biological invasions: towards an improved application of the concept

Boom‐bust dynamics – the rise of a population to outbreak levels, followed by a dramatic decline – have been associated with biological invasions and offered as a reason not to manage troublesome invaders. However, boom‐bust dynamics rarely have been critically defined, analyzed, or interpreted. Here, we define boom‐bust dynamics and provide specific suggestions for improving the application of the boom‐bust concept. Boom‐bust dynamics can arise from many causes, some closely associated with invasions, but others occurring across a wide range of ecological settings, especially when environmental conditions are changing rapidly. As a result, it is difficult to infer cause or predict future trajectories merely by observing the dynamic. We use tests with simulated data to show that a common metric for detecting and describing boom‐bust dynamics, decline from an observed peak to a subsequent trough, tends to severely overestimate the frequency and severity of busts, and should be used cautiously if at all. We review and test other metrics that are better suited to describe boom‐bust dynamics. Understanding the frequency and importance of boom‐bust dynamics requires empirical studies of large, representative, long‐term data sets that use clear definitions of boom‐bust, appropriate analytical methods, and careful interpretations.     

Potential changes to the biology and challenges to the management of invasive sea lamprey Petromyzon marinus in the Laurentian Great Lakes due to climate change

Control programs are implemented to mitigate the damage caused by invasive species worldwide. In the highly invaded Great Lakes, the climate is expected to become warmer with more extreme weather and variable precipitation, resulting in shorter iced‐over periods and variable tributary flows as well as changes to pH and river hydrology and hydrogeomorphology. We review how climate change influences physiology, behavior, and demography of a damaging invasive species, sea lamprey (Petromyzon marinus), in the Great Lakes, and the consequences for sea lamprey control efforts. Sea lamprey control relies on surveys to monitor abundance of larval sea lamprey in Great Lakes tributaries. The abundance of parasitic, juvenile sea lampreys in the lakes is calculated by surveying wounding rates on lake trout (Salvelinus namaycush), and trap surveys are used to enumerate adult spawning runs. Chemical control using lampricides (i.e., lamprey pesticides) to target larval sea lamprey and barriers to prevent adult lamprey from reaching spawning grounds are the most important tools used for sea lamprey population control. We describe how climate change could affect larval survival in rivers, growth and maturation in lakes, phenology and the spawning migration as adults return to rivers, and the overall abundance and distribution of sea lamprey in the Great Lakes. Our review suggests that Great Lakes sea lamprey may benefit from climate change with longer growing seasons, more rapid growth, and greater access to spawning habitat, but uncertainties remain about the future availability and suitability of larval habitats. Consideration of the biology of invasive species and adaptation of the timing, intensity, and frequency of control efforts is critical to the management of biological invasions in a changing world, such as sea lamprey in the Great Lakes.     

A multi‐basin approach determines variability in host fish suitability for unionids in tributaries of the Laurentian Great Lakes

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The role of long‐distance seed dispersal in the local population dynamics of an invasive plant species

Aim Long‐distance dispersal is important for plant population dynamics at larger spatial scales, but our understanding of this phenomenon is mostly based on computer modelling rather than field data. This paper, by combining field data and a simulation model, quantifies the fraction of the seed of the alien species Heracleum mantegazzianum that needs to disperse over a long distance for successful invasion. Location Central Europe, Czech Republic. Methods To assess the role of random dispersal in long‐term population dynamics of the studied species, we combined longitudinal data covering 50 years of the invasion of this plant from its very start, inferred from a series of aerial photographs of 60‐ha plots, with data on population dynamics at a fine scale of 10‐m² plots. Results A simulation model based on field data indicates that the fraction of seed that is dispersed from source plants not described by the short‐distance dispersal kernel ranges from 0.1 to 7.5% of the total seed set. The fraction of long‐distance dispersed seed that provides the best prediction of the observed spread was significantly negatively correlated with the percentage of habitats suitable for invasion. Main conclusions Our results indicate that the fraction of seeds that needed to be dispersed over long distances to account for the observed invasion dynamics decreased with increasing proportion of invasible habitats, indicating that the spatial pattern of propagule pressure differs in landscapes prone to invasion. Long‐distance dispersal is an important component of the population dynamics of an invasive species even at relatively small scales.     

Evaluating the effects of pollinator‐mediated interactions using pollen transfer networks: evidence of widespread facilitation in south Andean plant communities

Information about the relative importance of competitive or facilitative pollinator‐mediated interactions in a multi‐species context is limited. We studied interspecific pollen transfer (IPT) networks to evaluate quantity and quality effects of pollinator sharing among plant species on three high‐Andean communities at 1600, 1800 and 2000 m a.s.l. To estimate the sign of the effects (positive, neutral or negative), the relation between conspecific and heterospecific pollen deposited on stigmas was analysed with GLMMs. Network analyses showed that communities were characterised by the presence of pollen hub‐donors and receptors. We inferred that facilitative and neutral pollinator‐mediated interactions among plants prevailed over competition. Thus, the benefits from pollinator sharing seem to outweigh the costs (i.e. heterospecific deposition and conspecific pollen loss). The largest proportion of facilitated species was found at the highest elevation community, suggesting that under unfavourable conditions for the pollination service and at lower plant densities facilitation can be more common.     

Tracking biological invasions in space and time: elucidating the invasive history of the green alga Codium fragile using old DNA

With the advent of 'ancient DNA' studies on preserved material of extant and extinct species, museums and herbaria now represent an important although still underutilized resource in molecular ecology. The ability to obtain sequence data from archived specimens can reveal the recent history of cryptic species and introductions. We have analysed extant and herbarium samples of the highly invasive green alga Codium fragile , many over 100 years old, to identify cryptic accessions of the invasive strain known as C. fragile ssp. tomentosoides , which can be identified by a unique haplotype. Molecular characterization of specimens previously identified as native in various regions shows that the invasive tomentosoides strain has been colonizing new habitats across the world for longer than records indicate, in some cases nearly 100 years before it was noticed. It can now be found in the ranges of all the other native haplotypes detected, several of which correspond to recognized subspecies. Within regions in the southern hemisphere there was a greater diversity of haplotypes than in the northern hemisphere, probably as a result of dispersal by the Antarctic Circumpolar Current. The findings of this study highlight the importance of herbaria in preserving contemporaneous records of invasions as they occur, especially when invasive taxa are cryptic.     

Pollen transfer in fragmented plant populations: insight from the pollen loads of pollinators and stigmas in a mass‐flowering species

Pollinator and/or mate scarcity affects pollen transfer, with important ecological and evolutionary consequences for plant reproduction. However, the way in which the pollen loads transported by pollinators and deposited on stigmas are affected by pollination context has been little studied. We investigated the impacts of plant mate and visiting insect availabilities on pollen transport and receipt in a mass‐flowering and facultative autogamous shrub (Rhododendron ferrugineum). First, we recorded insect visits to R. ferrugineum in plant patches of diverse densities and sizes. Second, we analyzed the pollen loads transported by R. ferrugineum pollinators and deposited on stigmas of emasculated and intact flowers, in the same patches. Overall, pollinators (bumblebees) transported much larger pollen loads than the ones found on stigmas, and the pollen deposited on stigmas included a high proportion of conspecific pollen. However, comparing pollen loads of emasculated and intact flowers indicated that pollinators contributed only half the conspecific pollen present on the stigma. At low plant density, we found the highest visitation rate and the lowest proportion of conspecific pollen transported and deposited by pollinators. By contrast, at higher plant density and lower visitation rate, pollinators deposited larger proportion of conspecific pollen, although still far from sufficient to ensure that all the ovules were fertilized. Finally, self‐pollen completely buffered the detrimental effects on pollination of patch fragmentation and pollinator failure. Our results indicate that pollen loads from pollinators and emasculated flowers should be quantified for an accurate understanding of the relative impacts of pollinator and mate limitation on pollen transfer in facultative autogamous species.     

Using a climatic niche model to predict the direct and indirect impacts of climate change on the distribution of Douglas‐fir in New Zealand

Climate change is likely to have major impacts on the distribution of planted and natural forests. Herein, we demonstrate how a process‐based niche model (CLIMEX) can be extended to globally project the potential habitat suitable for Douglas‐fir. Within this distribution, we use CLIMEX to predict abundance of the pathogen P haeocryptopus gaeumannii and severity of its associated foliage disease, Swiss needle cast. The distribution and severity of the disease, which can strongly reduce growth rate of Douglas‐fir, is closely correlated with seasonal temperatures and precipitation. This model is used to project how climate change during the 2080s may alter the area suitable for Douglas‐fir plantations within New Zealand. The climate change scenarios used indicate that the land area suitable for Douglas‐fir production in the North Island will be reduced markedly from near 100% under current climate to 36–64% of the total land area by 2080s. Within areas shown to be suitable for the host in the North Island, four of the six climate change scenarios predict substantial increases in disease severity that will make these regions at best marginal for Douglas‐fir by the 2080s. In contrast, most regions in the South Island are projected to sustain relatively low levels of disease, and remain suitable for Douglas‐fir under climate change over the course of this century.     

Invasive plants in Minnesota are “joining the locals”: A trait‐based analysis

Questions

Predicting which newly arrived species will establish and become invasive is a problem that has long vexed researchers. In a study of cold temperate oak forest stands, we examined two contrasting hypotheses regarding plant functional traits to explain the success of certain non‐native species. Under the “join the locals” hypothesis, successful invaders are expected to share traits with resident species because they employ successful growth strategies under light‐limited understorey conditions. Instead, under the “try harder” hypothesis, successful invaders are expected to have traits different from native species in order to take advantage of unused niche space.

Location

Minnesota, USA.

Methods

We examined these two theories using 109 native and 11 non‐native plants in 68 oak forest stands. We focused on traits related to plant establishment and growth, including specific leaf area (SLA), leaf carbon‐to‐nitrogen ratio (C:N), wood density, plant maximum height, mycorrhizal type, seed mass and growth form. We compared traits of native and non‐native species using ordinations in multidimensional trait space and compared community‐weighted mean (CWM) trait values across sites.

Results

We found few differences between trait spaces occupied by native and non‐native species. Non‐native species occupied smaller areas of trait space than natives, yet were within that of the native species, indicating similar growth strategies. We observed a higher proportion of non‐native species in sites with higher native woody species CWM SLA and lower CWM C:N. Higher woody CWM SLA was observed in sites with higher soil pH, while lower CWM C:N was found in sites with higher light levels.

Conclusions

Non‐native plants in this system have functional traits similar to natives and are therefore “joining the locals.” However, non‐native plants may possess traits toward the acquisitive end of the native plant trait range, as evidenced by higher non‐native plant abundance in high‐resource environments.

     

Predicting socio‐economic and biodiversity impacts of invasive species: Red Imported Fire Ant in the developing western Pacific

Effective biosecurity and pest management are fundamental to sustainable development. Invasive ants pose significant risks to the environment and economy, which are well‐managed by biosecurity agencies in developed countries. However, a comprehensive view of the potential impacts of invasive ants in developing Pacific Island Countries and Territories (PICTs) is lacking. We quantified the potential costs of the Red Imported Fire Ant (Solenopsis invicta Buren) across multiple sectors using an extrapolation analysis. Overall, we estimated that the impacts of Red Imported Fire Ant on developing PICTs could amount to over USD 329 million annually, corresponding to approximately 0.7% of combined GDP. Over half of the costs were predicted to result from impacts on the agriculture sector, a major source of employment and subsistence. We found that over 350 highly threatened species could be at risk from Red Imported Fire Ant. We would expect countries with Least Developed Country status and relatively low GDP to be least able to respond to an incursion of these ants, and as a result the costs could be higher than we have extrapolated. Red Imported Fire Ant could therefore potentially have considerable impact on the on‐going development of the region.     

Living in two worlds: Evolutionary mechanisms act differently in the native and introduced ranges of an invasive plant

Identifying the factors that influence spatial genetic structure among populations can provide insights into the evolution of invasive plants. In this study, we used the common reed (Phragmites australis), a grass native in Europe and invading North America, to examine the relative importance of geographic, environmental (represented by climate here), and human effects on population genetic structure and its changes during invasion. We collected samples of P. australis from both the invaded North American and native European ranges and used molecular markers to investigate the population genetic structure within and between ranges. We used path analysis to identify the contributions of each of the three factors—geographic, environmental, and human‐related—to the formation of spatial genetic patterns. Genetic differentiation was observed between the introduced and native populations, and their genetic structure in the native and introduced ranges was different. There were strong effects of geography and environment on the genetic structure of populations in the native range, but the human‐related factors manifested through colonization of anthropogenic habitats in the introduced range counteracted the effects of environment. The between‐range genetic differences among populations were mainly explained by the heterogeneous environment between the ranges, with the coefficient 2.6 times higher for the environment than that explained by the geographic distance. Human activities were the primary contributor to the genetic structure of the introduced populations. The significant environmental divergence between ranges and the strong contribution of human activities to the genetic structure in the introduced range suggest that invasive populations of P. australis have evolved to adapt to a different climate and to human‐made habitats in North America.     

Seed set variation in wild Clarkia populations: teasing apart the effects of seasonal resource depletion,pollen quality,and pollen quantity

In habitats where resource availability declines during the growing season, selection may favor early‐flowering individuals. Under such ephemerally favorable conditions, late‐blooming species (and individuals) may be particularly vulnerable to resource limitation of seed production. In California, a region prone to seasonal drought, members of the annual genus Clarkia are among the last to flower in the spring. We compared pollen limitation (PL) of seed set and outcrossing rates between early‐ and late‐flowering individuals in two mixed‐mating Clarkia taxa to detect whether flowering time is associated with changes in seed set due to resource depletion, PL, or increased selfing. In 2008–2010, we hand‐pollinated one flower on a total of 1855 individual plants either Early (near the onset of flowering) or Late (near the end of flowering) in the flowering season and compared seed set to adjacent, open‐pollinated flowers on the same stem. To assess the contribution of pollen quality to reproduction, we first (2008) used allozymes to estimate outcrossing rates of seeds produced by Early and Late open‐pollinated flowers. Second (2009), we conducted an anther‐removal experiment to estimate self‐pollen deposition. Seed set in Clarkia unguiculata was not pollen‐limited. Clarkia xantiana ssp. xantiana was pollen‐limited in 2008 and 2010, but not 2009. PL did not differ between Early and Late treatments. In both taxa, seed set of Early flowers was greater than Late flowers, but not due to PL in the latter. Reproduction was generally pollinator‐dependent. Most pollen deposition was xenogamous, and outcrossing rates were >0.7 – and similar between Early and Late periods. These results suggest that pollen receipt and pollen quality remain seasonally consistent. By contrast, the resources necessary to provision seeds decline, reducing the fitness benefits associated with resource allocation to ovules.     

Simulating the effects of climate change on the distribution of an invasive plant,using a high resolution,local scale,mechanistic approach: challenges and insights

The growing economic and ecological damage associated with biological invasions, which will likely be exacerbated by climate change, necessitates improved projections of invasive spread. Generally, potential changes in species distribution are investigated using climate envelope models; however, the reliability of such models has been questioned and they are not suitable for use at local scales. At this scale, mechanistic models are more appropriate. This paper discusses some key requirements for mechanistic models and utilises a newly developed model (PSS[gt]) that incorporates the influence of habitat type and related features (e.g., roads and rivers), as well as demographic processes and propagule dispersal dynamics, to model climate induced changes in the distribution of an invasive plant (Gunnera tinctoria) at a local scale. A new methodology is introduced, dynamic baseline benchmarking, which distinguishes climate‐induced alterations in species distributions from other potential drivers of change. Using this approach, it was concluded that climate change, based on IPCC and C4i projections, has the potential to increase the spread‐rate and intensity of G. tinctoria invasions. Increases in the number of individuals were primarily due to intensification of invasion in areas already invaded or in areas projected to be invaded in the dynamic baseline scenario. Temperature had the largest influence on changes in plant distributions. Water availability also had a large influence and introduced the most uncertainty in the projections. Additionally, due to the difficulties of parameterising models such as this, the process has been streamlined by utilising methods for estimating unknown variables and selecting only essential parameters.