Comparative ecological niche models predict the invasive spread of variable-leaf milfoil (<Emphasis Type="Italic">Myriophyllum</Emphasis><Emphasis Type="Italic">heterophyllum</Emphasis>) and its potential impact on closely related native species

Invasive species can alter patterns of biodiversity by displacing closely related native species that occupy similar habitats. We used multivariate ecological niche modeling to determine the potential spread and displacement of native congeners by the invasive aquatic plant, variable-leaf milfoil (VLM, Myriophyllum heterophyllum) in New Hampshire, USA. We show that VLM occurs almost exclusively in “higher order” lakes characterized as large, low elevation systems with relatively high pH, alkalinity and conductivity. In contrast, native milfoils occur across a broad range of lake orders. The strong association between lake order and VLM invasions suggests that VLM is most likely to displace native milfoils in higher order lakes. However, the mechanism by which VLM occurs in higher order lakes—higher propagule pressure versus higher growth and survivorship—is unclear. We therefore caution that native species may ultimately be susceptible to displacement from lower order lakes if the current distribution of VLM reflects higher propagule pressures. Our model provides a valuable tool for the prioritization of monitoring efforts.     

Predicting spread of invasive macrophytes in New Zealand lakes using indirect measures of human accessibility

1. Models predicting invasive macrophyte spread between lakes provide an important tool for focusing proactive management efforts to lakes deemed susceptible to invasion. However, challenges to forecasting macrophyte spread include wide physiological tolerances of invasive macrophytes and a lack of information on the relative importance of the various human vectors (e.g. boating traffic). In New Zealand, three invasive species that reproduce vegetatively, Ceratophyllum demersum, Lagarosiphon major, Egeria densa, and a single species that reproduces sexually, Utricularia gibba, are currently spreading across the lake landscape at a great cost to the local ecology and economy. 2. In this study, we first examined whether variables that indirectly describe weed spread via human access and use, as well as a lake’s position in the landscape, could describe the distribution of these four species using a boosted regression trees (BRT) modelling approach. Then, as these invasive species have not reached their full invasion potential, we examined how giving more influence to infected lakes at the edge of the invasion front, and including all lakes across New Zealand as background samples, simulating ‘absences beyond the invasion front’, influenced our ability to forecast the potential for new lakes to be invaded. 3. The BRT models identified that variables characterising human access and use, as well as lake position, were associated with the occurrence of the three vegetatively reproducing macrophytes. Weed occurrence was more likely when there was a highway in the vicinity, human population density was high and if the lake was large (c. 55 km²). But in the single case of U. gibba, temperature was the variable that best explained occurrence. This is consistent with the suggestion that U. gibba is predominantly dispersed by waterbirds, rather than human activity. 4. But for all four species, the BRT models based on the recorded observations alone predicted observed invasions with low prediction probabilities and did not forecast further spread. By contrast, when observations at the edge of the invasion front were upweighted, and additional background lakes implemented into the model, recorded observations were predicted and additional lakes were forecast to be at risk, suggesting that these models better captured the current and potential distribution of these macrophyte species. 5. The use of variables that characterise weed spread could provide similar insights into other systems where survey information on the nature, strength and direction of invasion vectors is lacking. Furthermore, when weighting the data, many lakes across New Zealand were forecasted to be at risk of invasion. The advantage of weighing the presence data was that insights into the potential for a species to spread were obtained. The probabilistic estimates of risk, as derived from the models, together with other information for prioritising lakes, can be used to focus surveillance and protection efforts.     

In lakes but not in minds: stakeholder knowledge of invasive species in prairie lakes

Humans are key vectors in the spread and establishment of aquatic invasive species (AIS), and human behavior can exacerbate or help prevent further spread of non-native species. Therefore, stakeholders’ knowledge is critical to preventing establishment of AIS. However, stakeholders’ AIS knowledge in prairie lakes remains poorly understood. We used a survey questionnaire in Saskatchewan, Canada, to assess the state of AIS knowledge, identify predictors of knowledge, and optimize management strategies. Statistical analyses of the responses of 440 participants indicated a generally low level of AIS knowledge, suggesting low communication success. Respondents were generally more aware of non-native fishes than plants. Of concern was the observation of substantial knowledge gaps regarding non-native mussels and important preventative behaviors that may have devastating ecological, social, and economic consequences if left unaddressed. Better understanding of AIS issues was significantly associated with several trans-situational (age, sex and education), situational (recreational purpose and using multiple lakes), and lake-related knowledge (awareness of eutrophication) predictors. Exploitation of these predictors is recommended to improve effectiveness of outreach and communication efforts. Specifically, we propose that management strategies focus on improving communications by streamlining outreach messages, targeting low-knowledge groups (e.g., swimmers, cabin owners), and expanding education campaigns.     

Integrating spread dynamics and economics of timber production to manage Chinese tallow invasions in southern U.S. forestlands

Economic costs associated with the invasion of nonnative species are of global concern. We estimated expected costs of Chinese tallow (Triadica sebifera (L.) Small) invasions related to timber production in southern U.S. forestlands under different management strategies. Expected costs were confined to the value of timber production losses plus costs for search and control. We simulated management strategies including (1) no control (NC), and control beginning as soon as the percentage of invaded forest land exceeded (2) 60 (Low Control), (3) 25 (Medium Control), or (4) 0 (High Control) using a spatially-explicit, stochastic, bioeconomic model. With NC, simulated invasions spread northward and westward into Arkansas and along the Gulf of Mexico to occupy ≈1.2 million hectares within 20 years, with associated expected total costs increasing exponentially to ≈$300 million. With LC, MC, and HC, invaded areas reached ≈275, 34, and 2 thousand hectares after 20 years, respectively, with associated expected costs reaching ≈$400, $230, and $200 million. Complete eradication would not be cost-effective; the minimum expected total cost was achieved when control began as soon as the percentage of invaded land exceeded 5%. These results suggest the importance of early detection and control of Chinese tallow, and emphasize the importance of integrating spread dynamics and economics to manage invasive species.     

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 Battle Lost? Report on Two Centuries of Invasion and Management of Lantana camara L. in Australia,India and South Africa

Recent discussion on invasive species has invigorated the debate on strategies to manage these species. Lantana camara L., a shrub native to the American tropics, has become one of the worst weeds in recorded history. In Australia, India and South Africa, Lantana has become very widespread occupying millions of hectares of land. Here, we examine historical records to reconstruct invasion and management of Lantana over two centuries and ask: Can we fight the spread of invasive species or do we need to develop strategies for their adaptive management? We carried out extensive research of historical records constituting over 75% of records on invasion and management of this species in the three countries. The records indicate that governments in Australia, India and South Africa have taken aggressive measures to eradicate Lantana over the last two centuries, but these efforts have been largely unsuccessful. We found that despite control measures, the invasion trajectory of Lantana has continued upwards and that post-war land-use change might have been a possible trigger for this spread. A large majority of studies on invasive species address timescales of less than one year; and even fewer address timescales of >10 years. An understanding of species invasions over long time-scales is of paramount importance. While archival records may give only a partial picture of the spread and management of invasive species, in the absence of any other long-term dataset on the ecology of Lantana, our study provides an important insight into its invasion, spread and management over two centuries and across three continents. While the established paradigm is to expend available resources on attempting to eradicate invasive species, our findings suggest that in the future, conservationists will need to develop strategies for their adaptive management rather than fighting a losing battle.     

Invasions by ladybugs,ladybirds, and other predatory beetles

Species of predatory Coleoptera have become abundant in new geographic regions recently, raising concerns for invaded ecosystems. We address this topic by focusing on invasive alien ladybird beetles (Coccinellidae; known also as ladybugs). Humans appear directly or indirectly responsible for all or most ladybird invasions. Factors hypothesized to have promoted ladybird invasions include genetic diversity (e.g., for polymorphism), phenotypic plasticity, adaptation and genetic shift, generalized diet and habitat preferences, flexible life history and reproduction, large body size, and release from enemies. Factors such as climate, habitat and prey availability, and biotic resistance may sometimes prevent or slow ladybird invasions. Indigenous species (e.g., herbivores) may suffer from invasions, and biological control programs may be affected. Species of indigenous ladybirds throughout the world are reported to have declined in abundance following ladybird invasions, with increased competition and/or intraguild predation most often hypothesized or inferred. Similar recent studies especially of ground beetles (Carabidae) also make clear the potential of invasive alien predatory Coleoptera to disrupt invaded natural and agricultural ecosystems.     

High phenolic content fails to deter mesograzer consumption of <Emphasis Type="Italic">Myriophyllum spicatum</Emphasis> (Eurasian watermilfoil) in New England

Eurasian watermilfoil (Myriophyllum spicatum) is often considered one of the most aggressive macrophyte invaders in freshwater habitats throughout the USA. However, conditions leading to successful milfoil invasions are not well understood. This study sought to illuminate the role of herbivores in determining milfoil invasion success via the potential mechanisms of enemy release and biotic resistance. We determined feeding preferences of three herbivores native to the northeastern United States and measured macrophyte phenolic content, which may act as an herbivore feeding deterrent. We found that phenolic content in milfoil was more than two times higher than in the most abundant native macrophytes at our study sites, consistent with enemy release. However, laboratory feeding experiments demonstrated that milfoil phenolics did not deter amphipod (Hyalella azteca), snail (Physella sp.), or weevil (Euhrychiopsis lecontei) herbivory. Furthermore, amphipod consumption rates in our study were an order of magnitude higher than amphipod consumption rates reported in milfoil’s native range, contrary to the predictions of enemy release. Amphipods and snails from habitats invaded by milfoil consumed similar quantities of both milfoil and the low-phenolic native plant Elodea canadensis. In contrast, weevils consumed milfoil but not E. canadensis in choice experiments. Amphipods collected from milfoil-free habitats also readily consumed milfoil, and they consumed 2.5 times more milfoil than E. canadensis in a choice feeding trial. These results suggest that high phenolic levels do not prevent native herbivores from consuming invasive milfoil. Instead, native generalist grazers like amphipods and snails may limit milfoil proliferation and provide a measure of biotic resistance.     

Bridgehead Effect in the Worldwide Invasion of the Biocontrol Harlequin Ladybird

Recent studies of the routes of worldwide introductions of alien organisms suggest that many widespread invasions could have stemmed not from the native range, but from a particularly successful invasive population, which serves as the source of colonists for remote new territories. We call here this phenomenon the invasive bridgehead effect. Evaluating the likelihood of such a scenario is heuristically challenging. We solved this problem by using approximate Bayesian computation methods to quantitatively compare complex invasion scenarios based on the analysis of population genetics (microsatellite variation) and historical (first observation dates) data. We applied this approach to the Harlequin ladybird Harmonia axyridis (HA), a coccinellid native to Asia that was repeatedly introduced as a biocontrol agent without becoming established for decades. We show that the recent burst of worldwide invasions of HA followed a bridgehead scenario, in which an invasive population in eastern North America acted as the source of the colonists that invaded the European, South American and African continents, with some admixture with a biocontrol strain in Europe. This demonstration of a mechanism of invasion via a bridgehead has important implications both for invasion theory (i.e., a single evolutionary shift in the bridgehead population versus multiple changes in case of introduced populations becoming invasive independently) and for ongoing efforts to manage invasions by alien organisms (i.e., heightened vigilance against invasive bridgeheads).     

A genetic reconstruction of the invasion of the calanoid copepod <Emphasis Type="Italic">Pseudodiaptomus inopinus</Emphasis> across the North American Pacific Coast

The rate of aquatic invasions by planktonic organisms has increased considerably in recent decades. In order to effectively direct funding and resources to control the spread of such invasions, a methodological framework for identifying high-risk transport vectors, as well as ruling out vectors of lesser concern will be necessary. A number of estuarine ecosystems on the North American Pacific Northwest coast have experienced a series of high impact planktonic invasions that have slowly unfolded across the region in recent decades, most notably, that of the planktonic copepod crustacean Pseudodiaptomus inopinus. Although introduction of P. inopinus to the United States almost certainly occurred through the discharge of ballast water from commercial vessels originating in Asia (the species’ native range), the mechanisms and patterns of subsequent spread remain unknown. In order to elucidate the migration events shaping this invasion, we sampled the genomes of copepods from seven invasive and two native populations using restriction-site associated DNA sequencing. This genetic data was evaluated against spatially-explicit genetic simulation models to evaluate competing scenarios of invasion spread. Our results indicate that invasive populations of P. inopinus exhibit a geographically unstructured genetic composition, likely arising from infrequent and large migration events. This pattern of genetic patchiness was unexpected given the linear geographic structure of the sampled populations, and strongly contrasts with the clear invasion corridors observed in many aquatic systems.     

Offense and defense in landscape-level invasion control

Biological invasions are multi-stage processes comprising chance demographic events, species interactions, and dispersal. Despite this complexity, simple models can increase understanding of the invasion process. We model the spread of aquatic invasive species through a network of lakes to evaluate the effectiveness of two intervention strategies. The first, which we call offense, contains the invader at sources; the second, which we call defense, protects uninvaded destinations. Deterministic models reveal the effects of these intervention strategies on spread rates. Practical applications involve finite collections of uninvaded lakes, however, and we therefore also present a stochastic model to describe how these strategies affect expected times to important invasion milestones. When the goal is to reduce overall spread rates, both approaches agree that offense is better early in invasions, but that defense is better after 1/2 the lakes are invaded. When the goal is to protect areas of high conservation value, however, defensive site protection always provides lower per site introduction rates. Although we focus on lakes, our results are quite general, and could be applied to any discrete habitat patches including, for example, fragmented terrestrial habitats.     

An invasive fish promotes invasive plants in Minnesota lakes

1. Biological invasions can greatly alter ecological communities, affecting not only the diversity and abundance but also composition of invaded assemblages. This is because invaders’ impacts are mediated by characteristics of resident species: some may be highly sensitive to invader impacts while others are unaffected or even facilitated. In some cases, this can result in invasive species promoting further invasions; in particular, herbivory by introduced animals has been shown to disproportionately harm native plants, which can indirectly benefit non-native plants. Here, we investigated whether such patterns emerged through the effects of an invasive fish species on lake plant communities.
2. Specifically, we tested whether invasion of Minnesota (U.S.A.) lakes by Cyprinus carpio (common carp), an omnivorous, benthivorous fish known to reduce abundance and richness of aquatic plants, differentially affected native versus non-native plant species. We applied statistical models to a large, long-term monitoring dataset (206 macrophyte taxa recorded in 913 lakes over a 20-year time period) to test whether carp altered community composition, to identify which macrophyte species were most sensitive to carp and determine whether species characteristics predicted carp sensitivity, and to characterise consequences of carp invasion on lake-level vegetation attributes.
3. We found that carp exerted strong selective pressure on community composition. Native macrophytes, those with a more aquatic growth form, and those considered less tolerant of disturbance (i.e. higher coefficients of conservatism) were more sensitive to carp. Conversely, no introduced macrophytes exhibited sensitivity to carp and all had higher probabilities of occurrence as carp abundance increased. The net effect of carp invasion was a shift toward less species-rich plant communities characterised by more non-native and disturbance-tolerant species.
4. These results have several implications for conservation and management. First, they reinforce the need to prevent further spread of carp outside of their native range. Where carp have already established, their control should be incorporated into efforts to restore aquatic vegetation; this may be an essential step for recovering particular plant species of high conservation importance. Furthermore, reducing carp abundance could have ancillary benefits of reducing dominance by invasive plant species. Lastly, where carp cannot be eliminated, managers should target native macrophytes that are relatively tolerant of carp in shoreline plantings and other revegetation efforts.

     

Is Hybridization Responsible for Invasive Growth of Non-indigenous Water-milfoils?

Heterosis, or hybrid vigor, has recently been proposed as a factor promoting invasive growth of some non-indigenous aquatic plant species, particularly those capable of spreading rapidly within and among lakes through clonal reproduction. We tested this hypothesis for variable-leaf water milfoil (Myriophyllum heterophyllum), a non-indigenous aquatic plant that has become a major management and conservation concern in New England. Using nuclear ribosomal DNA, we looked for F₁ hybrid populations of invasive M. heterophyllum in 25 New Hampshire (NH) lakes. In contrast to a previous study that found F₁ hybrid lineages of invasive M. heterophyllum in Connecticut, we did not find hybrids in our study lakes. This result has two implications: (1) pure lineages of M. heterophyllum are also capable of invasive growth, and (2) the distribution of invasive M. heterophyllum lineages (hybrid vs. pure) may be spatially structured across New England. We stress the importance of more detailed distributional and ecological studies for understanding the invasive potential of this species.     

Exploring lake user and manager knowledge of aquatic invasive species in New Hampshire freshwater lake systems,USA

Aquatic invasive species (AIS) threaten freshwater ecosystem structure and function worldwide. Such changes trigger a variety of negative impacts on lake recreation and the economics of individuals, towns, and states. Recent studies suggest environmental educational efforts have been an effective tool in raising public awareness of AIS; however, we lack a more general understanding of how public versus manager knowledge of invasive species and their threats to freshwater ecosystems and human livelihoods. To fill this gap, we surveyed New Hampshire lake users and interviewed lake managers to 1) identify the key issues surrounding AIS management; and 2) assess public awareness of the AIS problem and their management at three lakes in New Hampshire. Our interviews with managers suggest that educational outreach is a key mechanism for combatting the AIS problem. The general public surveys further differentiated respondents into two groups that differed demography and in AIS knowledge and concern. Together, our results indicate that AIS management efforts depend heavily on funding, regional cooperation, and commitment of individual managers and lake-users. Blanket outreach campaigns have been effective in expanding the knowledge reach of AIS, but they remain too general to engage the public in AIS spread prevention and eradication. Instead, integrating practices of rapid response and appeals to responsibility norms among lake users is paramount for combatting the AIS problem in this region.     

An ounce of prevention or a pound of cure: bioeconomic risk analysis of invasive species

Numbers of non-indigenous species--species introduced from elsewhere - are increasing rapidly worldwide, causing both environmental and economic damage. Rigorous quantitative risk-analysis frameworks, however, for invasive species are lacking. We need to evaluate the risks posed by invasive species and quantify the relative merits of different management strategies (e.g. allocation of resources between prevention and control). We present a quantitative bioeconomic modelling framework to analyse risks from non-indigenous species to economic activity and the environment. The model identifies the optimal allocation of resources to prevention versus control, acceptable invasion risks and consequences of invasion to optimal investments (e.g. labour and capital). We apply the model to zebra mussels (Dreissena polymorpha), and show that society could benefit by spending up to US$324 000 year(-1) to prevent invasions into a single lake with a power plant. By contrast, the US Fish and Wildlife Service spent US$825 000 in 2001 to manage all aquatic invaders in all US lakes. Thus, greater investment in prevention is warranted.     

Biological invasions in rapidly urbanizing areas: a case study of Beijing, China

Urbanization is widely recognized as a major factor promoting biological invasions worldwide. In this article, we provide insights into the patterns of biological invasions in Beijing, one of the largest and quickly urbanizing cities of the world, by developing a comprehensive list of naturalized and invasive flora and their associate traits (e.g., distribution, life form, habitat, or geographic origin). One hundred and twelve naturalized (including 48 invasive) plants have been identified within the Beijing Municipality. Most of the naturalized and invasive plants belong to four families (Asteraceae, Poaceae, Amaranthaceae, and Euphorbiaceae) and are annual herbs that preferentially grow in disturbed sites. North and South America are the main contributors to the naturalized and invasive flora of Beijing. As expected, those Beijing districts that have recently experienced the highest human population growth, urban expansion, and the largest economic growth are also those with the highest number of naturalized and invasive species. Urban expansion is predicted to continue in the near term making additional invasions likely that will significantly increase the proportion of introduced species in Beijing’s flora. An integrated management strategy for the whole municipality is urgently needed that includes comprehensive scientific research that documents the extent of invasions and their effects on Beijing’s economy and environment.     

Multiplexed microsatellite recovery using massively parallel sequencing

Conservation and management of natural populations requires accurate and inexpensive genotyping methods. Traditional microsatellite, or simple sequence repeat (SSR), marker analysis remains a popular genotyping method because of the comparatively low cost of marker development, ease of analysis and high power of genotype discrimination. With the availability of massively parallel sequencing (MPS), it is now possible to sequence microsatellite-enriched genomic libraries in multiplex pools. To test this approach, we prepared seven microsatellite-enriched, barcoded genomic libraries from diverse taxa (two conifer trees, five birds) and sequenced these on one lane of the Illumina Genome Analyzer using paired-end 80-bp reads. In this experiment, we screened 6.1 million sequences and identified 356,958 unique microreads that contained di- or trinucleotide microsatellites. Examination of four species shows that our conversion rate from raw sequences to polymorphic markers compares favourably to Sanger- and 454-based methods. The advantage of multiplexed MPS is that the staggering capacity of modern microread sequencing is spread across many libraries; this reduces sample preparation and sequencing costs to less than $400 (USD) per species. This price is sufficiently low that microsatellite libraries could be prepared and sequenced for all 1373 organisms listed as 'threatened' and 'endangered' in the United States for under $0.5 M (USD).     

Habitat, predation, and coexistence between invasive and native crayfishes: prioritizing lakes for invasion prevention

Conditions fostering coexistence of native species with invasive species have received little attention in invasion biology, especially for closely related invasive and native species. We used long-term datasets on multiple replicate invasions to define conditions under which native virile crayfish (Orconectes virilis) can coexist with invasive rusty crayfish (O. rusticus). We examined multiple drivers of coexistence involving habitat use and predation at between-lake and within-lake scales to derive predictions that could guide prioritization efforts to prevent future introductions of rusty crayfish and mitigate impacts of existing invasions. Lakes in which native species persisted for many years had significantly less cobble and sand habitats, and significantly more vegetated habitats compared to lakes from which native crayfish have been displaced. In the presence of rusty crayfish, virile crayfish alter their habitat use to vegetated habitats relative to habitat use in the absence of rusty crayfish. Such vegetated habitats had greater plant standing crop, plant species richness, and sediment percent organic matter compared to vegetated sites occupied by rusty crayfish. Our results suggest that low abundance of cobble habitat and altered habitat use allows native crayfish to coexist with the rusty crayfish invader. At the within-lake scale, virile crayfish persist by escaping predation in the vegetated habitats, despite suboptimal abiotic conditions. By understanding these abiotic and biotic conditions that promote coexistence, managers could enhance native crayfish persistence by targeting high cobble lakes for efforts to prevent the introduction of invasive crayfish, and targeting vegetated habitats for protection in already invaded lakes.     

Physical control of Eurasian watermilfoil in an oligotrophic lake

The introduction of Eurasian watermilfoil (Myriophyllum spicatum) into oligotrophic waters of high water clarity in temperate zones of North America has produced growth in excess of 6 m depth and yearly biomass approaching 1000 g m^–2 dry weight. From its initial observation in Lake George, New York, USA in 1985, by 1993 milfoil had spread to 106 discrete locations within the lake. A 7-year study of one site having no management showed milfoil to grow expansively, suppressing native plant species from 20 in 1987 to 6 in 1993 with the average number of species m^–2 quadrat declining from 5.5 in 1987 to less than 2 in 1993. Management of milfoil by means of hand harvesting, suction harvesting and benthic barrier has reduced the number of unmanaged sites from 106 in 1993 to 11. One year post-treatment at sites utilizing suction harvesting, showed a greater number of native species at all sites than pretreatment with a substantial reduction in milfoil biomass. At sites where benthic barrier was removed 1–2 years after installation, milfoil had recolonized 44% of grid squares within 30 days. Ninety days after barrier removal 74% of grid squares contained milfoil and one year later 71% of the grids supported milfoil. During the first year following mat removal, the average number of species m^–2 peaked at 4.7 and stabilized at 4.5 during the second year. Hand harvesting by SCUBA in areas of limited milfoil growth (new sites of infestation and sites of former treatment) was found to reduce the number of milfoil plants present in subsequent years. Hand harvesting did not eliminate milfoil at any of the sites and regrowth/colonization necessitated reharvesting every 3 or more years. Results of evaluations of physical plant management techniques indicate that (1) an integrated program utilizing different techniques based on plant density reduced the growth of milfoil and (2) long term commitment to aquatic plant management is necessary since none of the techniques employed singly were found to eliminate milfoil.