Establishment Of Non-native Species Research Articles

Restoring plant species diversity and community composition in a ponderosa pine-bunchgrass ecosystem

Monitoring of ecological restoration treatments often focuses on changes in community structure and function. We suggest that long-term changes in community composition also need to be explicitly considered when evaluating the success of restoration treatments. In 1992, we initiated an experiment in a ponderosa pine-bunchgrass ecosystem to evaluate responses to restoration treatments: (a) thinning the overstory vegetation (‘thinning’), (b) thinning plus forest floor manipulation with periodic prescribed burning (‘composite’), and (c) untreated ‘control.’ Treatments were further stratified by forest patch type: presettlement tree clumps (trees that established prior to the onset of fire exclusion in 1876), patches of retained postsettlement trees, patches where all postsettlement trees were removed, and remnant grass openings. Species richness did not differ among treatments for 10 years, but was highest in the composite treatment in 11th and 12th year after initial treatment. Community composition diverged among treatments 5 years after initial treatment, and compositional changes were greatest in the composite treatment. Species richness and composition differed among patch types prior to treatment. Remnant grass patches were the most diverse and presettlement patches were the least diverse. Following treatment, species richness in the postsettlement removed and retained patches, gradually approached levels found in remnant grass patches. Compositional differences among patch types changed a little by 2005. Species richness at the 2 m2 scale increased only where the overstory was thinned and the understory was burned. However, these changes may not be detectable for many years, and can vary temporally in response to events such as severe droughts. Nonnative species establishment may be reduced by scheduling longer burn intervals or by refraining from burning where fuel loads are not hazardous, though these options may hinder goals of increasing diversity. Restoring species diversity and community composition continues to be more difficult than restoring ecosystem structure and function.

Landscape as a Model: The Importance of Geometry

In all models, but especially in those used to predict uncertain processes (e.g., climate change and nonnative species establishment), it is important to identify and remove any sources of bias that may confound results. This is critical in models designed to help support decisionmaking. The geometry used to represent virtual landscapes in spatially explicit models is a potential source of bias. The majority of spatial models use regular square geometry, although regular hexagonal landscapes have also been used. However, there are other ways in which space can be represented in spatially explicit models. For the first time, we explicitly compare the range of alternative geometries available to the modeller, and present a mechanism by which uncertainty in the representation of landscapes can be incorporated. We test how geometry can affect cell-to-cell movement across homogeneous virtual landscapes and compare regular geometries with a suite of irregular mosaics. We show that regular geometries have the potential to systematically bias the direction and distance of movement, whereas even individual instances of landscapes with irregular geometry do not. We also examine how geometry can affect the gross representation of real-world landscapes, and again show that individual instances of regular geometries will always create qualitative and quantitative errors. These can be reduced by the use of multiple randomized instances, though this still creates scale-dependent biases. In contrast, virtual landscapes formed using irregular geometries can represent complex real-world landscapes without error. We found that the potential for bias caused by regular geometries can be effectively eliminated by subdividing virtual landscapes using irregular geometry. The use of irregular geometry appears to offer spatial modellers other potential advantages, which are as yet underdeveloped. We recommend their use in all spatially explicit models, but especially for predictive models that are used in decisionmaking.

RAPID ASSESSMENT OF POSTFIRE PLANT INVASIONS IN CONIFEROUS FORESTS OF THE WESTERN UNITED STATES

Fire is a natural part of most forest ecosystems in the western United States, but its effects on nonnative plant invasion have only recently been studied. Also, forest managers are engaging in fuel reduction projects to lessen fire severity, often without considering potential negative ecological consequences such as nonnative plant species introductions. Increased availability of light, nutrients, and bare ground have all been associated with high-severity fires and fuel treatments and are known to aid in the establishment of nonnative plant species. We use vegetation and environmental data collected after wildfires at seven sites in coniferous forests in the western United States to study responses of nonnative plants to wildfire. We compared burned vs. unburned plots and plots treated with mechanical thinning and/or prescribed burning vs. untreated plots for nonnative plant species richness and cover and used correlation analyses to infer the effect of abiotic site conditions on invasibility. Wildfire was responsible for significant increases in nonnative species richness and cover, and a significant decrease in native cover. Mechanical thinning and prescribed fire fuel treatments were associated with significant changes in plant species composition at some sites. Treatment effects across sites were minimal and inconclusive due to significant site and site x treatment interaction effects caused by variation between sites including differences in treatment and fire severities and initial conditions (e.g., nonnative species sources). We used canonical correspondence analysis (CCA) to determine what combinations of environmental variables best explained patterns of nonnative plant species richness and cover. Variables related to fire severity, soil nutrients, and elevation explained most of the variation in species composition. Nonnative species were generally associated with sites with higher fire severity, elevation, percentage of bare ground, and lower soil nutrient levels and lower canopy cover. Early assessments of postfire stand conditions can guide rapid responses to nonnative plant invasions.

ZOOGEOGRAPHIC PATTERNS AND FAUNAL CHANGE OF SOUTH DAKOTA FISHES

We summarized historic and recent fish distributions in South Dakota and analyzed fish faunal similarity at 2 spatial scales (geomorphic province and river drainage) for both historic (native) and recent (post-1990) faunas. We quantified zoogeographic patterns between geomorphic provinces and among neighboring river drainages for historic and recent faunas. We also quantified faunal change (species losses and additions) between provinces and among drainages. Ninety-seven fishes were native to South Dakota, but 111 fishes were present in recent collections because 8 native species were missing, and 22 nonnatives were present. There was high β diversity among historic and recent river drainage fish faunas, but there was between 22% and 56% faunal change between periods. Recent faunas were homogenized compared to historic faunas at both provincial and river drainage spatial scales. Patterns of nonnative species establishment were geographically distinct from patterns of native species loss. Most nonnative species additions were in cold-water or human-made habitats of the Great Plains. Most native species declines stemmed from warm water streams of the Central Lowlands and Missouri River valley. Conservation of rare and declining native species and containment of nonnatives are both necessary to preserve historical patterns of fish biodiversity in South Dakota.

Reduced soil compaction enhances establishment of non-native plant species

Many studies have shown that soil disturbance facilitates establishment of invasive, non-native plant species, and a number of mechanisms have been isolated that contribute to the process. To our knowledge no studies have isolated the role of altered soil compaction, a likely correlate of many types of soil disturbance, in facilitating invasion. To address this, we measured the response of seeded non-native and native plant species to four levels of soil compaction in mesocosms placed in an abandoned agricultural field in the Methow Valley, Washington, USA. Soil compaction levels reflected the range of resistance to penetration (0.1–3.0 kg cm−2) measured on disturbed soils throughout the study system prior to the experiment. Percent cover of non-native species, namely Bromus tectorum and Centaurea diffusa, decreased by 34% from the least to the most compacted treatments, whereas percent cover of native species, mostly Pseudoroegneria spicata and Lupinus spp., did not respond to compaction treatments. Experimental results were supported by a survey of soil penetration resistance and percent cover by species in 18 abandoned agricultural fields. Percent cover of B. tectorum was negatively related to soil compaction levels, whereas none of the native species showed any response to soil compaction. These results highlight a potentially important, though overlooked, aspect of soil disturbance that may contribute to subsequent non-native plant establishment.

Occurrence and persistence of non-native Cichlasoma urophthalmus (family Cichlidae) in estuarine habitats of south-west Florida (USA): environmental controls and movement patterns

The geographic scope of species introductions has increased dramatically in the last 200 years. One of the greatest challenges to understanding the ecological consequences of non-native species is that it has been difficult to identify the attributes that allow them to become established outside their native range. This challenge is compounded by anthropogenic habitat alteration that often enhances the establishment and expansion of non-native species. Fishes in the family Cichlidae are notoriously adaptive, and contribute more non-native species to North America than any other fish family. This is especially true in Florida (USA), where 18 cichlid species have become established. The Mayan cichlid (Cichlasoma urophthalmus) was first documented in south Florida in 1983, and is present in many freshwater and estuarine habitats. This study reports a northward range expansion by C. urophthalmus, demonstrates the connectivity between altered freshwater habitats and tidal estuarine habitats, and suggests that the estuarine populations are ephemeral and dependent upon colonisation from populations in altered freshwater habitats.

Policy and management responses to earthworm invasions in North America

The introduction, establishment and spread of non-native earthworm species in North America have been ongoing for centuries. These introductions have occurred across the continent and in some ecosystems have resulted in considerable modifications to ecosystem processes and functions associated with above- and belowground foodwebs. However, many areas of North America have either never been colonized by introduced earthworms, or have soils that are still inhabited exclusively by native earthworm fauna. Although several modes of transport and subsequent proliferation of non-native earthworms have been identified, little effort has been made to interrupt the flow of new species into new areas. Examples of major avenues for introduction of earthworms are the fish-bait, horticulture, and vermicomposting industries. In this paper we examine land management practices that influence the establishment of introduced species in several ecosystem types, and identify situations where land management may be useful in limiting the spread of introduced earthworm species. Finally, we discuss methods to regulate the importation of earthworms and earthworm-containing media so that introduction of new exotic species can be minimized or avoided. Although our focus in this paper is necessarily North American, many of the management and policy options presented here could be applicable to the problem of earthworm invasions in other parts of the world.

Changes in stand structure and composition after restoration treatments in low elevation dry forests of northeastern Oregon

In many fire-dependent forests in the United States, changes occurring in the last century have resulted in overstory structures, conifer densities, down woody structure and understory plant communities that deviate from those described historically. With these changes, many forests are presumed to be unsustainable. Broad-scale treatments are proposed to promote stand development on trajectories toward more sustainable structures. Yet little research to date has identified the effects of these restoration treatments, especially in low elevation dry ponderosa pine ( Pinus ponderosa) and Douglas-fir ( Pseudotsuga menziesii) forests of northeastern Oregon. We report changes in tree structure, coarse woody debris (logs), and understory composition from an operational-scale, replicated ( N = 4), completely randomized experiment. Treatments included a single entry thin from below conducted in 1998, a late season burn conducted in 2000, a thin followed by burning (thin + burn), and a no action treatment which served as a control. Changes in live and dead tree structure and understory vascular plant community composition were compared between pre-treatment and 2004. Tree seedling density and composition and coarse woody debris structure were evaluated in 2004. Thin, burn, and thin + burn treatments reduced the density but not the basal area of live overstory trees. Thinning reduced the number of medium-diameter trees, and tended to decrease the abundance of shade tolerant, moist-site understory species yet increased the dominance of several rhizomatous species such as Calamagrostis rubescens, Symphoricarpos albus, and Spiraea betulifolia. Burning alone had little effect on large trees but reduced the number of small Douglas-fir and logs. Shade tolerant perennial species associated with fine textured soils such as S. albus, Spiraea betulifolia, C. rubescens, Carex geyeri, and Arnica cordifolia increased in frequency and average cover with burning. Conversely, cover of the bunch grass Festuca idahoensis was reduced while non-native invasive species establishment was little affected. Ordination scores suggested that burning increased the abundance of species representing greater shade tolerance and finer-textured soils. The thin + burn treatment left both ponderosa pine and Douglas-fir with modal or normal diameter distributions, and increased the abundance of understory species representing shallow, coarse texture soils. These results are discussed in the context of management options for restoration of ecosystem health in similar low elevation dry ponderosa pine and Douglas-fir forests.

Establishment of non-native plant species after wildfires: effects of fuel treatments, abiotic and biotic factors, and post-fire grass seeding treatments

Establishment and spread of non-native species following wildfires can pose threats to long-term native plant recovery. Factors such as disturbance severity, resource availability, and propagule pressure may influence where non-native species establish in burned areas. In addition, pre- and post-fire management activities may influence the likelihood of non-native species establishment. In the present study we examine the establishment of non-native species after wildfires in relation to native species richness, fire severity, dominant native plant cover, resource availability, and pre- and post-fire management actions (fuel treatments and post-fire rehabilitation treatments). We used an information-theoretic approach to compare alternative hypotheses. We analysed post-fire effects at multiple scales at three wildfires in Colorado and New Mexico. For large and small spatial scales at all fires, fire severity was the most consistent predictor of non-native species cover. Non-native species cover was also correlated with high native species richness, low native dominant species cover, and high seeded grass cover. There was a positive, but non-significant, association of non-native species with fuel-treated areas at one wildfire. While there may be some potential for fuels treatments to promote non-native species establishment, wildfire and post-fire seeding treatments seem to have a larger impact on non-native species.

Global Documentation of Fish Introductions: the Growing Crisis and Recommendations for Action

Fish provides 15% of the total animal protein in human diets. It is also the primary source of livelihood for 35 million people (30 M in Asia and 2.6 M in Africa). The increase in global population and demand for fish protein cannot be met by capture fisheries alone. Governments are turning towards aquaculture as the source of fish protein. However, it has also led to the introduction and establishment of non-native species in local ecosystems through their escapement from aquaculture facilities to the wild. In freshwater ecosystems with relatively high endemism, this has become a significant problem. Documenting the international movement of fish is one way of providing a general view of the magnitude of these movements and the existing and potential threat faced by ecosystems due to species invasiveness. Information, however, is limited and scattered in different journals and agency/project reports. Several agencies, both local and international, have databases that provide information on invasive species (terrestrial and aquatic, local, regional or international in scope). The critical challenge is for consolidation, common access through data sharing and development of risk assessment and management tools. This is proposed through the use of Internet technology, sharing of databases or having a gateway or portal to which all introduced and invasive fish species related databases link. The fusion of these information sources will allow access to updated and reliable information. The experience of the WorldFish Center in documenting these phenomena through developing the FishBase information system and global partnerships is presented with recommendations for harmonizing approaches.

Nurse effect of the native cushion plant Azorella monantha on the invasive non-native Taraxacum officinale in the high-Andes of central Chile

Positive interactions among native plant species are common in alpine habitats, particularly those where one species (nurse plant) generates microclimatic conditions that are more benign than the surrounding environment, facilitating the establishment of other species. Nonetheless, these microclimatic conditions could facilitate the establishment of non-native species as well. A conspicuous component of the alien alpine flora of the central Chilean Andes is the perennial herb Taraxacum officinale agg. (dandelion). In contrast to other alien species that are restricted to human-disturbed sites, T. officinale is frequently observed growing within native plant communities dominated by cushion plants. In this study we evaluated if T. officinale is positively associated with the cushion plant Azorella monantha. Via seedling survival experiments and gas-exchange measurements we also assessed the patterns of facilitation between cushions and dandelions, and explore the potential mechanisms of invasion by dandelions. T. officinale grows spatially positively associated with cushions of A. monantha. Survival of seedlings, as well as their net-photosynthetic rates and stomatal conductance, were higher within cushions than in open areas away from them, suggesting that the microclimatic modifications generated by this native cushion facilitates the establishment and performance of a non-native invasive species. Our results, as well as other recent studies, highlight the role of native communities in facilitating rather than constraining non-native plant invasions, particularly in stressful habitats such as alpine environments.

The influence of disturbance and habitat on the presence of non-native plant species along transport corridors

The impact from transportation corridors on surrounding habitat often reaches far beyond the edge of the corridor. The altered disturbance regime in plant communities along corridor edges and vehicle traffic facilitate the spread and establishment of invasive non-native plant species. We compared the frequency of non-native plant species along highways and railways and the ability of these species to invade grasslands and dense forests along these corridors. We measured the frequency of several non-native plant species along transects 0–150 m from the edge of highways and railways in grasslands and forests, as well as at control sites away from corridors. Both transportation corridors had higher frequency of non-native species than respective control sites. Grasslands had higher frequency of non-native species than forested habitats, but the frequency did not differ between the highways and the railways. The frequency of non-native species in grasslands along highways and railways was higher than at grassland control sites up to 150 m from the corridor edge, whereas the frequency in forested habitats along corridors was higher than at forested control sites up to only 10 m from the corridor edge. There was a significant decrease in the frequency of non-native species with increasing distance from both corridors in the forest, while grasslands showed no significant change in non-native species frequency with distance from corridors. This suggests that corridor edges and grassland habitats act as microhabitats for non-native species and are more prone to invasion than forests, especially if disturbed. Our results emphasize the importance of minimizing the disturbance of adjacent plant communities along highways and railways during construction and maintenance, particularly in grassland habitats and in areas sensitive to additional fragmentation and habitat loss.

Impact of Slash Pile Size and Burning on Soil Chemical Characteristics in Ponderosa Pine Forests

This study examines the impacts of slash pile size and burning on various soil chemical characteristics and nutrient concentrations. Changes in soil characteristics following slash pile burns associated with forest thinning treatments may affect native and non-native floral species establishment and have broad implications for ecosystem functions, processes, and management. We sampled soils from plots under various slash pile burning treatments at The Arboretum at Flagstaff, Arizona, to determine their chemical characteristics.The characteristics consist of concentrations of macro- and micro-nutrients (N, P, K, Ca, Mg, Na, Mn, Fe, Zn, Cu, B) and pH. We compared the amounts of nutrients in burned plots with those in unburned plots where slash was removed by chipping. The treatments were tested on large and small piles. Burned slash pile plots had higher soil nutrient levels overall. The levels of N, Ca, K, Zn, and Na best illustrate this fertilizing effect. Significantly different values of N and P existed between treatments; however, the results show no distinct patterns related to the burning of slash. Nevertheless, there is enough evidence for a forest management decision involving burning any part of the forest to consider the effect of fires on soil chemistry.

Monitoring Nonnative Plant Invasions Over Fifty Years in Wisconsin Forests1

Long-term ecological change is often hard to document without reliable baseline data. We obtained baseline data for and resurveyed 80 forest understory communities in Wisconsin first surveyed 50 yr ago. Sixty-two sites were in the forested region of northern Wisconsin, and 18 were in the agricultural region of southern Wisconsin. For 50 yr, the number of sites with nonnative plants increased from 4 to 62. Species richness in 1950 had little influence on nonnative species establishment by 2002, and nonnative species did not seem to influence rates of native species loss over time.Additional index words: Ecologically invasive, forest, monitoring, plant, species loss, Wisconsin.

Non-native plant invasions in managed and protected ponderosa pine/Douglas-fir forests of the Colorado Front Range

We examined patterns of non-native plant diversity in protected and managed ponderosa pine/Douglas-fir forests of the Colorado Front Range. Cheesman Lake, a protected landscape, and Turkey Creek, a managed landscape, appear to have had similar natural disturbance histories prior to European settlement and fire protection during the last century. However, Turkey Creek has experienced logging, grazing, prescribed burning, and recreation since the late 1800s, while Cheesman Lake has not. Using the modified-Whittaker plot design to sample understory species richness and cover, we collected data for 30 0.1 ha plots in each landscape. Topographic position greatly influenced results, while management history did not. At both Cheesman Lake and Turkey Creek, low/riparian plots had highest native and non-native species richness and cover; upland plots (especially east/west-facing, south-facing and flat, high plots) had the lowest. However, there were no significant differences between Cheesman Lake and Turkey Creek for native species richness, native species cover, non-native species richness, or non-native species cover for any topographic category. In general, non-native species richness and cover were highly positively correlated with native species richness and/or cover (among other variables). In total, 16 non-native species were recorded at Cheesman Lake and Turkey Creek; none of the 16 non-native species were more common at one site than another. These findings suggest that: (1) areas that are high in native species diversity also contain more non-native species; (2) both protected and managed areas can be invaded by non-native plant species, and at similar intensities; and (3) logging, grazing, and other similar disturbances may have less of an impact on non-native species establishment and growth than topographic position (i.e., in lowland and riparian zones versus upland zones).

Do human activities raise species richness? Contrasting patterns in United States plants and fishes

AbstractIn a given area, human activities usually cause the extinction of native species and the establishment of non‐native species. A key conservation issue is whether non‐native establishment tends to outpace native species extinction to produce a net gain in species richness. To determine this, empirical data must be accumulated at various scales. I show that, within the United States, the number of established non‐native plant species per state does tend to outpace the number of extinct and threatened species per state. The net gain in plant species is strongly and positively correlated with human population density. Continuation of this trend predicts substantial gains in net plant species richness for all states in the United States as human population grows. This contrasts with freshwater fishes, where most states show a net loss of species diversity as extinct and threatened species exceed established non‐native species. Changes in fish diversity do not correlate strongly with human population or non‐native species but are largely driven by the decline of native fish species.

Changes in fish communities of the Shoalhaven River 20 years after construction of Tallowa Dam, Australia

AbstractThis study was undertaken as part of a long‐term investigation of the ability of high‐level fishways to rehabilitate fish communities upstream of high dams. Effects of Tallowa Dam on fish of the Shoalhaven River system were studied by comparing species abundances, population size‐structures and the structure of fish communities above and below the dam. Fish were sampled twice yearly for two years at 12 sites throughout the catchment. Species richness was greater downstream of the dam, with 21 species, compared to 16 species upstream of the dam. Ten diadromous species are believed to be extinct above the dam because of obstructed fish passage. Another four migratory species capable of climbing the wall have reduced abundances upstream. Accumulations of fish, particularly juveniles, directly below the dam were evident for nine species. Fish communities upstream and downstream of the dam differed significantly, identifying the dam as a significant discontinuity in the available fish habitats within the system. Historical evidence suggests that before the dam was built, fish communities from the tidal limit to at least 130 m elevation were largely continuous. This study has demonstrated that Tallowa Dam is a major barrier to fish migration and has had adverse effects on the biodiversity of the system. The creation of Lake Yarrunga by Tallowa Dam has resulted in distinctive fish communities in riverine and lacustrine habitats. Populations of five species that occur both upstream and downstream of the dam have developed differences in their size structures. The fish community downstream of the dam also differs from its historical condition because of the virtual disappearance of Australian grayling (Prototroctes maraena) and the establishment of non‐native species.A high‐level fishway is now being designed for the dam to restore fish passage. Data from this study will serve as a baseline against which to assess the effectiveness of the fishway in rehabilitating fish communities of the river system. Copyright © 2002 John Wiley & Sons, Ltd.

Effects of human population, area, and time on non-native plant and fish diversity in the United States

Non-native species diversity of plants and fishes in the contiguous 48 United States is analyzed to measure the influence of human population size, time of modern settlement, area and native species diversity. Besides exotic (from outside USA) plants, four types of non-native fishes are examined: established exotic fishes, reported exotic fishes, US fishes not native to a state, and native state fishes moved to new locations in a state. Human population size is most highly correlated with exotic plant diversity ( r>70%) but is still significantly correlated with most types of non-native fish diversity. Time of modern settlement significantly increases non-native plant (but not most fish) diversity, even after the effects of current population size are removed. These patterns occur because most non-native plants are imported for landscaping, farming and other uses intimately linked to human settlements whereas, almost half of non-native fishes were released by state agencies for sport, often into large western states with relatively few humans. This also explains why state area is significantly correlated with all types of non-native fish diversity, but not non-native plant diversity where smaller eastern states have more people and more years of settlement which increase non-native plant diversity. Positive correlation of non-native plant diversity with native plant diversity is found, as humans tend to settle in states with high native species diversity. In contrast, negative correlation between non-native fish and native fish diversity is found. These findings may help predict non-native species diversity if past trends continue. They also imply that the most cost-effective way to slow non-native species impact may be to focus where human population is still small, because rate of establishment of non-native species decreases with increasing human population.

Impact of gull colonies on the flora of the Riou archipelago (Mediterranean islands of south-east France)

The effects of an increasingly large yellow-legged gull Larus cachinnans colony on the flora of a Mediterranean limestone archipelago (south-east France) were studied through the analysis of floristical changes which have occurred in the past 36 years. Island plant lists can be distinguished first according to the island area, but all the islands show a common pattern of historical changes in flora. This floristical change appears to be correlated with the gull density and the percentage of ruderal plant species. Disturbance by gulls favours the massive establishment of non-native plant species which has led to the extinction of some endangered taxa through this plant species ‘enrichment’. Small islets appear to be more affected than large islands. Some preliminary conservation measures are presented.

Management of the introduction and transfer of marine molluscs

1. The introduction and transfer of marine molluscs for fisheries and aquaculture will include a risk of transporting competitors, predators, parasites, pests and diseases which can compromise intended molluscan culture and wild fisheries. 2. Introductions as well as transfers, in the course of normal trade, particularly of half-grown oysters, have been responsible for the establishment of several harmful and nuisance non-native species. Once established at a new locality these may continue to be moved by various means or by natural expansions of their range. 3. Several taxa have been associated with molluscan introductions and range expansions of these can be rapid should more than one dispersal vector be involved. 4. A Code of Practice for the introduction and transfer of marine organisms, prepared by the International Council for the Exploration of the Sea, provides a procedure which enables responsible movement of commercial species. In the case of an introduction it requires examination of the intended species by assessment of its likely ecological and genetic interaction with native species in advance of its release. It also includes measures, which if adopted, considerably reduce the risk of introducing accompanied unwanted species. 5. Introduced and useful species themselves, intended for culture, have seldom caused subsequent difficulties. 6. In this account the management of movements of non-native species, to and within northern Europe, are discussed. ©1996 by John Wiley & Sons, Ltd.