Stream macrophytes increase invertebrate production and fish habitat utilization in a California stream

Stable flow and thermal regimes, coupled with geologically derived nutrients, are drivers of enhanced productivity in volcanic spring‐fed rivers. However, little information exists on biotic mechanisms or species interactions contributing to elevated productivity at higher trophic levels. In a California stream, juvenile steelhead trout were observed to preferentially select macrophyte habitat at a rate three times greater, on average, than five other habitat types. To understand the potential rearing benefits associated with macrophytes, we conducted a manipulative experiment to determine how macrophytes affect invertebrate prey availability and stream water velocity. Macrophytes supported up to nine times greater abundance of invertebrates than adjacent open gravel habitats. They also doubled invertebrate drift rates and reduced water velocity by up to 42‐fold. The results show that aquatic macrophytes are an important stream habitat feature that may be bioenergetically more favourable for rearing salmonids than more traditional lotic habitats. We suggest that macrophytes have the potential to enhance growth rates of juvenile salmonids when compared with other habitat types. Habitats that confer growth and size advantages may ultimately improve fitness and contribute to conservation of imperilled salmonids.     

Effects of transferring glacier-fed water to a clear-water mountain river on the production and food organisms of brown trout (Salmo trutta L.) in Southern Norway

The effects of transferring glacier-fed water to a clear-water river on production and food organisms of brown trout (Salmo trutta L.) were investigated in a mountain river, in southern Norway. Production in unregulated fluvial habitat was 271.5 g 100 m^?2 yr^?1 compared with 103.1 g 100 m^?2 yr^?1 in a glacier-fed reach of the river. This difference was due to low fish density and recruitment rate. The growth rate, at least for younger fish (2+ to 4+ age groups), was significantly higher in the regulated reach of the river. The main difference in food composition was the low abundance of crustaceans Eurycercus lamellatus and Gammarus lacustris in the regulated reach. Trichopterans were the main diet component in both sites.     

Application of GIS modelling to quantify fish habitats in lakes

Quantitative measures of fish habitat in lakes and impoundments have seldom been proposed. The availability of Geographic Information Systems (GIS), however, now provides a means for researchers to assess fish habitat on a whole-lake scale. GIS modelling was used in this study to quantify the impacts of stratification on the amount and distribution of brown trout ( Salmo trutta ) habitat in a large reservoir in Victoria, Australia. Species-specific temperature and dissolved oxygen tolerance ranges for brown trout were applied to the hydrological profile and lake bathymetry. The model predicted that brown trout habitat was reduced by 82% in the summer, and the model was validated with position data from acoustically tagged adult brown trout. This study demonstrated that a quantifiable assessment of fish habitat on a whole-lake spatial scale can be made. It also could be applied to a range of fish species, with applications in ecosystem monitoring and fisheries management.     

Periphyton and invertebrate response to wood placement in large pacific coastal rivers

Wood abundance in aquatic systems has been dramatically reduced compared to historical levels due to anthropogenic activities that led to wood removal and stream simplification. As a result, reintroduction of wood to aquatic systems is now a widely used and relatively well‐studied restoration technique for increasing habitat complexity. Although stream periphyton (biofilm) and invertebrates serve as food sources for a variety of predators including fish, birds and bats, data on how lower trophic levels respond to wood placement are relatively scarce. The purpose of this study was to test the hypothesis that periphyton biomass and aquatic invertebrate density were higher on Engineered Log Jams (ELJs) than on inorganic substrates in two large Pacific Northwest river systems. Among years and rivers, periphyton biomass and invertebrate densities were significantly higher on ELJs than on cobbles within the same reach. Invertebrate communities on ELJs were dominated by meiofauna (<500 µm), whereas cobbles were dominated by larger chironomids. We attribute these trophic level differences to substrate type, as we did not detect taxonomic differences between cobbles in reaches with and without ELJs. We show that adding wood to reaches with little or no naturally occurring wood increased overall habitat surface area and thereby the potential for increased productivity relative to reaches with low levels of wood. Finally, wood supports a unique community of invertebrates that are often overlooked in lotic system studies but may be contributing substantially to overall biological diversity. Published in 2009 by John Wiley & Sons, Ltd.     

Interactive effects of cover and hydraulics on brown trout habitat selection patterns

Habitat modelling results are extremely sensitive to the habitat suitability criteria (HSC) used in the simulations. HSCs are usually expressed as univariate habitat suitability curves, although such univariate approach has been long questioned, since overlooking interactions between hydraulic variables may misrepresent the complexity of fish behaviour in habitat selection. It could lead to adopt erroneous flow management decisions based on misleading results. Furthermore, the interactive effects of hydraulic variables on habitat selection may be driven by the structural features of the channel, which determine cover availability. Therefore, we compared brown trout habitat selection patterns through multivariate resource selection functions (RSFs) in structurally contrasting rivers to unveil the interactive effects of hydraulics and cover elements and their consequences in univariate HSC results. Microhabitat preferences of young‐of‐the‐year (0+) trout were similar across fast and slow waters, meanwhile juvenile (1+) and adult (>1+) preferences significantly changed. RSFs for young‐of‐the‐year trout were consistent with univariate results and did not differ among water types. However, RSFs for older trout varied among water types and revealed complex interactions among hydraulic variables and between hydraulics and structural elements, which were not described accurately by univariate curves. Therefore, results suggest that interactions between water depth and current velocity have a significant effect on habitat selection patterns in juvenile and adult brown trout, this effect being controlled by cover availability. Copyright © 2008 John Wiley & Sons, Ltd.     

PCB Concentrations of Lake Michigan Invertebrates: Reconstruction Based on PCB Concentrations of Alewives (Alosa pseudoharengus) and their Bioenergetics

Invertebrate PCB concentrations are a poorly quantified but crucial step in the trophic transfer of organochlorine contaminants to fishes. In fact, current attempts to quantify PCB fluxes in the Lake Michigan pelagic food web are hampered by poor knowledge of invertebrate PCB concentrations. Models exist that estimate PCB concentrations in fish based upon PCB concentrations in their food. We have used a complementary approach and estimated invertebrate PCB concentrations based upon historical records of alewife PCB concentrations in Lake Michigan. We first developed a model of total PCB accumulation in alewife (Alosa pseudoharengus) using a bioenergetics-based approach to growth. The PCB assimilation efficiency between invertebrate prey and alewife predators was estimated to be about 0.40. We then used the model to hindcast PCB concentrations in the invertebrates that comprised alewife prey. We estimated that median PCB concentrations of Lake Michigan invertebrates have dropped roughly 10-fold from 1976 to 1990; for example, the estimated Diporeia hoyi PCB concentration has dropped from ca. 0.48 mgkg^-1 wet weight to 0.04 mgkg^-1 wet weight over this time period. The biomagnification ratio (alewife PCB/zooplankton PCB) is about 16-fold (Diporeia hoyi) to 40-fold (copepods/cladocerans). The PCB concentrations in Lake Michigan invertebrates and alewife that we have estimated for 1993 and 1994 should be viewed as predictions, testable as data become available. Because historic data on invertebrate PCB concentrations in Lake Michigan are exceedingly scarce, our estimates should be useful for studies that attempt to quantify the Lake Michigan PCB fluxes or future modeling efforts that attempt to incorporate multiple levels of the Lake Michigan food web.     

Evaluating Methods to Establish Habitat Suitability Criteria: A Case Study in the Upper Delaware River Basin,USA

Defining habitat suitability criteria (HSC) of aquatic biota can be a key component to environmental flow science. HSC can be developed through numerous methods; however, few studies have evaluated the consistency of HSC developed by different methodologies. We directly compared HSC for depth and velocity developed by the Delphi method (expert opinion) and by two primary literature meta‐analyses (literature‐derived range and interquartile range) to assess whether these independent methods produce analogous criteria for multiple species (rainbow trout, brown trout, American shad, and shallow fast guild) and life stages. We further evaluated how these two independently developed HSC affect calculations of habitat availability under three alternative reservoir management scenarios in the upper Delaware River at a mesohabitat (main channel, stream margins, and flood plain), reach, and basin scale. In general, literature‐derived HSC fell within the range of the Delphi HSC, with highest congruence for velocity habitat. Habitat area predicted using the Delphi HSC fell between the habitat area predicted using two literature‐derived HSC, both at the basin and the site scale. Predicted habitat increased in shallow regions (stream margins and flood plain) using literature‐derived HSC while Delphi‐derived HSC predicted increased channel habitat. HSC generally favoured the same reservoir management scenario; however, no favoured reservoir management scenario was the most common outcome when applying the literature range HSC. The differences found in this study lend insight into how different methodologies can shape HSC and their consequences for predicted habitat and water management decisions. Published 2016. This article is a U.S. Government work and is in the public domain in the USA.     

A MODEL INCORPORATING DISTURBANCE AND RECOVERY PROCESSES IN BENTHIC INVERTEBRATE HABITAT—FLOW TIME SERIES

We describe and demonstrate a model (Benthic Invertebrate Time Series Habitat Simulation) for calculating the effect of changes to flow regimes on benthic invertebrate habitat and population dynamics. The following inputs are required: a hydrograph (discharge time series), habitat–discharge relationship, disturbance–discharge relationship, wetted width–discharge relationship and a recolonization time series. Habitat–discharge, disturbance–discharge and wetted width–discharge relationships are common outputs from instream hydraulic habitat models (e.g. Physical Habitat Simulation, River Hydraulic Habitat Simulation and River2D). Hydraulic habitat models calculate a combined habitat suitability index from physical habitat suitability curves for water depth, velocity and substrate composition and weight this by area to give a weighted usable area (WUA). Because conventional invertebrate habitat suitability curves are based on density estimates, the combined habitat suitability index can be treated as an index of density and WUA treated as an index of potential relative abundance (at the reach scale) in the absence of disturbance due to flow variation (flooding and drying) and biotic processes. Our approach begins with WUA and calculates realizable suitable habitat (i.e. relative abundance) by taking into account the resetting of benthic invertebrate densities by floods and drying and recovery (or accrual) rates and times. The approach is intended mainly to compare the relative amounts of productive invertebrate habitat sustained by natural and modified flow regimes, but it also has the potential for investigating the influence of flow variation on invertebrate population dynamics. We anticipate that the model will be particularly useful for assessing effects of changes in flow regimes caused by diversions, abstractions or water storage on annual benthic invertebrate productivity. Copyright © 2013 John Wiley & Sons, Ltd.     

Diet and trophic niche space and overlap of Lake Ontario salmonid species using stable isotopes and stomach contents

Lake Ontario supports a diversity of native and non-native salmonids which are managed largely through stocking practices. Ecological changes (e.g., invasive species) altering the food web structure accompanied with shifts in prey abundance, necessitate understanding the trophic niches of Lake Ontario salmonids to aid in management. The objectives of this study were to quantify salmonid (5 species) trophic niches and dietary proportions using stable isotope ratios (δ¹³C and δ¹⁵N) of a large sample set (adult fish (>300?mm; n?=?672) and key offshore prey (5 species, n?=?2037)) collected across Lake Ontario in 2013. Estimates of prey based on stable isotope ratios were similar to stomach contents. Based on stable isotope ratios, non-native prey dominated salmonid diet; in particular alewife (Alosa pseudoharengus) constituted the majority (0.31 to 0.93) of all salmonid diets, and round goby (Neogobius melanostomus) contributed 0.26 and 0.19 of brown trout (Salmo trutta) and lake trout (Salvelinus namaycush) diets, respectively. Trophic niche overlap was high between all salmonids, except lake trout. The largest trophic niche overlap occurred between Chinook (Oncorhynchus tshawytscha), coho (Oncorhynchus kisutch), and Atlantic salmon (Salmo salar), and their reliance on alewife infers a strong pelagic foraging strategy. Lake, brown and rainbow (Oncorhynchus mykiss) trout had larger and/or more distinct trophic niches indicative of a more variable diet across individuals and utilizing different foraging strategies and/or habitats. Overall, Lake Ontario salmonids maintained a high reliance on alewife, and their potential for plasticity in diet provides important information to management regarding population sustainability.     

Trophic interactions among algal blooms,macroinvertebrates, and brown trout: Implications for trout recovery in a restored river

Positive correlation between trout abundance and dissolved metal concentrations along the Upper Clark Fork River (UCFR; Montana, USA) have forced restoration practitioners to seek underlying causes of reduced fish density beyond heavy metal contamination. Throughout the river, nutrient enrichment and summer algal blooms may be hindering full recovery of trout populations. In this study, we evaluated the community structure and metal body burdens of benthic invertebrates and characterized existing trophic linkages between brown trout and dominant invertebrate taxa before and during summer algal blooms in a downstream reach of the UCFR where fish densities are low (20–30 trout/km), and where metal contamination is relevant but minimal compared with upstream. In spring, estimated invertebrate abundance was 1,727 ± 217 individuals/m² and dominated by Ephemerellidae and Baetidae families. During summer algal bloom, invertebrate abundance increased 15‐fold (20,580 ± 3,510 individuals/m²) mostly due to greater abundance of Chironomidae, Hydropsychidae, and Simulidae. Copper body burdens (130 ± 42 ppm) were higher than any other heavy metal regardless of season, but detectable concentrations of arsenic, cadmium, and lead were also found. A Bayesian mixing model combining metal burdens and stable isotopes showed that in the spring, trout of average size (355 ± 65 g) relied mostly on epibenthic taxa (Ephemerellidae and Hydropsychidae), contrasting with small (<100 g) and large (>400 g) trout relying heavily on Baetidae, a major component of invertebrate drift. Foraging segregation related to trout size did not occur during summer algal blooms, which may reflect increasing influence of benthic algal proliferation or indicate the indiscriminate use of pool habitats as thermal refugia over summer conditions by trout of different ages.     

Fatty acids reveal salmonine – prey relationships in Lake Michigan

Lake Michigan salmon and trout populations are important species for recreational fisheries and food web management, and are largely supported through stocking efforts, with varying degrees of natural recruitment. Ongoing fisheries management of these salmonine populations is dictated by relationships between predator and prey abundance as well as community structure within the lake. However, while prey fish biomass has declined, and species composition has changed in recent decades, knowledge of prey consumption by the salmonine community has lagged. Herein, we explore trophic relationships using fatty acids profiles, which offer insights into the foraging habits and energy pathways relied on over weeks to months prior to collection. Fatty acids of the prey base for salmonines in Lake Michigan indicate a gradient of foraging habits that range from pelagic (typified by alewife and rainbow smelt) versus benthic (i.e., slimy sculpin and round goby) resource use. Fatty acids implied that there was more variation in foraging habits among individual lake trout and brown trout compared to Chinook salmon, coho salmon and rainbow trout, which appeared to all rely almost exclusively on pelagic prey. Fatty acid profiles also indicated size-based shifts in foraging habits; for example, larger lake trout consuming a greater proportion of benthic prey than smaller individuals. Data herein suggest that Chinook and coho salmon, as well as rainbow trout, are more likely to experience competitive interactions during times of low pelagic prey-fish abundance in Lake Michigan, whereas brown and lake trout are able to utilize benthic resources to a greater degree.     

A probability-based model to quantify the impact of hydropeaking on habitat suitability in rivers

A negative effect of hydropower on river environment includes rapid changes in flow and habitat conditions. Any sudden flow change could force fish to move towards a refuge area in a short period of time, causing serious disturbances in the life cycle of the fish. A probability-based model was developed to quantify the impact of hydropeaking on habitat suitability for two fish species, brown trout (Salamo trutta) and Grayling (Thymallus thymallus). The model used habitat preference curves, river velocity and depth to develop the suitability maps. The suitability maps reveal that habitat suitability deteriorates as flow increases in the studied part of the river. The probability model showed that, on average, suitability indices are higher for adult grayling than juvenile trout in hydropeaking events in this part of the river. The method developed shows the potential to be used in river management and the evaluation of hydropeaking impacts in river systems affected by hydropower.     

Development and evaluation of bioenergetic-based habitat suitability criteria for trout

We constructed energetic models of habitat use for 82–322 g rainbow trout (Oncorhynchus mykiss) in a large regulated river, and 8–28 g Colorado River cutthroat trout (O. clarki pleuriticus) in a small headwater stream, to determine if observed summer habitat use by these species could be attributed to net energy acquisition, and to develop habitat suitability criteria based on net energy gain. Metabolic models of energy expenditure were derived from literature sources, but measurements of energy availability were site-specific. From the energy models, we assigned a suitability value of 1.0 to the entire range of velocities where positive net energy gains were predicted, and a suitability value of zero to velocities where negative net energy gains were predicted. Predicted net energy gain velocities were compared with observed velocities used by each species. For rainbow trout, the energetic model predicted energetically profitable velocities ranging from 5 to 45 cm s⁻¹. Predicted velocities were similar to velocities used by rainbow trout. This indicated that rainbow trout, as a group, were using energetically profitable stream locations, but some rainbow trout used non-profitable velocities. For Colorado River cutthroat trout, the energetic model predicted energetically profitable velocities ranging from 5 to 45 cm s⁻¹; however, Colorado River cutthroat trout used significantly lower velocities than predicted. The dissimilarity between velocities predicted and used by Colorado River cutthroat trout may be attributed to their inability to utilize energetically profitable velocities available in the stream because of depth restrictions The results suggest that the predictive abilities of energetic models vary between streams because of differences in depth and velocity availability. © 1997 John Wiley & Sons, Ltd.     

Production sources and food web of a macrophyte-dominated region in Lake Taihu,based on gut contents and stable isotope analyses

We estimated the food web linkages among primary producers, invertebrates and fish in a macrophyte-dominated region of a eutrophic lake (Lake Taihu) by analyzing gut contents and C and N stable isotope ratios. Observation of the gut contents reflected a variety of feeding modes among fish species that consume a diverse assortment of prey, with limited dietary overlap. Basal food sources were distinguishable based on their δ¹³C isotopic signatures, and wide seasonal variations of isotope values were observed in the lake biota with a general trend towards enriched δ¹³C and δ¹⁵N values in summer and depleted values in winter. This pattern could be explained by a combination of environmental (e.g., irradiance and nutrient inputs) and biotic (e.g., availability of food sources and plasticity in prey item choice) features. We adopted a paired (gut contents and stable isotopes) approach to reconstruct diets that used known food items as end members in the isotopic mixing model analysis and diagrammed the food web structure of Lake Taihu describing the organic matter pathways from primary producers to predators of the upper trophic levels. Our surveys also corroborate the finding that phytoplankton were the most important primary source of organic matter for fauna within this macrophyte-dominated lake region, whereas macrophytes also made a sizeable contribution for several invertebrate and fish species, especially in winter.     

An evaluation of inter-basin water transfers as a mechanism for augmenting salmonid and grayling habitat in the River Wear,North-East England

Transfers of water from the Kielder system have been used for 12 years to avoid low flow problems in the River Wear. Transfers are scheduled to avoid breaches of the river's statutory Minimum Maintained Flow (MMF). Despite this routine use, the role of transfers in augmenting instream habitat has never been evaluated. A physical habitat simulation (PHABSIM) study was undertaken in 1996 to investigate the influence of transfers and the MMF policy on brown trout Salmo trutta, Atlantic salmon Salmo salar and grayling Thymallus thymallus instream habitat at three sites on the Wear. Transfers support total habitat levels up to 10% greater than unregulated conditions. They impact usable instream habitat (weighted usable area) to a much greater extent. For salmon parr, the species/lifestage whose habitat is most limited by low flows, transfers have maintained relatively stable usable habitat levels during periods when otherwise they would have fallen by as much as 70%. The MMF policy results in minimum flow values which are higher than those which would have been set using the Montana Method and the availability of salmon parr usable habitat does not fall below 10% of its mean annual value. Judged in these terms, the MMF-based transfer regime has played a positive role in avoiding extreme habitat loss in the Wear. Simulations of four alternative Kielder transfer release policies indicate that near-optimum habitat levels could be maintained throughout summer low flow periods. However, this would result in unnatural temporal patterns of flow and habitat availability. © 1998 John Wiley & Sons, Ltd.     

The effect of discharge and predation on habitat use by wild and hatchery brown trout (Salmo Trutta)

The effects of trout stock, discharge and predation risk on habitat use by brown trout, Salmo trutta, were studied in four artificial streams. Trout stock had no effect on habitat use as both wild and hatchery fish used similar habitats. The presence of pike (Esox lucius) caused trout to decrease their use of pools, the habitat in which pike occurred, and increase their use of other habitats. Decreasing discharge reduced available area of the stream and resulted in fewer fish in the shallow margins. Both decreased flow and increased predation risk caused more overlap in habitat use, and thus increased the potential for intraspecific competition, predation and the use of poorer habitats. The results illustrate the danger of applying habitat use relationships obtained from one stream to all other streams where habitat availability and biotic interactions may differ.     

INFLUENCE OF THE MORPHOLOGICAL AND HYDRAULIC CHARACTERISTICS OF MOUNTAIN STREAMS ON FISH HABITAT SUITABILITY CURVES

Microhabitat preferences of adult brown trout (Salmo trutta m. fario) were monitored for the purpose of determining design parameters for river restoration. The habitat preferences were evaluated during the summer period of minimum flows. Since 1995, field measurements have been performed in 52 reaches in 43 mountain and piedmont streams. The relationship between hydraulic characteristics and the values of maximum habitat suitability derived from velocity and depth habitat suitability curves (HSCs) was statistically determined. Trout in natural stream reaches showed a strong degree of dependence on depths, but in regulated streams, they were dependent on velocities. The representative habitat suitability curves for four depth intervals were extrapolated. From these outputs, the optimum depths of a microhabitat for river restoration measures and/or assessment of the influence of water withdrawals can be derived. The influence of geological regions on the shape of HSCs has not been proved; therefore, it is conceivable that after verification, the generalized HSCs may also be valid in other mountain and piedmont regions. Copyright © 2011 John Wiley & Sons, Ltd.     

Gut contents from multiple morphs of lake trout (Salvelinus namaycush) at two offshore shoals in Lake Superior

Four lake trout, Salvelinus namaycush, Walbaum 1792 morphs occur in Lake Superior: lean, siscowet, humper, and redfin. Diets of lean and siscowet have been relatively well described. However, less is known about diets of humper and redfin, and overall few studies have been conducted at offshore shoals. We compared gut content data among mature (357–867 mm) sympatric lake trout morphs caught at two offshore shoals in Lake Superior, Stannard Rock and Superior Shoal, in 2013 and 2014 (total n = 416). All morphs were caught in shallow (<50 m), mid (50–100 m), and deep (>100 m) strata. Invertebrates made up a greater portion of the stomach contents than did fish for all morphs by both percent occurrence and proportional biomass, and Mysis was the primary invertebrate consumed by all morphs at both sites. Coregonus spp. and deepwater sculpin, Myoxocephalus thompsonii were the most commonly consumed fish. Humper had the highest average proportional biomass of deepwater sculpin and had no other identifiable species of fish in their guts. Biomass of fish in redfin guts was highest for Coregonus spp., followed by similar amounts of deepwater sculpin and burbot, Lota lota. Diet overlap among morphs was high, and differences in prey consumption between sites are likely related to prey availability. Additional study is needed to determine if differences in trophic ecology between humper and other morphs are sufficient to support concurrent stocking of multiple morphs, particularly in light of recent declines in native prey fishes, especially Coregonus spp., in the Laurentian Great Lakes.     

Evaluation of δD and δ18O as natural markers of invertebrate source environment and dispersal in the middle Mississippi River‐floodplain ecosystem

Movement of invertebrates among large rivers, tributaries, and floodplain lakes or dispersal of adult aquatic insects from riverine or floodplain habitats may provide important subsidies to food webs in receiving habitats. Intensive sampling at habitat interfaces and artificial labelling are two approaches to assess freshwater invertebrate dispersal, but these are difficult to implement at a landscape scale. Natural chemical tracers have been used to track dispersal of fishes and marine invertebrates, but the potential applicability of stable isotope ratios as natural tracers of invertebrate dispersal in freshwater environments has not been assessed. We evaluated stable hydrogen and oxygen isotopes (δD and δ¹⁸O) as natural markers of source environment and dispersal of macroinvertebrates in the middle Mississippi River, tributaries and floodplain wetlands. Water and invertebrates were collected from 12 sites during 2007–2008. Water δD and δ¹⁸O differed among the river, its tributaries, and floodplain wetlands and were strongly correlated with invertebrate δD and δ¹⁸O. Variability in invertebrate δ¹⁸O rendered it ineffective as an indicator of invertebrate source environment. Mean δD of Mississippi River invertebrates differed from δD of invertebrates from floodplain wetlands; δD distinguished invertebrates from these two environments with >80% accuracy. Neither δD nor δ¹⁸O of aquatic insects changed following emergence from their natal site. Preservation method (ethanol or freezing) did not affect invertebrate δD or δ¹⁸O. Invertebrate δD may be a useful natural tracer of natal environment and dispersal in the Mississippi River‐floodplain ecosystem and other freshwater systems where spatial variation in water δD is present. Copyright © 2010 John Wiley & Sons, Ltd.