From salmon to shad: Shifting sources of marine‐derived nutrients in the Columbia River Basin

Like Pacific salmon (Oncorhynchus spp.), nonnative American shad (Alosa sapidissima) have the potential to convey large quantities of nutrients between the Pacific Ocean and freshwater spawning areas in the Columbia River Basin (CRB). American shad are now the most numerous anadromous fish in the CRB, yet the magnitude of the resulting nutrient flux owing to the shift from salmon to shad is unknown. Nutrient flux models revealed that American shad conveyed over 15,000 kg of nitrogen (N) and 3,000 kg of phosphorus (P) annually to John Day Reservoir, the largest mainstem reservoir in the lower Columbia River. Shad were net importers of N, with juveniles and postspawners exporting just 31% of the N imported by adults. Shad were usually net importers of P, with juveniles and postspawners exporting 46% of the P imported by adults on average. American shad contributed <0.2% of the total annual P load into John Day Reservoir, but during June when most adult shad are migrating into John Day Reservoir, they contributed as much as 2.0% of the P load. Nutrient inputs by American shad were similar to current but far less than historical inputs of Pacific salmon owing to their smaller size. Given the relatively high background P levels and low retention times in lower Columbia River reservoirs, it is unlikely that shad marine‐derived nutrients affect nutrient balances or food web productivity through autotrophic pathways. However, a better understanding of shad spawning aggregations in the CRB is needed.     

Environmental DNA reveals the temporal and spatial extent of spawning migrations of European shad in a highly fragmented river basin

1. Artificial barriers on lowland rivers impede the spawning migrations of anadromous fishes, preventing access to historical spawning areas. In the cryptic European shads Alosa alosa and Alosa fallax (‘shad’ hereafter), this has resulted in population declines across their range. Conservation programmes aim to facilitate the passage of migrators over these barriers and so require baseline information on the spatial and temporal extent of current migrations.
2. Here, a shad-specific environmental DNA (eDNA) assay was used to quantify the spatial extent of shad spawning migrations in the River Severn basin, western England. This basin is characterized by the presence of multiple barriers in the lower catchment. In 2017, the eDNA assay was piloted in the River Teme, an important shad spawning tributary, and then applied in 2018 and 2019 across the lower Severn basin.
3. In all years, shad DNA was detected between mid-May and mid-June, with the maximum spatial extent of shad distribution being in early June when shad eDNA was detected upstream of weirs that were generally considered as impassable. In 2018, this included the detection of shad above the most upstream weir on the main River Severn that required individual fish to have passed six weirs.
4. Although barriers inhibit the spawning migrations of shad, this eDNA assay showed that some highly vagile individuals might be able to ascend these barriers and migrate considerable distances upstream. This suggests that efforts to increase the permeability of these barriers could result in relatively high numbers of migrating shad reaching upstream spawning areas. These results demonstrate that this eDNA assay could also be used across their range, to further quantify the spatial extent of their spawning, including in highly fragmented rivers and those where shad are believed to spawn only occasionally and are rarely observed.

     

Digestion of larval American shad by cyprinids

Cyprinids have pharyngeal teeth for grinding food before swallowing and a continuous gut with no discrete stomach. This digestive tract structure, as well as the feeding behavior traits shown by cyprinids, makes it difficult to identify and measure the amount of food consumed by these fishes. The relations among quantity of food in cyprinid gut, time after feeding and predator size were described by log-linear multiple regression. The number of intact American shad (Alosa sapidissima (Wilson)) larvae eaten in laboratory experiments was estimated by extrapolating the regression for the amount of food in the gut over time for predators of various size. The numbers of larval American shad in guts of fish captured in the Juniata River, Pennsylvania, were not significantly different from those estimated with the equations derived from laboratory data. In view of their abundance in rivers and their potential digestion rate for larval fish, cyprinids can be expected to have a marked influence on reducing the numbers of larval American shad.     

Large‐scale,seasonal habitat use and movements of yellow American eels in the St. Lawrence River revealed by acoustic telemetry

Large‐scale habitat use and movements of yellow American eels (Anguilla rostrata) from the St. Lawrence River were examined using acoustic telemetry from early summer to late fall in 2010 and 2011. Sixty‐seven eels were tagged, and their passage or presence was recorded using fixed acoustic arrays covering a 400 km distance along the St. Lawrence River and Estuary. Sixty‐four per cent of the 67 tagged eels were detected. Most eels were detected at only one array; the closest to their release location and at several occasions during the tracking period, suggesting a high proportion of freshwater residency in the upstream part of the St. Lawrence River. Downstream movements towards the brackish estuary (63–418 km distance) were demonstrated for 16.4% of the eels, particularly for those caught at the most downstream site that is close to the brackish estuary. Our results strongly suggest a lower activity of freshwater resident yellow eels during summer, a behaviour that may be related to day length, which defines time available for their nocturnal foraging. Indeed, yellow eels were detected primarily at night; no effect of moon phase was revealed. Movements in the vicinity of arrays (up to 116 km in the fluvial estuary) were suggested and smaller‐scale movements within Lac St. Louis were demonstrated, highlighting a yellow‐eel home range far more extensive than previously reported in smaller systems. Evidence for within‐season homing and site fidelity is also reported.     

Novel insights into the marine phase and river fidelity of anadromous twaite shad Alosa fallax in the UK and Ireland

1. Most research on anadromous fishes has been invested in their freshwater life‐phases, resulting in a relatively sparse understanding of their spatial ecology during marine life‐phases. However, understanding the marine dispersal of anadromous fishes is essential to identify threats and to implement conservation measures that fully encompass their lifecycle.
2. The twaite shad Alosa fallax is an anadromous fish increasingly imperilled across its range due to pollution, harvesting, and impediments to freshwater migration, but little is known about its distribution and movements during its marine life‐phase. Here, the application of acoustic telemetry provided novel insights into the coastal dispersal of twaite shad in the UK and Ireland during 2018–2019, and the freshwater entry of individuals during the 2019 spawning season.
3. Of 73 twaite shad acoustic‐tagged during their upstream migration in the River Severn in May 2018, 58 emigrated from the river. Twelve were subsequently detected 200 km to the south‐west at the Taw–Torridge Estuary between July 2018 and April 2019, where estuarine movements up to 5.8 km inland occurred in summer, winter, and spring. One was subsequently detected in the Munster Blackwater Estuary (Ireland) and then in the River Severn, indicating a minimum movement distance of 950 km. Thirty‐four (59%) of the emigrating individuals from 2018 re‐entered fresh water in the rivers Severn (n = 33) and Wye (n = 2) in April and May 2019.
4. These results suggest year‐round use of estuarine and nearshore habitats by at least a subset of the twaite shad population during their marine phase, providing evidence of potential range overlap between populations that spawn in different areas in the UK and Ireland, which may be facilitated by substantial dispersal. The results also highlight the potential of telemetry for estimating freshwater and marine mortality, and the benefits of sharing detection data across networks.

     

High variability in spawning migration of sea trout,Salmo trutta,in two northern Swedish rivers

Abstract Telemetry was used to examine spawning migration of sea trout, Salmo trutta L. (n = 126), in two rivers in northern Sweden. The spawning areas defined by radio‐tagged fish differed between the river systems. In the River Vindelälven, sea trout spawned in the main stem and 80% of tagged individuals returned to areas where hatchery‐reared juveniles had been previously stocked. In the River Piteälven, 74% of tagged sea trout ascended tributaries for spawning. Tagged fish were categorised into three groups of migratory pattern. cart (classification and regression tree) analysis indicated that distance from tagging location to spawning site (S_dist) explained the migratory patterns. Large S_dist separated fish with stepwise upstream migration from those with up‐ and downstream migrations and one‐directional direct migration. Fish tagged early in the season migrated the longest distance to spawning areas. Stocking locations and sex explained the large search behaviour up‐ and downstream in the rivers. The findings are important for the sustainable management of sea trout in the Gulf of Bothnia.     

Interannual differences in growth and hatch‐date distributions of early juvenile European anchovy in the Bay of Biscay: implications for recruitment

In order to understand better the recruitment variability in European anchovy in the Bay of Biscay, it is important to investigate the processes that affect survival during the early life stages. Anchovy juvenile growth trajectories and hatch‐date distributions were inferred over a 3‐year period based on otolith microstructure analysis. Otolith growth trajectories showed a characteristic shape depending on their hatch‐date timing. Earlier‐born juveniles had notably broader maximum increments than later born conspecifics, resulting in higher growth rates. This observation suggests that early hatching would be beneficial for larval and juvenile growth, and, therefore, survival. The estimated juvenile hatch‐date distributions were relatively narrow compared with the extended anchovy spawning season (March–August) in the Bay of Biscay and indicated that only individuals originated mainly from the summer months (June–August) survived until autumn. Hatch‐date distributions were markedly different among years and seemed to influence the interannual recruitment strength of anchovy. We conclude that years characterized by juvenile survivors originating from the peak spawning period (May and June) would lead to considerable recruitment success. Downwelling events during the peak spawning period seem to affect larval survival. Furthermore, size‐dependent overwinter mortality would be an additional process that regulates recruitment strength in the anchovy population in the Bay of Biscay.     

Environmental DNA as a non‐invasive sampling tool to detect the spawning distribution of European anadromous shads (Alosa spp.)

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Large‐bodied fish migration and residency in a flood basin of the Sacramento River,California, USA

River–floodplain complexes represent some of the most variable and diverse habitats on earth, yet they are among our planet's most threatened ecosystems. Use of these habitats by large‐bodied fishes is especially poorly understood, particularly in temperate regions. To provide insight into the factors that affect floodplain assemblages and migration, we sampled large‐bodied fishes with a fyke trap for 7 years in the Yolo Bypass, the primary flood basin of the Sacramento River, California. We collected a total of 18,336 individual fish comprised of 27 species, only 41% of which were native. Year‐round resident species white catfish Ameiurus catus, channel catfish Ictalurus punctatus and common carp Cyprinus carpio (all alien species) were the most abundant and comprised 74% of the total catch. Splittail Pogonichthys macrolepidotus (3.8%), white sturgeon Acipenser transmontanus (2.3%) and Sacramento sucker Catostomus occidentalis (1.1%) were the primary native species. We found that seasonal variation in water temperature and flood stage were important factors affecting the fish assemblage structure and the presence of migratory species. American shad Alosa sapidissima, an alien species, showed highest abundance during the early summer upstream migration, when temperatures were warmer. For native species, the abundances of white sturgeon, splittail, Sacramento pikeminnow Ptychocheilus grandis and Sacramento sucker were all highest during flood pulses. While our results suggest that flow alone is not sufficient to control alien species, the strong linkage between native fish migration and flow pulses highlights the importance of river–floodplain connectivity for the conservation of native fishes.     

Habitat use of age 0 Alabama shad in the Pascagoula River drainage, USA

Mickle PF, Schaefer JF, Adams SB, Kreiser BR. Habitat use of age 0 Alabama shad in the Pascagoula River drainage, USA. Ecology of Freshwater Fish 2010: 19: 107–115. © 2009 John Wiley & Sons A/S
Abstract – Alabama shad ( Alosa alabamae ) is an anadromous species that spawns in Gulf of Mexico drainages and is a NOAA Fisheries Species of Concern. Habitat degradation and barriers to migration are considered contributing factors to range contraction that has left just the Pascagoula River drainage population in Mississippi. We studied juvenile life history and autecology in three rivers within the drainage. We collected fish, habitat and physicochemical data in three habitat types (sandbar, open channel and bank) from June to October 2004–2006. Sandbar habitat was favoured by smaller individuals early in the year. Catch per unit effort (CPUE) decreased through the summer as larger fish began occupying bank and open channel habitat. The most parsimonious model of abundance included year and river variables, while patterns of presence and absence were best explained by river, habitat type and physiochemical variables. While all three rivers in the drainage contained Alabama shad, fish were less abundant and had lower condition values in the Chickasawhay River. Earlier work suggested the Alabama shad may gradually move downstream towards the Gulf of Mexico in their first year. However, we found no evidence of this and captured large fish high in the drainage late in the year.     

First report of a cystic malformation on the upper jaw of hatchery‐reared allis shad Alosa alosa

The anadromous allis shad Alosa alosa has suffered dramatic population declines throughout Europe and is currently considered as endangered throughout its entire distribution range. In order to reestablish allis shad in the River Rhine, which formerly housed one of the largest and most important populations, an EU‐LIFE Project ‘The re‐introduction of allis shad in the Rhine system’ was started in 2007. In course of the LIFE+ Projects, allis shad larvae bred from genitor fish of the Gironde–Garonne–Dordogne population in France were reared in a pilot ex situ stock plant pilot facility in Aßlar, Germany. At an age of 1–2 months, about 100% of these fish developed approximately 0.5‐ to 0.8‐cm large, fluid‐filled, transparent cysts in conjunction with the upper jaw. The performed microbiological, virological, parasitological and histological examinations did not detect any infectious agents. Possible causative agents are discussed with regard to environmental factors and the nutrition of larvae. In conclusion, the observed malformations are considered a sign for a severe health problem and therefore a risk for the successful breeding of allis shad in aquaculture.     

Anadromy and the dispersal of an invasive fish species (Oncorhynchus mykiss) in Eastern Quebec,as revealed by otolith microchemistry

Thibault I, Hedger RD, Dodson JJ, Shiao J‐C, Iizuka Y, Tzeng W‐N. Anadromy and the dispersal of an invasive fish species (Oncorhynchus mykiss) in Eastern Quebec, as revealed by otolith microchemistry. Ecology of Freshwater Fish 2010: 19: 348–360. © 2010 John Wiley & Sons A/S Abstract – Rainbow trout (Oncorhynchus mykiss) is invading rivers bordering the St. Lawrence Estuary (Quebec, Canada). Some rivers in Eastern Quebec support self‐sustaining populations while adult vagrants are frequently captured in rivers where no reproduction has been confirmed. We hypothesised that the development of anadromy has promoted the species dispersal. Otolith Sr:Ca analyses revealed that although all fish captured in the upstream stocking region were freshwater residents, both anadromous and freshwater resident phenotypes were found downstream in Eastern Quebec. The proportion of fish exhibiting the anadromous life cycle increased with the distance from the stocking zone. Eastern Quebec steelhead migrated to sea at the same age but at a larger size than steelhead within their native range. Age at first reproduction was similar to that observed in native populations. The development of the anadromous life cycle enables this species to colonise new rivers following long‐distance migrations along the St. Lawrence Estuary corridor.     

Interannual variation in the coastal distribution of a juvenile gadid in the northeast Pacific Ocean: The relevance of wind and effect on recruitment

Drift of propagules occurs within many populations inhabiting flow fields. This affects the number of propagules that rejoin their source population (recruitment) and plays a role in adaptive spatial redistribution. We focus on the cause and consequence of interannual variation in geographic distribution of population density among five cohorts of young‐of‐the‐year (age‐0) juvenile walleye pollock Gadus chalcogrammus in the western Gulf of Alaska (GOA). The coastal GOA is a wind‐driven advective system. Walleye pollock spawn during spring and their eggs and larvae drift southwestward; by late summer, age‐0 juveniles are variously distributed over the shelf. We found that high population densities of age‐0 juveniles (ca. 6 months old) near the southwestward exit of the Alaska Coastal Current from the GOA corresponded with high abundance of larvae from the major spawning area upstream, but did not translate into high abundance at older ages. Further, offshore and upwelling‐favorable winds were associated with the high downstream abundance and presumed export. In contrast, downwelling‐favorable (northeasterly) wind during and shortly after spawning (April–May) was associated with high recruitment at age 1. Finally, we found that recruitment also increased with apparent retention of age‐0 juveniles in favorable habitat upstream near the main spawning area. We hypothesize that wind‐related retention in superior upstream habitat favors recruitment. Our results argue for including wind‐driven transport in future walleye pollock recruitment models. We encourage more work on the juvenile stage of marine fishes aimed at understanding how transport and species‐specific habitat suitability interact to affect population response to large‐scale forcing.     

Solar and lunar influences on the spawning activity of Pacific bluefin tuna (Thunnus orientalis) in the south‐western North Pacific spawning ground

Pacific bluefin tuna (Thunnus orientalis) migrate to the south‐western North Pacific spawning grounds (area off Taiwan and the Ryukyu Islands) from mid‐April to early July. Reproductive traits of the species in this area have been studied for some time; however, temporal changes in spawning activity during this season are not well understood. In this study, the spawning fraction (proportion of females with post‐ovulatory follicles to total mature females) in relation to solar calendar date and lunar day was investigated using a generalized linear model. Sampling was conducted almost every day throughout the fishing season at Ishigaki Fishing Port (24°21′N, 124°09′E), southern Japan between 2012 and 2014, and a total of 212 ovaries were histologically observed. Spawning fraction continued to increase from the beginning to the end of the season. This indicates that the peak season of occurrence in the area and the peak of spawning activity did not co‐occur. Spawning fraction also changed with the lunar day and significantly increased around the new moon. Eggs and early larvae hatched around the new moon may have reduced the predation risk during night‐time. These findings improve the current knowledge of reproductive biology and resource evaluation of Pacific bluefin tuna and indicate that spawning activity is not constant, and has two or three peaks in each season.     

Modelling riverine habitat for robust redhorse: assessment for reintroduction of an imperilled species

A critical component of a species reintroduction is assessment of contemporary habitat suitability. The robust redhorse, Moxostoma robustum (Cope), is an imperilled catostomid that occupies a restricted range in the south‐eastern USA. A remnant population persists downstream of Blewett Falls Dam, the terminal dam in the Pee Dee River, North Carolina. Reintroduction upstream of Blewett Falls Dam may promote long‐term survival of this population. Tillery Dam is the next hydroelectric facility upstream, which includes a 30 rkm lotic reach. Habitat suitability indices developed in the Pee Dee River were applied to model suitable habitat for proposed minimum flows downstream of Tillery Dam. Modelling results indicate that the Tillery reach provides suitable robust redhorse habitat, with spawning habitat more abundant than non‐spawning habitat. Sensitivity analyses suggested that suitable water depth and substrate were limiting physical habitat variables. These results can inform decisions on flow regulation and guide planning for reintroduction of the robust redhorse and other species.     

Induced Spawning of Wild American Shad Alosa sapidissima Using Sustained Administration of Gonadotropin-Releasing Hormone Analog (GnRHa)

American shad Alosa supidissima broodstock were collected from the Susquehanna River during their spawning migration. Mean volume of expressible milt (± standard deviation) was 2.5 (±1.7) mL/kg body weight; mean spermatozoid count was 66.2 ± 10⁹ (±163 ± 10⁹) spermntozoa/mL; and duration of 50% motility was 36.5 (±10.3) see. Ovarian biopsies indicated the presence of oocytes of various sizes (200–2,000 μm in diameter) and stages of development. Fish were implanted with a delivery system loaded with gonadotropin-releasing hormone analog (GnRHa) and started spawning 2 d after treatment. Fertile eggs were collected daily for the next 9 d, for a total of 50,100 eggs/kg body weight with a mean fertilization success of 62%. Upon cessation of spawning, the ovaries of all females still contained large numbers of oocytes at various stages of development, as at the beginning of the experiment, but with a greater number of atretic oacytes. Our observations show that American shad have an asynchronous ovarian development, and treatment with a GnRHa delivery system is effective in inducing several successive spawns of fertile eggs.     

Dispersal and tributary immigration by juvenile coho salmon contribute to spatial expansion during colonisation

Anadromous fishes are frequently restricted by artificial barriers to movement such as dams and culverts, so measuring dispersal helps identify sites where improved connectivity could promote range expansion and population viability. We used a combination of DNA‐based parentage analysis and mark–recapture techniques to evaluate dispersal by juvenile coho salmon (Oncorhynchus kisutch) in a population in the initial stages of colonisation following installation of fish passage structures at a previously impassable dam on the Cedar River, WA, USA. The spatial distribution of individuals within maternal families revealed that dispersal was common. Among the offspring of radio‐tagged mothers, 28% were collected outside the spawning reach and dispersed up to 6.3 km (median = 1.5 km). Most juveniles captured in a tributary (Rock Creek, where few adults spawned) had immigrated from the Cedar River and represented many different families. Juvenile dispersal therefore provided a secondary phase of spatial expansion following initial colonisation by adults. Consistent with the condition‐dependent dispersal hypothesis, juveniles that dispersed farther upstream in the tributary were larger than fish collected near the tributary mouth. Overall, the results demonstrated widespread dispersal in a system with low coho salmon densities, and this might increase the rate of population growth if it reduces the effects of local density dependence. By implication, juveniles can take advantage of rearing habitats reconnected through barrier removal, even when such areas are located several kilometres from adult breeding grounds.     

Stability in reproductive timing and habitat usage of Chinook salmon across six years of varying environmental conditions and abundance

The spatial and temporal distribution of spawning activity by autumn‐run Chinook salmon Oncorhynchus tshawytscha (Walbaum) was examined across multiple years. The study period included two years of extreme drought conditions when water temperatures in the spawning reach of the Stanislaus River were considered sub‐optimal for spawning and egg incubation. Despite varying levels of superimposition, redd counts and densities remained stable and positively associated with river location, indicating that superimposition may be driven by habitat preference rather than the absence of suitable spawning locations. Spawning occurred slightly later during drought years (6–10 days later compared with wetter years). This delay was attributable to deferred migration instead of deferred spawning, and the majority of redds were constructed at water temperatures exceeding the optimal temperature range. As a consequence, estimates of juvenile production during 2014 and 2015 were among the lowest on record. These findings may be related in part to the high hatchery contribution to the population, above‐average temperatures during spawning and incubation, and superimposition rates. Management recommendations include adequate cold‐water storage in the upstream reservoir, refined spawning habitat restoration techniques in the light of superimposition rates and, on a broader system scale, actions that reduce the amount of stray hatchery Chinook salmon.     

美洲鲥(Alosa sapidissima)眼睛早期发育的组织学观察

应用石蜡切片及HE染色的方法对美洲鲥(Alosa sapidissima)早期发育过程中眼睛的发生、分化和形成过程进行了系统的观察.结果显示,受精后21 h 13 min,视泡出现;受精后26h 23 min,视泡发育成视杯;受精后35 h 44 min,原始视网膜和晶状体形成;受精后60h 15 min,角膜在视网膜前缘处形成,由单一的立方上皮构成;2日龄仔鱼,巩膜分化完成,由软骨组织和弹力纤维构成,脉络膜出现;3日龄仔鱼,虹膜出现,视网膜分化完全,由外向内的色素层、视觉细胞层、外界膜、外核层、外网膜层、内核层、内网膜层、视神经节细胞层、视神经纤维层和内界膜10层构成,此时视觉细胞层中视杆细胞出现;14日龄仔鱼,角膜分化完全,由自外向内的复层扁平上皮、前弹性层、基质层、后弹性层和内皮层5层构成;22日龄仔鱼,脉络膜腺出现;26日龄稚鱼脉络膜分化完全,由外向内依次为银膜层、血管层和色素层,虹膜也完全分化,由外向内依次为虹膜内皮层、前缘层、基质层、后缘层和色素层;45日龄幼鱼,视网膜内核层分化成两层水平细胞.此时美洲鲥眼睛的各个结构均发育完善.     

Habitat suitability analysis for lacustrine brown trout (Salmo trutta) in Lake Walchensee,Germany: implications for the conservation of an endangered flagship species

• 1. The lacustrine brown trout (Salmo trutta) is endangered and of high conservation importance. In the only spawning habitat of the population in the Bavarian Lake Walchensee, the River Obernach, a substantial decrease in spawning runs has been reported. In this study, the present ecological state of the spawning stream was analysed with the objective of identifying life‐stage specific limitations to successful recruitment attributable to deficiencies in (i) spawning migration, (ii) spawning habitat quality, and (iii) habitat quality for juveniles.
• 2. Structural stream analysis showed that discharge and several migration barriers — particularly near the river outlet into the lake — prevent successful spawning migrations at normal water levels. Migration barriers are probably the main limiting factor for reproduction of lacustrine brown trout, whereas structural variability of the Obernach meets the habitat requirements of both spawners and juveniles.
• 3. Spawning site quality was suitable for trout, as indicated by stream substratum texture and high exchange rates between free‐flowing water and the interstitial zone in physico‐chemical parameters (redox potential, dissolved oxygen, pH, temperature and conductivity).
• 4. Analyses of fish community structure revealed dominance of lithophilic species, in particular of riverine brown trout (Salmo trutta). Its density and intact demographic population structure suggest that spawning and juvenile habitat quality for salmonids is not limiting. Recapture of stocked lacustrine trout juveniles also indicates habitat suitability for the juvenile stage.
• 5. In conclusion, the results show that the methodology used in this study is suitable for the identification of life‐stage specific habitat deficiencies in lacustrine brown trout and other fish species. Availability of habitat data throughout the species' distribution range is a first crucial step for the development of an effective recovery plan. Copyright © 2009 John Wiley & Sons, Ltd.