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Temporal and Spatial Variation

Temporal and Spatial Variation

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Temporal and spatial variation in the energy intake of a brook trout ( 
Salvelinus fontinalis
 )population in an Appalachian watershed
Ryan M. Utz and Kyle J. Hartman
Stream-dwelling salmonids in eastern North America are often restricted to headwater watersheds, whereproductivity is low and thus feeding conditions are poor. We sought to quantify how energy intake varies with spatialand temporal variation by monitoring feeding rates in multiple sites over the course of two years. Daily rations werecalculated for 939 fish by examining stomach contents. Maintenance rations were compared with daily rations using abioenergetics model. Consumption peaked in spring, dropped substantially in summer, and remained low until the fol-lowing spring. A minority of fish fed at very high levels during all seasons, elevating the mean consumption of thepopulation. Fish occupying large sites with low trout densities consistently consumed more energy than fish in smallerstreams with high trout densities. A direct relationship between trout density and mean consumption was observed dur-ing summer, when feeding conditions were poorest. Our findings suggest that within a headwater watershed, largerreaches of streams where fewer trout are found act as important feeding areas and thus may be important habitat forbrook trout (
Salvelinus fontinalis
Résumé :
Les salmonidés des cours d’eau dans l’est de l’Amérique du nord sont souvent confinés aux bassins versantsd’amont où la productivité est faible et conséquemment les conditions alimentaires médiocres. Nous avons cherché àmesurer les variations spatiales et temporelles de l’ingestion d’énergie en suivant les taux d’alimentation à plusieurs si-tes durant deux années. L’analyse des contenus stomacaux de 939 poissons a permis de calculer les rations journalières.Un modèle bioénergétique a comparé les rations journalières aux rations de maintien. La consommation atteint unmaximum au printemps, diminue considérablement en été et reste faible jusqu’au printemps suivant. Un minorité depoissons se nourrit à un taux élevé en toutes saisons, ce qui augmente la consommation moyenne de la population. Lespoissons habitant de grands sites à faibles densités d’ombles consomment toujours plus d’énergie que les poissons dansles petits cours d’eau à forte densité d’ombles. On observe une relation directe entre la densité des ombles et laconsommation moyenne en été, quand les conditions d’alimentation sont les moins bonnes. Nos observations indiquentque, dans un bassin versant d’amont, les grandes sections habitées par un nombre réduit d’ombles sont des lieux im-portants d’alimentation et peuvent donc être un habitat vital pour l’omble de fontaine (
Salvelinus fontinalis
).[Traduit par la Rédaction]
Utz and Hartman2686
Brook trout (
Salvelinus fontinalis
) populations have de-clined in size and range as the result of a host of historic andpersistent anthropogenic impacts, including uncontrolledlogging, acidic precipitation, mine runoff, overfishing, andintroduced species (Larson and Moore 1985; Marschall andCrowder 1996; Wigington et al. 1996). A number of institu-tions consider the restoration of brook trout a managementgoal because of its potential value as a recreational fishery.Recent work has highlighted the importance of smallstreams with high alkalinity and low levels of fine sedimentin maintaining brook trout reproduction and populations(Petty et al. 2005; Hartman and Hakala 2006). However, be-cause of the low productivity of the watersheds that brook trout often occupy in Appalachia and elsewhere, factors otherthan reproductive success may be important in sustainingand expanding existing populations.Throughout eastern North America, brook trout are oftenrestricted to headwater watersheds that are naturally infer-tile. As a result, trout in these watersheds may be subjectedto prolonged periods of poor feeding conditions (Cada et al.1987; Ensign et al. 1990; Forrester et al. 1994), and foodabundance may affect trout production (Clarke and Scruton1999). Such episodes of reduced feeding usually occur dur-ing summer when aquatic insect activity is relatively low,although other seasons may present poor foraging opportuni-ties (Sweka and Hartman 2001). Although poor feeding has
Can. J. Fish. Aquat. Sci.
: 2675–2686(2006) doi:10.1139/F06-152 © 2006 NRC Canada
Received 22 February 2006. Accepted 2 August 2006. Published on the NRC Research Press Web site at http://cjfas.nrc.ca on25 November 2006.J19187
R.M. Utz
and K.J. Hartman.
WVU Wildlife and Fisheries Resources Program, Division of Forestry and Natural Resources,Davis College of Agriculture, Forestry, and Consumer Sciences, 320 Percival Hall, Morgantown, WV 26506-6125, USA.
Corresponding author (e-mail: rutz@al.umces.edu).
Present address: University of Maryland Appalachian Laboratory, 301 Braddock Road, Frostburg, MD 21532, USA.
not been shown to reduce populations under normal condi-tions, severe summer drought may cause environmental con-ditions to deteriorate (Hakala and Hartman 2004) along withdecreasing foraging rates, potentially resulting in mortality.A growing body of evidence suggests that lotic salmonidsoperate under density-dependent processes, which has impli-cations for feeding success. Fish density has been shown toaffect growth (Vøllestad et al. 2002; Imre et al. 2004;Lobón-Cerviá 2005) in that fish occupying reaches of streams with less fish per unit area experienced significantlyhigher growth rates in both natural and simulated settings.The territorial nature of salmonids likely drives this relation-ship. Density has been found to be positively correlated withemigration and mortality as a result of competitive interac-tions (Nakano 1995; Imre et al. 2004). Multiple studies haveconcluded that brook trout may be highly mobile (Gowanand Fausch 1996; Petty et al. 2005). The ability to movecoupled with the poor nature of high-density locations sug-gests that movement into low-density areas should be benefi-cial. Large adult brook trout have been shown to preferlarger reaches of streams where fish density is typically low(Petty et al. 2005). However, observational data pertaining toenergy intake at the watershed scale for stream salmonids re-main sparse in the literature.The concept that feeding intensity may be related to spa-tial variables has been established for salmonids with thegeneral conclusion that larger bodies of water offer greaterfeeding potential (Keeley and Grant 2001). However, the ex-tent at which this relationship exists within small watershedsremains to be described with detail. The relationship be-tween density, watershed position, and feeding intensity maybe of particular importance in eastern brook trout streamswhere food resources are often sparse. An understanding of temporal change in feeding intensity at the watershed scalemay provide insight into how brook trout use differentreaches of habitat to acquire energy. The recent developmentof a bioenergetics model for brook trout (Hartman and Sweka2001) facilitates the calculations of a number of feedingvariables, including daily ration and maintenance ration. Inthe current study, we sought to explain how consumptionchanges temporally and spatially by monitoring the ener-getic intake of a brook trout population throughout a head-water watershed over the course of two years.
Materials and methods
Study area
This study was conducted within the Middle Fork water-shed, a north-flowing tributary of the Tygart River in thecentral Appalachian Mountains of Randolph County, WestVirginia (Fig. 1). Most land cover within the watershed isthat of secondary-growth hardwood deciduous forest. Allsites in the study are located in the southernmost part of thewatershed and are of low order and high gradient. We chosenine 200 m sites encompassing a range of stream sizes (Ta-ble 1). All sites met the following criteria: each containedbrook trout, consistently held age-0 brook trout (indicatingwater quality was sufficient for spawning), and was devoidof fish barriers between other sites. Temperature regimens inthese reaches are suitable for trout as temperatures rarely ex-ceeded 20 °C through the duration of the study, determinedby analysis of temperature logger data conducted for thecurrent and previous studies. The West Virginia Division of Natural Resources (WVDNR) and the West Virginia Depart-ment of Environmental Protection have actively added lime-stone sand to riparian areas of streams within the watershedto remediate the effects of acid precipitation and acid minedrainage in the Middle Fork since the 1990s (WVDNR2001). This process is commonly used in the region and suc-cessfully increases pH, restores fish communities, and in-creases invertebrate abundance (Clayton et al. 1998), thoughinvertebrate productivity and diversity may not reach pre-acidification levels following treatment (McClurg 2004).Some sites selected in this study were not actively treatedwith lime. However, each site without a limestone treatmentretained the ability to support brook trout spawning andcarry fish populations. To quantify water quality in eachsite, water samples were taken in February, May, July, andOctober 2004 and tested for pH and alkalinity in a labora-tory at the WVDNR office in Elkins, West Virginia. Themeans of both measurements were calculated for each sitebased on the four seasonal measurements; this calculationprovided an estimate of average water quality across sea-sons.Fish diversity differed across sites but was typical of Ap-palachian headwater streams. The number of species en-countered increased with stream size. In nearly all sites, fishfauna was dominated by brook trout and mottled sculpin(
Cottus bairdii
). Other fish sampled include blacknose dace
© 2006 NRC Canada
2676 Can. J. Fish. Aquat. Sci. Vol. 63, 2006
Fig. 1.
The upper Middle Fork watershed and all sites describedfor this study.
 Rhynichthys obtusus
), longnose dace (
 Rhynichthys cata-ractae
), creek chub (
Semotilus atromaculatus
), white sucker(
Catostomus commersonii
), northern hog sucker (
 Hypen-telium nigricans
), and fantail darter (
 Etheostoma flabellare
Fish and stream size sampling
Fish sampling occurred 11 times over the course of twoyears (Table 2). Sampling was conducted between 0800 and1600, and the order of sites to sample was randomly chosenduring each day. A three-pass depletion electrofishing proce-dure (Platts and Nelson 1988) was used to estimate fish pop-ulations within each 200 m section. Before sampling, block nets were placed at the upstream and downstream ends of each section to restrict fish movement into or out of sectionsduring sampling. Sampling teams used an electrofishing unit(DC, 60 hz, 500–750 V; Smith-Root, Vancouver, Washing-ton) and dip nets to capture fish. Temperatures needed forbioenergetics modeling were recorded during fish samplingusing an electronic thermometer.Following collection, fish were processed at a streamsidestation. All fish were immobilized with a solution of cloveoil and 95% ethanol. Brook trout were weighed to the near-est 0.5 g, and total length was taken to the nearest milli-metre. We chose a subset of 10 brook trout per site permonth for stomach content removal. Only fish > 110 mmfork length (FL) were considered eligible for gut content re-moval because of gear restrictions; the gape of fish belowthis size was usually about the same size of the flushing tubediameter (7 mm). This size class generally represented age-1+ fish, which was apparent in length-frequency histograms(Utz 2005). An attempt was made to collect an equal rangeof fish sizes to analyze for gut contents at each site; asidefrom this, fish were chosen randomly. Stomach contentswere removed by directing a constant flow of stream waterinto the foregut (Twomey and Giller 1990). Gut items werefiltered through a 250
msieve and transferred to 95% etha-nol, a process that has proven effective (Light et al. 1983;Sweka 2003). All fish other than trout were counted andweighed. Each fish was allowed to recover from the cloveoil treatment and returned to the stream reach within 2 h. Asample of 10 trout, selected for gastric lavage, was randomlycollected from sites, frozen, and kept for dry weight analysisduring four select sampling periods. This resulted in a re-moval of 40 fish during the entire course of the study. Brook trout dry weights were needed to estimate fish energy den-sity, a necessary component of bioenergetics modeling.To determine if fish density was related to consumption atthe watershed scale, select habitat variables were calculatedat each site. Habitat measurements within each site were re-corded during base flow on 17 August 2005. Variables weremeasured on each 200 m reach of stream. Habitat unitswithin each site were classified as riffle or pool, and thelength of each unit was measured. Within each habitat unit,the wetted width and bankful width were taken along alinear transect perpendicular to the stream. The number of transects measured within a habitat unit depended on thelength of the unit, with measurements made approximatelyevery 20 m in riffles and every 2 m in pools. Pools weremore represented than riffles in mean width estimation be-cause of the tendency for large adults to use this habitat;however, riffles and pools did not differ markedly in widths.
© 2006 NRC Canada
Utz and Hartman 2677
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