Mercury toxicity in livers of northern pike (Esox lucius) from Isle Royale, USA

Many laboratory studies have documented that mercury can be toxic to fish, but it is largely unknown if mercury is toxic to fish in their natural environments. The objective of our study was to investigate the toxic effects of mercury on northern pike (Esox lucius) at Isle Royale, Michigan. In 124 northern pike from eight inland lakes, concentrations of total mercury in skin-on fillets ranged from 0.069 to 0.622 microg/g wet mass (wet wt). Concentrations of total mercury in livers increased exponentially compared with concentrations in fillets, to a maximum of 3.1 microg/g wet wt. Methylmercury constituted a majority of the mercury in livers with total mercury concentrations <0.5 microg/g wet wt, but declined to 28-51% of the mercury in livers with total mercury concentrations >0.5 microg/g wet wt. Liver color (absorbance at 400 nm) varied among northern pike and was positively related to liver total mercury concentration. The pigment causing variation in liver color was identified as lipofuscin, which results from lipid peroxidation of membranous organelles. An analysis of covariance revealed lipofuscin accumulation was primarily associated with mercury exposure, and this association obscured any normal accumulation from aging. We also documented decreased lipid reserves in livers and poor condition factors of northern pike with high liver total mercury concentrations. Our results suggest (i) northern pike at Isle Royale are experiencing toxicity at concentrations of total mercury common for northern pike and other piscivorous fish elsewhere in North America and (ii) liver color may be useful for indicating mercury exposure and effects in northern pike at Isle Royale and possibly other aquatic ecosystems and other fish species.     

Dynamics of a Boreal Lake Ecosystem during a Long-Term Manipulation of Top Predators

We assessed the long-term (16 years) effects of introducing piscivores (northern pike) into a small, boreal lake (Lake 221, Experimental Lakes Area) containing abundant populations of two planktivorous fish species. After the introduction, pearl dace were extirpated and yellow perch abundance was greatly reduced. Daphnia species shifted from D. galeata mendota to larger bodied Daphnia catawba, but the total zooplankton biomass did not increase, nor did the biomass of large grazers such as Daphnia. Phytoplankton biomass decreased after the northern pike introduction, but increased when northern pike were partially removed from the lake. Phosphorus (P) excretion by fish was ∼0.18 mg P m⁻² d⁻¹ before pike addition, declined rapidly to approximately 0.03–0.10 as planktivorous perch and dace populations were reduced by pike, and increased back to premanipulation levels after the pike were partially removed and the perch population recovered. When perch were abundant, P excretion by fish supported about 30% of the P demand by primary producers, decreasing to 6–14% when pike were abundant. Changes in phytoplankton abundance in Lake 221 appear to be driven by changes in P cycling by yellow perch, whose abundance was controlled by the addition and removal of pike. These results confirm the role of nutrient cycling in mediating trophic cascades and are consistent with previous enclosure experiments conducted in the same lake.     

Consumption of crucian carp (Carassius carassius L., 1758) by restocked pike (Esox lucius L., 1758) in a lake with frequent winter hypoxia

To assess the effects of stocked pike (Esox lucius L.) on crucian carp (Carassius carassius L.) biomasses, the annual consumption of pike was estimated and compared with removal fishery catches. The studied lake, Lake Savijärvi, is a small (40 ha), shallow and eutrophicated lake in southern Finland with frequent algal blooms during summers and fish kills during winters. Until the 1980s, the fish fauna consisted of pike, perch (Perca fluviatilis L.), roach [Rutilus rutilus (L.)], crucian carp, and tench (Tinca tinca L.). Since 2003, when crucian carp were abundant in the extreme and with only a few roach and tench individuals, the lake has been biomanipulated by removal seining. To enhance the effects of seining, piscivorous pike were restocked in the spring of 2008. During the 3‐year study, the catch of crucian carp decreased from 243 to 136 kg while the catch of pike increased from 0.1 to 5.7 kg per seining hectare due to their spawning in the lake as of 2009. The total crucian carp consumption by captured pike was 588 kg during the 2008–2010 study period. When applying the number of pike estimated with the mark‐recapture method, consumption estimate increased to 917 kg, or 22.9 kg per lake hectare in 2010. Thus, pike consumption of crucian carp was about 17% of the removal catch in 2010.     

What do the empty stomachs of northern pike (<Emphasis Type="Italic">Esox lucius</Emphasis>) reveal? Insights from carbon (δ<Superscript>13</Superscript>C) and nitrogen (δ<Superscript>15</Superscript>N) stable isotopes

The frequency of individuals with empty stomachs (FES) can vary greatly among northern pike populations. However, the FES has only seldom been analyzed in this species and its meaning is still not fully understood. It has been suggested that a high FES may reflect a strongly piscivorous behaviour while low FES could reflect a higher utilization of invertebrates. We compared the stomach contents and the trophic position of northern pike in 16 populations of individuals feeding mainly on fish or benthic invertebrates. We tested the hypothesis that northern pike with empty stomachs or with fish in their stomachs have a higher trophic position than individuals feeding on invertebrates. Carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope signatures were used to estimate the trophic position of individuals. We found no significant difference in the trophic position among piscivores, invertebrate feeders, and northern pike with empty stomachs. The average trophic position of northern pike was high (mean ± SD = 4.3 ± 0.4, n = 66) and was correlated with total length. These results indicate that, although invertebrates could be an important part of the diet of northern pike in Canadian Shield lakes, fish are still the dominant prey. Hence, feeding on invertebrates in our study lakes would reflect an opportunistic rather than a specialized feeding strategy.     

Individual specialization and trophic adaptability of northern pike (Esox lucius): an isotope and dietary analysis

Northern pike (Esox lucius) are often considered to be specialist piscivores, but under some circumstances will continue to eat invertebrates as adults. To examine effects of fish assemblage composition on the trophic ecology of pike, we combined stable isotope analysis (SIA) of carbon and nitrogen and stomach content analysis (SCA) on pike from five lakes in northern Alberta, three of which contain only pike (“pike-only”) and two that also contain yellow perch (Perca flavescens) or white sucker (Catostomus commersoni) (“pike-other”). Fish were more important as prey and empty stomachs, which often characterize piscivores, were significantly more frequent in pike-other than in pike-only lakes. However, even though invertebrates were more important for pike in pike-only lakes, SIA and SCA indicated that invertebrates were also an important component of pike diets in pike-other lakes. SIA and SCA also revealed considerable intrapopulation variation in trophic ecology, with individuals in some populations differing by as much as two trophic levels. Comparisons of stomach contents and isotope signatures of the same fish suggested that within these variable populations, specialization on invertebrates or fish was a long-term trait of some individuals. SIA indicated that trophic position increased and diets shifted to a greater importance of littoral prey as pike grew in pike-only lakes, but not in lakes with other fish present. Trophic adaptability in northern pike is expressed at both the population level, where the trophic ecology is sensitive to differences in prey regimes, and at the organismal level, in the form of intrapopulation variation and individual specialization. Received: 1 July 1998 / Accepted: 3 February 1999     

Mercury-binding capacity of organic and inorganic selenium in rat blood and liver

The mercury-binding capacity of seleno-DL-methionine and selenium dioxide was assessed in male Wistar rats. Mercury was supplied as fish loaves made of northern pike or rainbow trout. We used a selenium concentration of 3.4 mg/kg fish, about sixfold compared to the equivalent quantity of mercury. Seleno-DL-methionine had a tendency to increase both methyl mercury and total mercury in blood, although it also seemed to reduce the proportion of methyl mercury of total mercury. Selenium dioxide lowered mercury levels by 24–29% both in the blood and in the liver of rats that were fed with northern pike.     

Fish mercury levels in relation to characteristics of hydroelectric reservoirs in Newfoundland, Canada

Mercury levels in fish have been demonstrated to increase after impoundment with augmented levels of mercury predicted to decline as the reservoir ages. Previous research in Newfoundland predicted return rates in the order of 10 to 12 years for landlocked Atlantic salmon or ouananiche (Salmo salar) and 7 years for brook trout (Salvelinus fontinalis). In order to test the validity of these predictions on a broader spatial and temporal scale, and develop more generally predictive models, mercury levels in three fish species were studied in 16 older Newfoundland hydroelectric reservoirs of various age (32 to 95 years) and area flooded (21 to 13,000 ha).Mercury concentrations were standardized to fish length and correlated with physical, chemical, and biological characteristics of the sampling sites. Standard length mercury levels ranged from 0.23 to 0.86 ppm in ouananiche, 0.13 to 0.59 ppm in brook trout, and 0.22 to 0.72 in arctic charr (Salvelinus alpinus). Fish in excess of the Canadian Safety Limit (0.5 ppm) were collected from 14 of 16 sites for ouananiche, 8 of 17 sites for brook trout, and 3 of 7 sites for arctic charr, including control lakes. Standard length fish mercury levels were correlated with reservoir age and (log₁₀) area flooded for ouananiche and with pH for arctic charr. A multiple regression model was developed relating standard length mercury in ouananiche with reservoir age and log₁₀ of the flooded area. There were no apparent relationship between reservoir characteristics and brook trout mercury concentrations. Based on this analysis, it is not possible, at present, to develop generally predictive models for all species found in Newfoundland impoundments.     

Comparison of dietary mercury exposure in two sympatric top predator fishes, largemouth bass and northern pike: a bioenergetics modeling approach

Physical and ecologicalfactors, including lake temperature, fishphysiology, and diet, influence methylmercury(MeHg) exposure in fish. We employedbioenergetics modeling to compare dietary MeHgexposure in sympatric top predators, largemouthbass (Micropterus salmoides) and northernpike (Esox lucius). We comparedsimulations using field data to hypotheticalsimulations with (1) ± 25% change in meandaily lake temperature for juvenile and adultbass and pike; (2) ± 25% change inlong-term growth rate of pike; (3) adult bassdiet shift from generalist predator to strictpiscivore. Bass and pike MeHg exposures weresimilar in baseline simulations and reflectedpatterns in field tissue concentrations. Thisoccurred despite the fact that bass consumedhighly contaminated benthic invertebrates,while pike exclusively consumed lesscontaminated fish prey. Higher temperaturesincreased adult bass and pike MeHg exposures by35% and 27%, respectively. Shifting adultbass diets to 100% fish resulted in a 54%decrease in exposure, while increasing pikegrowth rates resulted in a 24% decrease. Bioenergetics modeling proved useful inunderstanding the influence of temperature,prey-base, and predator growth on differencesin Hg exposure across fish species.     

Length–weight relationships of four freshwater cyprinid species from a tributary of Ganga River Basin in North India

This paper provides length‐weight relationships (LWRs) for four freshwater cyprinid fish species, namely Barilius bendelisis (Hamilton, 1807), Bangana dero (Hamilton, 1807), Chagunius chagunio (Hamilton, 1807) and Labeo dyocheilus (McClelland, 1839) from the Kosi River, a tributary of the River Ganga in northern India. Altogether 848 specimens were caught bimonthly (for B. bendelisis) and quarterly for the other three species between March 2013 and December 2014 using cast nets (9 m length, 9 m width and 1/2 cm mesh size). LWRs for two of the four species (B. dero and C. chagunio) are recorded for the first time. The present study results will be useful for fishery researchers, management and conservation in this least‐explored tributary of the Ganga River Basin.     

Production and outmigration of young-of-year northern pike Esox lucius from natural and modified waterways connected to Lower Green Bay,Wisconsin

Production and outmigration of young-of-year (YOY) northern pike from natal sites in Lower Green Bay, WI, USA, were documented over three consecutive years (2013–2015). We tested the hypothesis that spawning success and outmigration characteristics of YOY northern pike would vary among natural and anthropogenically modified habitats. Sixteen focal study locations were surveyed, including a restored natural wetland, agricultural drainage ditches, a flooded forested wetland and several unimpounded tributaries. We collected 1469 YOY northern pike with most individuals (N = 1163) originating from a flooded forested wetland on the east shore. Most sites produced YOY in all years (range N = 2–1145 individuals among study years). Outmigration ranged between 1 and 40 days during 2013–2015. Greater production and extended outmigration times occurred at most sites in 2014 (range 17–40 days) when the region experienced a late spring with heavy precipitation. In contrast, the lowest production and shortest outmigration period occurred at most sites in 2015 (range 14–23 days) when environmental conditions reflected regional averages. Outmigration began nearly 3 weeks earlier in 2015 (5/8) than in other study years (8 June 2013 and 25 May 2014). Total length (TL) of outmigrating northern pike ranged between 17 and 138 mm. Total length of YOY was significantly different among sites in 2013 and 2014, with the smallest fish (17 mm TL) outmigrating from agricultural ditches in both years. There were no significant variations in size among sites in 2015 (range 21–95 mm TL). Our results indicate significant variation in YOY northern pike outmigration characteristics within Lower Green Bay that may reflect the interplay between adult spawning site selection and annual weather patterns. Our findings highlight the importance of quantifying overlooked habitats in regions of mixed development.     

Development of microsatellite markers for muskellunge (Esox masquinongy) and cross‐species amplification in two other esocids

We report the development of seven polymorphic microsatellite loci in muskellunge (Esox masquinongy) using an unenriched subgenomic library. Polymorphic loci exhibited 2–11 alleles with observed heterozygosities ranging from 0.190 to 0.917 (n = 24). All seven loci amplified by their respective primer pairs resulted in monomorphic products in northern pike (E. lucius) whereas three loci amplified monomorphic products in grass pickerel (E. americanus americanus); these results imply conservation of flanking sequence but loss of polymorphism between these closely related species. Only one of six microsatellite primers developed in a previous study in northern pike amplified polymorphic products in muskellunge.     

Metals (Hg,Pb, Cu,and Zn) Bioaccumulation in Sediment,Fish, and Human Scalp Hair: A Case Study from the City of Mersin Along the Southern Coast of Turkey

This study investigates mercury, lead, copper, and zinc concentrations in six most frequently consumed fish species (120 samples), sediments (20 samples) taken from Karaduvar Fishing Area where fish species live, and Mersin Port as a contrary region, and human scalp hair for people regularly consuming these fish species (50 samples) and non-fish-eaters (15 samples) in Mersin, Turkey. On taking living environment into account, the fish groups include pelagic species of Liza saliens, Liza aurata, and demersal species of Merluccius merluccius, Mullus barbatus, Upeneus moluccensis, and Solea solea. Total Hg (THg) was found to accumulate in muscle tissues at the lowest concentration (0.01 μg/g) in L. saliens and at the highest (2.66 μg/g) in S. solea. Pb was only detected at high concentrations of 1.86 μg/g in M. barbatus and of 2.16 μg/g in M. merluccius. Cu and Zn concentrations were below the detection limit within all fish species. In the sediment samples, Pb and Cu concentrations were persistently below their effect range–median (ERM) value, whereas this limiting value only maintained for 15% of THg concentrations. On the other hand, the effect range–low (ERL) of sediment exceeded at Pb in 15% of samples and Cu in 25% of samples. Zn remained below the detection limit for sediment samples. The metal concentrations at scalp hairs of regular consumers of these fish groups and non-fish eaters vary from the range 0.40–3.28 to 0.14–1.02 μg/g for THg, 11.16–107.84 to 8.00–22.38 μg/g for Pb, and 151.67–645.35 to 144.92–343.50 μg/g for Zn. An important finding of the present study is the significant adverse impact of sedimentary heavy metal bioaccumulation to human through the consumption of demersal fishes in the city of Mersin along the southern coast of Turkey.     

Restoration by biomanipulation in a small hypertrophic lake: first-year results

Biomanipulation was carried out in order to improve the water quality of the small hypertrophic Lake Zwemlust (1.5 ha; mean depth 1.5 m). In March 1987 the lake was drained to facilitate the elimination of fish. Fish populations were dominated by planktivorous and benthivorous species (total stock c. 1500 kg) and were collected by seine- and electro-fishing. The lake was subsequently re-stocked with 1500 northern pike fingerlings (Esox lucius L.) and a low density of adult rudd (Scardinius erythrophthalmus). The offspring of the rudd served as food for the predator pike. Stacks of Salix twigs, roots of Nuphar lutea and plantlets of Chara globularis were brought in as refuge and spawning grounds for the pike, as well as shelter for the zooplankton.The impact of this biomanipulation on the light penetration, phytoplankton density, macrophytes, zooplankton and fish communities and on nutrient concentrations was monitored from March 1987 onwards. This paper presents the results in the first year after biomanipulation.The abundance of phytoplankton in the first summer (1987) after this biomanipulation was very low, and consequently accompanied by increase of Secchi-disc transparency and drastic decline of chlorophyll a concentration.The submerged vegetation remained scarce, with only 5 % of the bottom covered by macrophytes at the end of the season.Zooplankters became more abundant and there was a shift from rotifers to cladocerans, comprised mainly of Daphnia and Bosmina species, the former including at least 3 species.The offspring of the stocked rudd was present in the lake from the end of August 1987. Only 19% of the stocked pike survived the first year.Bioassays and experiments with zooplankton community grazing showed that the grazing pressure imposed by the zooplankton community was able to keep chlorophyll a concentrations and algal abundance to low levels, even in the presence of very high concentrations of inorganic N and P. The total nutrient level increased after biomanipulation, probably due to increased release from the sediment by bioturbation, the biomass of chironomids being high.At the end of 1987 Lake Zwemlust was still in an unstable stage. A new fish population dominated by piscivores, intended to control the planktivorous and benthivorous fish, and the submerged macrophytes did not yet stabilize.     

Selenium and mercury concentrations in some fish species of the Madeira River, Amazon Basin, Brazil

Samples of 7 species of piscivorous, omnivorous, and herbivorous fish caught at 12 different sites on the Madeira River, Amazon Basin, were analyzed for selenium and mercury. Selenium was determined by anodic stripping voltammetry and mercury by cold vapor atomic absorption spectrophotometry. The means for selenium concentrations ranged from 0.49 to 3.11 nmol/g and for mercury from 0.41 to 6.66 nmol/g depending on the fish species. The molar ratios of Hg:Se increased according to the fish trophic level. Piscivorous species had the highest mean ratio (4.0) and herbivorous species the lowest (0.9). There was a positive and statistically significant correlation between selenium and mercury concentrations for the herbivorous species (r = 0.716;p = 0.0088) not seen for omnivororus and piscivorous species (r = -0.2032;p = 0.3407). These findings are significant for the fish-eating population of the Madeira River because the ingestion of mercury would always be in excess of selenium.     

Walleye, Stizostedion vitreum, and northern pike, Esox lucius, populations in three Alberta lakes

Results are presented from a survey of walleye and northern pike in Ethel, Marie and Wolf Lakes, Alberta conducted May-September, 1983. Three types of experimental gill net were fished, mainly inshore. Northern pike were abundant in all three lakes but white sucker and yellow perch also were common. Walleye were caught in largest numbers in Wolf Lake. Age and growth of both walleye and northern pike were determined from examination of the opercular bones. The walleye in the three lakes were represented by a few strong year classes but there was no synchronism between the lakes. The strong 1975 year class in Ethel and Wolf Lakes may have been correlated with high spring precipitation and heavy run-off. Although there were dominant year classes in the northern pike populations (especially the 1979 year class) the variability was not so marked as in the walleye. There were no significant differences between length-at-age for year classes of walleye in Ethel and Marie or northern pike in all three lakes. In Wolf Lake the 1978 and 1979 year classes grew faster than in previous years. Walleye and northern pike growth was well described by the von Bertalanffy growth model. Females of both species grew to a greater ultimate length and had a longer lifespan than the males. Fecundity data are presented for walleye from Wolf Lake (log₁₀ absolute fecundity = 0.856 + 2.441 log₁₀ length) and northern pike from Marie Lake (log₁₀ absolute fecundity =?2.671+4.052 log₁₀ length). The stomach contents of both species were examined during May and June. The majority of walleye had empty stomachs. Walleye and northern pike in Ethel and Wolf Lakes fed on a variety of fish and invertebrate species but both fed mainly on fish (in particular whitefish) in Marie Lake. The similarity of diet suggests competition for prey species. A walleye yield to commercial fishermen of c. 0.69 kg ha^?1 year^?1 in Wolf Lake has made up 8% of the total catch. Since 1970/71 when mesh size of nets was increased, the yield has been reduced to c. 0.30 kg ha^?1 year^?1. Anglers remove an additional c. 0.48 kg ha^?1 year^?1 walleye from Wolf Lake. Northern pike yields to commercial fishermen have been c. 2.00 kg ha^?1 year^?1 in Wolf Lake. Ethel and Marie Lakes have yielded only poor catches of walleye and northern pike. A yield model was used to illustrate that faster-growing northern pike have higher potential yields than walleye. Walleye produce higher yields in Wolf Lake than in Marie Lake, the reverse being true for northern pike. It is suggested that northern pike could be cropped at a higher rate in Marie and Wolf Lakes with a possible improvement in walleye stocks.     

Food consumption by fish populations in the Warta River,Poland, before and after impoundment

The impact of impoundment on the energy budget and gross ecological efficiency (K ₁) of fish populations in sites upstream from the backwater (AB) and downstream of the dam (CD) of a manmade reservoir (17.6 km²) located in the middle reaches of the lowland Warta River (808 km long), was investigated.Consumption (MJ ha^–1 yr^-1) in four years following the reservoir's impoundment was 34–74% lower in site AB, and 43–75% in site CD than before the impoundment. K ₁ of investigated species was higher before the impoundment and only pike exhibited an inverse efficiency trend. Significant differences were also observed in the diet of certain dominant fish species, which resulted from a change in the food supply as well as a reduction in their life span after the river was dammed.     

Mercury resistance and volatilization by oil utilizing haloarchaea under hypersaline conditions

The hydrocarbon utilizing haloarchaea, Haloferax (two strains), Halobacterium and Halococcus from a hypersaline coastal area of the Arabian Gulf, had the potential for resistance and volatilization of Hg²⁺. Individual haloarchaea resisted up to between 100 and 200 ppm HgCl₂ in hydrocarbon free media with salinities between 1 and 4 M NaCl, but only up to between 20 and 30 ppm in a mineral medium containing 3 M NaCl, with 0.5% (w/v) crude oil, as a sole source of carbon and energy. Halococcus and Halobacterium volatilized more mercury than Haloferax. The individual haloarchaea consumed more crude oil in the presence of 3 M NaCl than in the presence of 2 M NaCl. At both salinities, increasing the HgCl₂ concentration in the medium from 0 to 20 ppm resulted in decreasing the oil consumption values by the individual haloarchaea. However, satisfactory oil consumption still occurred in the presence of 10 ppm HgCl₂. It was concluded that haloarchaea with the combined potential for mercury resistance and volatilization and hydrocarbon consumption could be useful in removing toxic mercury forms effectively from oil free, mercury contaminated, hypersaline environments, and mercury and oil, albeit less effectively, from oily hypersaline environments.     

Mercury in waterfowl from a contaminated river in Virginia

Many bodies of water around the world are contaminated with mercury from historic industrial and mining activities or ongoing atmospheric deposition, resulting in numerous fish consumption advisories. However, concerns about mercury have only rarely led to consumption advisories on waterfowl. In contrast with fish, waterfowl frequently disperse long distances to new watersheds, so hunters and wildlife managers do not know whether waterfowl at a pristine site have spent time at a contaminated site elsewhere. We sampled tissue mercury concentrations of mallards (Anas platyrhynchos), wood ducks (Aix sponsa), and Canada geese (Branta canadensis) at a site contaminated with mercury, during the breeding and hunting seasons. We found that many mallards had bioaccumulated mercury to levels that had the potential to produce reproductive effects and exceeded consumption advisories set for fish by regulatory agencies, whereas this was true for only a few wood ducks and Canada geese. We also documented that mercury-exposed waterfowl from this contaminated site were harvested by hunters as far as 1,054 km away. Our results suggest the need for more proactive sampling of waterfowl for mercury, and likely other bioaccumulating contaminants, in order to allow hunters to make more informed choices about consumption of their harvest. © 2012 The Wildlife Society.     

Invasive pike establishment in Cook Inlet Basin lakes,Alaska: diet,native fish abundance and lake environment

Northern pike (Esox lucius) were introduced to the northern Susitna Basin of south-central Alaska in the 1950’s, and have since spread throughout the upper Cook Inlet Basin. Extirpations of several native fish populations have been documented in this area. It is hypothesized here that invasive pike remodel the ecology of lakes by removing vulnerable prey types and that these changes are reflected in the diet of invasive pike. Trends in pike diet suggest that pike switch to less desirable but more abundant macroinvertebrate prey as preferred fish prey are eliminated. The impacts of pike introduction were studied in detail for one species of resident fish, the threespine stickleback (Gasterosteus aculeatus). Stickleback abundance decreases as pike invasion progresses. Conductivity is a significant environmental predictor of stickleback abundance, with higher conductivity apparently mitigating population reduction. Higher conductivity water may lessen the physiological costs of developing more robust armor, which reduces vulnerability to predation. Maximum lake depth also appears to predict stickleback abundance, though this trend was only marginally significant. Deeper lakes may provide an open-water refuge from pike predation by allowing stickleback to exist outside of the pike inhabited littoral zone. These findings indicate the importance of diverse habitat types and certain chemical and physical characteristics to the outcome of invasion by fish predators.     

Ontogenetic niche shifts and resource partitioning in a subarctic piscivore fish guild

The feeding ecology of three piscivorous fish species (perch (Perca fluviatilis), pike (Esox lucius) and burbot (Lota lota)), was studied in the subarctic Pasvik watercourse (69 °N), northern Norway and Russia. All three species primarily occupied the benthic habitats in the watercourse. Perch and burbot exhibited distinct ontogenetic niche shifts in food resource use, perch changing from a dominance of zooplankton to zoobenthos to fish, and burbot from zoobenthos to fish. Fish prey dominated the diet of all the investigated size-classes of pike, but small-sized pike (<20 cm) were not represented in the sample. Fish prey size was positively related to predator size in all three species. Whitefish (Coregonus lavaretus) was the dominant prey of pike and large-sized burbot and perch. Nine-spined sticklebacks (Pungitius pungitius) was also an important prey and appeared to be a dietary stepping-stone enhancing the transition from invertebrate feeding to consumption of large-sized whitefish prey for all three predators. A cluster analysis separated the different size groups of the three predator species into five functional feeding groups, most of them containing two or all three species. Within these feeding groups, and especially among the piscivorous size-classes, there was a strong and significant interspecific overlap in prey selection, and the dietary similarities between the species were in general much larger than the intraspecific similarities between ontogenetic stages. All three piscivorous species are important top predators in the aquatic food web of the watercourse, and their ontogenetic diet shifts and resource partitioning patterns generate a substantial food web complexity in this subarctic ecosystem.