Introduced northern pike predation on salmonids in southcentral Alaska

Northern pike (Esox lucius) are opportunistic predators that can switch to alternative prey species after preferred prey have declined. This trophic adaptability allows invasive pike to have negative effects on aquatic food webs. In Southcentral Alaska, invasive pike are a substantial concern because they have spread to important spawning and rearing habitat for salmonids and are hypothesised to be responsible for recent salmonid declines. We described the relative importance of salmonids and other prey species to pike diets in the Deshka River and Alexander Creek in Southcentral Alaska. Salmonids were once abundant in both rivers, but they are now rare in Alexander Creek. In the Deshka River, we found that juvenile Chinook salmon (Oncorhynchus tshawytscha) and coho salmon (O. kisutch) dominated pike diets and that small pike consumed more of these salmonids than large pike. In Alexander Creek, pike diets reflected the distribution of spawning salmonids, which decrease with distance upstream. Although salmonids dominated pike diets in the lowest reach of the stream, Arctic lamprey (Lampetra camtschatica) and slimy sculpin (Cottus cognatus) dominated pike diets in the middle and upper reaches. In both rivers, pike density did not influence diet and pike consumed smaller prey items than predicted by their gape‐width. Our data suggest that (1) juvenile salmonids are a dominant prey item for pike, (2) small pike are the primary consumers of juvenile salmonids and (3) pike consume other native fish species when juvenile salmonids are less abundant. Implications of this trophic adaptability are that invasive pike can continue to increase while driving multiple species to low abundance.     

Food consumption of northern pike (Esox lucius L.), estimated with a bioenergetics model

Abstract– A bioenergetics model was used to estimate the food consumption of northern pike ( Esox lucius L.) in two lakes in northeastern Finland, and the results compared with those obtained by a method based on stomach contents and gastric evacuation rates. The annual specific food consumption was 7–5 g per one gram pike for one- and two-year-old pike, and 4–3 g g⁻¹ in age groups 3–6 years old, according to the bioenergetics model. The proportion of ingested food used for growth was 10–30%, and was highest in the youngest age groups. The food consumption rates estimated by the stomach contents method were considerably lower, only about half those produced by the bioenergetics model. The method based on stomach contents tends to underestimate the food consumption. The bioenergetics model was considered more valid for estimating food consumption, but some of the parameter values for northern pike may differ in populations from different latitudes. According to a sensitivity analysis, the food consumption estimate of the bioenergetics model was most sensitive to parameters in the allometric function for respiration.     

Getting cosy in freshwater: Assumed to be brackish pike are not so brackish after all

Pike (Esox lucius) occupy coastal streams and rivers of the Baltic Sea, where they attain large sizes (>5 kg). These large sizes are perhaps due to the fact that they can tolerate relatively high salinities and can thus forage in the nearby more productive brackish environments. In an attempt to quantify the extent to which pike utilise brackish environments, and to provide some insight into the underlying causes for brackish water migrations, we tagged 30 pike from a western Baltic river with acoustic transmitters and were able to track 21 individuals for 1 year. Based on experienced from local anglers, this population was assumed to be brackish in nature, where individuals underwent freshwater migrations to spawn. Our findings however suggest that the smallest and most active individuals make short exits into brackish waters and do so on rare occasions. Our results further indicate that neither sex nor size is related to activity level. We suggest that these patterns reflect two distinct behaviours—active and passive—and that large pike can be supported by the food availability in the river, without the need to venture into coastal zones, thus defying the conventional view that Baltic pike are all brackish in nature.     

Hatching time effect on the intra‐spawning larval morphology and growth in Northern pike (Esox luciusL.)

Based on the analysis of 11 morphometric variables of body (total length, body area and perimeter, myotome height and eye diameter) and yolk sac (length, height, area, perimeter and volume) of pike larvae, the aim of this study was to evaluate how larval size at hatching and growth of larvae hatched from single egg batches vary according to three hatching times: early, mid and late. Hatching time structures strongly pike larval morphometrics. Early hatched larvae have smaller body sizes at hatch, faster growth and higher yolk use efficiency than late hatched ones. Early hatched larvae seem to be premature and hatch at precocious developmental stage whereas late hatched individuals continue their growth within the egg shell and hatch at larger size but with lower reserves (yolk). A compensatory growth phase was observed for the early hatching pike larvae particularly during the first 5 days post hatch. Consequently, no significant difference in body parameters was recorded from day 10 post hatching whatever the hatching time. The higher growth accomplished by early hatched larvae may be related to a particular metabolic activity that converts more efficiently yolk into body tissues.     

Effects of angling and manual handling on pike behaviour investigated by high‐resolution positional telemetry

Human disturbances such as angling and manual handling may have long‐term effects on the behaviour of pike, Esox lucius L., an ecologically important species. Using continuous high‐resolution positional telemetry, this study compared the swimming activity of handled and unhandled pike in a small lake. Pike pre‐equipped with acoustic transmitters were angled and exposed to a handling protocol including measurements of length and mass. Pike not recaptured constituted an unhandled control group. Results demonstrated that the handling protocol caused temperature‐dependent changes in pike activity, with higher temperatures leading to lower activity of the recaptured pike. The effects, however, were transitory and not detectable after 48‐h post‐release. These findings indicate that pike are relatively resilient to handling and quickly resume pre‐handling activity.     

Pacific herring (Clupea pallasii) consumption by marine birds during winter in Prince William Sound,Alaska

Following the 1989 MV Exxon Valdez oil spill (EVOS) and subsequent herring population collapse in Alaska's Prince William Sound (PWS), the Pacific herring (Clupea pallasii) fishery was closed. In the 25 yr since EVOS, herring and several herring‐dependent marine bird species have failed to reach pre‐spill population levels. One hypothesis is that intense predation pressure may be inhibiting herring recovery. To inform herring modeling efforts, this study estimated marine bird consumption of juvenile and adult herring in PWS for 10 winters over an 18‐yr period (1989–90 through 2006–2007). Total estimated herring consumption by wintering marine birds averaged 2409 ± 950 t, indicating that avian consumption represents a substantial and inter‐annually variable source of herring mortality. Common murre (Uria aalge) consumed the greatest portion (16–80%) of herring in all years among marine bird species. Juvenile herring biomass consumed annually by common murre was greater than murre consumption of adult herring biomass. Time lag analyses showed that marine bird consumption of adult herring is negatively correlated with the amount of herring spawn observed in subsequent years, but such effects were not observed more than 2 yr. Our models indicate that during years in which herring recruitment is low or bird populations are particularly large, marine birds can consume up to 10% of the annual adult herring biomass. Our results highlight the importance of herring to wintering PWS birds. We propose that future management of herring stocks seeks to reduce negative impacts on marine birds that prey on herring.     

Meta‐analysis of growth rates for a circumpolar fish,the northern pike ( Esox lucius), with emphasis on effects of continent,climate and latitude

For circumpolar species, little is known on how somatic growth rates can vary at large, transcontinental spatial scales. In this study, a meta‐analysis of growth rates was conducted for northern pike ( Esox lucius) across North America and Eurasia. Growth rates of northern pike did not differ between North American and ‘coastal Eurasian’ pike (e.g., UK, Ireland, Sweden), while growth rates for both of these groups were significantly higher compared to ‘inland Eurasian’ pike (mainly in Russia). There was no difference in growth between lentic and lotic habitats on either continent. In North America, pike growth was positively correlated with temperature, but in Eurasia, pike growth correlated poorly with most climatic variables. Similarly, maximum longevity in pike populations was significantly predicted by latitude in North America, but not in Eurasia. After standardising annual pike growth by the thermal opportunity for growth, a highly significant countergradient growth relationship was found for North American pike, while a significant, but considerably less predictive countergradient growth relationship was found for Eurasian pike. This study provides novel insights into the ecology of a circumpolar species and how populations function at extraordinarily large spatial scales. First, pike appear to be cosmopolitan across hydrologic habitats having fast or slow growth in either lentic or lotic environments. Secondly, continental‐scale differences in pike growth rates are suggestive of major genetic and life‐history differences. Finally, variable climate–growth relationships and countergradient growth patterns indicate that global climate change is likely to affect circumpolar fishes like pike in complex, nonlinear ways.     

Climate effects and bottom‐up controls on growth and size‐at‐age of Pacific halibut (Hippoglossus stenolepis) in Alaska (USA)

Pacific halibut (Hippoglossus stenolepis) are an ecologically, commercially, and culturally important Alaskan groundfish species. Commercial harvest of halibut dates back to the late 19th century and has been managed by the International Pacific Halibut Commission (IPHC) since 1921. IPHC surveys have revealed declining trends in survey biomass in multiple regions and region‐specific declines in mean size‐at‐age (size‐at‐age) over the past two decades (>50% in some areas). Changes in size‐at‐age can arise from a variety of physical, ecological, sampling, and fishery effects, including size‐dependent fishery or predation mortality, alteration in growth from variability in prey quality or quantity, and changes in temperature‐dependent metabolic demands. Here, we develop and apply a bioenergetics model for halibut using survey‐based diet and temperature data for Alaska to evaluate potential environmental drivers of size‐at‐age. In general, juvenile (<40 cm fork length) foraging rates were highest in the Gulf of Alaska concomitant with higher potential growth and elevated basal metabolic demands during warm summer conditions. In contrast, adult (40–120 cm FL) potential growth was highest in the Eastern Bering Sea, potentially reflecting lower metabolic costs and higher rates of prey consumption in that region. We additionally find evidence for interannual variation in potential growth, with a higher frequency of reduced growth potential in the last decade, particularly in the Eastern Bering Sea in 2015 and 2016 for both juvenile and adult halibut. These results suggest the potential for patterns in size‐at‐age to arise from trophic and environmental constraints that collectively limit growth in some regions and years.     

Saving large fish through harvest slots outperforms the classical minimum‐length limit when the aim is to achieve multiple harvest and catch‐related fisheries objectives

We address the problem of optimal size‐selective exploitation in an age‐structured fish population model by systematically examining how density and size dependency in growth, mortality and fecundity affect optimal harvesting patterns when judged against a set of fisheries objectives. The study offers five key insights. First, while minimum‐length limits often maximize the biomass yield, exploitation using harvest slots (i.e. regulations that protect both immature and very large individuals) can generate within 95% of maximum yield; harvest slots also generally maximize the number of fish that are harvested. Second, density dependence in growth and size‐dependent mortality predict more liberal optimal size limits than those derived under assumptions of no density and size dependence. Third, strong density dependence in growth maximizes the catch of trophy fish only when modest harvest is introduced; the same holds for numbers harvested, when the stock–recruitment function follows the Ricker type. Fourth, the inclusion of size‐dependent maternal effects on fecundity or egg viability has only limited effects on optimal size limits, unless the increase in fecundity with mass (“hyperallometry”) is very large. However, large hyperallometry in fecundity shifts the optimal size limit for biomass yield from the traditional minimum‐length limit to a harvest slot. Fifth, harvest slots generally provide the best compromises among multiple objectives. We conclude that harvest slots, or more generally dome‐shaped selectivity to harvest, can outperform the standard minimum‐length selectivity. The exact configuration of optimal size limits crucially depends on objectives, local fishing pressure, the stock–recruitment function, and the density and size dependency of growth, mortality and fecundity.     

Habitat partitioning among three predatory fish in a temperate reservoir

The diversity of predatory species plays a key role in ecosystem functioning but our understanding of the mechanisms underlying their coexistence is limited, particularly in freshwater ecosystems. Northern pike Esox lucius, European perch Perca fluviatilis and pikeperch Sander lucioperca are three widespread predatory species in European lakes, where they often coexist. As potential competitors, we hypothesised that partitioning habitat is a determinant of species coexistence. This was tested by quantifying the variability of their habitat use in tracking adult individuals in the Bariousses reservoir (France, 86.6 ha, mean depth 7.1 m). Specifically, we investigated their distribution along the littoral–pelagic and depth axes along the daily cycle and across seasons. From littoral to pelagic waters were first found pike, then perch and finally pikeperch. Pike was the closest to the surface while pikeperch was the deepest. This general pattern was, however, variable across seasons with the three species located in the upper layer in summer during reservoir stratification. Individuals were more evenly distributed along the littoral–pelagic axis and closer to the bottom when water was mixing (autumn, winter). In summer, perch used more intensively in the pelagic zone during the daytime. Other species did not show any diel change of habitats. Our results highlighted that species coexistence is associated with habitat partitioning among these three predators, with perch showing a more variable behaviour regarding habitat characteristics. Now more than ever, in the context of global change which modifies habitats, it is of crucial importance to understand the coexistence mechanisms of species that shape ecosystems.     

Spawning movements of humpback whitefish in interior Alaska rivers

This study was conducted to describe the spawning movements and identify spawning areas for humpback whitefish Coregonus pidschian in the Minto Flats–Chatanika River complex, Alaska, during 2008 and 2009. Radio transmitters were surgically implanted in humpback whitefish in 2008 (N = 60) and 2009 (N = 100), and fish positions were determined through a combination of boat and aerial surveys and fixed receiving stations. Two spawning areas were identified: one in the Chatanika River downstream of the Elliot Highway Bridge and the other in the Tanana River near Fairbanks. Humpback whitefish dispersed from the wetland complex of Minto Flats in June, moved upstream through late August, arrived at the spawning areas in early September and began moving downstream in early October. In 2009, spatially segregated movements were observed when approximately 40% of the radio‐tagged humpback whitefish moved to the Tanana River, suggesting that humpback whitefish in Minto Flats are comprised of mixed spawning stocks. These study results provide a complete account of humpback whitefish movements and their associated spawning habitats, which will allow for better‐informed management strategies.     

Interactions between body mass and water temperature cause energetic bottlenecks in juvenile steelhead

Across taxa, it is generally accepted that there are fitness advantages to rapid growth early in life. For stream‐dwelling salmonids, however, high temperatures and associated energetic costs during the summer growing season might offset or even prevent the competitive advantage of large body size. Our overall objective was to understand the relative importance of factors that can cause variation in growth rates in an age‐0 cohort of wild steelhead (Oncorhynchus mykiss) in Idaho, USA, where temperatures approach, and temporarily exceed, their tolerance level. For individually tagged fish inhabiting the same stream reach, we found that growth rates were negatively related to fish mass (slopes of the two best approximating models were both ?0.024). Comparing growth rates from 16 different stream reaches throughout the watershed, we found that temperature‐induced metabolic cost was the single best approximating model (AIC w_i = 1.0) of the variation in individual growth rates. The bioenergetic model showed that mass‐specific metabolic costs decreased with mass, but the absolute energetic demands increased over the same size range. Because temperature had a multiplicative effect on metabolic cost, our results suggest that the effect of food limitation increased with fish size. We conclude that high water temperatures pose energetic bottlenecks and can be a potentially strong mechanism limiting growth in juvenile salmonids in summer, particularly as streams in the region experience warming trends.     

Growth and production of Pacific ocean perch (Sebastes alutus) in nursery habitats of the Gulf of Alaska

Nursery areas for juvenile fishes are often important for determining recruitment in marine populations by providing habitats that can maximize growth and thereby minimize mortality. Pacific ocean perch (POP, Sebastes alutus) have an extended juvenile period where they inhabit rocky nursery habitats. We examined POP nursery areas to link growth potential to recruitment. Juvenile POP were captured from nursery areas in 2004 and 2008, and estimated growth rates ranged from ?0.19 to 0.60 g day^?1 based on differences in size between June and August. Predicted growth rates from a bioenergetics model ranged from 0.05 to 0.49 g day^?1 and were not significantly different than observed. Substrate preferences and the distribution of their preferred habitats were utilized to predict the extent of juvenile POP nursery habitat in the Gulf of Alaska. Based on densities of fish observed on underwater video transects and the spatial extent of nursery areas, we predicted 278 and 290 million juvenile POP were produced in 2004 and 2008. Growth potential for juvenile POP was reconstructed using the bioenergetics model, spring zooplankton bloom timing and duration and bottom water temperature for 1982–2008. When a single outlying recruitment year in 1986 was removed, growth potential experienced by juvenile POP in nursery areas was significantly correlated to the recruitment time‐series from the stock assessment, explaining ～30% of the variability. This research highlights the potential to predict recruitment using habitat‐based methods and provides a potential mechanism for explaining some of the POP recruitment variability observed for this population.     

温度对白斑狗鱼早期发育和生长的影响

为了能给白斑狗鱼耐高温性状的改良提供有效的分子标记,本研究基于白斑狗鱼热敏感组 (109尾)和耐高温组 (103尾)进行简化基因组测序,对25个染色体中的InDel标记,以前两个PCA为协变量,利用MLM模型(Masked Language Mode)与白斑狗鱼的耐热性状进行了关联分析。结果显示,大部分InDel分布在内含子 (63.69%),外显子分布的InDel位点较少 (1.30%)。通过GWAS分析,发现5个位点与白斑狗鱼耐热性状显著关联,分别位于safb基因、未知基因LOC117593903和CLSTN2基因内含子中。其中9N del、4N del-1和4N del-2三个InDel位点均位于CLSTN2基因第3内含子,这3个位点在212尾个体中基因型分布高度连锁。本研究中发现的5个InDel突变可能会对白斑狗鱼的耐热性状产生显著的影响,可作为白斑狗鱼耐热性状改良的候选分子标记。进一步在验证群体中利用KASP技术对部分位点进行了验证。发现9N del位点DD基因型个体在热敏感组中占优势,DI基因型个体在耐高温组中占优势,与简化基因组测序结果基本一致。位于CLSTN2基因第2内含子的3个InDel位点可能影响CLSTN2基因的转录,该基因可能是白斑狗鱼耐高温性状相关的重要候选基因。研究结果为白斑狗鱼分子标记辅助育种提供了理论依据,为白斑狗鱼耐热性状的改良提供了候选分子标记。

     

Food consumption and digestive enzyme activity of Clarias batrachus exposed to various temperatures

The effect of temperature on the food consumption rate and the digestive enzyme activities of Clarias batrachus (80.60 ± 5.34 g) were evaluated. Fish were exposed to six different temperatures of 10, 15, 20, 25, 30 and 35 °C following an acclimation temperature of 25 °C. The rate of temperature change was 2 °C day^?1. Highest food consumption was recorded at 25 °C. It gradually reduced with decreasing water temperature. Food consumption rate was significantly (P < 0.05) lower at 10 °C compared with other treatments. Hence, 46.67, 8.20–23.58 and 1.02–6.15% reduced food consumptions were recorded in groups exposed at 10, 15 and 20 °C temperatures, respectively, compared with the 25 °C. The consumption rate was not affected in fish exposed at 30 and 35 °C. Total protease, trypsin and chymotrypsin activities were significantly (P < 0.05) higher in fish exposed at 25 °C compared with others. Lipase activity was significantly (P < 0.05) higher in fish exposed at 30 °C compared with others. Lowest enzyme activities were recorded at 10 °C. Water temperature below 25 °C affected the food consumption and digestive enzyme activities in fish that served as indicators of stress in fish.     

Microsatellite polymorphism in Italian populations of northern pike (Esox lucius L.)

Northern pike (Esox lucius) is not considered an endangered species in Italy, but since recent studies indicate the decline of this population, conservation and management strategies based on the genetic differentiation of natural northern pike populations are needed. In this paper, genetic diversity was analysed in 10 Italian and 2 East European northern pike populations by means of seven microsatellite loci. Data indicated an appreciable genetic differentiation, in spite of a low genetic variation, and agreed with the low level of genetic polymorphism already observed for this species in North America and North Europe. Results of statistical tests revealed genetic peculiarities of the Italian populations, even though signals of recent contact between populations were found and discussed in relation to anthropic impacts, particularly to the stocking practice. This investigation represents the first approach to the knowledge of the genetic variability of Italian pike populations using microsatellite markers, and reported results could be of interest for future management and conservation programmes of this species in Italy.     

Spawning distribution and abundance of a northern Chinook salmon population

Chinook salmon, Oncorhynchus tshawytscha (Walbaum), is an important biological and cultural resource in Alaska, but knowledge about Chinook salmon ecology is limited in many regions. From 2009 to 2012, spawning distribution and abundance of a northern Chinook salmon population on the Togiak River in south‐west Alaska were assessed. Chinook salmon preferred deeper mainstem channel spawning habitat, with 12% (14 of 118 tags in 2009) to 21% (22 of 106 tags in 2012) of radio‐tagged fish spawning in smaller order tributaries. Tributary spawners tended to have earlier run timing than mainstem spawners. Chinook salmon exhibited extended holding and backout (entering freshwater but returning to saltwater before completing anadromous migration) behaviours near the mouth of Togiak River, potentially prolonging their exposure to fishery harvest. Mark–recapture total annual run estimates (2010–2012) ranged from 11 240 (2011) to 18 299 (2012) fish. Exploitation of Chinook salmon ranged from 36% (2012) to 55% (2011) during the study period, with incidental fishery catches near the mouth of the river comprising the largest source of harvest.     

Thermal ecology of subadult and adult muskellunge in a thermally enriched reservoir

The movement of adult muskellunge, Esox masquinongy Mitchill, has been investigated in a variety of systems, but temperature selection by muskellunge has not been examined where well‐oxygenated waters were available over a range of temperatures for much of the year. Thirty subadult and adult muskellunge tagged internally with temperature‐sensing radio tags were tracked from March 2010 to March 2011 in a Tennessee reservoir. Mean tag temperatures were 18.9 °C in spring (March to May), 22.1 °C in summer (June to August), 16.5 °C in autumn and 9.8 °C in winter (December to February). When the greatest range in water temperatures was available (7.1–33.3 °C; May to early August 2010), their realised thermal niche (mean ± 1 SD) was 22.3 °C ± 1.8; the realised thermal niche was affected by fish size (smaller fish selected slightly warmer temperatures) but not sex. An electric generating steam plant discharging warm water resumed operation in January 2011, and most (86%) tagged fish occupied the plume where temperatures were ≈10 °C warmer than ambient water temperatures. No mortalities were observed 15 days later when plant operations ceased. Their affinity for the heated plume prompted concerns that muskellunge will be too easily exploited when the plant operates during winter.     

Spat availability of commercial bivalve species recruited on artificial collectors from the northern Gulf of California. Seasonal changes in species composition

This study reports a year‐round recruitment of spat of four commercial bivalve species; Pteria sterna, Euvola vogdesi, Pinctada mazatlanica and Pinna rugosa collected in the region of Puerto Peñasco, north‐eastern coast of the Gulf of California. Bimonthly recruitment of commercial bivalve spat on netlon^® collectors was evaluated for six sites from June 2007 to August 2008. To describe spat recruitment abundances with environmental parameters, sea surface temperature (°C) and surface chlorophyll a concentration (mg m^?3) were characterized by means of monthly Aqua/MODIS satellite data. For each species a repeated measures anova was used to evaluate differences in the number of spat between months, sites and depths. Maximum sea surface temperature was recorded in August–September (~31.5°C) and the minimum in January–February (~15°C), while the minimum surface chlorophyll a was observed in June–September (mean range = 1.5–2 mg m^?3) and the maximum in January–March (mean range = 2–5 mg m^?3). Spat recruitment showed distinct patterns; P. sterna can be characterized as having a Winter–Spring pattern, E. vogdesi a winter pattern, while P. mazatlanica and P. rugosa a summer spat recruitment pattern. This information constitutes part of the fundamental data needed for the development of aquaculture and conservation initiatives in the region based on wild spat supply.     

Effects of protein and carbohydrate levels on survival,consumption and gonad index in adult sea urchin Strongylocentrotus purpuratus (Stimpson 1857) from Baja California,Mexico

A greater understanding of dietary protein and carbohydrate levels with regard to gonad production in Strongylocentrotus purpuratus would increase our nutritional knowledge of this sea urchin and guide the development of formulated diets for such aquaculture target species. A total of 255 purple sea urchins were captured from Ensenada Bay, Mexico, and maintained in 200‐L tanks for 9 weeks. Formulated diets that contained 30%, 26%, 23%, 20% and 17% of protein and 42%, 46%, 50%, 54% and 58% carbohydrates were offered ad libitum. Survival was affected by diets; urchins that were fed high‐protein–low‐carbohydrate diet experienced decreased survival. No significant differences were found in gonad index, but gonad production efficiency was lower in urchins that were fed a medium‐low‐protein–medium‐high‐carbohydrate diet. Urchins that were fed high carbohydrate levels utilized protein more efficiently and showed better digestibility of the diet and protein. These data suggest that all of our diets support gonad growth, but in terms of consumption, a diet that contains protein levels of 17% and 23% with carbohydrate levels of 50% and 58% are beneficial for S. purpuratus.