Daniel Yule | Utah State University (original) (raw)

Papers by Daniel Yule

Journal of Great Lakes Research

The ability of planktivorous fishes to exert top-down control on Bythotrephes potentially has far... more The ability of planktivorous fishes to exert top-down control on Bythotrephes potentially has far-reaching impacts on aquatic food-webs, given previously described effects of Bythotrephes on zooplankton communities. We estimated consumption of Bythotrephes by planktivorous and benthivorous fishes, using bioenergetics and daily ration models at nearshore (18 m), intermediate (46 m), and offshore (110 m) depths along one western Lake Superior transect (April, and September–November) and two northern Lake Michigan transects (April, July, and September). In Lake Superior, consumption (primarily by cisco Coregonus artedi) exceeded Bythotrephes production at all offshore sites in September–November (up to 396% of production consumed) and at the intermediate site in November (842%) with no evidence of consumption nearshore. By comparing Bythotrephes biomass following months of excessive consumption, we conservatively concluded that top-down control was evident only at the offshore site dur...

North American Journal of Fisheries Management, 2000

... 2 Daniel L. Yule a , Roy A. Whaley a , Paul H. Mavrakis a , Dirk D. Miller b & St... more ... 2 Daniel L. Yule a , Roy A. Whaley a , Paul H. Mavrakis a , Dirk D. Miller b & Steve A. Flickinger c pages 10-18. ... Oikos , 51: 220–226. [CrossRef], [Web of Science ®], [CSA] View all references; Johannes et al. 19899. Johannes, MRS, McQueen, DJ, Stewart, TJ and Post, JR 1989. ...

Acoustic (AC) methods are used to estimate the density of pelagic prey fish in large lakes, with ... more Acoustic (AC) methods are used to estimate the density of pelagic prey fish in large lakes, with midwater trawling (MT) used to apportion these estimates to species. Accurate apportionment of species composition is a challenge because only a small fraction of water sampled acoustically is sampled with MT gear, and catch composition can be biased because different fish species (and sizes) have different capture probabilities. We demonstrate a new method of apportionment that uses classification tree modeling to identify variables important for classifying AC targets to species.During the summer of 2011, we conducted a spatially-balance lake-wide AC and MT survey of Lake Superior that included 51 sites in four bathymetric strata (0-30 m, 30-100 m, 100-200 m and > 200 m). Using a data set of 83 MT samples, we developed tree-based models for each stratum. Fish length was the most important variable for classifying species in all strata. Using previously published data on AC target st...

Ecological Modelling, 2014

ABSTRACT The relative importance of predation and food availability as contributors to larval cis... more ABSTRACT The relative importance of predation and food availability as contributors to larval cisco (Coregonus artedi) mortality in Lake Superior were investigated using a visual foraging model to evaluate potential predation pressure by rainbow smelt (Osmerus mordax) and a bioenergetic model to evaluate potential starvation risk. The models were informed by observations of rainbow smelt, larval cisco, and zooplankton abundance at three Lake Superior locations during the period of spring larval cisco emergence and surface-oriented foraging. Predation risk was highest at Black Bay, ON, where average rainbow smelt densities in the uppermost 10 m of the water column were >1000 ha−1. Turbid conditions at the Twin Ports, WI-MN, affected larval cisco predation risk because rainbow smelt remained suspended in the upper water column during daylight, placing them alongside larval cisco during both day and night hours. Predation risk was low at Cornucopia, WI, owing to low smelt densities (<400 ha−1) and deep light penetration, which kept rainbow smelt near the lakebed and far from larvae during daylight. In situ zooplankton density estimates were low compared to the values used to develop the larval coregonid bioenergetics model, leading to predictions of negative growth rates for 10 mm larvae at all three locations. The model predicted that 15 mm larvae were capable of attaining positive growth at Cornucopia and the Twin Ports where low water temperatures (2–6 °C) decreased their metabolic costs. Larval prey resources were highest at Black Bay but warmer water temperatures there offset the benefit of increased prey availability. A sensitivity analysis performed on the rainbow smelt visual foraging model showed that it was relatively insensitive, while the coregonid bioenergetics model showed that the absolute growth rate predictions were highly sensitive to input parameters (i.e., 20% parameter perturbation led to order of magnitude differences in model estimates). Our modelling indicated that rainbow smelt predation may limit larval cisco survival at Black Bay and to a lesser extent at Twin Ports, and that starvation may be a major source of mortality at all three locations. The framework we describe has the potential to further our understanding of the relative importance of starvation and predation on larval fish survivorship, provided information on prey resources available to larvae are measured at sufficiently fine spatial scales and the models provide a realistic depiction of the dynamic processes that the larvae experience.

Science of The Total Environment, 2014

We determined whole-fish polychlorinated biphenyl (PCB) concentrations of 25 male and 25 female a... more We determined whole-fish polychlorinated biphenyl (PCB) concentrations of 25 male and 25 female age-7 ciscoes (Coregonus artedi) captured from a spawning aggregation in Thunder Bay, Lake Superior, during November 2010. We also determined PCB concentrations in the ovaries and somatic tissue of five additional female ciscoes (ages 5-22). All 55 of these ciscoes were in ripe or nearly ripe condition. Bioenergetics modeling was used to determine the contribution of the growth dilution effect toward a difference in PCB concentrations between the sexes, as females grew substantially faster than males. Results showed that the PCB concentration of males (mean = 141 ng/g) was 43% greater than that of females (mean = 98 ng/g), and this difference was highly significant (P<0.0001). Mean PCB concentrations in the ovaries and the somatic tissue of the five females were 135 and 100 ng/g, respectively. Based on these PCB determinations for the ovaries and somatic tissue, we concluded that release of eggs by females at previous spawnings was not a contributing factor to the observed difference in PCB concentrations between the sexes. Bioenergetics modeling results indicated that the growth dilution effect could explain males being higher than females in PCB concentration by only 3-7%. We concluded that the higher PCB concentration in males was most likely due to higher rate of energy expenditure, originating from greater activity and a higher resting metabolic rate. Mean PCB concentration in the cisco eggs was well below the U. S. Food and Drug Administration and Ontario Ministry of Environment guidelines of 2000 and 844 ng/g, respectively, and this finding may have implications for the cisco roe fishery currently operating in Lake Superior.

ABSTRACT Cisco (Coregonus artedi Leseur, formerly lake herring Leucichthys artedi Leseur) populat... more ABSTRACT Cisco (Coregonus artedi Leseur, formerly lake herring Leucichthys artedi Leseur) populations in each of the Laurentian Great Lakes collapsed between the late 1920s and early 1960s following a multitude of stressors, and never recovered in Lakes Michigan, Erie and Ontario. Prior to their collapse, Koelz (1929) studied Leucichthys spp. in the Great Lakes basin and provided a description of their diversity. Three cisco morphotypes were described; a 'slim terete' morphotype (L. artedi artedi), a 'deep compressed' morphotype (L. artedi albus), and a deep-bodied form resembling tullibee in western Canadian lakes (L. artedi manitoulinus). Based on body measurements of 159 individuals (Koelz 1929), we used discriminant function analysis (DFA) to discriminate historic morphotypes. Shapes of historic morphotypes were found to vary significantly (Pillai's trace = 1.16, P < 0.0001). The final DFA model used nine body measurements and correctly classified 90% of the historic cisco. Important discriminating measurements included body depth, eye diameter, and dorsal fin base and height. Between October-November of 2007-2011, we sampled cisco from 16 Great Lakes sites collecting digital photographs of over 1,700 individuals. We applied the DFA model to their body measurements and classified each individual to a morphotype. Contemporary cisco from Lakes Superior, Ontario and Michigan were predominantly classified as artedi, while the most common classifications from northern Lake Huron were albus and manitoulinus. Finding historic morphotypes is encouraging because it suggests that the morphological variation present prior to their collapse still exists. We conclude that contemporary cisco having shapes matching the missing historic morphotypes in the lower lakes warrant special consideration as potential donor populations in reestablishment efforts.

Diel migration patterns of fishes in nearshore (15-80 m depth) and offshore (>80 m) waters of Lak... more Diel migration patterns of fishes in nearshore (15-80 m depth) and offshore (>80 m) waters of Lake Superior were examined to assess the potential for diel migration to link benthic and pelagic, and nearshore and offshore habitats. In our companion article, we described three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no diel migration. DVM was expressed by fishes migrating from benthopelagic to pelagic positions and DBM was expressed by fishes migrating horizontally from deep to shallow waters at night. Fishes not exhibiting diel migration typically showed increased activity by moving from benthic to benthopelagic positions within demersal habitat. The distribution and biomass of fishes in Lake Superior was characterized by examining 704 bottom trawl samples collected between 2001 and 2008 from four depth zones: ≤40, 41-80, 81-160, and >160 m. Diel migration behaviors of fishes described in our companion article were applied to estimates of areal biomass (kg ha −1 ) for each species by depth zone. The relative strength of diel migrations were assessed by applying lake area to areal biomass estimates for each species by depth zone to yield estimates of lake-wide biomass (metric tonnes). Overall, species expressing DVM accounted for 83%, DBM 6%, and non-migration 11% of the total lake-wide community biomass. In nearshore waters, species expressing DVM represented 74% of the biomass, DBM 25%, and non-migration 1%. In offshore waters, species expressing DVM represented 85%, DBM 1%, and non-migration 14% of the biomass. Of species expressing DVM, 83% of total biomass occurred in offshore waters. Similarly, 97% of biomass of non-migrators occurred in offshore waters while 83% of biomass of species expressing DBM occurred in nearshore waters. A high correlation (R 2 = 0.996) between lake area and community biomass by depth zone resulted in 81% of the lake-wide biomass occurring in offshore waters. Accentuating this nearshore-offshore trend was one of increasing estimated total areal biomass of the fish community with depth zone, which ranged from 13.71 kg ha −1 at depths ≤40 m to 18.81 kg ha −1 at depths >160 m, emphasizing the importance of the offshore fish community to the lake ecosystem. The prevalence of diel migration expressed by Lake Superior fishes increases the potential of fish to link benthic and pelagic and shallow and deepwater habitats. These linkages enhance the potential for habitat coupling, a condition where habitats become interconnected and interdependent through transfers of energy and nutrients. Habitat coupling facilitates energy and nutrient flow through a lake ecosystem, thereby increasing productivity, especially in large lakes where benthic and pelagic, and nearshore and offshore habitats are often well separated. We propose that the application of biomass estimates to patterns of diel migration in fishes can serve as a useful metric for assessing the potential for habitat linkages and habitat coupling 355 356 Gorman et al. / Aquatic Ecosystem Health and Management 15 (2012) [355][356][357][358][359][360][361][362][363][364][365][366][367][368] in lake ecosystems, and provide an important indicator of ecosystem health and function. The decline of native Lake Trout and ciscoes and recent declines in exotic Alewife and Rainbow Smelt populations in other Great Lakes have likely reduced the capacity for benthic-pelagic coupling in these systems compared to Lake Superior. We recommend comparing the levels and temporal changes in diel migration in other Great Lakes as a means to assess changes in the relative health and function of these ecosystems.

Diel patterns of distribution of fishes in nearshore (15-80 m depth) and offshore (>80 m) waters ... more Diel patterns of distribution of fishes in nearshore (15-80 m depth) and offshore (>80 m) waters of the Apostle Islands region of Lake Superior were described using bottom trawls, mid-water trawls, and acoustic gear during day and night sampling. These data revealed three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no migration. DVM was expressed by fishes migrating from benthopelagic to pelagic strata and DBM was expressed by fishes migrating horizontally from deeper waters in the day to shallower waters at night while remaining within the benthopelagic stratum. Most fishes that did not exhibit diel migration showed increased nighttime densities as a result of increased activity and movement from benthic to benthopelagic strata. Rainbow Smelt (Osmerus mordax), Cisco (Coregonus artedi), Bloater (C. hoyi), Kiyi (C. kiyi), juvenile Trout-Perch (Percopsis omiscomaycus), and adult siscowet (Salvelinus namaycush siscowet) exhibited DVM. Lake Whitefish (C. clupeaformis), lean Lake Trout (Salvelinus namaycush namaycush), and juvenile siscowet exhibited DBM. Adult Trout-Perch and adult Pygmy Whitefish (Prosopium coulteri) exhibited a mixture of DBM and DVM. Burbot (Lota lota), Slimy Sculpin (Cottus cognatus), Spoonhead Sculpin (C. ricei), and Deepwater Sculpin (Myoxocephalus thompsonii) did not exhibit diel migration, but showed evidence of increased nocturnal activity. Ninespine Stickleback (Pungitius pungitius) exhibited a mixture of DVM and non-migration. Juvenile Pygmy Whitefish did not show a diel change in density or depth distribution. Species showing ontogenetic shifts in depth distribution with larger, adult life stages occupying deeper waters included, Rainbow Smelt, lean and siscowet Lake Trout, Lake Whitefish, Pygmy Whitefish, Ninespine Stickleback and Trout-Perch. Of these species, siscowet also showed an ontogenetic shift from primarily DBM as juveniles to primarily DVM as adults. Across all depths, fishes expressing DVM accounted for 73% of the total estimated community areal biomass (kg ha −1 ) while those expressing DBM accounted for 25% and non-migratory species represented 2% of the biomass. The proportion of total community biomass exhibiting DVM increased with depth, from 59% to 95% across ≤30 m to >90 m depth zones. Along the same depth gradient, the proportion of total community biomass exhibiting DBM declined from 40% to 1%, while non-migrators increased from 1% to 4%. These results indicate that DVM and DBM behaviors are pervasive in the Lake Superior fish community and potentially provide strong linkages that effect coupling of benthic and pelagic and nearshore and offshore habitats.

The Great Lakes Science Center has conducted daytime nearshore bottom trawl surveys of Lake Super... more The Great Lakes Science Center has conducted daytime nearshore bottom trawl surveys of Lake Superior (15-80 m bathymetric depth zone) each spring since 1978 to provide long-term trends of relative abundance and biomass of the fish community. Between 19 May and 20 June 2011, 82 stations distributed around the perimeter of the lake were sampled with a 12-m Yankee bottom trawl towed cross-contour. The 2011 estimate of fish community biomass was 3.63 kg/ha, seventh lowest in the 34-year survey history, but up from 1.37 kg/ha observed in the 2010 survey. The distribution of biomass across jurisdictions was uneven; mean biomass in Canada East, Canada West, Michigan, Minnesota and Wisconsin waters were 2.23, 3.64, 2.07, 0.01, and 7.73 kg/ha, respectively. Dominant species in the catch, in order of relative biomass, were lake whitefish, rainbow smelt, bloater, cisco, and longnose sucker. Compared to 2010 levels, biomass of cisco, bloater, shortjaw cisco, lake whitefish, rainbow smelt, and lean and siscowet lake trout increased. Year-class strengths for the 2010 cisco and bloater cohorts were well below average and ranked as the ninth and twelfth weakest year-classes, respectively, in the past 34 years. The 2011 cisco age structure was dominated by age-2 fish (2009 year-class), which accounted for 91% of the ciscoes captured. Remaining ciscoes captured were composed mostly of adults from the 2005, 2003 and 1998 year-classes. Year-class strength of rainbow smelt was the fifth weakest in the survey record, continuing a decline that began in 2008.

and sharing with colleagues.

ABSTRACT 1. We hypothesised that the autumn spawning migration of Lake Superior cisco (Coregonus ... more ABSTRACT 1. We hypothesised that the autumn spawning migration of Lake Superior cisco (Coregonus artedi) provides a resource subsidy, in the form of energy-rich cisco eggs, from the offshore pelagic to the nearshore benthic community over winter, when alternate prey production is likely to be low. 2. We tested this hypothesis using fish and macroinvertebrate surveys, fish population demographics, diet and stable isotope analyses, and bioenergetics modelling. 3. The benthic, congeneric lake whitefish (C. clupeaformis) was a clear beneficiary of cisco spawning. Cisco eggs represented 16% of lake whitefish annual consumption in terms of biomass, but 34% of energy (because of their high energy density: >10 kJ g wet mass 1). Stable isotope analyses were consistent with these results and suggest that other nearshore fish species may also rely on cisco eggs. 4. The lipid content of lake whitefish liver almost doubled from 26 to 49% between November and March, while that of muscle increased from 14 to 26% over the same period, suggesting lake whitefish were building, rather than depleting, lipid reserves during winter. 5. In the other Laurentian Great Lakes, where cisco populations remain very low and rehabilitation efforts are underway, the offshore-to-nearshore ecological link apparent in Lake Superior has been replaced by non-native planktivorous species. These non-native species spawn in spring have smaller eggs and shorter incubation periods. The rehabilitation of cisco in these systems should reinstate the onshore subsidy as it has in Lake Superior.

Fisheries Research, 2015

ABSTRACT We examined the spatial scale of recruitment variability for disparate cisco (Coregonus ... more ABSTRACT We examined the spatial scale of recruitment variability for disparate cisco (Coregonus artedi) populations in the Great Lakes (n = 8) and Minnesota inland lakes (n = 4). We found that the scale of synchrony was approximately 400 km when all available data were utilized; much greater than the 50-km scale suggested for freshwater fish populations in an earlier global analysis. The presence of recruitment synchrony between Great Lakes and inland lake cisco populations supports the hypothesis that synchronicity is driven by climate and not dispersal. We also found synchrony in larval densities among three Lake Superior populations separated by 25–275 km, which further supports the hypothesis that broad-scale climatic factors are the cause of spatial synchrony. Among several candidate climate variables measured during the period of larval cisco emergence, maximum wind speeds exhibited the most similar spatial scale of synchrony to that observed for cisco. Other factors, such as average water temperatures, exhibited synchrony on broader spatial scales, which suggests they could also be contributing to recruitment synchrony. Our results provide evidence that abiotic factors can induce synchronous patterns of recruitment for populations of cisco inhabiting waters across a broad geographic range, and show that broad-scale synchrony of recruitment can occur in freshwater fish populations as well as those from marine systems.

Freshwater Biology, 2014

1. In large lakes around the world, depth-based changes in the abundance and distribution of inve... more 1. In large lakes around the world, depth-based changes in the abundance and distribution of invertebrate and fish species suggest that there may be concomitant changes in patterns of resource allocation. Using Lake Superior of the Laurentian Great Lakes as an example, we explored this idea through stable isotope analyses of 13 major fish taxa. 2. Patterns in carbon and nitrogen isotope ratios revealed use of both littoral and profundal benthos among populations of most taxa analysed regardless of the depth of their habitat, providing evidence of nearshore-offshore trophic linkages in the largest freshwater lake by area in the world. 3. Isotope-mixing model results indicated that the overall importance of benthic food-web pathways to fish was highest in nearshore species, whereas the importance of planktonic pathways increased in offshore species. These characteristics, shared with the Great Lakes of Africa, Russia and Japan, appear to be governed by two key processes: high benthic production in nearshore waters and the prevalence of diel vertical migration (DVM) among offshore invertebrate and fish taxa. DVM facilitates use of pelagic food resources by deep-water biota and represents an important process of trophic linkage among habitats in large lakes. 4. Support of whole-lake food webs through trophic linkages among pelagic, profundal and littoral habitats appears to be integral to the functioning of large lakes. These linkages can be disrupted though ecosystem disturbance such as eutrophication or the effects of invasive species and should be considered in native species restoration efforts.

Canadian Journal of Fisheries and Aquatic Sciences, 2014

We synthesized data from multiple sampling programs and years to describe the Lake Superior pelag... more We synthesized data from multiple sampling programs and years to describe the Lake Superior pelagic biomass size structure. Data consisted of Coulter counts for phytoplankton, optical plankton counts for zooplankton, and acoustic surveys for pelagic prey fish. The size spectrum was stable across two time periods separated by 5 years. The primary scaling or overall slope of the normalized biomass size spectra for the combined years was −1.113, consistent with a previous estimate for Lake Superior (−1.10). Periodic dome structures within the overall biomass size structure were fit to polynomial regressions based on the observed sub-domes within the classical taxonomic positions (algae, zooplankton, and fish). This interpretation of periodic dome delineation was aligned more closely with predator-prey size relationships that exist within the zooplankton (herbivorous, predacious) and fish (planktivorous, piscivorous) taxonomic positions. Domes were spaced approximately every 3.78 log 10 units along the axis and with a decreasing peak magnitude of −4.1 log 10 units. The relative position of the algal and herbivorous zooplankton domes predicted well the subsequent biomass domes for larger predatory zooplankton and planktivorous prey fish.

Journal of Great Lakes Research

The ability of planktivorous fishes to exert top-down control on Bythotrephes potentially has far... more The ability of planktivorous fishes to exert top-down control on Bythotrephes potentially has far-reaching impacts on aquatic food-webs, given previously described effects of Bythotrephes on zooplankton communities. We estimated consumption of Bythotrephes by planktivorous and benthivorous fishes, using bioenergetics and daily ration models at nearshore (18 m), intermediate (46 m), and offshore (110 m) depths along one western Lake Superior transect (April, and September–November) and two northern Lake Michigan transects (April, July, and September). In Lake Superior, consumption (primarily by cisco Coregonus artedi) exceeded Bythotrephes production at all offshore sites in September–November (up to 396% of production consumed) and at the intermediate site in November (842%) with no evidence of consumption nearshore. By comparing Bythotrephes biomass following months of excessive consumption, we conservatively concluded that top-down control was evident only at the offshore site dur...

North American Journal of Fisheries Management, 2000

... 2 Daniel L. Yule a , Roy A. Whaley a , Paul H. Mavrakis a , Dirk D. Miller b & St... more ... 2 Daniel L. Yule a , Roy A. Whaley a , Paul H. Mavrakis a , Dirk D. Miller b & Steve A. Flickinger c pages 10-18. ... Oikos , 51: 220–226. [CrossRef], [Web of Science ®], [CSA] View all references; Johannes et al. 19899. Johannes, MRS, McQueen, DJ, Stewart, TJ and Post, JR 1989. ...

Acoustic (AC) methods are used to estimate the density of pelagic prey fish in large lakes, with ... more Acoustic (AC) methods are used to estimate the density of pelagic prey fish in large lakes, with midwater trawling (MT) used to apportion these estimates to species. Accurate apportionment of species composition is a challenge because only a small fraction of water sampled acoustically is sampled with MT gear, and catch composition can be biased because different fish species (and sizes) have different capture probabilities. We demonstrate a new method of apportionment that uses classification tree modeling to identify variables important for classifying AC targets to species.During the summer of 2011, we conducted a spatially-balance lake-wide AC and MT survey of Lake Superior that included 51 sites in four bathymetric strata (0-30 m, 30-100 m, 100-200 m and > 200 m). Using a data set of 83 MT samples, we developed tree-based models for each stratum. Fish length was the most important variable for classifying species in all strata. Using previously published data on AC target st...

Ecological Modelling, 2014

ABSTRACT The relative importance of predation and food availability as contributors to larval cis... more ABSTRACT The relative importance of predation and food availability as contributors to larval cisco (Coregonus artedi) mortality in Lake Superior were investigated using a visual foraging model to evaluate potential predation pressure by rainbow smelt (Osmerus mordax) and a bioenergetic model to evaluate potential starvation risk. The models were informed by observations of rainbow smelt, larval cisco, and zooplankton abundance at three Lake Superior locations during the period of spring larval cisco emergence and surface-oriented foraging. Predation risk was highest at Black Bay, ON, where average rainbow smelt densities in the uppermost 10 m of the water column were >1000 ha−1. Turbid conditions at the Twin Ports, WI-MN, affected larval cisco predation risk because rainbow smelt remained suspended in the upper water column during daylight, placing them alongside larval cisco during both day and night hours. Predation risk was low at Cornucopia, WI, owing to low smelt densities (<400 ha−1) and deep light penetration, which kept rainbow smelt near the lakebed and far from larvae during daylight. In situ zooplankton density estimates were low compared to the values used to develop the larval coregonid bioenergetics model, leading to predictions of negative growth rates for 10 mm larvae at all three locations. The model predicted that 15 mm larvae were capable of attaining positive growth at Cornucopia and the Twin Ports where low water temperatures (2–6 °C) decreased their metabolic costs. Larval prey resources were highest at Black Bay but warmer water temperatures there offset the benefit of increased prey availability. A sensitivity analysis performed on the rainbow smelt visual foraging model showed that it was relatively insensitive, while the coregonid bioenergetics model showed that the absolute growth rate predictions were highly sensitive to input parameters (i.e., 20% parameter perturbation led to order of magnitude differences in model estimates). Our modelling indicated that rainbow smelt predation may limit larval cisco survival at Black Bay and to a lesser extent at Twin Ports, and that starvation may be a major source of mortality at all three locations. The framework we describe has the potential to further our understanding of the relative importance of starvation and predation on larval fish survivorship, provided information on prey resources available to larvae are measured at sufficiently fine spatial scales and the models provide a realistic depiction of the dynamic processes that the larvae experience.

Science of The Total Environment, 2014

We determined whole-fish polychlorinated biphenyl (PCB) concentrations of 25 male and 25 female a... more We determined whole-fish polychlorinated biphenyl (PCB) concentrations of 25 male and 25 female age-7 ciscoes (Coregonus artedi) captured from a spawning aggregation in Thunder Bay, Lake Superior, during November 2010. We also determined PCB concentrations in the ovaries and somatic tissue of five additional female ciscoes (ages 5-22). All 55 of these ciscoes were in ripe or nearly ripe condition. Bioenergetics modeling was used to determine the contribution of the growth dilution effect toward a difference in PCB concentrations between the sexes, as females grew substantially faster than males. Results showed that the PCB concentration of males (mean = 141 ng/g) was 43% greater than that of females (mean = 98 ng/g), and this difference was highly significant (P<0.0001). Mean PCB concentrations in the ovaries and the somatic tissue of the five females were 135 and 100 ng/g, respectively. Based on these PCB determinations for the ovaries and somatic tissue, we concluded that release of eggs by females at previous spawnings was not a contributing factor to the observed difference in PCB concentrations between the sexes. Bioenergetics modeling results indicated that the growth dilution effect could explain males being higher than females in PCB concentration by only 3-7%. We concluded that the higher PCB concentration in males was most likely due to higher rate of energy expenditure, originating from greater activity and a higher resting metabolic rate. Mean PCB concentration in the cisco eggs was well below the U. S. Food and Drug Administration and Ontario Ministry of Environment guidelines of 2000 and 844 ng/g, respectively, and this finding may have implications for the cisco roe fishery currently operating in Lake Superior.

ABSTRACT Cisco (Coregonus artedi Leseur, formerly lake herring Leucichthys artedi Leseur) populat... more ABSTRACT Cisco (Coregonus artedi Leseur, formerly lake herring Leucichthys artedi Leseur) populations in each of the Laurentian Great Lakes collapsed between the late 1920s and early 1960s following a multitude of stressors, and never recovered in Lakes Michigan, Erie and Ontario. Prior to their collapse, Koelz (1929) studied Leucichthys spp. in the Great Lakes basin and provided a description of their diversity. Three cisco morphotypes were described; a 'slim terete' morphotype (L. artedi artedi), a 'deep compressed' morphotype (L. artedi albus), and a deep-bodied form resembling tullibee in western Canadian lakes (L. artedi manitoulinus). Based on body measurements of 159 individuals (Koelz 1929), we used discriminant function analysis (DFA) to discriminate historic morphotypes. Shapes of historic morphotypes were found to vary significantly (Pillai's trace = 1.16, P < 0.0001). The final DFA model used nine body measurements and correctly classified 90% of the historic cisco. Important discriminating measurements included body depth, eye diameter, and dorsal fin base and height. Between October-November of 2007-2011, we sampled cisco from 16 Great Lakes sites collecting digital photographs of over 1,700 individuals. We applied the DFA model to their body measurements and classified each individual to a morphotype. Contemporary cisco from Lakes Superior, Ontario and Michigan were predominantly classified as artedi, while the most common classifications from northern Lake Huron were albus and manitoulinus. Finding historic morphotypes is encouraging because it suggests that the morphological variation present prior to their collapse still exists. We conclude that contemporary cisco having shapes matching the missing historic morphotypes in the lower lakes warrant special consideration as potential donor populations in reestablishment efforts.

Diel migration patterns of fishes in nearshore (15-80 m depth) and offshore (>80 m) waters of Lak... more Diel migration patterns of fishes in nearshore (15-80 m depth) and offshore (>80 m) waters of Lake Superior were examined to assess the potential for diel migration to link benthic and pelagic, and nearshore and offshore habitats. In our companion article, we described three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no diel migration. DVM was expressed by fishes migrating from benthopelagic to pelagic positions and DBM was expressed by fishes migrating horizontally from deep to shallow waters at night. Fishes not exhibiting diel migration typically showed increased activity by moving from benthic to benthopelagic positions within demersal habitat. The distribution and biomass of fishes in Lake Superior was characterized by examining 704 bottom trawl samples collected between 2001 and 2008 from four depth zones: ≤40, 41-80, 81-160, and >160 m. Diel migration behaviors of fishes described in our companion article were applied to estimates of areal biomass (kg ha −1 ) for each species by depth zone. The relative strength of diel migrations were assessed by applying lake area to areal biomass estimates for each species by depth zone to yield estimates of lake-wide biomass (metric tonnes). Overall, species expressing DVM accounted for 83%, DBM 6%, and non-migration 11% of the total lake-wide community biomass. In nearshore waters, species expressing DVM represented 74% of the biomass, DBM 25%, and non-migration 1%. In offshore waters, species expressing DVM represented 85%, DBM 1%, and non-migration 14% of the biomass. Of species expressing DVM, 83% of total biomass occurred in offshore waters. Similarly, 97% of biomass of non-migrators occurred in offshore waters while 83% of biomass of species expressing DBM occurred in nearshore waters. A high correlation (R 2 = 0.996) between lake area and community biomass by depth zone resulted in 81% of the lake-wide biomass occurring in offshore waters. Accentuating this nearshore-offshore trend was one of increasing estimated total areal biomass of the fish community with depth zone, which ranged from 13.71 kg ha −1 at depths ≤40 m to 18.81 kg ha −1 at depths >160 m, emphasizing the importance of the offshore fish community to the lake ecosystem. The prevalence of diel migration expressed by Lake Superior fishes increases the potential of fish to link benthic and pelagic and shallow and deepwater habitats. These linkages enhance the potential for habitat coupling, a condition where habitats become interconnected and interdependent through transfers of energy and nutrients. Habitat coupling facilitates energy and nutrient flow through a lake ecosystem, thereby increasing productivity, especially in large lakes where benthic and pelagic, and nearshore and offshore habitats are often well separated. We propose that the application of biomass estimates to patterns of diel migration in fishes can serve as a useful metric for assessing the potential for habitat linkages and habitat coupling 355 356 Gorman et al. / Aquatic Ecosystem Health and Management 15 (2012) [355][356][357][358][359][360][361][362][363][364][365][366][367][368] in lake ecosystems, and provide an important indicator of ecosystem health and function. The decline of native Lake Trout and ciscoes and recent declines in exotic Alewife and Rainbow Smelt populations in other Great Lakes have likely reduced the capacity for benthic-pelagic coupling in these systems compared to Lake Superior. We recommend comparing the levels and temporal changes in diel migration in other Great Lakes as a means to assess changes in the relative health and function of these ecosystems.

Diel patterns of distribution of fishes in nearshore (15-80 m depth) and offshore (>80 m) waters ... more Diel patterns of distribution of fishes in nearshore (15-80 m depth) and offshore (>80 m) waters of the Apostle Islands region of Lake Superior were described using bottom trawls, mid-water trawls, and acoustic gear during day and night sampling. These data revealed three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no migration. DVM was expressed by fishes migrating from benthopelagic to pelagic strata and DBM was expressed by fishes migrating horizontally from deeper waters in the day to shallower waters at night while remaining within the benthopelagic stratum. Most fishes that did not exhibit diel migration showed increased nighttime densities as a result of increased activity and movement from benthic to benthopelagic strata. Rainbow Smelt (Osmerus mordax), Cisco (Coregonus artedi), Bloater (C. hoyi), Kiyi (C. kiyi), juvenile Trout-Perch (Percopsis omiscomaycus), and adult siscowet (Salvelinus namaycush siscowet) exhibited DVM. Lake Whitefish (C. clupeaformis), lean Lake Trout (Salvelinus namaycush namaycush), and juvenile siscowet exhibited DBM. Adult Trout-Perch and adult Pygmy Whitefish (Prosopium coulteri) exhibited a mixture of DBM and DVM. Burbot (Lota lota), Slimy Sculpin (Cottus cognatus), Spoonhead Sculpin (C. ricei), and Deepwater Sculpin (Myoxocephalus thompsonii) did not exhibit diel migration, but showed evidence of increased nocturnal activity. Ninespine Stickleback (Pungitius pungitius) exhibited a mixture of DVM and non-migration. Juvenile Pygmy Whitefish did not show a diel change in density or depth distribution. Species showing ontogenetic shifts in depth distribution with larger, adult life stages occupying deeper waters included, Rainbow Smelt, lean and siscowet Lake Trout, Lake Whitefish, Pygmy Whitefish, Ninespine Stickleback and Trout-Perch. Of these species, siscowet also showed an ontogenetic shift from primarily DBM as juveniles to primarily DVM as adults. Across all depths, fishes expressing DVM accounted for 73% of the total estimated community areal biomass (kg ha −1 ) while those expressing DBM accounted for 25% and non-migratory species represented 2% of the biomass. The proportion of total community biomass exhibiting DVM increased with depth, from 59% to 95% across ≤30 m to >90 m depth zones. Along the same depth gradient, the proportion of total community biomass exhibiting DBM declined from 40% to 1%, while non-migrators increased from 1% to 4%. These results indicate that DVM and DBM behaviors are pervasive in the Lake Superior fish community and potentially provide strong linkages that effect coupling of benthic and pelagic and nearshore and offshore habitats.

The Great Lakes Science Center has conducted daytime nearshore bottom trawl surveys of Lake Super... more The Great Lakes Science Center has conducted daytime nearshore bottom trawl surveys of Lake Superior (15-80 m bathymetric depth zone) each spring since 1978 to provide long-term trends of relative abundance and biomass of the fish community. Between 19 May and 20 June 2011, 82 stations distributed around the perimeter of the lake were sampled with a 12-m Yankee bottom trawl towed cross-contour. The 2011 estimate of fish community biomass was 3.63 kg/ha, seventh lowest in the 34-year survey history, but up from 1.37 kg/ha observed in the 2010 survey. The distribution of biomass across jurisdictions was uneven; mean biomass in Canada East, Canada West, Michigan, Minnesota and Wisconsin waters were 2.23, 3.64, 2.07, 0.01, and 7.73 kg/ha, respectively. Dominant species in the catch, in order of relative biomass, were lake whitefish, rainbow smelt, bloater, cisco, and longnose sucker. Compared to 2010 levels, biomass of cisco, bloater, shortjaw cisco, lake whitefish, rainbow smelt, and lean and siscowet lake trout increased. Year-class strengths for the 2010 cisco and bloater cohorts were well below average and ranked as the ninth and twelfth weakest year-classes, respectively, in the past 34 years. The 2011 cisco age structure was dominated by age-2 fish (2009 year-class), which accounted for 91% of the ciscoes captured. Remaining ciscoes captured were composed mostly of adults from the 2005, 2003 and 1998 year-classes. Year-class strength of rainbow smelt was the fifth weakest in the survey record, continuing a decline that began in 2008.

and sharing with colleagues.

ABSTRACT 1. We hypothesised that the autumn spawning migration of Lake Superior cisco (Coregonus ... more ABSTRACT 1. We hypothesised that the autumn spawning migration of Lake Superior cisco (Coregonus artedi) provides a resource subsidy, in the form of energy-rich cisco eggs, from the offshore pelagic to the nearshore benthic community over winter, when alternate prey production is likely to be low. 2. We tested this hypothesis using fish and macroinvertebrate surveys, fish population demographics, diet and stable isotope analyses, and bioenergetics modelling. 3. The benthic, congeneric lake whitefish (C. clupeaformis) was a clear beneficiary of cisco spawning. Cisco eggs represented 16% of lake whitefish annual consumption in terms of biomass, but 34% of energy (because of their high energy density: >10 kJ g wet mass 1). Stable isotope analyses were consistent with these results and suggest that other nearshore fish species may also rely on cisco eggs. 4. The lipid content of lake whitefish liver almost doubled from 26 to 49% between November and March, while that of muscle increased from 14 to 26% over the same period, suggesting lake whitefish were building, rather than depleting, lipid reserves during winter. 5. In the other Laurentian Great Lakes, where cisco populations remain very low and rehabilitation efforts are underway, the offshore-to-nearshore ecological link apparent in Lake Superior has been replaced by non-native planktivorous species. These non-native species spawn in spring have smaller eggs and shorter incubation periods. The rehabilitation of cisco in these systems should reinstate the onshore subsidy as it has in Lake Superior.

Fisheries Research, 2015

ABSTRACT We examined the spatial scale of recruitment variability for disparate cisco (Coregonus ... more ABSTRACT We examined the spatial scale of recruitment variability for disparate cisco (Coregonus artedi) populations in the Great Lakes (n = 8) and Minnesota inland lakes (n = 4). We found that the scale of synchrony was approximately 400 km when all available data were utilized; much greater than the 50-km scale suggested for freshwater fish populations in an earlier global analysis. The presence of recruitment synchrony between Great Lakes and inland lake cisco populations supports the hypothesis that synchronicity is driven by climate and not dispersal. We also found synchrony in larval densities among three Lake Superior populations separated by 25–275 km, which further supports the hypothesis that broad-scale climatic factors are the cause of spatial synchrony. Among several candidate climate variables measured during the period of larval cisco emergence, maximum wind speeds exhibited the most similar spatial scale of synchrony to that observed for cisco. Other factors, such as average water temperatures, exhibited synchrony on broader spatial scales, which suggests they could also be contributing to recruitment synchrony. Our results provide evidence that abiotic factors can induce synchronous patterns of recruitment for populations of cisco inhabiting waters across a broad geographic range, and show that broad-scale synchrony of recruitment can occur in freshwater fish populations as well as those from marine systems.

Freshwater Biology, 2014

1. In large lakes around the world, depth-based changes in the abundance and distribution of inve... more 1. In large lakes around the world, depth-based changes in the abundance and distribution of invertebrate and fish species suggest that there may be concomitant changes in patterns of resource allocation. Using Lake Superior of the Laurentian Great Lakes as an example, we explored this idea through stable isotope analyses of 13 major fish taxa. 2. Patterns in carbon and nitrogen isotope ratios revealed use of both littoral and profundal benthos among populations of most taxa analysed regardless of the depth of their habitat, providing evidence of nearshore-offshore trophic linkages in the largest freshwater lake by area in the world. 3. Isotope-mixing model results indicated that the overall importance of benthic food-web pathways to fish was highest in nearshore species, whereas the importance of planktonic pathways increased in offshore species. These characteristics, shared with the Great Lakes of Africa, Russia and Japan, appear to be governed by two key processes: high benthic production in nearshore waters and the prevalence of diel vertical migration (DVM) among offshore invertebrate and fish taxa. DVM facilitates use of pelagic food resources by deep-water biota and represents an important process of trophic linkage among habitats in large lakes. 4. Support of whole-lake food webs through trophic linkages among pelagic, profundal and littoral habitats appears to be integral to the functioning of large lakes. These linkages can be disrupted though ecosystem disturbance such as eutrophication or the effects of invasive species and should be considered in native species restoration efforts.

Canadian Journal of Fisheries and Aquatic Sciences, 2014

We synthesized data from multiple sampling programs and years to describe the Lake Superior pelag... more We synthesized data from multiple sampling programs and years to describe the Lake Superior pelagic biomass size structure. Data consisted of Coulter counts for phytoplankton, optical plankton counts for zooplankton, and acoustic surveys for pelagic prey fish. The size spectrum was stable across two time periods separated by 5 years. The primary scaling or overall slope of the normalized biomass size spectra for the combined years was −1.113, consistent with a previous estimate for Lake Superior (−1.10). Periodic dome structures within the overall biomass size structure were fit to polynomial regressions based on the observed sub-domes within the classical taxonomic positions (algae, zooplankton, and fish). This interpretation of periodic dome delineation was aligned more closely with predator-prey size relationships that exist within the zooplankton (herbivorous, predacious) and fish (planktivorous, piscivorous) taxonomic positions. Domes were spaced approximately every 3.78 log 10 units along the axis and with a decreasing peak magnitude of −4.1 log 10 units. The relative position of the algal and herbivorous zooplankton domes predicted well the subsequent biomass domes for larger predatory zooplankton and planktivorous prey fish.