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David M. Warner

Publications and source records attributed to David M. Warner.

33 records · Page 2Linked to original sources

A multispecies statistical age-structured model to assess predator-prey balance: application to an intensively managed Lake Michigan pelagic fish community

Using a Bayesian model fitting approach, we developed a multispecies statistical catch-at-age model to assess trade-offs between predatory demands and prey productivities, focusing on the Lake Michigan pelagic fish community. We assessed these trade-offs in terms of predation mortalities and productivities of alewife (Alosa pseudoharengus) and rainbow smelt (Osmerus mordax) and functional responses of salmonines. Our predation mortality estimates suggest that salmonine consumption has been a major driver of historical fluctuations in prey abundance, with sharp declines in alewife abundance in the 1980s and 2000s coinciding with estimated increases in predation mortalities. While Chinook salmon (Oncorhynchus tshawytscha) were food limited during periods of low alewife abundance, other salmonines appeared to maintain a (near) maximum per-predator consumption across all observed prey densities, suggesting that feedback mechanisms are unlikely to help maintain a balance between predator consumption and prey productivity in Lake Michigan. This study demonstrates that a multispecies modeling approach that combines stock assessment methods with explicit consideration of predator–prey interactions could provide the basis for tactical decision-making from a broader ecosystem perspective.

Lake Michigan

Nearshore energy subsidies support Lake Michigan fishes and invertebrates following major changes in food web structure

Aquatic food webs that incorporate multiple energy channels (e.g. nearshore benthic or pelagic) with varying productivity and turnover rates convey stability to biological communities by providing multiple independent energy sources. Within the Lake Michigan food web, invasive dreissenid mussels have caused rapid changes to food web structure and potentially altered the channels through which consumers acquire energy. We used stable C and N isotopes to determine how Lake Michigan food web structure has changed in the past decade, coincident with the expansion of dreissenid mussels, decreased pelagic phytoplankton production and increased nearshore benthic algal production. Fish and invertebrate samples collected from sites around Lake Michigan were analyzed to determine taxa-specific 13C:12C (delta 13C) and 15N:14N (delta 15N) ratios. Sampling took place during two distinct periods, 2002-2003 and 2010-2012, that spanned the period of dreissenid expansion, and included nearshore, pelagic and profundal fish and invertebrate taxa. Magnitude and direction of the 13C shift indicated significantly greater reliance upon nearshore benthic energy sources among nearly all fish taxa as well as profundal invertebrates. Although the mechanisms underlying this 13C shift likely varied among species, possible causes include the transport of benthic algal production to offshore waters and an increased reliance on nearshore prey items. Delta 15N shifts were more variable and of smaller magnitude across taxa although declines in delta 15N among some pelagic fishes may indicate a shift to alternative prey resources. Lake Michigan fishes and invertebrates appear to have responded to dreissenid induced changes in nutrient and energy pathways by switching from pelagic to alternative nearshore energy subsidies. Although large shifts in energy allocation (i.e. pelagic to nearshore benthic) resulting from invasive species appear to have affected total production at upper trophic levels, changes in trophic structure and utilization of novel energy pathways may help to stabilize food webs following species invasions.

Lake Michigan

Status of pelagic prey fishes in Lake Michigan, 2013

Acoustic surveys were conducted in late summer/early fall during the years 1992-1996 and 2001-2013 to estimate pelagic prey fish biomass in Lake Michigan. Midwater trawling during the surveys as well as target strength provided a measure of species and size composition of the fish community for use in scaling acoustic data and providing species-specific abundance estimates. The 2013 survey consisted of 27 acoustic transects (546 km total) and 31 midwater trawl tows. Mean prey fish biomass was 6.1 kg/ha (relative standard error, RSE = 11%) or 29.6 kilotonnes (kt = 1,000 metric tons), which was similar to the estimate in 2012 (31.1 kt) and 23.5% of the long-term (18 years) mean. The numeric density of the 2013 alewife year class was 6% of the time series average and this year-class contributed 4% of total alewife biomass (5.2 kg/ha, RSE = 12%). Alewife ≥age-1 comprised 96% of alewife biomass. In 2013, alewife comprised 86% of total prey fish biomass, while rainbow smelt and bloater were 4 and 10% of total biomass, respectively. Rainbow smelt biomass in 2013 (0.24 kg/ha, RSE = 17%) was essentially identical to the rainbow smelt biomass in 2012 and was 6% of the long term mean. Bloater biomass in 2013 was 0.6 kg/ha, only half the 2012 biomass, and 6% of the long term mean. Mean density of small bloater in 2013 (29 fish/ha, RSE = 29%) was lower than peak values observed in 2007-2009 and was 23% of the time series mean. In 2013, pelagic prey fish biomass in Lake Michigan was similar to Lake Huron, but pelagic community composition differs in the two lakes, with Lake Huron dominated by bloater.

Lake Michigan

Status and trends of pelagic prey fish in Lake Huron, 2013

The USGS Great Lakes Science Center (GLSC) conducted acoustic and midwater trawl surveys of Lake Huron during 1997 and annually during 2004-2013. The 2013 survey was conducted during September and October and included transects in Lake Huron’s main basin, Georgian Bay, and North Channel. Pelagic fish density was 1,033 fish/ha in 2013 and increased 62% over the 2012 estimate. Total biomass in 2013 (6.07 kg/ha) was similar to the 2012 estimate (6.97 kg/ha). Mean numeric density of alewife Alosa pseudoharengus was substantially greater in 2013 than in 2012, but the 2013 estimate has low precision. Age-0 rainbow smelt Osmerus mordax abundance increased from 2012, whereas age-1+ rainbow smelt decreased. Age-0 bloater Coregonus hoyi abundance increased over 2012 estimates. Density and biomass of large bloater in 2013 was similar to 2012 levels. Emerald shiner Notropis atherinoides density and biomass increased during 2013. Two adult cisco Coregonus artedi were captured in Georgian Bay. Based on comparable biomass estimates during 2012 and 2013, prey fish availability during 2014 will likely be similar to 2013. Lake Huron has pelagic fish biomass similar to that observed in recent lake-wide acoustic surveys of Lake Michigan and Lake Superior, but species composition differs in the three lakes. There is an increasing gradient of diversity and native species occurrence from Lake Michigan to Lake Superior, with Lake Huron being intermediate in the prevalence of native fish species like coregonines and emerald shiner.

Lake Huron

Lake Michigan offshore ecosystem structure and food web changes from 1987 to 2008

Ecosystems undergo dynamic changes owing to species invasions, fisheries management decisions, landscape modifications, and nutrient inputs. At Lake Michigan, new invaders (e.g., dreissenid mussels ( Dreissena spp.), spiny water flea ( Bythotrephes longimanus ), round goby ( Neogobius melanostomus )) have proliferated and altered energy transfer pathways, while nutrient concentrations and stocking rates to support fisheries have changed. We developed an ecosystem model to describe food web structure in 1987 and ran simulations through 2008 to evaluate changes in biomass of functional groups, predator consumption, and effects of recently invading species. Keystone functional groups from 1987 were identified as Mysis , burbot ( Lota lota ), phytoplankton, alewife ( Alosa pseudoharengus ), nonpredatory cladocerans, and Chinook salmon (Oncorhynchus tshawytscha ). Simulations predicted biomass reductions across all trophic levels and predicted biomasses fit observed trends for most functional groups. The effects of invasive species (e.g., dreissenid grazing) increased across simulation years, but were difficult to disentangle from other changes (e.g., declining offshore nutrient concentrations). In total, our model effectively represented recent changes to the Lake Michigan ecosystem and provides an ecosystem-based tool for exploring future resource management scenarios.

Lake Michigan

Changing ecosystem dynamics in the Laurentian Great Lakes: Bottom-up and top-down regulation

Understanding the relative importance of top-down and bottom-up regulation of ecosystem structure is a fundamental ecological question, with implications for fisheries and water-quality management. For the Laurentian Great Lakes, where, since the early 1970s, nutrient inputs have been reduced, whereas top-predator biomass has increased, we describe trends across multiple trophic levels and explore their underlying drivers. Our analyses revealed increasing water clarity and declines in phytoplankton, native invertebrates, and prey fish since 1998 in at least three of the five lakes. Evidence for bottom-up regulation was strongest in Lake Huron, although each lake provided support in at least one pair of trophic levels. Evidence for top-down regulation was rare. Although nonindigenous dreissenid mussels probably have large impacts on nutrient cycling and phytoplankton, their effects on higher trophic levels remain uncertain. We highlight gaps for which monitoring and knowledge should improve the understanding of food-web dynamics and facilitate the implementation of ecosystem-based management.

Illinois, Indiana, Michigan, Minnesota, new York,

Chinook salmon foraging patterns in a changing Lake Michigan

Since Pacific salmon stocking began in Lake Michigan, managers have attempted to maintain salmon abundance at high levels within what can be sustained by available prey fishes, primarily Alewife Alosa pseudoharengus . Chinook Salmon Oncorhynchus tshawytscha are the primary apex predators in pelagic Lake Michigan and patterns in their prey selection (by species and size) may strongly influence pelagic prey fish communities in any given year. In 1994–1996, there were larger Alewives, relatively more abundant alternative prey species, fewer Chinook Salmon, and fewer invasive species in Lake Michigan than in 2009–2010. The years 2009–2010 were instead characterized by smaller, leaner Alewives, fewer alternative prey species, higher abundance of Chinook Salmon, a firmly established nonnative benthic community, and reduced abundance of Diporeia , an important food of Lake Michigan prey fish. We characterized Chinook Salmon diets, prey species selectivity, and prey size selectivity between 1994–1996 and 2009–2010 time periods. In 1994–1996, Alewife as prey represented a smaller percentage of Chinook Salmon diets than in 2009–2010, when alewife comprised over 90% of Chinook Salmon diets, possibly due to declines in alternative prey fish populations. The size of Alewives eaten by Chinook Salmon also decreased between these two time periods. For the largest Chinook Salmon in 2009–2010, the average size of Alewife prey was nearly 50 mm total length shorter than in 1994–1996. We suggest that changes in the Lake Michigan food web, such as the decline in Diporeia , may have contributed to the relatively low abundance of large Alewives during the late 2000s by heightening the effect of predation from top predators like Chinook Salmon, which have retained a preference for Alewife and now forage with greater frequency on smaller Alewives.

Transactions of the American Fisheries Society

Sensitivity of fish density estimates to standard analytical procedures applied to Great Lakes hydroacoustic data

Standardized methods of data collection and analysis ensure quality and facilitate comparisons among systems. We evaluated the importance of three recommendations from the Standard Operating Procedure for hydroacoustics in the Laurentian Great Lakes (GLSOP) on density estimates of target species: noise subtraction; setting volume backscattering strength (S v ) thresholds from user-defined minimum target strength (TS) of interest (TS-based S v threshold); and calculations of an index for multiple targets (N v index) to identify and remove biased TS values. Eliminating noise had the predictable effect of decreasing density estimates in most lakes. Using the TS-based Sv threshold decreased fish densities in the middle and lower layers in the deepest lakes with abundant invertebrates (e.g., Mysis diluviana). Correcting for biased in situ TS increased measured density up to 86% in the shallower lakes, which had the highest fish densities. The current recommendations by the GLSOP significantly influence acoustic density estimates, but the degree of importance is lake dependent. Applying GLSOP recommendations, whether in the Laurentian Great Lakes or elsewhere, will improve our ability to compare results among lakes. We recommend further development of standards, including minimum TS and analytical cell size, for reducing the effect of biased in situ TS on density estimates.

Great Lakes

Evaluating analytical approaches for estimating pelagic fish biomass using simulated fish communities

Pelagic fish assessments often combine large amounts of acoustic-based fish density data and limited midwater trawl information to estimate species-specific biomass density. We compared the accuracy of five apportionment methods for estimating pelagic fish biomass density using simulated communities with known fish numbers that mimic Lakes Superior, Michigan, and Ontario, representing a range of fish community complexities. Across all apportionment methods, the error in the estimated biomass generally declined with increasing effort, but methods that accounted for community composition changes with water column depth performed best. Correlations between trawl catch and the true species composition were highest when more fish were caught, highlighting the benefits of targeted trawling in locations of high fish density. Pelagic fish surveys should incorporate geographic and water column depth stratification in the survey design, use apportionment methods that account for species-specific depth differences, target midwater trawling effort in areas of high fish density, and include at least 15 midwater trawls. With relatively basic biological information, simulations of fish communities and sampling programs can optimize effort allocation and reduce error in biomass estimates.

Canadian Journal of Fisheries and Aquatic Sciences

Relationship between mid-water trawling effort and catch composition uncertainty in two large lakes (Huron and Michigan) dominated by alosines, osmerids, and coregonines

Because it is not possible to identify species with echosounders alone, trawling is widely used as a method for collecting species and size composition data for allocating acoustic fish density estimates to species or size groups. In the Laurentian Great Lakes, data from midwater trawls are commonly used for such allocations. However, there are no rules for how much midwater trawling effort is required to adequately describe species and size composition of the pelagic fish communities in these lakes, so the balance between acoustic sampling effort and trawling effort has been unguided. We used midwater trawl data collected between 1986 and 2008 in lakes Michigan and Huron and a variety of analytical techniques to develop guidance for appropriate levels of trawl effort. We used multivariate regression trees and re-sampling techniques to i. identify factors that influence species and size composition of the pelagic fish communities in these lakes, ii. identify stratification schemes for the two lakes, iii. determine if there was a relationship between uncertainty in catch composition and the number of tows made, and iv. predict the number of tows required to reach desired uncertainty targets. We found that depth occupied by fish below the surface was the most influential explanatory variable. Catch composition varied between lakes at depths <38.5 m below the surface, but not at depths ≥38.5 m below the surface. Year, latitude, and bottom depth influenced catch composition in the near-surface waters of Lake Michigan, while only year was important for Lake Huron surface waters. There was an inverse relationship between RSE [relative standard error = 100 × (SE/mean)] and the number of tows made for the proportions of the different size and species groups. We found for the fifth (Lake Huron) and sixth (Lake Michigan) largest lakes in the world, 15–35 tows were adequate to achieve target RSEs (15% and 30%) for ubiquitous species, but rarer species required much higher, and at times, impractical effort levels to reach these targets.

Fisheries Research

Influences on Bythotrephes longimanus life-history characteristics in the Great Lakes

We compared Bythotrephes population demographics and dynamics to predator (planktivorous fish) and prey (small-bodied crustacean zooplankton) densities at a site sampled through the growing season in Lakes Michigan, Huron, and Erie. Although seasonal average densities of Bythotrephes were similar across lakes (222/m 2 Erie, 247/m 2 Huron, 162/m 2 Michigan), temporal trends in abundance differed among lakes. In central Lake Erie where Bythotrephes ' prey assemblage was dominated by small individuals (60%), where planktivorous fish densities were high (14,317/ha), and where a shallow water column limited availability of a deepwater refuge, the Bythotrephes population was characterized by a small mean body size, large broods with small neonates, allocation of length increases mainly to the spine rather than to the body, and a late summer population decline. By contrast, in Lake Michigan where Bythotrephes ' prey assemblage was dominated by large individuals (72%) and planktivorous fish densities were lower (5052/ha), the Bythotrephes population was characterized by a large mean body size (i.e., 37–55% higher than in Erie), small broods with large neonates, nearly all growth in body length occurring between instars 1 and 2, and population persistence into fall. Life-history characteristics in Lake Huron tended to be intermediate to those found in Lakes Michigan and Erie, reflecting lower overall prey and predator densities (1224/ha) relative to the other lakes. Because plasticity in life history can affect interactions with other species, our findings point to the need to understand life-history variation among Great Lakes populations to improve our ability to model the dynamics of these ecosystems.

Great Lakes

Status of pelagic prey fishes in Lake Michigan, 2012

Acoustic surveys were conducted in late summer/early fall during the years 1992-1996 and 2001-2012 to estimate pelagic prey fish biomass in Lake Michigan. Midwater trawling during the surveys as well as target strength provided a measure of species and size composition of the fish community for use in scaling acoustic data and providing species-specific abundance estimates. The 2012 survey consisted of 26 acoustic transects (576 km total) and 31 midwater tows. Mean total prey fish biomass was 6.4 kg/ha (relative standard error, RSE = 15%) or 31 kilotonnes (kt = 1,000 metric tons), which was 1.5 times the estimate for 2011 and 22% of the long-term mean. The increase from 2011 resulted from increased biomass of age-0 alewife, age-1 or older alewife, and large bloater. The abundance of the 2012 alewife year class was similar to the average, and this year-class contributed 35% of total alewife biomass (4.9 kg/ha, RSE = 17%), while the 2010 alewife year-class contributed 58%. The 2010 year class made up 89% of age-1 or older alewife biomass. In 2012, alewife comprised 77% of total prey fish biomass, while rainbow smelt and bloater were 4 and 19% of total biomass, respectively. Rainbow smelt biomass in 2012 (0.25 kg/ha, RSE = 17%) was 40% of the rainbow smelt biomass in 2011 and 5% of the long term mean. Bloater biomass was much lower (1.2 kg/ha, RSE = 12%) than in the 1990s, and mean density of small bloater in 2012 (191 fish/ha, RSE = 24%) was lower than peak values observed in 2007-2009. In 2012, pelagic prey fish biomass in Lake Michigan was similar to Lake Superior and Lake Huron. Prey fish biomass remained well below the Fish Community Objectives target of 500-800 kt, and key native species remain absent or rare.

Lake Michigan

Status and trends of pelagic prey fishes in Lake Huron, 2012

The USGS Great Lakes Science Center (GLSC) conducted acoustic/midwater trawl surveys of Lake Huron during 1997 and annually during 2004-2012. The 2012 survey was conducted during September and October, and included transects in Lake Huron’s Main Basin, Georgian Bay, and North Channel. Pelagic fish density (638 fish/ha) was lower in 2012 compared to 2011, with density in 2012 only 34% of 2011. Total biomass in 2012 was 74% of the 2011 value. Alewife Alosa pseudoharengus remained nearly absent, and only one cisco Coregonus artedi was captured. Rainbow smelt Osmerus mordax density was only 31% of the 2011 density. Bloater Coregonus hoyi density was less than half the 2011 density, mostly as a result of lower density of small bloater. Density and biomass of large bloater in 2012 were similar to 2011 levels. During 2012 we observed significantly higher fish biomass in North Channel than in the Main Basin or Georgian Bay. Prey availability during 2013 will likely be similar to 2012. Lake Huron now has pelagic fish biomass similar to that observed in recent lakewide acoustic surveys of Lake Michigan and Lake Superior, but species composition differs in the three lakes. There is an increasing diversity and prevalence of native species gradient from Lake Michigan to Lake Superior, with Lake Huron being intermediate in the prevalence of native fish species like coregonines and emerald shiner Notropis atherinoides.

Report

Resurgence of emerald shiners Notropis atherinoides in Lake Huron's main basin

Emerald shiners Notropis atherinoides were formerly common in Lakes Huron and Michigan, but declined during the 1960s as the exotic alewife Alosa pseudoharengus proliferated. The Lake Huron emerald shiner population was chronically depressed through 2004; however, we detected resurgence in emerald shiner density and biomass in Lake Huron during acoustic and midwater trawl surveys conducted during 2004–2006. Emerald shiners were not found during 2004, but by 2006 main basin density exceeded 500 fish/ha, biomass estimates exceeded 0.5 kg/ha, and emerald shiners contributed more to pelagic biomass than alewives or rainbow smelt Osmerus mordax . Length frequency distributions suggested that increased density was the result of two consecutive strong year classes in 2005 and 2006. Emerald shiner distributions also expanded from a focus in western Lake Huron in 2005 to a lakewide distribution in 2006. Emerald shiners occurred offshore, but were nearly always associated with epilimnetic surface waters warmer than 19°C. Resurgence of emerald shiners was likely a consequence of reduced alewife abundance, as they declined concurrently with alewife proliferation during the early 1960s. Return of this species may benefit native nearshore piscivores; however, benefits to Pacific salmonids Oncorhynchus spp. are uncertain because emerald shiners are smaller and still less abundant than historically important prey species, and they may be thermally segregated from salmonines.

Journal of Great Lakes Research

Changes in seasonal nearshore zooplankton abundance patterns in Lake Ontario following establishment of the exotic predator Cercopagis pengoi

Cercopagis pengoi, a zooplanktivore first discovered in Lake Ontario in 1998, may reduce availability of prey for planktivorous fish. Cercoapgis pengoi is most abundant in late summer and fall. Therefore, we hypothesized that abundance of small zooplankton (bosminids and cyclopoids) species would decrease at that time. To determine if the establishment of C. pengoi was followed by changes in the zooplankton community, seasonal patterns in nearshore zooplankton collected from May to October 1995&ndash;2000 were examined. Early summer density of small zooplankton was similar in all years while late summer and fall densities were significantly lower in 1998&ndash;2000 than in 1995&ndash;1997. The declines of small zooplankton coincided seasonally with the peak in C. pengoi density. Other possible causes for the observed changes in small zooplankton are less likely. High levels of fish predation should have resulted in smaller zooplankton in 1998&ndash;2000 than in 1995&ndash;1997 and larger declines in Daphnia than other groups. This was not observed. There was no significant decline in chlorophyll-a concentrations or changes in temperature between 1995&ndash;1997 and 1998&ndash;2000. Therefore, the declines in density of small zooplankton were most likely the result of C. pengoi predation. The effect of C. pengoi establishment on alewives is increased competition for zooplankton prey but C. pengoi has replaced a portion of the zooplankton biomass and adult alewife diet formerly dominated by Diacyclops thomasi and Bosmina longirostris .

Journal of Great Lakes Research