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Spatial and temporal genetic diversity of lake whitefish (Coregonus clupeaformis (Mitchill)) from Lake Huron and Lake Erie

Lake whitefish (Coregonus clupeaformis (Mitchill)) are important commercially, culturally, and ecologically in the Laurentian Great Lakes. Stocks of lake whitefish in the Great Lakes have recovered from low levels of abundance in the 1960s. Reductions in abundance, loss of habitat and environmental degradation can be accompanied by losses of genetic diversity and overall fitness that may persist even as populations recover demographically. Therefore, it is important to be able to identify stocks that have reduced levels of genetic diversity. In this study, we investigated patterns of genetic diversity at microsatellite DNA loci in lake whitefish collected between 1927 and 1929 (historical period) and between 1997 and 2005 (contemporary period) from Lake Huron and Lake Erie. Genetic analysis of lake whitefish from Lakes Huron and Erie shows that the amount of population structuring varies from lake to lake. Greater genetic divergences among collections from Lake Huron may be the result of sampling scale, migration patterns and demographic processes. Fluctuations in abundance of lake whitefish populations may have resulted in periods of increased genetic drift that have resulted in changes in allele frequencies over time, but periodic genetic drift was not severe enough to result in a significant loss of genetic diversity. Migration among stocks may have decreased levels of genetic differentiation while not completely obscuring stock boundaries. Recent changes in spatial boundaries to stocks, the number of stocks and life history characteristics of stocks further demonstrate the potential of coregonids for a swift and varied response to environmental change and emphasise the importance of incorporating both spatial and temporal considerations into management plans to ensure that diversity is preserved.

Lake Erie, Lake Huron

Morphometric variation among spawning cisco aggregations in the Laurentian Great Lakes: are historic forms still present?

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.

Lake Michigan;Lake Erie;Lake Ontario;Lake Huron;La

Export of pelagic fish larvae from a large Great Lakes connecting channel

The St. Clair-Detroit River System is located in the heart of the North American Laurentian Great Lakes, connecting lakes Huron and Erie, contributing over 90% of the inflow to Lake Erie, and providing spawning habitat for many fishes including walleye (Sander vitreus), yellow perch (Perca flavescens), and lake whitefish (Coregonus clupeaformis). Dredging and channelization have greatly altered the Detroit River thereby reducing available spawning habitat and redirecting a majority of river discharge through deep canal-like channels to offshore areas of western Lake Erie at high velocities. Fish reproduction has been measured in the Detroit River using larval surveys for over 10 years. However, quantitative estimates of contributions from spawning in the system to Lake Erie fish populations have not been determined. We use long-term biological sampling and physical habitat surveys to characterize larval drift patterns and a Bayesian approach to quantify larval export from the Detroit River. Larval export from the Detroit River into Lake Erie varied by species, across years (2006–2015), and spatially among the multiple navigational channels in the lower Detroit River. Total annual export of walleye larvae ranged from 12 million in 2013 to 114 million in 2014, yellow perch larvae ranged from 319 million in 2014 to 690 million in 2013, and lake whitefish larvae ranged from 29 million in 2010 to 84 million in 2011. Given the widespread spawning, large numbers of larvae produced, and continued system-wide water quality and aquatic habitat improvements, the Detroit River provides valuable habitat and added resilience to Lake Erie fisheries despite suffering from severe anthropogenic disturbances.

Michigan, Ontario

Pelagic forage versus abiotic factors as drivers of walleye growth in northern Wisconsin lakes

Understanding ecological relationships among fishes and their environments are important for informing management policies. We conducted a statewide assessment of cisco (Coregonus artedi) in inland lakes of Wisconsin to better understand the status of this pelagic, coldwater forage fish. We then used long-term (2005–2014), standardized walleye (Sander vitreus) survey data from the Ceded Territory of Wisconsin (CTWI) to test for the influence of cisco (present, extirpated, or never present) and several abiotic factors on walleye growth trajectories described using sex-specific asymptotic lengths (L∞), Brody growth coefficients (K), and time in years required to attain common length limits used to manage harvest of walleye in the recreational fishery (381 and 457 mm). Despite being top predators in many north-temperate waters, walleye growth was highly variable among lakes, suggesting that forage base and abiotic factors may be important drivers. Growth characteristics of 160 CTWI walleye populations revealed that females reached greatest L∞ in lakes with cisco compared to those where cisco were never present or those lakes where cisco have been extirpated; however, differences were not statistically significant. Male walleye L∞ did not differ based on cisco presence. Brody growth coefficients (K) for female walleye were positively correlated with growing degree days and Secchi depth; K for males was positively correlated with Secchi depth. Average time to attain 381 and 457 mm were lowest in lakes where cisco have been extirpated. Our results suggest that cooler water temperatures and lower water clarity may be more important drivers of walleye maximum growth potential in northern Wisconsin lakes than the presence of cisco.

Wisconsin

Short-term survival of lake whitefish following surgical implantation of acoustic transmitters using chemical anesthesia and electroimmobilization

The recreational, tribal, and commercial importance of lake whitefish ( Coregonus clupeaformis ) has prompted interest in conducting large-scale telemetry studies to evaluate movement patterns, stock structure, and spatial distribution of fish relative to fishing effort in the Laurentian Great Lakes. However, there is a lack of knowledge concerning the postoperative effects of intracoelomic transmitter implantation and the use of different anesthetic agents and immobilization techniques during surgery on the survival of lake whitefish. Consequently, we evaluated 48-h survival of lake whitefish following intracoelomic implantation of acoustic transmitters using chemical anesthesia (i.e., Aqui-S 20E®) and electroimmobilization through a series of field surgery trials during November 2016 and 2017. Lake whitefish were collected using electrofishing and assigned to treatment groups including: 1) a loop tag; 2) a loop tag and acoustic transmitter with no anesthesia or immobilization; 3) a loop tag and transmitter using Aqui-S 20E®; 4) a loop tag and transmitter using electroimmobilization; and 5) reference fish. In treatments involving anesthesia or immobilization, induction and recovery times were recorded, and all fish were monitored for postoperative mortality. After 48 h, all fish that underwent anesthesia or immobilization during surgery were alive, but 4 of 25 (16%) fish that underwent surgery without anesthesia or immobilization died within 48 h. Induction and recovery times were substantially shorter for fish immobilized with electricity (induction time: instantaneous; recovery time < 20 s) than for those anesthetized with Aqui-S 20E® (induction time > 200 s; recovery time > 500 s). Results indicate that survival of lake whitefish following surgical implantation of acoustic transmitters was high when surgeries were conducted with Aqui-S 20E® or electroimmobilization. Compared to using Aqui-S 20E®, electroimmobilization can provide shorter induction and recovery times and may be a preferred option for field-based tagging activities with limitations on time and fish holding capacity.

Great Lakes

Spatial extent of contemporary lake whitefish spawning in western Lake Erie

Degradation of lake whitefish (Coregonus clupeaformis) spawning areas in the Detroit River, Maumee Bay, and western Lake Erie reefs and shoals has been identified as a contributing factor to Lake Erie’s population collapse in the 1950s. This decline prompted the United States and Canada to take steps to improve the aquatic ecosystem of Lake Erie. A recent increase in commercial fish harvest and catch of early-life stages of lake whitefish in the western basin of Lake Erie provide evidence that lake whitefish are once again spawning in some of their historical spawning areas, however the extent of use is unknown. To investigate the contemporary distribution of lake whitefish spawning within western Lake Erie, 31 potential spawning locations were sampled for eggs in 2016 and 2017. A subset of sites within Maumee Bay and an open lake reef complex were sampled repeatedly each year to determine the onset of spawning and 17 sites outside these two areas were each sampled once to describe the spatial extent of spawning. Spawned eggs were first detected on 22 November in 2016 and on 18 November in 2017. Viable eggs were collected at 27 of 31 sampled locations, verifying that lake whitefish spawned in Maumee Bay, on the mid-lake reefs, and other locations in 2016 and 2017. These findings confirm that lake whitefish are using historical spawning areas and available spawning habitat in western Lake Erie. The extensive use of available spawning habitat adds spawning stock diversity that is vital for generating population stability and resilience.

Lake Erie

Lake Ontario cisco population dynamics based on long-term surveys

Prior to European settlement, cisco (Coregonus artedi) were likely one of Lake Ontario’s most abundant fishes but currently represent a small portion of the fish community. To understand how the population has changed over the past 70 years we compared trends in annual catch rates from gillnet and bottom trawl surveys and commercial fishery landings. In surveys, cisco were generally rare, and represented 0.2, 0.4, and 0.001% of all fish caught in two gillnet surveys and bottom trawl surveys. Cisco catch rates in gillnets and trawls were positively correlated and correlations increased when gillnet catches two years later were compared to trawls since trawls tended to capture smaller, juvenile-sized cisco relative to gillnets. Survey catch rates suggest recruitment is generally low, but discrete periods of relatively greater recruitment in the 1980s and mid-2010s suggest reproductive conditions for cisco vary temporally. Trawl surveys were the most spatially extensive survey and illustrated catch rates were highest in northeastern Lake Ontario. Greater cisco abundance in this region may be related to more-abundant embayment spawning habitat, greater distance from winter aggregations of nonnative planktivores, or more appropriate environmental conditions during spawing. At the basin scale, Lake Ontario bottom trawl catch per unit effort (CPUE) was positively correlated to Lake Superior trawl CPUE suggesting a regional driver, such as climate, may be similarly impacting both populations. Concurrent patterns across Lake Ontario surveys support the idea that cisco are currently a small portion of the fish community, recruitment remains inconsistent, and habitats in northeastern Lake Ontario appear critical to the remnant populations.

Lake Ontario

Historic coregonine habitat use and assessment of larval nursery locations in Lake Erie

Coregonine fishes (Coregonus spp.) are important components of Great Lake food webs and support lucrative commercial and recreational fisheries. Due to a combination of several factors including habitat loss, over-exploitation, and introduction of exotic species, the distribution and abundance of coregonines have been reduced. Examples of these declines are evident in Lake Erie where cisco (C. artedi) have been nearly extirpated, and lake whitefish (C. clupeaformis), while still abundant, are declining. To identify key habitat locations of coregonines in Lake Erie, we conducted a literature review of historical spawning, nursery, and adult habitat sites where coregonines have been observed. We used these sites as a reference for larval sampling at six locations across the southern shore of Lake Erie during spring 2017. Paired bongo samplers were used to collect larvae, and average densities were calculated for comparison across sites. Larval coregonine (46 visually identified as lake whitefish; 8 classified as coregonines) densities were highest at Huron, OH (0.880/1,000 m3 ± 1.61), followed by Sandusky, OH (0.426/1,000 m3 ± 1.05), Dunkirk, NY (0.208/1,000 m3 ± 0.703), Fairport, OH (0.185/1,000 m3 ± 0.680), Erie, PA (0.120/1,000 m3 ± 0.532), and Conneaut, OH (0.1196/1,000 m3 ± 0.528). Using contemporary sampling data coupled with historical spawning locations, we identified sites that are currently being used as nursery locations by lake whitefish. By validating the contemporary use of historic spawning and nursery sites, this study identifies locations where habitat protection and restoration or future stocking of coregonids could be conducted in Lake Erie in efforts to recover populations and improve fishery production.

Lake Erie

Genetic structure of lake whitefish, Coregonus clupeaformis, populations in the northern main basin of Lake Huron

Genetic analysis of spawning lake whitefish ( Coregonus clupeaformis ) from six sites in the main basin of Lake Huron was conducted to determine population structure. Samples from fisheryindependent assessment surveys in the northwest main basin were analyzed to determine the relative contributions of lake whitefish genetic populations. Genetic population structure was identified using data from seven microsatellite DNA loci. One population was identified at Manitoulin Island, one to two were observed in the east-central main basin (Fishing Island and Douglas Point), and one to two populations were found in the northwest (Thunder Bay and Duncan Bay). The genetic identity of collections from Duncan Bay and Thunder Bay was not consistent among methods used to analyze population structure. Low genetic distances suggested that they comprised one population, but genic differences indicated that they may constitute separate populations. Simulated data indicated that the genetic origins of samples from a mixed-fishery could be accurately identified, but accuracy could be improved by incorporating additional microsatellite loci. Mixture analysis and individual assignment tests performed on mixed-stock samples collected from the western main basin suggested that genetic populations from the east-central main basin contributed less than those from the western main basin and that the proportional contribution of each baseline population was similar in each assessment sample. Analysis of additional microsatellite DNA loci may be useful to help improve the precision of the estimates, thus increasing our ability to manage and protect this valuable resource.

Michigan, Ontario

Successional change in the Lake Superior fish community: Population trends in ciscoes, rainbow smelt, and lake trout, 1958-2008

The Lake Superior fish community underwent massive changes in the second half of the 20th century. Those changes are largely reflected in changes in abundance of the adults of principal prey species, the ciscoes ( Coregonus spp. ), the invasive rainbow smelt ( Osmerus mordax ), and the principal predator, lake trout ( Salvelinus namaycush ). To better understand changes in species abundances, a comprehensive series of gillnet and bottom trawl data collected from 1958 to 2008 were examined. In the late 1950s/early 1960s, smelt abundance was at its maximum, wild lake trout was at its minimum, and an abundance of hatchery lake trout was increasing rapidly. The bloater ( Coregonus hoyi ) was the prevalent cisco in the lake; abundance was more than 300% greater than the next most abundant cisco, shortjaw cisco ( C. zenithicus ), followed by kiyi ( C. kiyi ) and lake cisco ( C. artedi ). By the mid-1960s, abundance of hatchery lake trout was nearing maximum, smelt abundance was beginning to decline, and abundances of all ciscoes declined, but especially that of shortjaw cisco and kiyi. By the late 1970s, recovery of wild lake trout stocks was well underway and abundances of hatchery lake trout and smelt were declining and the ciscoes were reaching their nadir. During 1980–1990, the fish community underwent a dramatic shift in organization and structure. The rapid increase in abundance of wild lake trout, concurrent with a rapid decline in hatchery lake trout, signaled the impending recovery. Rainbow smelt abundance dropped precipitously and within four years, lake cisco and bloater populations rebounded on the heels of a series of strong recruitment events. Kiyi populations showed signs of recovery by 1989, and shortjaw by 2000, though well below historic maximum abundances. High abundance of adult smelt prior to 1980 appears to be the only factor linked to recruitment failure in the ciscoes. Life history traits of the cisco species were examined to better understand their different responses to conditions of low and high predator levels, i.e., late 1950s–early 1960s vs. post 1980. Bloaters are most likely to become the predominant cisco in the absence of strong predation and the least abundant under prolonged predation; smelt share this pattern. Conversely, the lake cisco and shortjaw cisco fare better when predator abundance is high. The recovery of lake trout in Lake Superior reestablished a strong top-down influence on the fish community and its present structure and organization appears to be approaching an equilibrium that reflects a more natural state. If lake trout recovery is sustained, shortjaw cisco abundance is expected to increase and join lake cisco and kiyi as dominant cisco species, and bloater and smelt will oscillate at lower abundances.

Lake Superior

Can migration mitigate the effects of ecosystem change? Patterns of dispersal, energy acquisition and allocation in Great Lakes lake whitefish (Coregonus clupeaformis)

Migration can be a behavioural response to poor or declining home range habitat quality and can occur when the costs of migration are overcome by the benefi ts of encountering higher-quality resources elsewhere. Despite dramatic ecosystem-level changes in the benthic food web of the Laurentian Great Lakes since the colonization of dreissenid mussels, coincident changes in condition and growth rates among benthivorous lake whitefi sh populations have been variable. We hypothesized that this variation could be in part mitigated by differences in migratory habits among populations, where increased migration distance can result in an increased probability of encountering high-quality habitat (relative to the home range). Results from four Great Lakes populations support this hypothesis; relative growth rates increased regularly with migration distance. The population with the largest average migration distance also had the least reduction in size-at-age during a period of signifi cant ecosystem change and among the highest estimated consumption and activity rates. In comparison, the population with the greatest declines in size-at-age was among the least mobile, demonstrating only moderate rates of consumption and activity. The least mobile population of lake whitefi sh was supported by a remnant Diporeia population and has experienced only moderate temporal growth declines. Our study provides evidence for the potential role of migration in mitigating the effects of ecosystem change on lake whitefi sh populations.

Great Lakes

Life history characteristics of a recovering lake whitefish Coregonus clupeaformis stock in the Detroit River, North America

The Detroit River is part of a channel connecting Lakes Huron and Erie and was once a prolific spawning area for lake whitefish, Coregonus clupeaformis . Large numbers of lake whitefish migrated into the river to spawn where they were harvested by commercial fisheries and for fish culture operations. Prior to our study, the last lake whitefish was landed from the Detroit River in 1925. Loss of spawning habitat during shipping channel construction and over-fishing, likely reduced lake whitefish spawning runs. Because lake whitefish are recovering in Lake Erie, and spawning in the western basin, we suspected they may also be spawning in the Detroit River. We sampled in the Detroit River for lake whitefish adults and eggs in October–December 2005–07 and for larvae during March–May 2006–08. A total of 15 spawning-ready lake whitefish from 4 to 18 years old, were collected. Viable eggs were collected during mid-November 2006–07; highest egg densities were found mid-river. Sac-fry whitefish larvae were collected in the river and near the river mouth. No whitefish larvae were retained in the river. Because high numbers of larvae were collected from mid- and downstream river sites, reproduction of lake whitefish in the Detroit River could contribute substantially to the Lake Erie lake whitefish metapopulation.

Michigan, Ontario

Repeat surveys of spawning cisco (Coregonus artedi) in western Lake Superior: Timing, distribution and composition of spawning stocks

Acoustic (AC) and midwater trawl (MT) surveys of spawning cisco ( Coregonus artedi ) in Lake Superior have been combined with commercial yield to estimate exploitation. To time surveys properly, it is important to understand when adults typically arrive at spawning grounds and how numbers change as the spawning season progresses. We conducted repeat autumn surveys during nighttime hours at coastal sites where commercial roe fisheries occur. Spawner densities increased significantly from October to mid-November, but differences measured at sites sampled from mid- to late-November were comparatively small. Spawners occupied the upper 20–30 m of the water column during mid-November before utilizing a wider range of depths by late-November. We compared repeat AC densities to temporal trends of catch-per-unit-effort (CPUE) in suspended commercial gillnets and found good agreement within sites. Because different gillnet mesh sizes were used in each roe fishery. CPUE and AC density were poorly correlated among sites. We recommend that future surveys be conducted between mid- and late-November, and that MT gear be used to measure cisco densities in the uppermost 10 m of the water column where AC estimates may be conservative. Given the short temporal window for assessing spawner density, we believe both AC-MT and gillnet surveys will be needed to ensure that harvest of different stocks is kept at a sustainable level.

Lake Superior

Mitochondrial DNA variability among Lake Baikal omul Coregonus autumnalis migratorius (Georgi)

Omul, Coregonus autumnalis, are a commercially important coregonine fish from Lake Baikal, Siberia, Russia. In Lake Baikal, three morphotypes recognized by fishery experts occupy different zones in the lake: they are referred to as "littoral," "pelagic," and "benthic". Expressed character divergence was supported by whole-body morphometric analysis, but it is not known whether discrete genetic differences accompany the observed morphological variation. This study was designed to assess the genetic variation of three different omul morphotypes sampled from different locations in Lake Baikal that were segregated by morphotype in multivariate analysis. We surveyed genetic variation with restriction fragment length polymorphism analysis of specific gene loci amplified with the polymerase chain reaction. Sequence variation was localized in the mitochondrial control region. Though no discrete genetic markers were found, there is evidence of reproductive segregation by geographic location that corroborates geographic variation in morphological characters.

Advances in Limnology

Population dynamics of bloaters (Coregonus hoyi) in Lake Michigan, 1973-1998

Past research has demonstrated that the recruitment pattern of Lake Michigan bloaters (Coregonus hoyi) was best represented by a Beverton-Holt function during 1977-1983. In this present study, we re-assessed relations in bloater recruitment and investigated trends in sex ratio and mean length at age as they related to recent changes in bloater abundance. Bloaters were collected from 1973 to 1997 during lake-wide bottom-trawl surveys and in western Lake Michigan from 1980 to 1998 during graded-mesh gill-net surveys. Length, weight, age, sex, and maturity of bloaters were recorded. The relative abundance of females in the population increased throughout the past 10-15 years to more than 80% in 1997. Relative abundance of females in the population was inversely related to overall abundance during periods of increasing female dominance. Female dominance may be related to an increasingly older population, which resulted from low recruitment, and to the greater longevity of female bloaters. Mean length at age decreased steadily in the 1980s with increased bloater abundance. Bloater recruitment was density dependent and followed a Ricker-type stock-recruitment relation. Future research will focus on identifying other factors that influence bloater recruitment such as the effect of female dominance.

Advances in Limnology

Diversity in shortjaw cisco (Coregonus zenithicus) in North America

Shortjaw cisco (Coregonus zenithicus) exhibit morphological variability across their geographic range in North America and could comprise more than one distinct morph or taxon. To investigate this, principal components analysis was applied to a data set that consisted of four variables from nine localities. All data were obtained from digital images of the specimens and the excised first gill arch. Confidence ellipses (95%) about the means of bivariate distributions of the principal components revealed that some populations were distinct from the others, but a continuity of overlap clouded understanding of pattern among the variation. Most populations had more and longer gillrakers than shortjaw cisco from George Lake (Manitoba) and Basswood Lake (Ontario) that had fewer and shorter gillrakers. This analysis supports the existence of a short- and few-rakered morph and a long- and many-rakered morph. However, most populations of shortjaw cisco from the Great Lakes across Canada to the Arctic share a similar morphology and likely represent a single, widespread species.

Advances in Limnology

The growth-temperature relation and preferred temperatures of juvenile lake herring

Lake herring, Coregonus artedi, were once extremely abundant in the Great Lakes where they functioned as a major trophic integrator, directly linking the planktonic crustacean resource to lake trout. Salvelinus namaycush. Lake herring populations in the Great Lakes collapsed during the middle third of the 20th century due to overfishing, degradation of critical habitat in major production areas, and interaction with exotic species. Fishery and habitat impediments to the recovery of lake herring have been removed, and it may now be possible to reestablish the species in its former habitats in the Great Lakes if adverse interactions with exotic species can be controlled within acceptable limits. To determine the potential for thermal niche overlap and adverse interaction with exotic fishes, juvenile (age-0) lake herring were held in the laboratory at 5, 10, 15, 18, and 21 C, and fed ad libitum for 54 days. The optimum temperature for growth in weight was about 14.5 C, indicating the fundamental thermal niche was 12.5-16.5 C. Fish used in the growth study were also tested in a vertical thermal gradient tank to measure their final preferendum. The final preferendum, 16.5 C, was in close agreement with the optimum temperature for growth and within the fundamental thermal niche. Both the optimum temperature for growth and the final preferendum have been used as measures of thermal niche, but this is the first time both measures were made on the same group of fish. Published information on the fundamental thermal niche, preferred temperatures, thermal habitat use, and feeding habits of alewives, rainbow smelt, and ruffe, indicates they will co-occur in spring, or summer with age-0 lake herring and that collectively they pose a predation threat to small, age-0 lake herring.

Advances in Limnology

One-hundred fundamental, open questions to integrate methodological approaches in lake ice research

The rate of technological innovation within aquatic sciences outpaces the collective ability of individual scientists within the field to make appropriate use of those technologies. The process of in situ lake sampling remains the primary choice to comprehensively understand an aquatic ecosystem at local scales; however, the impact of climate change on lakes necessitates the rapid advancement of understanding and the incorporation of lakes on both landscape and global scales. Three fields driving innovation within winter limnology that we address here are autonomous real-time in situ monitoring, remote sensing, and modeling. The recent progress in low-power in situ sensing and data telemetry allows continuous tracing of under-ice processes in selected lakes as well as the development of global lake observational networks. Remote sensing offers consistent monitoring of numerous systems, allowing limnologists to ask certain questions across large scales. Models are advancing and historically come in different types (process-based or statistical data-driven), with the recent technological advancements and integration of machine learning and hybrid process-based/statistical models. Lake ice modeling enhances our understanding of lake dynamics and allows for projections under future climate warming scenarios. To encourage the merging of technological innovation within limnological research of the less-studied winter period, we have accumulated both essential details on the history and uses of contemporary sampling, remote sensing, and modeling techniques. We crafted 100 questions in the field of winter limnology that aim to facilitate the cross-pollination of intensive and extensive modes of study to broaden knowledge of the winter period.

Water Resources Research