Given the rapidly increasing visitation levels to Fraser Island, there is increasing concern that tourist activities may threaten the long-term ecological health of the region's unique dune lakes. This project aimed to investigate the consequences of tourist use of Fraser Island's dune lakes and to develop appropriate monitoring tools and management objectives in light of the projected increases in visitation levels in the foreseeable future. The initial phase of this research aimed to identify the relative importance of some of the most popular dune lakes on the island as key destinations for tourists. Tourist surveys, in conjunction with the development of a Tourist Pressure Index (TPI), which quantifies logistic, social and natural variables, identified Lakes McKenzie, Allom and Birrabeen as the lakes most at risk from excessive tourist use. In addition, analyses of water quality in 15 lakes on Fraser Island aimed to determine the current trophic status of dune lakes on Fraser Island and the ecological implications of tourist use of these systems. Detailed comparisons of nutrient and chlorophyll a concentrations in five popular dune lakes in February 1990 (data from Arthington et al. 1990) and February 1999 suggested that productivity has increased significantly in the past decade. More detailed examinations of nutrient and algal variables in five popular perched dune lakes revealed that while ambient nutrient and phytoplankton chlorophyll a concentrations remained relatively stable, periphyton chlorophyll a concentrations increased over the course of the 1999-2000 summer in most lakes. Significantly, these increases were found only in heavily visited (disturbed) sites in the clear lakes examined (McKenzie and Birrabeen). In these lakes, where algal growth is likely to be only limited by nutrient availability, tourist nutrient additions may stimulate excessive periphyton production. Experimental algal bioassays identified that phytoplankton and to a lesser degree periphyton growth was stimulated by nutrient additions in all five perched dune lakes. However, the degree to which growth was stimulated was both lake and nutrient (nitrogen versus phosphorus versus nitrogen + phosphorus) dependent, highlighting the variable nature of systems within a relatively small geographic range. Since periphyton biomass was higher in heavily visited areas of lakes and was likely to be stimulated by nutrient additions by tourists, stable isotope analyses of littoral zone food webs were conducted to quantify the percent contribution of periphyton to consumer diets. There was a trend towards higher periphyton contributions in systems identified as key tourist locations (on the basis of their TPI scores) and this indicates that increasing visitation may increase the contribution of periphyton to littoral zone food webs, both via increases in the quantity and quality of periphyton as a food resource. To further explore the contribution of periphyton in littoral zone food webs of heavily visited lakes, a 15N-tracer addition experiment was conducted to establish the fate of nutrient additions within the littoral zone. Nutrients were added in quantities that mimicked those likely from tourists, to enable a realistic appraisal of the fate of tourist additions. As expected, periphyton rapidly assimilated the added 15N-tracer and was found to be the first and most significant sink for nutrients entering the littoral zone. Finally, the results from this research were used to develop a conceptual model of nutrient enrichment for perched dune lakes on Fraser Island. The model indicates that although nutrient additions from tourists may lead to undesirable increases in periphyton biomass, the degree to which this is deemed to be a detrimental ecological outcome is likely to be mediated by water level fluctuations and the consumptive capacity of grazers. Given that excessive periphyton growth is likely to be seen as negative impact of tourism, regular periphyton monitoring (biomass and percent contribution to littoral zone food webs) should be built into an updated monitoring program for this series of dune lakes. Whilst the implementation of periphyton monitoring is likely to enable the early detection of deleterious impacts of excessive tourist use, it is likely that the long-term conservation of the region will, in the future, require the implementation of strict visitation level guidelines, to ensure that the irreversible consequences of long term additions of nutrients are ameliorated.

Es gibt bereits viele Hinweise dafür, dass Seen sehr sensibel auf die anthropogen verursachte Klimaerwärmung reagiert haben. Bis jetzt haben sich die Studien der Klimafolgenforschung hauptsächlichst auf die Auswirkungen der Erwärmung im Winter und Frühling konzentriert. Über den Einfluss der Klimaerwärmung auf Seen in den gemäßigten Breiten im Sommer ist weniger bekannt. In der vorliegenden Doktorarbeit habe ich einige Faktoren, welche die Reaktion von Seen auf die Erwärmung im Sommer vermutlich stark mitbestimmt haben, untersucht. Der Schwerpunkt lag dabei auf klimatisch induzierten Auswirkungen auf die thermische Charakteristik und die Phänologie und Abundanz des Planktons eines flachen und polymiktischen Sees (Müggelsee, Berlin). Zuerst wurde der Einfluss der Klimaerwärmung auf die Phänologie und Abundanz des Planktons in verschiedenen Jahreszeiten untersucht. Das schnellwachsende Phyto- und Zooplankton (Daphnia) im Frühjahr hat sich vorwiegend synchron vorverschoben, wohingegen Veränderungen des Sommerzooplanktons deutlich artspezifisch und nicht synchron waren. Die Phänologie oder Abundanz einiger Sommercopepoden hat sich entsprechend der individuellen thermischen Anforderungen innerhalb bestimmter Entwicklungsstufen, wie zum Beispiel der Emergenz von der Diapause im Frühling, verändert. Die Studie unterstreicht, dass nicht nur der Grad der Erwärmung, sondern auch dessen Zeitpunkt innerhalb des Jahres von großer ökologischer Bedeutung ist. Um die Auswirkungen des Klimawandels auf die thermischen Eigenschaften des Sees zu erforschen, habe ich die Langzeitentwicklung der täglichen epilimnischen Temperaturextrema während des Sommers untersucht. Durch diese Studie wurde zum ersten Mal für Seen gezeigt, dass die täglichen epilimnischen Minima (Nacht) stärker angestiegen sind als die Maxima (Tag), wodurch sich der tägliche epilimnische Temperaturbereich deutlich verringert hat. Diese Tag-Nacht-Asymmetrie in der epilimnischen Temperatur wurde durch eine erhöhte Emission von Langwellenstrahlung aus der Atmosphäre während der Nacht verursacht. Dies unterstreicht, dass nicht nur Erhöhungen der Lufttemperatur, sondern auch Änderungen anderer meteorologischer Variablen wie der Windgeschwindigkeit, der Luftfeuchte und der Bewölkung eine wichtige Rolle bei der Bestimmung der Seetemperatur im Hinblick auf weitere Klimaveränderungen spielen werden. Zudem wurde eine Kurzzeitanalyse zum Schichtungsverhalten des polymiktischen Sees durchgeführt, um die Häufigkeit und Dauer von Schichtungsereignissen und deren Einfluss auf den gelösten Sauerstoff, die gelösten Nährstoffe und das Phytoplankton zu untersuchen. Selbst während der längsten Schichtungsereignisse (Hitzewellen 2003 und 2006) unterschieden sich die Auswirkungen auf den See von denen, welche in flachen dimiktischen Seen während der kontinuierlichen Sommerschichtung auftreten. Die hypolimnische Temperatur war höher, was die Sauerstoffzehrung und die Akkumulation von gelösten Nährstoffen begünstigt hat. Die thermische Schichtung wird in Zukunft sehr wahrscheinlich zunehmen. Dies lässt darauf schließen, dass polymiktische Seen sehr anfällig gegenüber Änderungen im Hinblick auf projizierte Klimaveränderungen sein werden. Abschließend wurde eine Studie über Lang- und Kurzzeitveränderungen in der Entwicklung der planktischen Larven der Muschel Dreissena polymorpha durchgeführt, um den Einfluss der Veränderungen im thermischen und trophischen Regime des Sees zu analysieren. Die Klimaerwärmung und die Verringerung in der externen Nährstofffracht haben die Abundanz der Larven stark beeinflusst indem sie jeweils auf bestimmte Entwicklungsphasen dieser Art während der warmen Jahreszeiten gewirkt haben. Der Anstieg in der Abundanz und der Länge der Larven stand im Zusammenhang mit dem Rückgang der Nährstofffracht und der Veränderung der Phytoplanktonzusammensetzung. Die Hitzewellen in den Jahren 2003 und 2006 haben diesen positiven Effekt auf die Larvenabundanz jedoch durch ungünstige Sauerstoffkonzentrationen während der sehr langen Schichtung aufgehoben. Die Klimaerwärmung kann demzufolge entgegenwirkende Effekte in produktiven flachen Seen, in welchen die externe Nährstofffracht reduziert wurde, auslösen. Aus diesen Ergebnissen schließe ich, dass nicht nur die Art des Klimawandels und damit der Zeitpunkt der Erwärmung und das Auftreten von Extremen wie Hitzewellen, sondern auch standortspezifische Bedingungen wie Schichtungsverhalten und Trophiegrad entscheidende Faktoren sind, welche die Auswirkungen der Klimaerwärmung auf interne Seeprozesse im Sommer bestimmen. Somit sollte sich die weiterführende Klimafolgenforschung für Seen darauf konzentrieren, wie verschiedene Seetypen auf die komplexen Umweltveränderungen im Sommer reagieren, damit ein umfassenderes Verständnis über den Einfluss von anthropogen verursachten Veränderungen auf Seen der gemäßigten Breiten erreicht wird. / There is already strong evidence that temperate lakes have been highly vulnerable to human induced climate warming during the last century. Hitherto climate impact studies have mainly focussed on the impacts of the recent long-term warming in winter and spring and little is known on the influence of climate warming on temperate lakes in summer. In the present thesis, I studied some aspects, which may have been strongly involved in determining the response of a lake to climate warming in summer. Thereby I have focussed on climate induced impacts on the thermal characteristics and the phenology and abundance of summer plankton in a shallow polymictic lake (Müggelsee, Germany). First, the influence of climate warming on the phenology and abundance of the lake plankton was investigated across seasons. Fast-growing spring phytoplankton and zooplankton (Daphnia) advanced largely synchronously, whereas long-term changes in the phenology of slow-growing summer zooplankton were clearly species-specific and not synchronised. The phenology and/or abundance of several summer copepod species changed according to their individual thermal requirements at decisive developmental stages such as emergence from diapause in spring. The study emphasises that not only the degree of warming, but also its timing within the annual cycle is of great ecological importance. To analyse the impact of climate change on the thermal characteristics of the lake, I examined the long-term development of the daily epilimnetic temperature extrema during summer. The study demonstrated for the first time for lakes that the daily epilimnetic minima (during nighttime) have increased more rapidly than the daily epilimnetic maxima (during daytime), resulting in a distinct decrease in the daily epilimnetic temperature range. This day-night asymmetry in epilimnetic temperature was likely caused by an increased nighttime emission of long-wave radiation from the atmosphere. This underlines that not only increases in air temperature, but also changes in other meteorological variables such as wind speed, relative humidity and cloud cover may play an important role in determining the lake temperature with respect to further climate change. Furthermore, a short-term analysis on the mixing regime of the polymictic lake was conducted to examine the frequency and duration of stratification events and their impacts on dissolved oxygen, dissolved nutrients and summer phytoplankton. Even during the longest stratification events (heatwaves in 2003 and 2006) the thermal characteristics of the lake differed from those typically found in shallow dimictic lakes, which exhibit a continuous stratification during summer. Particularly, hypolimnetic temperatures were higher, favouring the depletion of oxygen and the accumulation of dissolved nutrient in the hypolimnion. Thermal stratification will be very likely amplified in the future, thus, I conclude that polymictic lakes will be very vulnerable to alterations in the thermal regime with respect to projections of further climate change during summer. Finally, a long-term case study on the long and short-term changes in the development of the planktonic larvae of the freshwater mussel Dreissena polymorpha was performed to analyse the impacts of simultaneous changes in the thermal and in the trophic regime of the lake. Both the climate warming and the decrease in external nutrient load were important in determining the abundance of the pelagic larvae by affecting different features of the life-history of this species throughout the warm season. The long-term increase in the abundance and length of larvae was related to the decrease in external nutrient loading and the change in phytoplankton composition. However, the recent heatwaves in 2003 and 2006 have offset this positive effect on larval abundance, due to unfavourable low oxygen concentrations that had resulted from extremely long stratification events, mimicking the effects of nutrient enrichment. Climate warming may thus induce counteracting effects in productive shallow lakes that underwent lake restoration through a decrease in external nutrient loading. I conclude that not only the nature of climate change and thus the timing of climate warming throughout the seasons and the occurrence of climatic extremes as heatwaves, but also site-specific lake conditions as the thermal mixing regime and the trophic state are crucial factors governing the impacts of climate warming on internal lake processes during summer. Consequently, further climate impact research on lake functioning should focus on how the different lake types respond to the complex environmental forcing in summer, to allow for a comprehensive understanding of human induced environmental changes in lakes.

The Earths climate is changing at a higher rate, i.e between 1861 and 1994 the annual mean temperature in Scandinavia increased with 0,68º C and according to recent climate models the annual mean temperature is likely to rise with another 3º C during this century. A warmer climate in many ways is associated with changing conditions for lake ecosystems. An expected higher water temperature and a stronger summer stratification of the water column increases the risk of anoxic conditions at the lake bottom. Thus anoxic conditions are likely to cause a phosphate leakage from the sediment, i.e. a higher internal loading of phosphate. In this project, the extremely warm summer of 2002 has been used as an example for a possible scenario for a future climate. By comparing levels of phosphorus in the summer of 2002 with a ten-year median value, a phosphorus related sensitivity to climate change has been analyzed for 55 Swedish lakes. This sensitivity has then been related to several parameters of which in particular the lake morphometry and the land use in the catchment of the lake influenced the climatic sensitivity of the lake to climatic change. / Jordens klimat förändras i en allt snabbare takt. Mellan 1861 och 1994 steg årsmedeltemperaturen i Skandinavien med 0,68º C. Enligt aktuella klimatmodeller förväntas årsmedeltemperaturen i Skandinavien öka med ytterligare 3º C det närmaste seklet. Ett varmare klimat innebär på flera sätt nya förutsättningar för ekosystemen. Genom höjda vattentemperaturer och en starkare stratifikation sommartid ökar risken för syrefria förhållanden i sjöar. Då sedimentet under syrefria förhållanden kan läcka fosfat innebär detta en ökad internbelastning av fosfor. I detta projekt har den extremt varma sommaren 2002 använts som ett möjligt framtida klimat. Genom att jämföra fosforhalter sommaren 2002 med ett medianvärde för 10 år har den fosforrelaterade känsligheten för klimatförändringar kunnat analyseras för 55 svenska sjöar. Denna känslighet har sedan relaterats till diverse parametrar så som sjöns morfometri och avrinningsområdets sammansättning.

Phosphorus (P) is a limiting nutrient regulating productivity in both freshwater and marine ecosystems. A full knowledge of the sources and pathways of the P cycle is essential for understanding aquatic ecosystem function and for managing eutrophication. However, two significant pathways are poorly understood or remain uncharacterized. First, aquatic metazoans represent a significant internal regenerative pathway of P through the mineralization, translocation (i.e., benthic pelagic coupling) and excretion of nutrients. Rates of P excreted are expected to vary across taxa (i.e., zooplankton vs. mussels vs. benthic macroinvertebrates vs. fish), yet the significance of any one group of taxa in supplying P to bacteria and algae is unknown. Therefore, I developed the first comprehensive set of empirical models of nutrient release for aquatic metazoans (zooplankton, mussels, other benthic macroinvertebrates, and detritivorous and non-detritivorous fish) and compared inter-taxonomic differences in P excretion. I demonstrated that detritivorous fish excrete P at rates greater than all other taxa (as a function of individual organism mass); whereas, mussels generally excreted P at rates less than other taxa. Significant differences in the rate of P excretion between zooplankton and non-detritivorous fish were not observed [i.e., the allometry of P excretion was similar between zooplankton and non-detritivorous fish (as a function of individual body mass)]. I subsequently applied the models to assemblage biomass and abundance data to examine and compare the relative contribution of each taxa to the internal supply of P, and to examine the turnover time of P bound in metazoan biomass. I clearly demonstrated a hierarchy in the contribution by different metazoan assemblages to P cycling (zooplankton > benthic macroinvertebrates > mussels > fish) and clarified the significance of different metazoan taxa in P cycling. Moreover, I demonstrated that the slow turnover time of P bound in fish biomass (relative to other metazoans) indicates that fish are important as sinks rather than sources of P. A second potentially significant P pathway is through the influence of ultraviolet radiation (UVR) on P cycling. UVR may alter P cycling abiotically through changes in P availability and biotically through changes in the acquisition and regeneration of dissolved P by plankton. However, the significance of P released from the photodecomposition of dissolved organic P compounds (DOP), and the effect of UVR on the uptake and regeneration of dissolved P, the turnover of particulate P, and on ambient phosphate (PO43-) concentration has not been investigated and remains unknown. Therefore, my initial experiments applied the novel use of radiophosphate uptake assays to quantify the significance of the photodecomposition of DOP to PO43-. I concluded that the liberation of PO43- through the photodecomposition of DOP is not a significant pathway. However, the photochemical liberation of PO43- from suspended sediments was evident and should be an important pathway supplying PO43- to plankton in shallow polymictic lakes. This represents the first study to identify this P pathway in lakes. The turnover time of the PO43- pool increased under UVR irradiance (i.e., uptake of P by plankton decreased), while the regeneration rate of dissolved P and turnover rate of planktonic P were generally not affected. The net effect of UVR was an increase in steady state PO43- concentration (ssPO43-). Alkaline phosphatase activity (APA) in the dissolved and particulate fractions was significantly reduced in UVR treatments, but unrelated to changes in P uptake as proposed in the literature. This is the first study to comprehensively investigate the biotic effects of UVR on P cycling and represents a major advancement in the field of photobiology. In summary, I have characterized several poorly understood pathways in the P cycle of lakes. With the models I have developed, aquatic metazoans can now be integrated into the P cycle of lakes, for example, with other internal and external sources of P (e.g., from inlets, lake sediments and the atmosphere). This will advance our knowledge of P cycling, and will provide researchers with a better understanding of the nutrient pathways supporting primary production.

Along the Toronto shoreline, small coastal embayments (0.4 – 32 ha) are being constructed or modified to restore warmwater fish habitat. I describe how Lake Ontario (hereafter the Lake) alters the thermal regime of these small coastal embayments, how the altered thermal regimes affect growth and survival of age-0 warmwater fishes, and how the thermal habitat quality for such fishes can be improved by altering embayment design. During the warming period of the ice-free season, embayments warm faster than the Lake and so are cooled by exchanges with the Lake. Later in the year Lake exchange warms the rapidly cooling embayments, but the net effect of Lake-embayment exchange is cooling. The degree of cooling in Toronto’s small coastal embayments varies; many have temperatures near that of the Lake, and a few warm as much as local ponds. Age-0 pumpkinseed (Lepomis gibbosus) can fail to reach sufficient size to survive the winter in cooler embayments because their spawning is delayed and their growth is slowed. Most embayments along the Toronto shoreline are too cold to produce age-0 fish that can survive the winter, but all embayments are occupied by age >1 pumpkinseed, suggesting movement from warmer to cooler embayments. Using otolith microchemistry to identify natal embayments of fish, I confirm that age-0 and age-1 pumpkinseed, as well as age-0 largemouth bass (Micropterus salmoides) and age-0 yellow perch (Perca flavescens), exist in metapopulations. Embayment bathymetry is a poor predictor of temperature because almost all embayment flushing rates are very fast, usually 1-2 days. Warmer embayments are located in protected areas of Lake Ontario and receive waters that have already heated substantially. Cold embayments are located along the exposed shoreline of Lake Ontario. To protect embayments from cold lake waters, the cross-sectional area of embayment channels need to be reduced to 1-10% of their current size.

This research examines the perceptions and use of environmental indicators and state of the environment reports by local government and Conservation Authority decision makers and practitioner’s within the Ontario portion of the Great Lakes and St. Lawrence basin. Participants describe their information needs and how indicators and SOER are used at the local level; and what limitations or challenges they face to bridge the gap between monitoring information and policy. A multi-method approach including a web-based survey and follow-up telephone interviews was the primary data collection method used. Indicator and SOER knowledge and information are further explored to determine information exchange amongst different levels of governance. To review the dissemination of indicator and SOER information from a higher spatial scale down to the local level, the State of the Great Lakes environmental indicators and SOER, developed by the governments of Canada and the United States served as a case study.

This research examines the perceptions and use of environmental indicators and state of the environment reports by local government and Conservation Authority decision makers and practitioner’s within the Ontario portion of the Great Lakes and St. Lawrence basin. Participants describe their information needs and how indicators and SOER are used at the local level; and what limitations or challenges they face to bridge the gap between monitoring information and policy. A multi-method approach including a web-based survey and follow-up telephone interviews was the primary data collection method used. Indicator and SOER knowledge and information are further explored to determine information exchange amongst different levels of governance. To review the dissemination of indicator and SOER information from a higher spatial scale down to the local level, the State of the Great Lakes environmental indicators and SOER, developed by the governments of Canada and the United States served as a case study.

This study was designed to provide information about the juvenile life history of coho salmon, Oncorhynchus kisutch, in the Tenmile Lakes in Oregon by 1) classifying scales of spawning fish and outgoing smolts (1+), 2) assessing the contribution of juvenile life history types to the returns of adults (3₂) and jacks (2₂), and 3) comparing growth rates and survival of different types. Scales of outmigrating smolts from one brood year, jacks from two brood years, and scales of adults from four brood years were classified into juvenile life histories and examined for several scale characters. A linear discriminant function analysis was used to show the separation of the types and to determine the most distinguishing scale characters. Length of smolts at ocean entry back-calculated from scales of jacks and adults was compared with the length of the observed group of emigrating smolts. Based on variation in numbers and spacing of circuli and the size of the freshwater scale zone, believed to represent different residence time in the tributaries and the lakes, four types of juvenile life histories were defined. Fish classified as type 1, stream-reared, are believed to have reared in the tributaries until migrating as smolts in the following year. Type 2, stream-lake-reared, fish are thought to have reared in the tributaries for almost a year but then moved down to the lakes, where rearing continued until smolt migration in spring. Type 3, stream-lake- reared, are believed to have reared for a short time in the tributaries, then moved down to the lakes sometime in their first year of life. Rearing continued in the lakes until spring of the following year. Type 4, lake-reared, are thought to have moved down to the lakes shortly after emergence from the gravel, where they reared until migrating as smolts in the following spring. Scales of smolts, jacks, and adults were sorted into these four types. Presently, coho salmon fry and yearlings appear to be moving out of tributaries in late spring and from March to beginning of May, respectively. Migration of smolts out of the lake system to the ocean occurs mostly within the month of May. Recent habitat surveys show that dramatic seasonal changes occur in use of rearing habitat by juvenile coho salmon in the Tenmile Lakes tributaries from summer to winter. In winter juveniles appear to be using more low gradient, low velocity, off-channel areas than in summer. Fish of type 4 represented 90%, 43%, and 74% of the returns of adults in 1957- 58, 1963-64, and 1971-72, respectively, and 90% of the returns of jacks in 1962-63 and 1970-71. However, no type 4 fish was represented in the returns of adults in 1985-86, whereas type 1 fish represented 89% of the returns. Type 4 appeared to have grown better in fresh water, reached a larger size at outmigration, and have a greater relative survival than fish of type 2, when compared among the observed group of smolts, returning jacks, and adults. The large proportion of the escapement returning as jacks for some of the years may indicate good growing condition for juvenile coho salmon in fresh water. According to the classification of jack scales a large proportion of fish returning as jacks were lake-reared and were found to be larger at migration to ocean than fish returning as adults. This may suggest that fish that reared well in the lakes and reached a large size at outmigration had the tendency to mature early and return as jacks. Based on the analysis of scales, lake-reared juvenile coho salmon formerly contributed well to adult returns. The former high returns of jacks and adults reflect the importance of the lake habitat for the coho salmon populations of this system. In order to enhance this stock to a higher level, management strategies should be focused on making the lake habitat available to juvenile coho salmon for at least some part of their freshwater life. / Graduation date: 1992

Along the Toronto shoreline, small coastal embayments (0.4 – 32 ha) are being constructed or modified to restore warmwater fish habitat. I describe how Lake Ontario (hereafter the Lake) alters the thermal regime of these small coastal embayments, how the altered thermal regimes affect growth and survival of age-0 warmwater fishes, and how the thermal habitat quality for such fishes can be improved by altering embayment design. During the warming period of the ice-free season, embayments warm faster than the Lake and so are cooled by exchanges with the Lake. Later in the year Lake exchange warms the rapidly cooling embayments, but the net effect of Lake-embayment exchange is cooling. The degree of cooling in Toronto’s small coastal embayments varies; many have temperatures near that of the Lake, and a few warm as much as local ponds. Age-0 pumpkinseed (Lepomis gibbosus) can fail to reach sufficient size to survive the winter in cooler embayments because their spawning is delayed and their growth is slowed. Most embayments along the Toronto shoreline are too cold to produce age-0 fish that can survive the winter, but all embayments are occupied by age >1 pumpkinseed, suggesting movement from warmer to cooler embayments. Using otolith microchemistry to identify natal embayments of fish, I confirm that age-0 and age-1 pumpkinseed, as well as age-0 largemouth bass (Micropterus salmoides) and age-0 yellow perch (Perca flavescens), exist in metapopulations. Embayment bathymetry is a poor predictor of temperature because almost all embayment flushing rates are very fast, usually 1-2 days. Warmer embayments are located in protected areas of Lake Ontario and receive waters that have already heated substantially. Cold embayments are located along the exposed shoreline of Lake Ontario. To protect embayments from cold lake waters, the cross-sectional area of embayment channels need to be reduced to 1-10% of their current size.

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