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Cold-water lakes situated in high latitudes and altitudes have pivotal socio-ecological importance both globally and locally. However, they are increasingly threatened by multiple anthropogenic stressors, such as climate change, hydropower and invasive species. The development of efficient management strategies is therefore urgently needed and requires a comprehensive understanding of the factors influencing the biodiversity and ecological processes of these ecosystems. We provide a holistic knowledge base for informed future research and management by addressing the interplay between local and global environmental drivers of food webs in Arctic charr (Salvelinus alpinus, Salmonidae) and brown trout (Salmo trutta, Salmonidae) dominated cold-water lakes in Fennoscandia. The trophic niche and population dynamics of these generalist top consumers provide extensive insights into the effects of natural and anthropogenic drivers on food webs in intensively studied Fennoscandian cold-water lakes, covering marked biogeographical gradients in abiotic and biotic conditions. Drawing on a synthesis of existing literature, our focus is on three pivotal drivers: (1) lake location and connectivity, (2) lake area and morphometry and (3) fish community composition. These drivers significantly influence the complexity and the origin and flow of energy in lake food webs, and ultimately the size structure of the charr and trout populations. Furthermore, we highlight ongoing environmental changes in Fennoscandian cold-water lakes caused by hydropower and invasive species. Finally, we identify crucial knowledge gaps and propose management actions for improving the future state of Fennoscandian cold-water lake ecosystems and their charr and trout populations.

Cold-water lakes situated in high latitudes and altitudes have pivotal socio-ecological importance both globally and locally. However, they are increasingly threatened by multiple anthropogenic stressors, such as climate change, hydropower and invasive species. The development of efficient management strategies is therefore urgently needed and requires a comprehensive understanding of the factors influencing the biodiversity and ecological processes of these ecosystems. We provide a holistic knowledge base for informed future research and management by addressing the interplay between local and global environmental drivers of food webs in Arctic charr (Salvelinus alpinus, Salmonidae) and brown trout (Salmo trutta, Salmonidae) dominated cold-water lakes in Fennoscandia. The trophic niche and population dynamics of these generalist top consumers provide extensive insights into the effects of natural and anthropogenic drivers on food webs in intensively studied Fennoscandian cold-water lakes, covering marked biogeographical gradients in abiotic and biotic conditions. Drawing on a synthesis of existing literature, our focus is on three pivotal drivers: (1) lake location and connectivity, (2) lake area and morphometry and (3) fish community composition. These drivers significantly influence the complexity and the origin and flow of energy in lake food webs, and ultimately the size structure of the charr and trout populations. Furthermore, we highlight ongoing environmental changes in Fennoscandian cold-water lakes caused by hydropower and invasive species. Finally, we identify crucial knowledge gaps and propose management actions for improving the future state of Fennoscandian cold-water lake ecosystems and their charr and trout populations.

Cold-water lakes situated in high latitudes and altitudes have pivotal socio-ecological importance both globally and locally. However, they are increasingly threatened by multiple anthropogenic stressors, such as climate change, hydropower and invasive species. The development of efficient management strategies is therefore urgently needed and requires a comprehensive understanding of the factors influencing the biodiversity and ecological processes of these ecosystems. We provide a holistic knowledge base for informed future research and management by addressing the interplay between local and global environmental drivers of food webs in Arctic charr (Salvelinus alpinus, Salmonidae) and brown trout (Salmo trutta, Salmonidae) dominated cold-water lakes in Fennoscandia. The trophic niche and population dynamics of these generalist top consumers provide extensive insights into the effects of natural and anthropogenic drivers on food webs in intensively studied Fennoscandian cold-water lakes, covering marked biogeographical gradients in abiotic and biotic conditions. Drawing on a synthesis of existing literature, our focus is on three pivotal drivers: (1) lake location and connectivity, (2) lake area and morphometry and (3) fish community composition. These drivers significantly influence the complexity and the origin and flow of energy in lake food webs, and ultimately the size structure of the charr and trout populations. Furthermore, we highlight ongoing environmental changes in Fennoscandian cold-water lakes caused by hydropower and invasive species. Finally, we identify crucial knowledge gaps and propose management actions for improving the future state of Fennoscandian cold-water lake ecosystems and their charr and trout populations.

Cold-water lakes situated in high latitudes and altitudes have pivotal socio-ecological importance both globally and locally. However, they are increasingly threatened by multiple anthropogenic stressors, such as climate change, hydropower and invasive species. The development of efficient management strategies is therefore urgently needed and requires a comprehensive understanding of the factors influencing the biodiversity and ecological processes of these ecosystems. We provide a holistic knowledge base for informed future research and management by addressing the interplay between local and global environmental drivers of food webs in Arctic charr (Salvelinus alpinus, Salmonidae) and brown trout (Salmo trutta, Salmonidae) dominated cold-water lakes in Fennoscandia. The trophic niche and population dynamics of these generalist top consumers provide extensive insights into the effects of natural and anthropogenic drivers on food webs in intensively studied Fennoscandian cold-water lakes, covering marked biogeographical gradients in abiotic and biotic conditions. Drawing on a synthesis of existing literature, our focus is on three pivotal drivers: (1) lake location and connectivity, (2) lake area and morphometry and (3) fish community composition. These drivers significantly influence the complexity and the origin and flow of energy in lake food webs, and ultimately the size structure of the charr and trout populations. Furthermore, we highlight ongoing environmental changes in Fennoscandian cold-water lakes caused by hydropower and invasive species. Finally, we identify crucial knowledge gaps and propose management actions for improving the future state of Fennoscandian cold-water lake ecosystems and their charr and trout populations.

Climate change is expected to alter global temperature and precipitation patterns resulting in complex environmental impacts. The proposed higher precipitation in northern Scandinavia would increase runoff from land, hence increase the inflow of terrestrial dissolved organic matter (tDOM) in coastal regions. This could promote heterotrophic bacterial production and shift the food web structure, by favoring the microbial food web. The altered climate is also expected to affect transport and availability of organic micropollutants (MPs), with downstream effects on exposure and accumulation in biota. This study aimed to assess climate-induced changes in a Bothnian Sea food web structure as well as bioaccumulation patterns of MPs. We performed a mesocosms-study, focusing on aquatic food webs with fish as top predator. Alongside increased temperature, mesocosm treatments included tDOM and MP addition. The tDOM addition affected nutrient availability and boosted both phytoplankton and heterotrophic bacteria in our fairly shallow mesocosms. The increased tDOM further benefitted flagellates, ciliates and mesozooplankton, while the temperature increase and MP addition had minor effect on those organism groups. Temperature, on the other hand, had a negative impact on fish growth and survival, whereas tDOM and MP addition only had minor impact on fish. Moreover, there were indications that bioaccumulation of MPs in fish either increased with tDOM addition or decreased at higher temperatures. If there was an impact on bioaccumulation, moderately lipophilic MPs (log Kow 3.6 - 4.6) were generally affected by tDOM addition and more lipophilic MPs (log Kow 3.8 to 6.4) were generally affected by increased temperature. This study suggest that both increased temperatures and addition of tDOM likely will affect bioaccumulation patterns of MPs in shallow coastal regions, albeit with counteracting effects.

Climate change is expected to alter global temperature and precipitation patterns resulting in complex environmental impacts. The proposed higher precipitation in northern Scandinavia would increase runoff from land, hence increase the inflow of terrestrial dissolved organic matter (tDOM) in coastal regions. This could promote heterotrophic bacterial production and shift the food web structure, by favoring the microbial food web. The altered climate is also expected to affect transport and availability of organic micropollutants (MPs), with downstream effects on exposure and accumulation in biota. This study aimed to assess climate-induced changes in a Bothnian Sea food web structure as well as bioaccumulation patterns of MPs. We performed a mesocosms-study, focusing on aquatic food webs with fish as top predator. Alongside increased temperature, mesocosm treatments included tDOM and MP addition. The tDOM addition affected nutrient availability and boosted both phytoplankton and heterotrophic bacteria in our fairly shallow mesocosms. The increased tDOM further benefitted flagellates, ciliates and mesozooplankton, while the temperature increase and MP addition had minor effect on those organism groups. Temperature, on the other hand, had a negative impact on fish growth and survival, whereas tDOM and MP addition only had minor impact on fish. Moreover, there were indications that bioaccumulation of MPs in fish either increased with tDOM addition or decreased at higher temperatures. If there was an impact on bioaccumulation, moderately lipophilic MPs (log Kow 3.6 - 4.6) were generally affected by tDOM addition and more lipophilic MPs (log Kow 3.8 to 6.4) were generally affected by increased temperature. This study suggest that both increased temperatures and addition of tDOM likely will affect bioaccumulation patterns of MPs in shallow coastal regions, albeit with counteracting effects.

Globally, lakes are warming and browning with ongoing climate change. These changes significantly impact a lake's biogeochemical properties and all organisms, including invertebrate consumers. The effects of these changes are essential to understand, especially during critical periods after and before the growing season, that is, autumn and spring, which can determine the composition of the invertebrate consumer community. In this study, we used a large-scale experimental pond system to test the combined effect of warming (+3°C) and increased input of terrestrial and coloured dissolved organic carbon (gradient of 1.6–8.8 mg/L in the ambient and 1.6–9.3 mg/L in the warm)—which causes browning—on zooplankton and benthic macroinvertebrate biomass and composition during the autumn and the following spring. Total zooplankton biomass decreased with warming and increased with browning, while total zoobenthos did not respond to either treatment. Warming and browning throughout the autumn had no overall interactive effects on zooplankton or zoobenthos. Autumnal warming decreased total pelagic consumer biomass, caused by a decrease in both Rotifera and Copepoda. In contrast, there was no effect on overall benthic consumer biomass, with only Asellus sp. biomass showing a negative response to warming. An autumnal increase in dissolved organic carbon led to increased total pelagic consumer biomass, which was related to increases in Daphnia sp. biomass but did not affect zoobenthos biomass. While we expected zooplankton and zoobenthos biomass to follow responses in primary and bacterial production to treatments, we did not find any relationship between consumer groups and these estimates of resource production. Our results suggest that consumer responses to warming and browning during autumn may lead to less overarching general changes in consumer biomass, and responses are mostly taxon-specific. This study gives novel insights into the effects of warming and browning on consumer

Globally, lakes are warming and browning with ongoing climate change. These changes significantly impact a lake's biogeochemical properties and all organisms, including invertebrate consumers. The effects of these changes are essential to understand, especially during critical periods after and before the growing season, that is, autumn and spring, which can determine the composition of the invertebrate consumer community. In this study, we used a large-scale experimental pond system to test the combined effect of warming (+3°C) and increased input of terrestrial and coloured dissolved organic carbon (gradient of 1.6–8.8 mg/L in the ambient and 1.6–9.3 mg/L in the warm)—which causes browning—on zooplankton and benthic macroinvertebrate biomass and composition during the autumn and the following spring. Total zooplankton biomass decreased with warming and increased with browning, while total zoobenthos did not respond to either treatment. Warming and browning throughout the autumn had no overall interactive effects on zooplankton or zoobenthos. Autumnal warming decreased total pelagic consumer biomass, caused by a decrease in both Rotifera and Copepoda. In contrast, there was no effect on overall benthic consumer biomass, with only Asellus sp. biomass showing a negative response to warming. An autumnal increase in dissolved organic carbon led to increased total pelagic consumer biomass, which was related to increases in Daphnia sp. biomass but did not affect zoobenthos biomass. While we expected zooplankton and zoobenthos biomass to follow responses in primary and bacterial production to treatments, we did not find any relationship between consumer groups and these estimates of resource production. Our results suggest that consumer responses to warming and browning during autumn may lead to less overarching general changes in consumer biomass, and responses are mostly taxon-specific. This study gives novel insights into the effects of warming and browning on consumer

Globally, lakes are warming and browning with ongoing climate change. These changes significantly impact a lake's biogeochemical properties and all organisms, including invertebrate consumers. The effects of these changes are essential to understand, especially during critical periods after and before the growing season, that is, autumn and spring, which can determine the composition of the invertebrate consumer community. In this study, we used a large-scale experimental pond system to test the combined effect of warming (+3°C) and increased input of terrestrial and coloured dissolved organic carbon (gradient of 1.6–8.8 mg/L in the ambient and 1.6–9.3 mg/L in the warm)—which causes browning—on zooplankton and benthic macroinvertebrate biomass and composition during the autumn and the following spring. Total zooplankton biomass decreased with warming and increased with browning, while total zoobenthos did not respond to either treatment. Warming and browning throughout the autumn had no overall interactive effects on zooplankton or zoobenthos. Autumnal warming decreased total pelagic consumer biomass, caused by a decrease in both Rotifera and Copepoda. In contrast, there was no effect on overall benthic consumer biomass, with only Asellus sp. biomass showing a negative response to warming. An autumnal increase in dissolved organic carbon led to increased total pelagic consumer biomass, which was related to increases in Daphnia sp. biomass but did not affect zoobenthos biomass. While we expected zooplankton and zoobenthos biomass to follow responses in primary and bacterial production to treatments, we did not find any relationship between consumer groups and these estimates of resource production. Our results suggest that consumer responses to warming and browning during autumn may lead to less overarching general changes in consumer biomass, and responses are mostly taxon-specific. This study gives novel insights into the effects of warming and browning on consumer

Climate change is expected to alter global temperature and precipitation patterns resulting in complex environmental impacts. The proposed higher precipitation in northern Scandinavia would increase runoff from land, hence increase the inflow of terrestrial dissolved organic matter (tDOM) in coastal regions. This could promote heterotrophic bacterial production and shift the food web structure, by favoring the microbial food web. The altered climate is also expected to affect transport and availability of organic micropollutants (MPs), with downstream effects on exposure and accumulation in biota. This study aimed to assess climate-induced changes in a Bothnian Sea food web structure as well as bioaccumulation patterns of MPs. We performed a mesocosms-study, focusing on aquatic food webs with fish as top predator. Alongside increased temperature, mesocosm treatments included tDOM and MP addition. The tDOM addition affected nutrient availability and boosted both phytoplankton and heterotrophic bacteria in our fairly shallow mesocosms. The increased tDOM further benefitted flagellates, ciliates and mesozooplankton, while the temperature increase and MP addition had minor effect on those organism groups. Temperature, on the other hand, had a negative impact on fish growth and survival, whereas tDOM and MP addition only had minor impact on fish. Moreover, there were indications that bioaccumulation of MPs in fish either increased with tDOM addition or decreased at higher temperatures. If there was an impact on bioaccumulation, moderately lipophilic MPs (log Kow 3.6 - 4.6) were generally affected by tDOM addition and more lipophilic MPs (log Kow 3.8 to 6.4) were generally affected by increased temperature. This study suggest that both increased temperatures and addition of tDOM likely will affect bioaccumulation patterns of MPs in shallow coastal regions, albeit with counteracting effects.

Climate change is expected to alter global temperature and precipitation patterns resulting in complex environmental impacts. The proposed higher precipitation in northern Scandinavia would increase runoff from land, hence increase the inflow of terrestrial dissolved organic matter (tDOM) in coastal regions. This could promote heterotrophic bacterial production and shift the food web structure, by favoring the microbial food web. The altered climate is also expected to affect transport and availability of organic micropollutants (MPs), with downstream effects on exposure and accumulation in biota. This study aimed to assess climate-induced changes in a Bothnian Sea food web structure as well as bioaccumulation patterns of MPs. We performed a mesocosms-study, focusing on aquatic food webs with fish as top predator. Alongside increased temperature, mesocosm treatments included tDOM and MP addition. The tDOM addition affected nutrient availability and boosted both phytoplankton and heterotrophic bacteria in our fairly shallow mesocosms. The increased tDOM further benefitted flagellates, ciliates and mesozooplankton, while the temperature increase and MP addition had minor effect on those organism groups. Temperature, on the other hand, had a negative impact on fish growth and survival, whereas tDOM and MP addition only had minor impact on fish. Moreover, there were indications that bioaccumulation of MPs in fish either increased with tDOM addition or decreased at higher temperatures. If there was an impact on bioaccumulation, moderately lipophilic MPs (log Kow 3.6 - 4.6) were generally affected by tDOM addition and more lipophilic MPs (log Kow 3.8 to 6.4) were generally affected by increased temperature. This study suggest that both increased temperatures and addition of tDOM likely will affect bioaccumulation patterns of MPs in shallow coastal regions, albeit with counteracting effects.

Globally, lakes are warming and browning with ongoing climate change. These changes significantly impact a lake's biogeochemical properties and all organisms, including invertebrate consumers. The effects of these changes are essential to understand, especially during critical periods after and before the growing season, that is, autumn and spring, which can determine the composition of the invertebrate consumer community. In this study, we used a large-scale experimental pond system to test the combined effect of warming (+3°C) and increased input of terrestrial and coloured dissolved organic carbon (gradient of 1.6–8.8 mg/L in the ambient and 1.6–9.3 mg/L in the warm)—which causes browning—on zooplankton and benthic macroinvertebrate biomass and composition during the autumn and the following spring. Total zooplankton biomass decreased with warming and increased with browning, while total zoobenthos did not respond to either treatment. Warming and browning throughout the autumn had no overall interactive effects on zooplankton or zoobenthos. Autumnal warming decreased total pelagic consumer biomass, caused by a decrease in both Rotifera and Copepoda. In contrast, there was no effect on overall benthic consumer biomass, with only Asellus sp. biomass showing a negative response to warming. An autumnal increase in dissolved organic carbon led to increased total pelagic consumer biomass, which was related to increases in Daphnia sp. biomass but did not affect zoobenthos biomass. While we expected zooplankton and zoobenthos biomass to follow responses in primary and bacterial production to treatments, we did not find any relationship between consumer groups and these estimates of resource production. Our results suggest that consumer responses to warming and browning during autumn may lead to less overarching general changes in consumer biomass, and responses are mostly taxon-specific. This study gives novel insights into the effects of warming and browning on consumer

Globally, lakes are warming and browning with ongoing climate change. These changes significantly impact a lake's biogeochemical properties and all organisms, including invertebrate consumers. The effects of these changes are essential to understand, especially during critical periods after and before the growing season, that is, autumn and spring, which can determine the composition of the invertebrate consumer community. In this study, we used a large-scale experimental pond system to test the combined effect of warming (+3°C) and increased input of terrestrial and coloured dissolved organic carbon (gradient of 1.6–8.8 mg/L in the ambient and 1.6–9.3 mg/L in the warm)—which causes browning—on zooplankton and benthic macroinvertebrate biomass and composition during the autumn and the following spring. Total zooplankton biomass decreased with warming and increased with browning, while total zoobenthos did not respond to either treatment. Warming and browning throughout the autumn had no overall interactive effects on zooplankton or zoobenthos. Autumnal warming decreased total pelagic consumer biomass, caused by a decrease in both Rotifera and Copepoda. In contrast, there was no effect on overall benthic consumer biomass, with only Asellus sp. biomass showing a negative response to warming. An autumnal increase in dissolved organic carbon led to increased total pelagic consumer biomass, which was related to increases in Daphnia sp. biomass but did not affect zoobenthos biomass. While we expected zooplankton and zoobenthos biomass to follow responses in primary and bacterial production to treatments, we did not find any relationship between consumer groups and these estimates of resource production. Our results suggest that consumer responses to warming and browning during autumn may lead to less overarching general changes in consumer biomass, and responses are mostly taxon-specific. This study gives novel insights into the effects of warming and browning on consumer

Globally, the length of tributaries to lakes varies from 0 to more than 15,000 km, but scaling relationships describing this aspect of lake-river connectivity are lacking. In this study, we describe a simple theoretical scaling relationship for tributary length based on the principle of line intercepts of topographic features, and test this theory using data from Scandinavia. Tributary length increases by 73% for each doubling of lake area. This pattern reflects the relationship between catchment and lake area, and is modified by inlet frequency, junction angle, and lake shape—factors related to specific geologic and hydrologic processes. The theory is precise (r2 = 0.74), with low bias (mean error is 14% of mean tributary length) when the characteristic junction angle (∼76°) is estimated statistically. Our study bridges the gap between geomorphic and large-scale statistical relationships to provide simple rules for understanding complex patterns of lake-river connectivity.

Gross primary production (GPP) by benthic microalgae growing on soft sediments is an important contributor to lake productivity in many lakes world-wide. As benthic microalgae have access to nutrients in the sediment they have been regarded as primarily controlled by light, while the role of CO2 as a limiting factor for benthic GPP in lake ecosystems is largely unknown. In this study, we experimentally tested for CO2 limitation of benthic GPP by collecting littoral surface sediments, with associated benthic microalgae, from a typical boreal lake. Intact sediment cores were incubated at different depths (light conditions) after addition of dissolved inorganic (bicarbonate) or organic (DOC; glucose) carbon as direct and indirect sources of CO2, respectively. Benthic microalgal GPP was stimulated by both dissolved inorganic carbon and DOC additions at high, but not at low, light levels. This study shows that benthic microalgal GPP can be CO2-limited when light is not limiting and suggests that both direct (e.g., via groundwater inflow) and indirect (via mineralisation of DOC) CO2 supply can stimulate benthic GPP.

Gross primary production (GPP) by benthic microalgae growing on soft sediments is an important contributor to lake productivity in many lakes world-wide. As benthic microalgae have access to nutrients in the sediment they have been regarded as primarily controlled by light, while the role of CO2 as a limiting factor for benthic GPP in lake ecosystems is largely unknown. In this study, we experimentally tested for CO2 limitation of benthic GPP by collecting littoral surface sediments, with associated benthic microalgae, from a typical boreal lake. Intact sediment cores were incubated at different depths (light conditions) after addition of dissolved inorganic (bicarbonate) or organic (DOC; glucose) carbon as direct and indirect sources of CO2, respectively. Benthic microalgal GPP was stimulated by both dissolved inorganic carbon and DOC additions at high, but not at low, light levels. This study shows that benthic microalgal GPP can be CO2-limited when light is not limiting and suggests that both direct (e.g., via groundwater inflow) and indirect (via mineralisation of DOC) CO2 supply can stimulate benthic GPP.

Gross primary production (GPP) by benthic microalgae growing on soft sediments is an important contributor to lake productivity in many lakes world-wide. As benthic microalgae have access to nutrients in the sediment they have been regarded as primarily controlled by light, while the role of CO2 as a limiting factor for benthic GPP in lake ecosystems is largely unknown. In this study, we experimentally tested for CO2 limitation of benthic GPP by collecting littoral surface sediments, with associated benthic microalgae, from a typical boreal lake. Intact sediment cores were incubated at different depths (light conditions) after addition of dissolved inorganic (bicarbonate) or organic (DOC; glucose) carbon as direct and indirect sources of CO2, respectively. Benthic microalgal GPP was stimulated by both dissolved inorganic carbon and DOC additions at high, but not at low, light levels. This study shows that benthic microalgal GPP can be CO2-limited when light is not limiting and suggests that both direct (e.g., via groundwater inflow) and indirect (via mineralisation of DOC) CO2 supply can stimulate benthic GPP.

Globally, the length of tributaries to lakes varies from 0 to more than 15,000 km, but scaling relationships describing this aspect of lake-river connectivity are lacking. In this study, we describe a simple theoretical scaling relationship for tributary length based on the principle of line intercepts of topographic features, and test this theory using data from Scandinavia. Tributary length increases by 73% for each doubling of lake area. This pattern reflects the relationship between catchment and lake area, and is modified by inlet frequency, junction angle, and lake shape—factors related to specific geologic and hydrologic processes. The theory is precise (r2 = 0.74), with low bias (mean error is 14% of mean tributary length) when the characteristic junction angle (∼76°) is estimated statistically. Our study bridges the gap between geomorphic and large-scale statistical relationships to provide simple rules for understanding complex patterns of lake-river connectivity.

Ecological theory predicts that the relative distribution of primary production across habitats influence fish size structure and biomass production. In this study, we assessed individual, population, and community-level consequences for brown trout (Salmo trutta) and Arctic char (Salvelinus alpinus) of variation in estimated habitat specific (benthic and pelagic) and total whole lake (GPPwhole) gross primary production in 27 northern oligotrophic lakes. We found that higher contribution of benthic primary production to GPPwhole was associated with higher community biomass and larger maximum and mean sizes of fish. At the population level, species-specific responses differed. Increased benthic primary production (GPPBenthic) correlated to higher population biomass of brown trout regardless of being alone or in sympatry, while Arctic char responded positively to pelagic primary production (GPPPelagic) in sympatric populations. In sympatric lakes, the maximum size of both species was positively related to both GPPBenthic and the benthic contribution to GPPWhole. In allopatric lakes, brown trout mean and maximum size and Arctic char mean size were positively related to the benthic proportion of GPPWhole. Our results highlight the importance of light-controlled benthic primary production for fish biomass production in oligotrophic northern lakes. Our results further suggest that consequences of ontogenetic asymmetry and niche shifts may cause the distribution of primary production across habitats to be more important than the total ecosystem primary production for fish size, population biomass, and production. Awareness of the relationships between light availability and asymmetric resource production favoring large fish and fish production may allow for cost-efficient and more informed management actions in northern oligotrophic lakes.

The shoreline development index—The ratio of a lake’s shore length to the circumference of a circle with the lake’s area—Is a core metric of lake morphometry used in Earth and planetary sciences. In this paper, we demonstrate that the shoreline development index is scale-dependent and cannot be used to compare lakes with different areas. We show that large lakes will have higher shoreline development index measurements than smaller lakes of the same characteristic shape, even when mapped at the same scale. Specifically, the shoreline development index increases by about 14% for each doubling of lake area. These results call into question previously reported patterns of lake shape. We provide several suggestions to improve the application of this index, including a bias-corrected formulation for comparing lakes with different surface areas.

Ecological theory predicts that the relative distribution of primary production across habitats influence fish size structure and biomass production. In this study, we assessed individual, population, and community-level consequences for brown trout (Salmo trutta) and Arctic char (Salvelinus alpinus) of variation in estimated habitat specific (benthic and pelagic) and total whole lake (GPPwhole) gross primary production in 27 northern oligotrophic lakes. We found that higher contribution of benthic primary production to GPPwhole was associated with higher community biomass and larger maximum and mean sizes of fish. At the population level, species-specific responses differed. Increased benthic primary production (GPPBenthic) correlated to higher population biomass of brown trout regardless of being alone or in sympatry, while Arctic char responded positively to pelagic primary production (GPPPelagic) in sympatric populations. In sympatric lakes, the maximum size of both species was positively related to both GPPBenthic and the benthic contribution to GPPWhole. In allopatric lakes, brown trout mean and maximum size and Arctic char mean size were positively related to the benthic proportion of GPPWhole. Our results highlight the importance of light-controlled benthic primary production for fish biomass production in oligotrophic northern lakes. Our results further suggest that consequences of ontogenetic asymmetry and niche shifts may cause the distribution of primary production across habitats to be more important than the total ecosystem primary production for fish size, population biomass, and production. Awareness of the relationships between light availability and asymmetric resource production favoring large fish and fish production may allow for cost-efficient and more informed management actions in northern oligotrophic lakes.

The shoreline development index—The ratio of a lake’s shore length to the circumference of a circle with the lake’s area—Is a core metric of lake morphometry used in Earth and planetary sciences. In this paper, we demonstrate that the shoreline development index is scale-dependent and cannot be used to compare lakes with different areas. We show that large lakes will have higher shoreline development index measurements than smaller lakes of the same characteristic shape, even when mapped at the same scale. Specifically, the shoreline development index increases by about 14% for each doubling of lake area. These results call into question previously reported patterns of lake shape. We provide several suggestions to improve the application of this index, including a bias-corrected formulation for comparing lakes with different surface areas.

Ecological theory predicts that the relative distribution of primary production across habitats influence fish size structure and biomass production. In this study, we assessed individual, population, and community-level consequences for brown trout (Salmo trutta) and Arctic char (Salvelinus alpinus) of variation in estimated habitat specific (benthic and pelagic) and total whole lake (GPPwhole) gross primary production in 27 northern oligotrophic lakes. We found that higher contribution of benthic primary production to GPPwhole was associated with higher community biomass and larger maximum and mean sizes of fish. At the population level, species-specific responses differed. Increased benthic primary production (GPPBenthic) correlated to higher population biomass of brown trout regardless of being alone or in sympatry, while Arctic char responded positively to pelagic primary production (GPPPelagic) in sympatric populations. In sympatric lakes, the maximum size of both species was positively related to both GPPBenthic and the benthic contribution to GPPWhole. In allopatric lakes, brown trout mean and maximum size and Arctic char mean size were positively related to the benthic proportion of GPPWhole. Our results highlight the importance of light-controlled benthic primary production for fish biomass production in oligotrophic northern lakes. Our results further suggest that consequences of ontogenetic asymmetry and niche shifts may cause the distribution of primary production across habitats to be more important than the total ecosystem primary production for fish size, population biomass, and production. Awareness of the relationships between light availability and asymmetric resource production favoring large fish and fish production may allow for cost-efficient and more informed management actions in northern oligotrophic lakes.

The shoreline development index—The ratio of a lake’s shore length to the circumference of a circle with the lake’s area—Is a core metric of lake morphometry used in Earth and planetary sciences. In this paper, we demonstrate that the shoreline development index is scale-dependent and cannot be used to compare lakes with different areas. We show that large lakes will have higher shoreline development index measurements than smaller lakes of the same characteristic shape, even when mapped at the same scale. Specifically, the shoreline development index increases by about 14% for each doubling of lake area. These results call into question previously reported patterns of lake shape. We provide several suggestions to improve the application of this index, including a bias-corrected formulation for comparing lakes with different surface areas.

Globally, the length of tributaries to lakes varies from 0 to more than 15,000 km, but scaling relationships describing this aspect of lake-river connectivity are lacking. In this study, we describe a simple theoretical scaling relationship for tributary length based on the principle of line intercepts of topographic features, and test this theory using data from Scandinavia. Tributary length increases by 73% for each doubling of lake area. This pattern reflects the relationship between catchment and lake area, and is modified by inlet frequency, junction angle, and lake shape—factors related to specific geologic and hydrologic processes. The theory is precise (r2 = 0.74), with low bias (mean error is 14% of mean tributary length) when the characteristic junction angle (∼76°) is estimated statistically. Our study bridges the gap between geomorphic and large-scale statistical relationships to provide simple rules for understanding complex patterns of lake-river connectivity.

The shoreline development index—The ratio of a lake’s shore length to the circumference of a circle with the lake’s area—Is a core metric of lake morphometry used in Earth and planetary sciences. In this paper, we demonstrate that the shoreline development index is scale-dependent and cannot be used to compare lakes with different areas. We show that large lakes will have higher shoreline development index measurements than smaller lakes of the same characteristic shape, even when mapped at the same scale. Specifically, the shoreline development index increases by about 14% for each doubling of lake area. These results call into question previously reported patterns of lake shape. We provide several suggestions to improve the application of this index, including a bias-corrected formulation for comparing lakes with different surface areas.

Ecological theory predicts that the relative distribution of primary production across habitats influence fish size structure and biomass production. In this study, we assessed individual, population, and community-level consequences for brown trout (Salmo trutta) and Arctic char (Salvelinus alpinus) of variation in estimated habitat specific (benthic and pelagic) and total whole lake (GPPwhole) gross primary production in 27 northern oligotrophic lakes. We found that higher contribution of benthic primary production to GPPwhole was associated with higher community biomass and larger maximum and mean sizes of fish. At the population level, species-specific responses differed. Increased benthic primary production (GPPBenthic) correlated to higher population biomass of brown trout regardless of being alone or in sympatry, while Arctic char responded positively to pelagic primary production (GPPPelagic) in sympatric populations. In sympatric lakes, the maximum size of both species was positively related to both GPPBenthic and the benthic contribution to GPPWhole. In allopatric lakes, brown trout mean and maximum size and Arctic char mean size were positively related to the benthic proportion of GPPWhole. Our results highlight the importance of light-controlled benthic primary production for fish biomass production in oligotrophic northern lakes. Our results further suggest that consequences of ontogenetic asymmetry and niche shifts may cause the distribution of primary production across habitats to be more important than the total ecosystem primary production for fish size, population biomass, and production. Awareness of the relationships between light availability and asymmetric resource production favoring large fish and fish production may allow for cost-efficient and more informed management actions in northern oligotrophic lakes.

The shoreline development index—The ratio of a lake’s shore length to the circumference of a circle with the lake’s area—Is a core metric of lake morphometry used in Earth and planetary sciences. In this paper, we demonstrate that the shoreline development index is scale-dependent and cannot be used to compare lakes with different areas. We show that large lakes will have higher shoreline development index measurements than smaller lakes of the same characteristic shape, even when mapped at the same scale. Specifically, the shoreline development index increases by about 14% for each doubling of lake area. These results call into question previously reported patterns of lake shape. We provide several suggestions to improve the application of this index, including a bias-corrected formulation for comparing lakes with different surface areas.

Ecological theory predicts that the relative distribution of primary production across habitats influence fish size structure and biomass production. In this study, we assessed individual, population, and community-level consequences for brown trout (Salmo trutta) and Arctic char (Salvelinus alpinus) of variation in estimated habitat specific (benthic and pelagic) and total whole lake (GPPwhole) gross primary production in 27 northern oligotrophic lakes. We found that higher contribution of benthic primary production to GPPwhole was associated with higher community biomass and larger maximum and mean sizes of fish. At the population level, species-specific responses differed. Increased benthic primary production (GPPBenthic) correlated to higher population biomass of brown trout regardless of being alone or in sympatry, while Arctic char responded positively to pelagic primary production (GPPPelagic) in sympatric populations. In sympatric lakes, the maximum size of both species was positively related to both GPPBenthic and the benthic contribution to GPPWhole. In allopatric lakes, brown trout mean and maximum size and Arctic char mean size were positively related to the benthic proportion of GPPWhole. Our results highlight the importance of light-controlled benthic primary production for fish biomass production in oligotrophic northern lakes. Our results further suggest that consequences of ontogenetic asymmetry and niche shifts may cause the distribution of primary production across habitats to be more important than the total ecosystem primary production for fish size, population biomass, and production. Awareness of the relationships between light availability and asymmetric resource production favoring large fish and fish production may allow for cost-efficient and more informed management actions in northern oligotrophic lakes.

Gross primary production (GPP) by benthic microalgae growing on soft sediments is an important contributor to lake productivity in many lakes world-wide. As benthic microalgae have access to nutrients in the sediment they have been regarded as primarily controlled by light, while the role of CO2 as a limiting factor for benthic GPP in lake ecosystems is largely unknown. In this study, we experimentally tested for CO2 limitation of benthic GPP by collecting littoral surface sediments, with associated benthic microalgae, from a typical boreal lake. Intact sediment cores were incubated at different depths (light conditions) after addition of dissolved inorganic (bicarbonate) or organic (DOC; glucose) carbon as direct and indirect sources of CO2, respectively. Benthic microalgal GPP was stimulated by both dissolved inorganic carbon and DOC additions at high, but not at low, light levels. This study shows that benthic microalgal GPP can be CO2-limited when light is not limiting and suggests that both direct (e.g., via groundwater inflow) and indirect (via mineralisation of DOC) CO2 supply can stimulate benthic GPP.

Globally, the length of tributaries to lakes varies from 0 to more than 15,000 km, but scaling relationships describing this aspect of lake-river connectivity are lacking. In this study, we describe a simple theoretical scaling relationship for tributary length based on the principle of line intercepts of topographic features, and test this theory using data from Scandinavia. Tributary length increases by 73% for each doubling of lake area. This pattern reflects the relationship between catchment and lake area, and is modified by inlet frequency, junction angle, and lake shape—factors related to specific geologic and hydrologic processes. The theory is precise (r2 = 0.74), with low bias (mean error is 14% of mean tributary length) when the characteristic junction angle (∼76°) is estimated statistically. Our study bridges the gap between geomorphic and large-scale statistical relationships to provide simple rules for understanding complex patterns of lake-river connectivity.

Gross primary production (GPP) by benthic microalgae growing on soft sediments is an important contributor to lake productivity in many lakes world-wide. As benthic microalgae have access to nutrients in the sediment they have been regarded as primarily controlled by light, while the role of CO2 as a limiting factor for benthic GPP in lake ecosystems is largely unknown. In this study, we experimentally tested for CO2 limitation of benthic GPP by collecting littoral surface sediments, with associated benthic microalgae, from a typical boreal lake. Intact sediment cores were incubated at different depths (light conditions) after addition of dissolved inorganic (bicarbonate) or organic (DOC; glucose) carbon as direct and indirect sources of CO2, respectively. Benthic microalgal GPP was stimulated by both dissolved inorganic carbon and DOC additions at high, but not at low, light levels. This study shows that benthic microalgal GPP can be CO2-limited when light is not limiting and suggests that both direct (e.g., via groundwater inflow) and indirect (via mineralisation of DOC) CO2 supply can stimulate benthic GPP.

Globally, the length of tributaries to lakes varies from 0 to more than 15,000 km, but scaling relationships describing this aspect of lake-river connectivity are lacking. In this study, we describe a simple theoretical scaling relationship for tributary length based on the principle of line intercepts of topographic features, and test this theory using data from Scandinavia. Tributary length increases by 73% for each doubling of lake area. This pattern reflects the relationship between catchment and lake area, and is modified by inlet frequency, junction angle, and lake shape—factors related to specific geologic and hydrologic processes. The theory is precise (r2 = 0.74), with low bias (mean error is 14% of mean tributary length) when the characteristic junction angle (∼76°) is estimated statistically. Our study bridges the gap between geomorphic and large-scale statistical relationships to provide simple rules for understanding complex patterns of lake-river connectivity.

Climate changes are predicted to influence gross primary production (GPP) of lakes directly through warming and indirectly through increased loads of allochthonous coloured dissolved organic matter (cDOM) from surrounding landscapes. However, few studies have investigated this combined effect. Here we tested the effects of warming (elevated 3celcius) and cDOM input (three levels of humic river water addition) on GPP in autumn (2 months including open water and ice-covered periods) in experimental pond ecosystems. The cDOM input decreased whole-ecosystem GPP at natural temperature conditions mainly as a result of lower benthic GPP not fully counteracted by an increase in pelagic GPP, while warming increased whole-ecosystem GPP due to a positive response of mainly pelagic GPP at all levels of cDOM input. Warming delayed autumn ice cover formation by 2 weeks but did not affect light availability in the water column compared to ambient ice-covered treatments. Gross primary production during this period was still affected by warming and cDOM. The results stress the importance of accounting for multiple climate drivers and habitats when predicting lake GPP responses to climate change. We conclude that climate change may shift whole-ecosystem GPP through different responses of habitat-specific GPP to increasing cDOM inputs and warming.

Climate changes are predicted to influence gross primary production (GPP) of lakes directly through warming and indirectly through increased loads of allochthonous coloured dissolved organic matter (cDOM) from surrounding landscapes. However, few studies have investigated this combined effect. Here we tested the effects of warming (elevated 3celcius) and cDOM input (three levels of humic river water addition) on GPP in autumn (2 months including open water and ice-covered periods) in experimental pond ecosystems. The cDOM input decreased whole-ecosystem GPP at natural temperature conditions mainly as a result of lower benthic GPP not fully counteracted by an increase in pelagic GPP, while warming increased whole-ecosystem GPP due to a positive response of mainly pelagic GPP at all levels of cDOM input. Warming delayed autumn ice cover formation by 2 weeks but did not affect light availability in the water column compared to ambient ice-covered treatments. Gross primary production during this period was still affected by warming and cDOM. The results stress the importance of accounting for multiple climate drivers and habitats when predicting lake GPP responses to climate change. We conclude that climate change may shift whole-ecosystem GPP through different responses of habitat-specific GPP to increasing cDOM inputs and warming.

The littoral zone varies in size among lakes from ∼3% to 100% of lake surface area. In this paper, we derive a simple theoretical scaling relationship that explains this variation, and test this theory using bathymetric data across the size spectra of freshwater lakes (surface area = 0.01–82,103 km2, maximum depth = 2–1,741 m). Littoral area primarily reflects the ratio of the maximum depth of photosynthesis to maximum lake depth. However, lakes that are similar in these characteristics can have different relative littoral areas because of variation in basin shape. Hypsometric (area-elevation) models that describe these patterns for individual lakes can be generalized among lakes to accurately predict the relative size of littoral habitat when there is incomplete bathymetric information. Collectively, our results provide simple rules for understanding patterns of littoral habitat size at the regional and global scales.

The littoral zone varies in size among lakes from ∼3% to 100% of lake surface area. In this paper, we derive a simple theoretical scaling relationship that explains this variation, and test this theory using bathymetric data across the size spectra of freshwater lakes (surface area = 0.01–82,103 km2, maximum depth = 2–1,741 m). Littoral area primarily reflects the ratio of the maximum depth of photosynthesis to maximum lake depth. However, lakes that are similar in these characteristics can have different relative littoral areas because of variation in basin shape. Hypsometric (area-elevation) models that describe these patterns for individual lakes can be generalized among lakes to accurately predict the relative size of littoral habitat when there is incomplete bathymetric information. Collectively, our results provide simple rules for understanding patterns of littoral habitat size at the regional and global scales.

The littoral zone varies in size among lakes from ∼3% to 100% of lake surface area. In this paper, we derive a simple theoretical scaling relationship that explains this variation, and test this theory using bathymetric data across the size spectra of freshwater lakes (surface area = 0.01–82,103 km2, maximum depth = 2–1,741 m). Littoral area primarily reflects the ratio of the maximum depth of photosynthesis to maximum lake depth. However, lakes that are similar in these characteristics can have different relative littoral areas because of variation in basin shape. Hypsometric (area-elevation) models that describe these patterns for individual lakes can be generalized among lakes to accurately predict the relative size of littoral habitat when there is incomplete bathymetric information. Collectively, our results provide simple rules for understanding patterns of littoral habitat size at the regional and global scales.

Climate changes are predicted to influence gross primary production (GPP) of lakes directly through warming and indirectly through increased loads of allochthonous coloured dissolved organic matter (cDOM) from surrounding landscapes. However, few studies have investigated this combined effect. Here we tested the effects of warming (elevated 3celcius) and cDOM input (three levels of humic river water addition) on GPP in autumn (2 months including open water and ice-covered periods) in experimental pond ecosystems. The cDOM input decreased whole-ecosystem GPP at natural temperature conditions mainly as a result of lower benthic GPP not fully counteracted by an increase in pelagic GPP, while warming increased whole-ecosystem GPP due to a positive response of mainly pelagic GPP at all levels of cDOM input. Warming delayed autumn ice cover formation by 2 weeks but did not affect light availability in the water column compared to ambient ice-covered treatments. Gross primary production during this period was still affected by warming and cDOM. The results stress the importance of accounting for multiple climate drivers and habitats when predicting lake GPP responses to climate change. We conclude that climate change may shift whole-ecosystem GPP through different responses of habitat-specific GPP to increasing cDOM inputs and warming.

Measures of environmental DNA (eDNA) concentrations in water samples have the potential to be both a cost-efficient and a nondestructive method to estimate fish population abundance. However, the inherent temporal and spatial variability in abiotic and biotic conditions in aquatic systems have been suggested to be a major obstacle to determine relationships between fish eDNA concentrations and fish population abundance. Moreover, once water samples are collected, methodological biases are common, which introduces additional sources of variation to potential relationships between eDNA concentrations and fish population abundance. Here, we evaluate the performance of applying the droplet digital PCR (ddPCR) method to estimate fish population abundance in experimental enclosures. Using large-scale enclosure ecosystems that contain populations of nine-spined stickleback (Pungitius pungitius), we compared the concentrations of fish eDNA (COI mitochondrial region, 134 bp) obtained with the ddPCR method with high precision estimates of fish population abundance (i.e., number of individuals) and biomass. To evaluate the effects of contrasted concentrations of humic substances (potential PCR inhibitors) on the performance of ddPCR assays, we manipulated natural dissolved organic carbon (DOC) concentrations (range 4–11 mg/L) in the enclosures. Additionally, water temperature (+2°C) was manipulated in half of the enclosures. Results showed positive relationships between eDNA concentration and fish abundance and biomass estimates although unexplained variation remained. Still and importantly, fish eDNA estimates from high DOC enclosures were not lowered by potential inhibitory effects with our procedure. Finally, water temperature (although only 2°C difference) was neither detected as a significant factor influencing fish eDNA estimates. Altogether, our work highlights that ddPCR-based eDNA is a promising method for future quantification of fish population abundance in natural

Scaling relationships provide simple rules for understanding complex hydrographic patterns. Globally, river inlet abundance varies among lakes by about three orders of magnitude, but few scaling relationships describe this aspect of lake-river connectivity. In this study, we describe a simple theoretical scaling relationship between lake surface area and river inlet abundance, and test this theory using data from Scandinavia. On average, the number of inlets increases by 67% for each doubling of lake area. However, lakes of vastly different areas can have the same number of inlets with relatively small variations of drainage density, lake shape, or junction angle - characteristics that can often be linked to specific geological processes. Our approach bridges the gap between the detailed understanding of geomorphic processes and large-scale statistical relationships, and engenders predictions about additional patterns including the relationship between lake area and water residence time.

Measures of environmental DNA (eDNA) concentrations in water samples have the potential to be both a cost-efficient and a nondestructive method to estimate fish population abundance. However, the inherent temporal and spatial variability in abiotic and biotic conditions in aquatic systems have been suggested to be a major obstacle to determine relationships between fish eDNA concentrations and fish population abundance. Moreover, once water samples are collected, methodological biases are common, which introduces additional sources of variation to potential relationships between eDNA concentrations and fish population abundance. Here, we evaluate the performance of applying the droplet digital PCR (ddPCR) method to estimate fish population abundance in experimental enclosures. Using large-scale enclosure ecosystems that contain populations of nine-spined stickleback (Pungitius pungitius), we compared the concentrations of fish eDNA (COI mitochondrial region, 134 bp) obtained with the ddPCR method with high precision estimates of fish population abundance (i.e., number of individuals) and biomass. To evaluate the effects of contrasted concentrations of humic substances (potential PCR inhibitors) on the performance of ddPCR assays, we manipulated natural dissolved organic carbon (DOC) concentrations (range 4–11 mg/L) in the enclosures. Additionally, water temperature (+2°C) was manipulated in half of the enclosures. Results showed positive relationships between eDNA concentration and fish abundance and biomass estimates although unexplained variation remained. Still and importantly, fish eDNA estimates from high DOC enclosures were not lowered by potential inhibitory effects with our procedure. Finally, water temperature (although only 2°C difference) was neither detected as a significant factor influencing fish eDNA estimates. Altogether, our work highlights that ddPCR-based eDNA is a promising method for future quantification of fish population abundance in natural

Scaling relationships provide simple rules for understanding complex hydrographic patterns. Globally, river inlet abundance varies among lakes by about three orders of magnitude, but few scaling relationships describe this aspect of lake-river connectivity. In this study, we describe a simple theoretical scaling relationship between lake surface area and river inlet abundance, and test this theory using data from Scandinavia. On average, the number of inlets increases by 67% for each doubling of lake area. However, lakes of vastly different areas can have the same number of inlets with relatively small variations of drainage density, lake shape, or junction angle - characteristics that can often be linked to specific geological processes. Our approach bridges the gap between the detailed understanding of geomorphic processes and large-scale statistical relationships, and engenders predictions about additional patterns including the relationship between lake area and water residence time.

Scaling relationships provide simple rules for understanding complex hydrographic patterns. Globally, river inlet abundance varies among lakes by about three orders of magnitude, but few scaling relationships describe this aspect of lake-river connectivity. In this study, we describe a simple theoretical scaling relationship between lake surface area and river inlet abundance, and test this theory using data from Scandinavia. On average, the number of inlets increases by 67% for each doubling of lake area. However, lakes of vastly different areas can have the same number of inlets with relatively small variations of drainage density, lake shape, or junction angle - characteristics that can often be linked to specific geological processes. Our approach bridges the gap between the detailed understanding of geomorphic processes and large-scale statistical relationships, and engenders predictions about additional patterns including the relationship between lake area and water residence time.

Measures of environmental DNA (eDNA) concentrations in water samples have the potential to be both a cost-efficient and a nondestructive method to estimate fish population abundance. However, the inherent temporal and spatial variability in abiotic and biotic conditions in aquatic systems have been suggested to be a major obstacle to determine relationships between fish eDNA concentrations and fish population abundance. Moreover, once water samples are collected, methodological biases are common, which introduces additional sources of variation to potential relationships between eDNA concentrations and fish population abundance. Here, we evaluate the performance of applying the droplet digital PCR (ddPCR) method to estimate fish population abundance in experimental enclosures. Using large-scale enclosure ecosystems that contain populations of nine-spined stickleback (Pungitius pungitius), we compared the concentrations of fish eDNA (COI mitochondrial region, 134 bp) obtained with the ddPCR method with high precision estimates of fish population abundance (i.e., number of individuals) and biomass. To evaluate the effects of contrasted concentrations of humic substances (potential PCR inhibitors) on the performance of ddPCR assays, we manipulated natural dissolved organic carbon (DOC) concentrations (range 4–11 mg/L) in the enclosures. Additionally, water temperature (+2°C) was manipulated in half of the enclosures. Results showed positive relationships between eDNA concentration and fish abundance and biomass estimates although unexplained variation remained. Still and importantly, fish eDNA estimates from high DOC enclosures were not lowered by potential inhibitory effects with our procedure. Finally, water temperature (although only 2°C difference) was neither detected as a significant factor influencing fish eDNA estimates. Altogether, our work highlights that ddPCR-based eDNA is a promising method for future quantification of fish population abundance in natural

Climate changes are predicted to influence gross primary production (GPP) of lakes directly through warming and indirectly through increased loads of allochthonous coloured dissolved organic matter (cDOM) from surrounding landscapes. However, few studies have investigated this combined effect. Here we tested the effects of warming (elevated 3celcius) and cDOM input (three levels of humic river water addition) on GPP in autumn (2 months including open water and ice-covered periods) in experimental pond ecosystems. The cDOM input decreased whole-ecosystem GPP at natural temperature conditions mainly as a result of lower benthic GPP not fully counteracted by an increase in pelagic GPP, while warming increased whole-ecosystem GPP due to a positive response of mainly pelagic GPP at all levels of cDOM input. Warming delayed autumn ice cover formation by 2 weeks but did not affect light availability in the water column compared to ambient ice-covered treatments. Gross primary production during this period was still affected by warming and cDOM. The results stress the importance of accounting for multiple climate drivers and habitats when predicting lake GPP responses to climate change. We conclude that climate change may shift whole-ecosystem GPP through different responses of habitat-specific GPP to increasing cDOM inputs and warming.

Climate changes are predicted to influence gross primary production (GPP) of lakes directly through warming and indirectly through increased loads of allochthonous coloured dissolved organic matter (cDOM) from surrounding landscapes. However, few studies have investigated this combined effect. Here we tested the effects of warming (elevated 3celcius) and cDOM input (three levels of humic river water addition) on GPP in autumn (2 months including open water and ice-covered periods) in experimental pond ecosystems. The cDOM input decreased whole-ecosystem GPP at natural temperature conditions mainly as a result of lower benthic GPP not fully counteracted by an increase in pelagic GPP, while warming increased whole-ecosystem GPP due to a positive response of mainly pelagic GPP at all levels of cDOM input. Warming delayed autumn ice cover formation by 2 weeks but did not affect light availability in the water column compared to ambient ice-covered treatments. Gross primary production during this period was still affected by warming and cDOM. The results stress the importance of accounting for multiple climate drivers and habitats when predicting lake GPP responses to climate change. We conclude that climate change may shift whole-ecosystem GPP through different responses of habitat-specific GPP to increasing cDOM inputs and warming.

The littoral zone varies in size among lakes from ∼3% to 100% of lake surface area. In this paper, we derive a simple theoretical scaling relationship that explains this variation, and test this theory using bathymetric data across the size spectra of freshwater lakes (surface area = 0.01–82,103 km2, maximum depth = 2–1,741 m). Littoral area primarily reflects the ratio of the maximum depth of photosynthesis to maximum lake depth. However, lakes that are similar in these characteristics can have different relative littoral areas because of variation in basin shape. Hypsometric (area-elevation) models that describe these patterns for individual lakes can be generalized among lakes to accurately predict the relative size of littoral habitat when there is incomplete bathymetric information. Collectively, our results provide simple rules for understanding patterns of littoral habitat size at the regional and global scales.

Measures of environmental DNA (eDNA) concentrations in water samples have the potential to be both a cost-efficient and a nondestructive method to estimate fish population abundance. However, the inherent temporal and spatial variability in abiotic and biotic conditions in aquatic systems have been suggested to be a major obstacle to determine relationships between fish eDNA concentrations and fish population abundance. Moreover, once water samples are collected, methodological biases are common, which introduces additional sources of variation to potential relationships between eDNA concentrations and fish population abundance. Here, we evaluate the performance of applying the droplet digital PCR (ddPCR) method to estimate fish population abundance in experimental enclosures. Using large-scale enclosure ecosystems that contain populations of nine-spined stickleback (Pungitius pungitius), we compared the concentrations of fish eDNA (COI mitochondrial region, 134 bp) obtained with the ddPCR method with high precision estimates of fish population abundance (i.e., number of individuals) and biomass. To evaluate the effects of contrasted concentrations of humic substances (potential PCR inhibitors) on the performance of ddPCR assays, we manipulated natural dissolved organic carbon (DOC) concentrations (range 4–11 mg/L) in the enclosures. Additionally, water temperature (+2°C) was manipulated in half of the enclosures. Results showed positive relationships between eDNA concentration and fish abundance and biomass estimates although unexplained variation remained. Still and importantly, fish eDNA estimates from high DOC enclosures were not lowered by potential inhibitory effects with our procedure. Finally, water temperature (although only 2°C difference) was neither detected as a significant factor influencing fish eDNA estimates. Altogether, our work highlights that ddPCR-based eDNA is a promising method for future quantification of fish population abundance in natural

Scaling relationships provide simple rules for understanding complex hydrographic patterns. Globally, river inlet abundance varies among lakes by about three orders of magnitude, but few scaling relationships describe this aspect of lake-river connectivity. In this study, we describe a simple theoretical scaling relationship between lake surface area and river inlet abundance, and test this theory using data from Scandinavia. On average, the number of inlets increases by 67% for each doubling of lake area. However, lakes of vastly different areas can have the same number of inlets with relatively small variations of drainage density, lake shape, or junction angle - characteristics that can often be linked to specific geological processes. Our approach bridges the gap between the detailed understanding of geomorphic processes and large-scale statistical relationships, and engenders predictions about additional patterns including the relationship between lake area and water residence time.