Thermal Refugia Research Articles

Timing and frequency of high temperature events bend the onset of behavioural thermoregulation in Atlantic salmon (Salmo salar).

The role of temperature on biological activities and the correspondent exponential relationship with temperature has been known for over a century. However, lacking to date is knowledge relating to (a) the recovery of ectotherms subjected to extreme temperatures in the wild, and (b) the effects repeated extreme temperatures have on the temperatures that induce behavioural thermoregulation (aggregations). We examined these questions by testing the hypothesis that thermal thresholds which initiate aggregations in juvenile Atlantic salmon (AS) (Salmo salar) are not static, but are temporally dynamic across a summer and follow a hysteresis loop. To test our hypothesis, we deployed custom-made underwater camera (UWC) systems in known AS thermal refuges to observe the timing of aggregation events in a natural system and used these data to develop and test models that predict the temperatures that induce thermal aggregations. Consistent with our hypothesis our UWC observations revealed a range of aggregation onset temperatures (AOT) ranging from 24.2°C to 27.1°C, thus confirming our hypothesis that AOTs are dynamic across summer. Our models suggest it take ~ 11days of non-thermally taxing temperatures for the AOT to rebound in the study river. Conversely, we found that as the frequency of events increased, the AOT declined, from 27.1°C to 24.2°C. Integrating both model components led to more robust model performance. Further, when these models were tested against an independent data set from the same river, the results remained robust. Our findings illustrate the complexity underlying behavioural thermoregulation in AS-a complexity that most likely extends to other salmonids. The frequency of extreme heat events is predicted to increase, and this has the capacity to decrease AOT thresholds in AS, ultimately reducing their resilience to extreme temperature events.

Quality of thermal refuges influences use by the cold intolerant Florida manatee

Quality of thermal refuges influences use by the cold intolerant Florida manatee

Evaluating annual severe coral bleaching risk for marine protected areas across Indonesia

Coral reefs face an uncertain future under global climate change, with thermal-induced bleaching increasing in frequency such that corals will soon experience annual severe bleaching (ASB). Marine Protected Areas (MPAs) are therefore becoming increasingly important as a conservation tool. Here we evaluate (i) Indonesia’s coral reefs’ spatial variation in ASB, (ii) whether reefs projected to have a later onset of ASB (i.e. possible climate refugia) are protected within MPAs, and (iii) the ASB risk profiles for reefs related to MPAs receiving priority investments. Our results highlight considerable variability across Indonesia’s reefs being at risk of ASB. The ASB risk before 2028 is greater for coral reefs protected by MPAs versus those outside MPA boundaries. The ASB risk before 2025 is greater for coral reefs protected by priority MPAs versus those protected by non-priority MPAs. Overall, our results show that only ∼45% of the coral reef areas that are currently located within MPAs will likely act as thermal refugia (ASB > 2044). This is unsurprising given that the MPA network in Indonesia has been established over many decades, with most MPAs designated before suitable bleaching risk projections were available to inform MPA placement. Our results highlight the scope to further incorporate potential climate refugia for reefs into new MPA designations. This study also provides strategic information, which can support the development of Indonesia’s long-term MPA and coral reef conservation strategy to effectively manage, mitigate, and adapt to the impacts of climate change on coral reefs.

Snow cover dynamics: an overlooked yet important feature of winter bird occurrence and abundance across the United States

Snow cover dynamics (i.e. depth, duration and variability) are dominant drivers of ecological processes during winter. For overwintering species, changes and gradients in snow cover may impact survival and population dynamics (e.g. facilitating survival via thermal refugia or limiting survival via reduced resource acquisition). However, snow cover dynamics are rarely used in species distribution modelling, especially for overwintering birds. Currently, we lack understanding of how snow cover gradients affect overwintering bird distributions and which functional traits drive these associations at regional and continental scales. Using observations from eBird, a global community science network, we explored the effects of snow cover dynamics on continental patterns of occurrence and counts for 150 bird species. We quantified the relative importance, species‐specific responses and trait‐based relationships of bird occurrence and abundance patterns to ecologically relevant snow cover dynamics across the United States. Snow cover dynamics were important environmental predictors in species distributions models, ranking within the top three predictors for most species occurrence (> 90%) and count (> 79%) patterns across the contiguous United States. Species exhibited a gradient of responses to snow cover from snow association to snow avoidance, yet most birds were limited by long, persistent snow seasons. Duration of winter and percent frozen ground without snow structured species distributions in the eastern USA, whereas snow cover variability was a stronger driver in the western USA. Birds associated with long, persistent snow seasons had traits associated with greater dispersal capacity and dietary diversity, whereas birds inhabiting regions with variable snow cover were generally habitat generalists. Our results suggest that various snow cover dynamics are important ecological filters of species distributions during winter. Global climate change is rapidly degrading key characteristics of seasonal snow cover. A changing cryosphere may elicit variable distributional changes for many overwintering birds, potentially accelerating range shifts and novel community assemblages.

Cooler, bigger; warmer, smaller: Fine‐scale thermal heterogeneity maps age class and species distribution in behaviourally thermoregulating salmonids

AbstractBehavioural thermoregulation is a survival strategy that occurs in response to an exceedance of thermal stress‐inducing thresholds. When salmonids experience exceedance of these thresholds, they seek regions of colder water, known as thermal refuges. During an extreme temperature event of summer 2021 (main stem ~31.5°C) a large aggregation of Atlantic salmon (Salmo salar—all age classes) and brook trout (Salvelinus fontinalis—>20 cm) was observed on the Little Southwest Miramichi River in New Brunswick, Canada. Using a drone‐mounted thermal infrared (TIR) sensor, fine‐scale TIR imagery of the occupied refuge was acquired. Polarized glasses were worn by an onshore observer to make visual observations. Constructing maps from these data we examined the spatial distribution of fish, and the corresponding temperature of the areas they occupied. Salmonids were found to be distributed by age class and species, with the distribution driven by the mosaic of temperatures in the refuge. Young of the year (YOY), 1+, 2+ and adult Atlantic salmon occupied areas with average temperatures ~30.1, 28.8, 25.7 and 21.9°C, respectively; whilst mature brook trout occupied areas ~21.8°C. Noteworthy is the observation of thermally aggregating young of the year Atlantic salmon, and the range in temperatures they occupied (~8°C). One isolated, shallow, cold water patch (~22°C) exclusively held YOY Atlantic salmon. Our findings highlight the importance of several different thermal characteristics of thermal refuges and their link to salmonid occupancy and can aid the design of ecologically meaningful thermal refuge augmentation/restoration projects.

Tough places and safe spaces: Can refuges save salmon from a warming climate?

The importance of thermal refuges in a rapidly warming world is particularly evident for migratory species, where individuals encounter a wide range of conditions throughout their lives. In this study, we used a spatially explicit, individual-based simulation model to evaluate the buffering potential of cold-water thermal refuges for anadromous salmon and trout (Oncorhynchus spp.) migrating upstream through a warm river corridor that can expose individuals to physiologically stressful temperatures. We considered upstream migration in relation to migratory phenotypes that were defined in terms of migration timing, spawn timing, swim speed, and use of cold-water thermal refuges. Individuals with different migratory phenotypes migrated upstream through riverine corridors with variable availability of cold-water thermal refuges and mainstem temperatures. Use of cold-water refuges (CWRs) decreased accumulated sublethal exposures to physiologically stressful temperatures when measured in degree-days above 20, 21, and 22°C. The availability of CWRs was an order of magnitude more effective in lowering accumulated sublethal exposures under current and future mainstem temperatures for summer steelhead than fall Chinook Salmon. We considered two emergent model outcomes, survival and percent of available energy used, in relation to thermal heterogeneity and migratory phenotype. Mean percent energy loss attributed to future warmer mainstem temperatures was at least two times larger than the difference in energy used in simulations without CWRs for steelhead and salmon. We also found that loss of CWRs reduced the diversity of energy-conserving migratory phenotypes when we examined the variability in entry timing and travel time outside of CWRs in relation to energy loss. Energy-conserving phenotypic space contracted by 7%-23% when CWRs were unavailable under the current thermal regime. Our simulations suggest that, while CWRs do not entirely mitigate for stressful thermal exposures in mainstem rivers, these features are important for maintaining a diversity of migration phenotypes. Our study suggests that the maintenance of diverse portfolios of migratory phenotypes and cool- and cold-water refuges might be added to the suite of policies and management actions presently being deployed to improve the likelihood of Pacific salmonid persistence into a future characterized by climate change.

Foundation plant species provide resilience and microclimatic heterogeneity in drylands

Climate change profoundly influences plants and animals in all ecosystems including drylands such as semi-arid and arid scrublands and grasslands. At the peak of an extended megadrought in the Southwestern USA, the microclimatic refuges provided by foundation plant species and through associated vegetation were examined. Shrubs and open interstitial spaces without a canopy but with annual plants were instrumented in 2016 and the wet season of 2017 in the central drylands of California. In both years and all seasons tested, vegetation significantly mediated fine-scale near-surface air temperature and relative soil moisture content—defined here as microclimate. The foundation species with other vegetation provided the most significant thermal refuge potential capacity for other plants and animals, but there was variation by growing season. Soil moisture content was frequently increased by the direct canopy effects of shrubs. This evidence suggests that the climate many plants and animals experience, even during an extended megadrought, is mediated by the local plants in highly impacted drylands with anthropogenic disturbance and significant water-induced challenges. Foundation species such as shrubs in drylands function as a potent starting point in examining the ecological relevance of climate at scales germane to many species locally. An ecological framework for climate resilience using shrubs will improve conservation and restoration planning in drylands.

Model software and input for simulation of thermal refuge effects on stream trout populations

Model software and input for simulation of thermal refuge effects on stream trout populations

Range expansion and establishment of a non-indigenous tunicate (Diplosoma listerianum)in thermal refugia is mediated by environmental variability in changing coastal environments

Coastal marine non-indigenous species (NIS) have expanded their range into Atlantic Canada under more variable and anomalously warmer sea surface temperature (SST) conditions observed since 2012. In this study we apply species distribution modeling (SDM), in combination with empirical validation using annual field monitoring surveys, to assess the influence of anomalously warmer ocean conditions on the range expansion of NIS into coastal Nova Scotia using the recently established colonial tunicate Diplosoma listerianum as a test case. SDM accurately predicted D. listerianum’s introduction to Atlantic Canada, including its range expansion, contraction, and long-term persistence within thermal refugia as local environmental conditions generally warmed between 2012 and 2019. This tunicate’s ability to survive the winter was the primary constraint on its range expansion into Nova Scotia prior to 2012. The identification of local thermal refugia in which D. listerianum spread under anomalously warm winter conditions and persisted during colder winters provides new insights and a mechanism for range expansion and establishment of NIS. Ultimately, interannual modeling and long-term projections employed here can be applied to predict, monitor, and manage transient range expansion and long-term persistence of NIS within changing environments.

The impact of coastal upwelling on coral reef ecosystem under anthropogenic influence: Coral reef community and its response to environmental factors

Mass coral bleaching is a severe threat to coral reefs in the era of global warming, and upwelling areas may serve as thermal refugia for coral reefs by mitigating the effects of heating anomalies. This study seeks to examine the impact of the Qiongdong upwelling (QDU) on the coastal coral reefs of the Hainan Island. The environmental variables (i.e., physiochemical parameters, coastal land use) differed significantly in upwelling and non-upwelling areas. In particular, the presence of colder and more saline water was the important indicator of coastal upwelling. In the QDU, turf algae and macroalgae dominated the algae communities, but habitat-specific environment and coastal development contributed to lower calcified algae, coral larvae replenishment and live corals cover. Lower abundance of coral reefs fishes could be attributed to the disappearance of live corals, increase of algae coverage, low habitat complexity and strong wave exposure at upwelling stations. Upwelling-related environmental parameters, wave exposure, land use and potential local stressors were revealed as major driving factors related to significant spatial changes in stony coral and fish communities. Overall, the present findings suggest that the Qiongdong upwelling has the potential to provide thermal refuge for coral reefs, but is severely affected by local anthropogenic activities.

Present and future thermal regimes of intertidal groundwater springs in a threatened coastal ecosystem

Abstract. In inland settings, groundwater discharge thermally modulates receiving surface water bodies and provides localized thermal refuges; however, the thermal influence of intertidal springs on coastal waters and their thermal sensitivity to climate change are not well studied. We addressed this knowledge gap with a field- and model-based study of a threatened coastal lagoon ecosystem in southeastern Canada. We paired analyses of drone-based thermal imagery with in situ thermal and hydrologic monitoring to estimate discharge to the lagoon from intertidal springs and groundwater-dominated streams in summer 2020. Results, which were generally supported by independent radon-based groundwater discharge estimates, revealed that combined summertime spring inflows (0.047 m3 s−1) were comparable to combined stream inflows (0.050 m3 s−1). Net advection values for the streams and springs were also comparable to each other but were 2 orders of magnitude less than the downwelling shortwave radiation across the lagoon. Although lagoon-scale thermal effects of groundwater inflows were small compared to atmospheric forcing, spring discharge dominated heat transfer at a local scale, creating pronounced cold-water plumes along the shoreline. A numerical model was used to interpret measured groundwater temperature data and investigate seasonal and multi-decadal groundwater temperature patterns. Modelled seasonal temperatures were used to relate measured spring temperatures to their respective aquifer source depths, while multi-decadal simulations forced by historic and projected climate data were used to assess long-term groundwater warming. Based on the 2020–2100 climate scenarios (for which 5-year-averaged air temperature increased up to 4.32∘), modelled 5-year-averaged subsurface temperatures increased 0.08–2.23∘ in shallow groundwater (4.2 m depth) and 0.32–1.42∘ in the deeper portion of the aquifer (13.9 m), indicating the depth dependency of warming. This study presents the first analysis of the thermal sensitivity of groundwater-dependent coastal ecosystems to climate change and indicates that coastal ecosystem management should consider potential impacts of groundwater warming.

Behavioural temperature regulation is a low priority in a coral reef fish (Plectropomus leopardus): insights from a novel behavioural thermoregulation system

Current understanding of behavioural thermoregulation in aquatic ectotherms largely stems from systems such as 'shuttle boxes', which are generally limited in their capacity to test large-bodied species. Here, we introduce a controlled system that allows large aquatic ectotherms to roam freely in a tank at sub-optimal temperatures, using thermal refuges to increase body temperature to their thermal optimum as desired. Of the 10 coral grouper (Plectropomus leopardus; length ∼400 mm) implanted with thermal loggers, three fish maintained themselves at the ambient tank temperature of 17.5-20.5°C for the entire 2-4 days of the trial. Of the other seven fish, body temperature never exceeded ∼21.5°C, which was well below the temperature available in the thermal refuges (∼31°C) and below the species' optimal temperature of ∼27°C. This study adds to a growing literature documenting an unexpected lack of behavioural thermoregulation in aquatic ectotherms in controlled, heterothermal environments.

Identification of Thermal Refuges and Water Temperature Patterns in Salmonid-Bearing Subarctic Rivers of Northern Quebec

In summer, salmonids can experience thermal stress during extreme weather conditions. This may affect their growth and even threaten their survival. Cool water zones in rivers constitute thermal refuges, allowing fish to be more comfortable to grow and survive in extreme events. Therefore, identifying and understanding the spatiotemporal variability of discrete thermal refuges and larger scale cooling zones in rivers is of fundamental interest. This study analyzes thermal refuges as well as cooling zones in two salmonid rivers in a subarctic climate by use of thermal infrared (TIR) imagery. The two studied rivers are the Koroc and Berard Rivers, in Nunavik, Quebec, Canada. On the 17 km studied section of the Berard River, four thermal refuges and five cooling zones were detected, covering 46% of the surveyed section of the river. On the 41 km section studied for the Koroc River, 67 thermal refuges and five cooling zones were identified which represent 32% of the studied section of the river. 89% of identified thermal refuges and about 60% of cooling zones are groundwater-controlled. Continuity of permafrost and shape of the river valley were found to be the main parameters controlling the distribution of refuges and cooling zones. These data provide important insights into planning and conservation measures for the salmonid population of subarctic Nunavik rivers.

Upwelling enhances seaweed nutrient quality, altering feeding behavior and growth rates in an intertidal sea urchin, Loxechinus albus

Upwelling systems deliver nutrient-rich water into coastal ecosystems, influencing primary productivity and potentially altering seaweed-herbivore interactions. Upwelling bottom-up effects on distinct trophic levels are well-known. However, their influence on seaweed biomolecules and on algae-herbivore interactions and growth are less known. The aim of this study was threefold: i) to compare physical-chemical characteristics and nutrient levels in the water of upwelling (U) and downwelling (DU) zones, ii) to quantify their influence on the content of protein and carbohydrates in seaweed tissues of representative U and DU locations, and iii) to experimentally assess their effect on the feeding behavior and growth of a prominent intertidal herbivore, the sea urchin Loxechinus albus. Waters from U zones showed lower temperatures and pH, and higher phosphate concentrations than those from downwelling zones. Similarly, the tissue of seaweeds from a U location had significantly more proteins and carbohydrates than those from a DU location. The origin location of the sea urchins had a significant influence on consumption and growth rates: in general, those coming from a site with U conditions consumed and grew more than those coming from DU conditions. The quality of the algae was a significant factor on consumption rates, although in the case of preference trials, this factor interacted with sea urchin origin location. Our results show that the availability and quality of the food in upwelling zones has an influence on herbivore-seaweed direct interactions. However, these interactions and the growth of the sea urchins were also related to the coastal site and conditions from which the sea urchins came from. These results are relevant considering the expected impact of climate change on the world's oceans, and the importance of U zones as thermal (cold water) refuges for marine ectotherms.

Seasonal Movement Patterns and Summertime Use of Thermal Refuge Areas by Muskellunge in the Nontidal Potomac River, Maryland

AbstractThe nontidal Potomac River is Maryland's most popular freshwater fishery, largely due to the multiple angling opportunities and accessibility. Increasing popularity of the Muskellunge Esox masquinongy fishery has prompted further investigation of Muskellunge seasonal movement patterns and summer habitat use during thermally stressful periods. Radiotelemetry was used to investigate seasonal movement and summertime habitat use by Muskellunge in the Potomac River from 2017 to 2020. Movement was greatest during spring, with reduced movement during summer, fall, and winter. Potomac River main‐stem temperatures during summer often exceeded levels considered stressful to Muskellunge. During thermally stressful periods, Muskellunge sought thermal refugia and movements ceased. A generalized linear mixed model predicted that (1) when daily average temperatures reached 24.0°C, over 50% of the Muskellunge habitually occupied thermal refugia and (2) at 26.0°C, greater than 90% of tagged Muskellunge were located in refuge sites. Aggregation of Muskellunge in thermal refugia during summer has implications for increased angling vulnerability and fishing mortality in a warming climate, which could potentially limit the quality of these sport fisheries in southern river systems.

Microclimate predicts kelp forest extinction in the face of direct and indirect marine heatwave effects.

Marine heatwaves threaten the persistence of kelp forests globally. However, the observed responses of kelp forests to these events have been highly variable on local scales. Here, we synthesize distribution data from an environmentally diverse region to examine spatial patterns of canopy kelp persistence through an unprecedented marine heatwave. We show that, although often overlooked, temperature variation occurring at fine spatial scales (i.e., a few kilometers or less) can be a critical driver of kelp forest persistence during these events. Specifically, though kelp forests nearly all persisted toward the cool outer coast, inshore areas were >3°C warmer at the surface and experienced extensive kelp loss. Although temperatures remained cool at depths below the thermocline, kelp persistence in these thermal refugia was strongly constrained by biotic interactions, specifically urchin populations that increased during the heatwave and drove transitions to urchin barrens in deeper rocky habitat. Urchins were, however, largely absent from mixed sand and cobble benthos, leading to an unexpected association between bottom substrate and kelp forest persistence at inshore sites with warm surface waters. Our findings demonstrate both that warm microclimates increase the risk of habitat loss during marine heatwaves and that biotic interactions modified by these events will modulate the capacity of cool microclimates to serve as thermal refugia.

Experimental Investigation of a Prototype Thermal Refuge for Trout

Recent years have seen an increase in surface water temperatures in several streams and fisheries, which has a detrimental effect on cold-water species such as trout. One possibility to resolve this issue is to create localized refugia of colder water generated through active cooling. The current work focuses on a prototype thermal refugia design and field testing. Various configurations of the prototype thermal refugia were tested in a stream, which could benefit from additional refugia regions. The prototypes featured a staggered, tube-bundle heat exchanger placed inside an enclosure with an aperture. The results demonstrate that in remote locations, man-made refugia can be provided. While the base enclosure (91.5 cm × 91.5 cm × 45.8 cm) allowed for excess mixing with the warmer free stream and resulted in low performance (dimensionless temperature difference of θ¯avg= 0.07), additional modifications improved performance. By utilizing a panel or an extension, the dimensionless temperature difference quadrupled (θ¯avg= 0.26) while the average heat transfer per dimensionless temperature difference was reduced to approximately one-fifth (1.92 kWth/θ¯avg) of the base enclosure. But these configurations increased the standard deviation of the temperature differences inside the refugia due to localized cooling. The combination of the panel and the extension did not further increase the standard deviation but resulted in an even higher dimensionless temperature difference (θ¯avg= 0.55) and a lower heat transfer per dimensionless temperature difference (0.81 kW/θ¯avg). This suggests that the enclosure design can be used to achieve a desirable temperature differential while maintaining a reasonable spatial fluctuation in that temperature difference and power requirement.

Overwinter survival of Corbicula fluminea in a central Minnesota lake.

Although Corbicula fluminea has been one of the more prolific freshwater invasive species in the world, previous studies have suggested a low probability for overwinter survival in northern latitudes without an artificially created thermal refuge. The discovery of live C. fluminea in a central Minnesota lake absent any known thermal refuge in 2020 presented an opportunity to further evaluate the overwinter survival and population structure of C. fluminea at the presumed edge of their potential range. The population was monitored from December 2020 through September 2021 alongside water temperature to better understand at which temperatures C. fluminea survived and if the population structure suggested reproduction occurring in the lake. We documented live C. fluminea in temperatures as low as 0.3°C. Shell size of recovered individuals suggested multiple cohorts, and the appearance of a new cohort at the end of the study, indicating active reproduction in the lake and suggesting the population had likely been present in the lake for at least two winters by the conclusion of the study period. Our findings provide evidence of the survival below historically documented lower lethal temperature limits and suggests adaptations to modeling predicting suitable habitat, both present and in a changing climate, are necessary to better assess risk of invasion by this species.

Selective deep water coral bleaching occurs through depth isolation

Climate change is degrading coral reefs around the world. Mass coral bleaching events have become more frequent in recent decades, leading to dramatic declines in coral cover. Mesophotic coral ecosystems (30–150 m depth) comprise an estimated 50–80 % of global coral reef area. The potential for these to act as refuges from climate change is unresolved. Here, we report three mesophotic-specific coral bleaching events in the northern Red Sea over the course of eight years. Over the last decade, faster temperature increases at mesophotic depths resulted in ~50 % decline in coral populations, while the adjacent shallow coral reefs remained intact. Further, community structure shifted from hard coral dominated to turf algae dominated throughout these recurrent bleaching events. Our results do not falsify the notion of the northern Red Sea as a thermal refuge for shallow coral reefs, but question the capacity of mesophotic ecosystems to act as a universal tropical refuge.

On Internal Tides Driving Residual Currents and Upwelling on an Island

AbstractWe present a modeling study of an idealized island with sloping sides in a stratified ocean with internal tides propagating through the domain, which transform into nonlinear internal waves as they encounter shallow depths. In the base condition, representative of a mid‐latitude island with relatively steep super critical slopes, the nonlinear internal waves shoaling on the island wrap around the island, scatter, and create an asymmetric wavefield. Radiation stress gradients from this wavefield and rotation are balanced by pressure gradients and turbulent vertical diffusion in the momentum equations which drive residual mean currents around the island. We explore the effects of different physical parameters including no rotation, higher Froude number, subcritical slope, addition of barotropic tides, larger excursion number, and addition of large‐scale mean currents. All simulations showed formation of residual currents of varying intensity, upwelled deep waters in shallow regions, and cooler waters on the side of the island consistent with the direction of incoming internal tides. While the no rotation and strong mean flow conditions created some upwelling, of all the modeled conditions, the subcritical slope has the greatest potential for creating favorable conditions for benthic organisms through enhanced upwelling. Areas of the world's oceans with stratified waters sufficient to sustain internal tide motions, strong internal tide energy, and local subcritical bottom slopes are likely to have cooler waters and more upwelled deep waters than the surrounding areas, and potentially serve as thermal refugia from future ocean warming.