Climate tracking by freshwater fishes suggests that fish diversity in temperate lakes may be increasingly threatened by climate warming

Aim

Many freshwater fishes are migrating poleward to more thermally suitable habitats in response to warming climates. In this study, we aimed to identify which freshwater fishes are most sensitive to climatic changes and asked: (i) how fast are lakes warming? (ii) how fast are fishes moving? and (iii) are freshwater fishes tracking climate?

Location

Ontario, Canada.

Methods

We assembled a database containing time series data on climate and species occurrence data from 10,732 lakes between 1986 and 2017. We calculated the rate of lake warming and climate velocity for these lakes. Climate velocities were compared with biotic velocities, specifically the rate at which the northernmost extent of each species shifted north.

Results

Lakes in Ontario warmed by 0.2°C decade⁻¹ on average, at a climate velocity of 9.4 km decade⁻¹ between 1986 and 2017. In response, some freshwater fishes have shifted their northern range boundaries with considerable interspecific variation ranging from species moving southwards at a rate of −58.9 km decade⁻¹ to species ranges moving northwards at a rate of 83.6 km decade⁻¹ over the same time period. More freshwater fish species are moving into northern lakes in Ontario than those being lost. Generally, predators are moving their range edges northwards, whereas prey fishes are being lost from northern lakes.

Main Conclusions

The concurrent loss of cooler refugia, combined with antagonistic competitive and predatory interactions with the range expanding species, has resulted in many commercially important predators moving their range edges northwards, whereas prey species have contracted their northern range edge boundaries. Trophic partitioning of range shifts highlights a previously undocumented observation of the loss of freshwater fishes from lower trophic levels in response to climate-driven migrations.     

Species traits and climate velocity explain geographic range shifts in an ocean‐warming hotspot

Species' ranges are shifting globally in response to climate warming, with substantial variability among taxa, even within regions. Relationships between range dynamics and intrinsic species traits may be particularly apparent in the ocean, where temperature more directly shapes species' distributions. Here, we test for a role of species traits and climate velocity in driving range extensions in the ocean‐warming hotspot of southeast Australia. Climate velocity explained some variation in range shifts, however, including species traits more than doubled the variation explained. Swimming ability, omnivory and latitudinal range size all had positive relationships with range extension rate, supporting hypotheses that increased dispersal capacity and ecological generalism promote extensions. We find independent support for the hypothesis that species with narrow latitudinal ranges are limited by factors other than climate. Our findings suggest that small‐ranging species are in double jeopardy, with limited ability to escape warming and greater intrinsic vulnerability to stochastic disturbances.     

The combined effects of climate change and river fragmentation on the distribution of Andean Amazon fishes

Upstream range shifts of freshwater fishes have been documented in recent years due to ongoing climate change. River fragmentation by dams, presenting physical barriers, can limit the climatically induced spatial redistribution of fishes. Andean freshwater ecosystems in the Neotropical region are expected to be highly affected by these future disturbances. However, proper evaluations are still missing. Combining species distribution models and functional traits of Andean Amazon fishes, coupled with dam locations and climatic projections (2070s), we (a) evaluated the potential impacts of future climate on species ranges, (b) investigated the combined impact of river fragmentation and climate change and (c) tested the relationships between these impacts and species functional traits. Results show that climate change will induce range contraction for most of the Andean Amazon fish species, particularly those inhabiting highlands. Dams are not predicted to greatly limit future range shifts for most species (i.e., the Barrier effect). However, some of these barriers should prevent upstream shifts for a considerable number of species, reducing future potential diversity in some basins. River fragmentation is predicted to act jointly with climate change in promoting a considerable decrease in the probability of species to persist in the long‐term because of splitting species ranges in smaller fragments (i.e., the Isolation effect). Benthic and fast‐flowing water adapted species with hydrodynamic bodies are significantly associated with severe range contractions from climate change.     

The abiotic and biotic factors limiting establishment of predatory fishes at their expanding northern range boundaries in Ontario,Canada

There is a poor understanding of the importance of biotic interactions in determining species distributions with climate change. Theory from invasion biology suggests that the success of species introductions outside of their historical ranges may be either positively (biotic acceptance) or negatively (biotic resistance) related to native biodiversity. Using data on fish community composition from two survey periods separated by approximately 28 years during which climate was warming, we examined the factors influencing the establishment of three predatory centrarchids: Smallmouth Bass (Micropterus dolomieu), Largemouth Bass (M. salmoides), and Rock Bass (Ambloplites rupestris) in lakes at their expanding northern range boundaries in Ontario. Variance partitioning demonstrated that, at a regional scale, abiotic factors play a stronger role in determining the establishment of these species than biotic factors. Pairing lakes within watersheds where each species had established with lakes sharing similar abiotic conditions where the species had not established revealed both positive and negative relationships between the establishment of centrarchids and the historical presence of other predatory species. The establishment of these species near their northern range boundaries is primarily determined by abiotic factors at a regional scale; however, biotic factors become important at the lake‐to‐lake scale. Studies of exotic species invasions have previously highlighted how spatial scale mediates the importance of abiotic vs. biotic factors on species establishment. Our study demonstrates how concepts from invasion biology can inform our understanding of the factors controlling species distributions with changing climate.     

Asymmetric boundary shifts of tropical montane Lepidoptera over four decades of climate warming

Aim To estimate whether species have shifted at equal rates at their leading edges (cool boundaries) and trailing edges (warm boundaries) in response to climate change. We provide the first such evidence for tropical insects, here examining elevation shifts for the upper and lower boundaries shifts of montane moths. Threats to species on tropical mountains are considered. Location Mount Kinabalu, Sabah, Malaysia. Methods We surveyed Lepidoptera (Geometridae) on Mount Kinabalu in 2007, 42 years after the previous surveys in 1965. Changes in species upper and lower boundaries, elevational extents and range areas were assessed. We randomly subsampled the data to ensure comparable datasets between years. Estimated shifts were compared for endemic versus more widespread species, and for species that reached their range limits at different elevations. Results Species that reached their upper limits at 2500–2700 m (n= 28 species, 20% of those considered) retreated at both their lower and upper boundaries, and hence showed substantial average range contractions (?300 m in elevational extent and ?45 km² in estimated range area). These declines may be associated with changes in cloud cover and the presence of ecological barriers (geological and vegetation transitions) which impede uphill movement. Other than this group, most species (n= 109, 80% of the species considered) expanded their upper boundaries upwards (by an average of 152 m) more than they retreated at their lower boundaries (77 m). Main conclusions Without constraints, leading margins shifted uphill faster than trailing margins retreated, such that many species increased their elevational extents. However, this did not result in increases in range area because the area of land available declines with increasing elevation. Species close to a major ecological/geological transition zone on the mountain flank declined in their range areas. Extinction risk may increase long before species reach the summit, even when undisturbed habitats are available.     

Biological traits predict shifts in geographical ranges of freshwater invertebrates during climatic warming and drying

Aim To test the ability of biological traits to predict climate‐related changes in geographical ranges of running‐water invertebrates. Location The Australian state of New South Wales and the Australian Capital Territory. Methods I analysed data from 8928 biomonitoring samples collected during a 16‐year period of generally rising air temperatures and declining precipitation. I used quantile regression to test for expansions and contractions on the climatically cooler, warmer, drier and wetter edges of the ranges of 120 invertebrate taxa, and correlated these shifts with the traits of thermophily (degree of preference for high versus low temperature) and rheophily (preference for flowing versus still water). Results The most commonly inferred range shifts were cool‐edge expansion plus warm‐edge contraction (71 taxa) and wet‐edge expansion plus dry‐edge contraction (71), but contractions from both cool and warm extremes (36) and from both dry and wet extremes (28) were also frequent. High‐temperature preference was associated with cool‐edge expansion and low‐temperature preference with wet‐edge expansion and contraction from all other extremes. A preference for flow was associated with wet‐edge expansion and dry‐edge contraction. Main conclusions Trait analysis has potential for predicting which species will expand their ranges and which will contract, but needs to be coupled with assessment of how the landscape provides each species with opportunities to track or avoid climate change. Improved quantification of climatically relevant traits and integration of trait analysis with species distribution modelling are likely to be beneficial.     

Interactions between range‐expanding tropical fishes and the northern Gulf of Mexico red snapper Lutjanus campechanus

Experimental investigation of the intensity of potential competitive interactions among increasingly abundant tropically‐associated grey Lutjanus griseus and lane snapper Lutjanus synagris and resident northern Gulf of Mexico (nGOM) red snapper Lutjanus campechanus was undertaken in large outdoor mesocosms. In pair‐wise interaction trials, compared with L. synagris, L. campechanus demonstrated significantly increased roving behaviour and predatory activity. While no significant difference in these activities was observed between L. campechanus and L. griseus, when all three snappers (Lutjanidae) were grouped together L. campechanus swimming activity significantly decreased in the presence of both tropically‐associated species. Overall, L. campechanus were more active and aggressive predators and appear to be competitively resistant to L. griseus and L. synagris. As lower latitude species have continued to become increasingly prevalent in nGOM habitats and regional warming continues to affect resident reef‐associated fishes, these findings contribute to the assessment of the effects of warming‐related species shifts upon nGOM fishes and document current partial resilience of L. campechanus to climate‐related expansions of tropical confamilials.     

Trophic niche segregation allows range‐extending coral reef fishes to co‐exist with temperate species under climate change

Changing climate is forcing many terrestrial and marine species to extend their ranges poleward to stay within the bounds of their thermal tolerances. However, when such species enter higher latitude ecosystems, they engage in novel interactions with local species, such as altered predator–prey dynamics and competition for food. Here, we evaluate the trophic overlap between range‐extending and local fish species along the east coast of temperate Australia, a hotspot for ocean warming and species range extensions. Stable isotope ratios (δ¹⁵N and δ¹³C) of muscle tissue and stomach content analysis were used to quantify overlap of trophic niche space between vagrant tropical and local temperate fish communities along a 730 km (6°) latitudinal gradient. Our study shows that in recipient temperate ecosystems, sympatric tropical and temperate species do not overlap significantly in their diet—even though they forage on broadly similar prey groups—and are therefore unlikely to compete for trophic niche space. The tropical and temperate species we studied, which are commonly found in shallow‐water coastal environments, exhibited moderately broad niche breadths and local‐scale dietary plasticity, indicating trophic generalism. We posit that because these species are generalists, they can co‐exist under current climate change, facilitating the existence of novel community structures.     

Reduced variability in range‐edge butterfly populations over three decades of climate warming

Populations at the high latitude edge of species’ geographical ranges are thought to show larger interannual population fluctuations, with subsequent higher local extinction risk, than those within the ‘core’ climatic range. As climate envelopes shift northward under climate warming, however, we would expect populations to show dampened variability. We test this hypothesis using annual abundance indices from 19 butterfly species across 79 British monitoring sites between 1976 and 2009, a period of climatic warming. We found that populations in the latter (warmer) half of the recording period show reduced interannual population variability. Species with more southerly European distributions showed the greatest dampening in population variability over time. Our results suggest that increases in population variability occur towards climatic range boundaries. British sites, previously existing at the margins of suitable climate space, now appear to fall closer to the core climatic range for many butterfly species.     

Dietary generalism accelerates arrival and persistence of coral‐reef fishes in their novel ranges under climate change

Climate change is redistributing marine and terrestrial species globally. Life‐history traits mediate the ability of species to cope with novel environmental conditions, and can be used to gauge the potential redistribution of taxa facing the challenges of a changing climate. However, it is unclear whether the same traits are important across different stages of range shifts (arrival, population increase, persistence). To test which life‐history traits most mediate the process of range extension, we used a 16‐year dataset of 35 range‐extending coral‐reef fish species and quantified the importance of various traits on the arrival time (earliness) and degree of persistence (prevalence and patchiness) at higher latitudes. We show that traits predisposing species to shift their range more rapidly (large body size, broad latitudinal range, long dispersal duration) did not drive the early stages of redistribution. Instead, we found that as diet breadth increased, the initial arrival and establishment (prevalence and patchiness) of climate migrant species in temperate locations occurred earlier. While the initial incursion of range‐shifting species depends on traits associated with dispersal potential, subsequent establishment hinges more on a species’ ability to exploit novel food resources locally. These results highlight that generalist species that can best adapt to novel food sources might be most successful in a future ocean.     

Projecting marine species range shifts from only temperature can mask climate vulnerability

Climate change is causing range shifts in many marine species, with implications for biodiversity and fisheries. Previous research has mainly focused on how species' ranges will respond to changing ocean temperatures, without accounting for other environmental covariates that could affect future distribution patterns. Here, we integrate habitat suitability modeling approaches, a high‐resolution global climate model projection, and detailed fishery‐independent and ‐dependent faunal datasets from one of the most extensively monitored marine ecosystems—the U.S. Northeast Shelf. We project the responses of 125 species in this region to climate‐driven changes in multiple oceanographic factors (e.g., ocean temperature, salinity, sea surface height) and seabed characteristics (i.e., rugosity and depth). Comparing model outputs based on ocean temperature and seabed characteristics to those that also incorporated salinity and sea surface height (proxies for primary productivity and ocean circulation features), we explored how an emphasis on ocean temperature in projecting species' range shifts can impact assessments of species' climate vulnerability. We found that multifactor habitat suitability models performed better in explaining and predicting species historical distribution patterns than temperature‐based models. We also found that multifactor models provided more concerning assessments of species' future distribution patterns than temperature‐based models, projecting that species' ranges will largely shift northward and become more contracted and fragmented over time. Our results suggest that using ocean temperature as a primary determinant of range shifts can significantly alter projections, masking species' climate vulnerability, and potentially forestalling proactive management.     

Macro-zooplankter responses to simulated climate warming in experimental freshwater microcosms

1. We report data collected from 48 replicated microcosm communities created to mimic plant‐dominated shallow lake and pond environments. Over a 2‐year period, the microcosms were subjected to warming treatments (continuous 3 °C above ambient and 3 °C above ambient during summer only), a nutrient addition treatment and the presence or absence of fish. We tracked macro‐zooplankter dynamics, censusing cladoceran populations at the species level, copepods at the order level and ostracods as a class. 2. Responses to warming were subtle. Cladoceran diversity and overall abundance were not significantly affected by warming, although measures of community evenness increased. Warming effects on patterns of population trajectories tended to be strongly seasonal and most apparent during periods of pronounced increase. Populations of the prevalent cladocerans, Chydorus sphaericus and Simocephalus vetulus, displayed idiosyncratic patterns, with evidence in the case of S. vetulus for a negative relationship between warming and body‐size at maturity. Copepod populations were reduced in size by warming, but those of ostracods increased. 3. The effects of the nutrient addition and fish treatments were strong and consistent, interacting little with warming effects in statistical models. Zooplankter abundance tended to be the highest in the fish‐free microcosms receiving additional nutrient inputs and lowest when fish were present and no nutrients were added. Both treatments reduced cladoceran diversity and community evenness. 4. We suggest that warming, independently, is unlikely to supplant the effects of changing nutrient loading and fish predation as the major driver of zooplankter dynamics in shallow lakes and ponds. Moreover, in the situations where warming was of significant influence in our experiment, the distinction between summer‐only warming and year‐around warming was blurred. This suggests that warming effects were most pervasive during the summer, at the upper end of the temperature spectrum.