Colin T. Kremer

A Global Pattern of Thermal Adaptation in Marine Phytoplankton

Local Optima for Plankton Recent ocean warming has changed the seasonality and composition of the marine phytoplankton. Thomas et al. (p. 1085, published online 25 October) investigated the direct effect of temperature on phytoplankton. By fitting published data to reaction temperature norms for 194 phytoplankton strains isolated from a wide latitudinal range, the resultant temperature-related traits (maximum growth rate, temperature optimum, and thermal niche width) reveal latitudinal trends in temperature optima and diversity. Biogeographical differences indicate an increased susceptibility of tropical strains to further warming, and modeling predicts poleward shifts of tropical strains and a loss of phytoplankton diversity in the tropics within the next hundred years. Optimal growth rates for phytoplankton correlate with local temperature and predict movement toward the poles with warming. Rising ocean temperatures will alter the productivity and composition of marine phytoplankton communities, thereby affecting global biogeochemical cycles. Predicting the effects of future ocean warming on biogeochemical cycles depends critically on understanding how existing global temperature variation affects phytoplankton. Here we show that variation in phytoplankton temperature optima over 150 degrees of latitude is well explained by a gradient in mean ocean temperature. An eco-evolutionary model predicts a similar relationship, suggesting that this pattern is the result of evolutionary adaptation. Using mechanistic species distribution models, we find that rising temperatures this century will cause poleward shifts in species’ thermal niches and a sharp decline in tropical phytoplankton diversity in the absence of an evolutionary response.

Global biogeochemical impacts of phytoplankton: a trait‐based perspective

Summary Phytoplankton are key players in the global carbon cycle, contributing about half of global primary productivity. Within the phytoplankton, functional groups (characterized by distinct traits) have impacts on other major biogeochemical cycles, such as nitrogen, phosphorus and silica. Changes in phytoplankton community structure, resulting from the unique environmental sensitivities of these groups, may significantly alter elemental cycling from local to global scales. We review key traits that distinguish major phytoplankton functional groups, how they affect biogeochemistry and how the links between community structure and biogeochemical cycles are modelled. Finally, we explore how global environmental change will affect phytoplankton communities, from the traits of individual species to the relative abundance of functional groups, and how that, in turn, may alter biogeochemical cycles. Synthesis. We can increase our mechanistic understanding of the links between the community structure of primary producers and biogeochemistry by focusing on traits determining functional group responses to the environment (response traits) and their biogeochemical functions (effect traits). Identifying trade-offs including allometric and phylogenetic constraints among traits will help parameterize predictive biogeochemical models, enhancing our ability to anticipate the consequences of global change.

Synchronous dynamics of zooplankton competitors prevail in temperate lake ecosystems

Although competing species are expected to exhibit compensatory dynamics (negative temporal covariation), empirical work has demonstrated that competitive communities often exhibit synchronous dynamics (positive temporal covariation). This has led to the suggestion that environmental forcing dominates species dynamics; however, synchronous and compensatory dynamics may appear at different length scales and/or at different times, making it challenging to identify their relative importance. We compiled 58 long-term datasets of zooplankton abundance in north-temperate and sub-tropical lakes and used wavelet analysis to quantify general patterns in the times and scales at which synchronous/compensatory dynamics dominated zooplankton communities in different regions and across the entire dataset. Synchronous dynamics were far more prevalent at all scales and times and were ubiquitous at the annual scale. Although we found compensatory dynamics in approximately 14% of all combinations of time period/scale/lake, there were no consistent scales or time periods during which compensatory dynamics were apparent across different regions. Our results suggest that the processes driving compensatory dynamics may be local in their extent, while those generating synchronous dynamics operate at much larger scales. This highlights an important gap in our understanding of the interaction between environmental and biotic forces that structure communities.

Coexistence in a variable environment: Eco-evolutionary perspectives

A central question in community ecology is the means by which species coexist. Models of coexistence often assume that species have fixed trait values and consider questions such as how tradeoffs and environmental variation influence coexistence and diversity. However, species traits can be dynamic, varying between populations and individuals and changing over time as species adapt and evolve, at rates that are relevant to ecological processes. Consequently, adding evolution to ecological coexistence models may modify their predictions and stability in complex or unexpected ways. We extend a well-studied coexistence mechanism depending on resource fluctuations by allowing evolution along a tradeoff between maximum growth rate and competitive ability. Interactions between favorable season length and the period of fluctuations constrain coexistence, with two species coexistence favored by intermediate season length and arising through evolutionary branching or non-local invasion. However, these results depend on the relative rates of ecological and evolutionary processes: rapid evolution leads to a complete breakdown of otherwise stable coexistence. Other coexistence mechanisms should be evaluated from an evolutionary perspective to examine how evolutionary forces may alter predicted ecological dynamics.

Rapid thermal adaptation in a marine diatom reveals constraints and trade-offs

Rapid evolution in response to environmental change will likely be a driving force determining the distribution of species across the biosphere in coming decades. This is especially true of microorganisms, many of which may evolve in step with warming, including phytoplankton, the diverse photosynthetic microbes forming the foundation of most aquatic food webs. Here we tested the capacity of a globally important, model marine diatom Thalassiosira pseudonana, for rapid evolution in response to temperature. Selection at 16 and 31°C for 350 generations led to significant divergence in several temperature response traits, demonstrating local adaptation and the existence of trade-offs associated with adaptation to different temperatures. In contrast, competitive ability for nitrogen (commonly limiting in marine systems), measured after 450 generations of temperature selection, did not diverge in a systematic way between temperatures. This study shows how rapid thermal adaptation affects key temperature and nutrient traits and, thus, a populations long-term physiological, ecological, and biogeographic response to climate change.

Incomplete loss of a conserved trait: function, latitudinal cline, and genetic constraints

Retention of nonfunctional traits over evolutionary time is puzzling, because the cost of trait production should drive loss. Indeed, several studies have found nonfunctional traits are rapidly eliminated by selection. However, theory suggests that complex genetic interactions and a lack of genetic variance can constrain evolution, including trait loss. In the mustard family Brassicaceae the conserved floral condition includes four long and two short stamens, but we show that short stamens in the highly self‐pollinating mustard Arabidopsis thaliana do not significantly increase selfed seed set, suggesting that the trait has lost most or all of its function after the transition to selfing. We find that short stamen loss is common in native populations. Loss is incomplete and decreases with increasing latitude, a cline unexplained by correlations with flowering time or ovule count (which also vary with latitude). Using recombinant inbred lines derived from a cross between plants at the latitudinal extremes of the native range, we found three QTLs affecting short stamen number, with epistasis among them constraining stamen loss. Constraints on stamen loss from both epistasis and low genetic variance may be augmented by high selfing rates, suggesting that these kinds of constraints may be common in inbred species.