Thermal adaptation and ecological speciation

Ecological speciation is defined as the emergence of reproductive isolation as a direct or indirect consequence of divergent ecological adaptation. Several empirical examples of ecological speciation have been reported in the literature which very often involve adaptation to biotic resources. In this review, we investigate whether adaptation to different thermal habitats could also promote speciation and try to assess the importance of such processes in nature. Our survey of the literature identified 16 animal and plant systems where divergent thermal adaptation may underlie (partial) reproductive isolation between populations or may allow the stable coexistence of sibling taxa. In many of the systems, the differentially adapted populations have a parapatric distribution along an environmental gradient. Isolation often involves extrinsic selection against locally maladapted parental or hybrid genotypes, and additional pre- or postzygotic barriers may be important. Together, the identified examples strongly suggest that divergent selection between thermal environments is often strong enough to maintain a bimodal genotype distribution upon secondary contact. What is less clear from the available data is whether it can also be strong enough to allow ecological speciation in the face of gene flow through reinforcement-like processes. It is possible that intrinsic features of thermal gradients or the genetic basis of thermal adaptation make such reinforcement-like processes unlikely but it is equally possible that pertinent systems are understudied. Overall, our literature survey highlights (once again) the dearth of studies that investigate similar incipient species along the continuum from initial divergence to full reproductive isolation and studies that investigate all possible reproductive barriers in a given system.     

Non-climatic thermal adaptation: implications for species' responses to climate warming

There is considerable interest in understanding how ectothermic animals may physiologically and behaviourally buffer the effects of climate warming. Much less consideration is being given to how organisms might adapt to non-climatic heat sources in ways that could confound predictions for responses of species and communities to climate warming. Although adaptation to non-climatic heat sources (solar and geothermal) seems likely in some marine species, climate warming predictions for marine ectotherms are largely based on adaptation to climatically relevant heat sources (air or surface sea water temperature). Here, we show that non-climatic solar heating underlies thermal resistance adaptation in a rocky–eulittoral-fringe snail. Comparisons of the maximum temperatures of the air, the snail''s body and the rock substratum with solar irradiance and physiological performance show that the highest body temperature is primarily controlled by solar heating and re-radiation, and that the snail''s upper lethal temperature exceeds the highest climatically relevant regional air temperature by approximately 22°C. Non-climatic thermal adaptation probably features widely among marine and terrestrial ectotherms and because it could enable species to tolerate climatic rises in air temperature, it deserves more consideration in general and for inclusion into climate warming models.     

Effects of drought and phenology on GPP in Pinus sylvestris: a simulation study along a geographical gradient

1. A simple canopy model was developed for Scots Pine ( Pinus sylvestris L.) and applied to a transect of six meteorological stations in Europe. The model accounts for possible genetic adaptation of phenology of photosynthesis to the local climate and to decreases of gas exchange owing to drought.
2. Simulations accounting for adaptation of phenology to the local climate differed up to 20% from simulations using the same phenology parameter values for all locations.
3. A temperature increase of 3°C and a doubling of the CO₂ concentration, while adjusting the photosynthesis parameters to give approximately the observed changed photosynthesis of +30%, also increased the length of the growing season by 23–42%. Combination of increases in the rate of photosynthesis and the length of the growing season resulted in increases of yearly Gross Primary Productivity (GPP) from 72 to 101%. Increases in transpiration were smaller.
4. A decrease of the precipitation by 25% reduced this increase to 54–64%.
5. The relative magnitude of the simulated increases in GPP was similar for locations representing boreal, temperate and mediterranean climates.     

Genome-wide evolutionary response to a heat wave in Drosophila

Extreme climatic events can substantially affect organismal performance and Darwinian fitness. In April 2011, a strong heat wave struck extensive geographical areas of the world, including Western Europe. At that time, we happened to resume and extend a long-term time series of seasonal genetic data in the widespread fly Drosophila subobscura, which provided a unique opportunity to quantify the intensity of the genetic perturbation caused by the heat wave. We show that the spring 2011 genetic constitution of the populations transiently shifted to summer-like frequencies, and that the magnitude of the genetic anomaly quantitatively matched the temperature anomaly. The results provide compelling evidence that direct effects of rising temperature are driving adaptive evolutionary shifts, and also suggest a strong genetic resilience in this species.     

Breeding for the future: what are the potential impacts of future frost and heat events on sowing and flowering time requirements for Australian bread wheat (Triticum aestivium) varieties?

Extreme climate, especially temperature, can severely reduce wheat yield. As global warming has already begun to increase mean temperature and the occurrence of extreme temperatures, it has become urgent to accelerate the 5–20 year process of breeding for new wheat varieties, to adapt to future climate. We analyzed the patterns of frost and heat events across the Australian wheatbelt based on 50 years of historical records (1960–2009) for 2864 weather stations. Flowering dates of three contrasting‐maturity wheat varieties were simulated for a wide range of sowing dates in 22 locations for ‘current’ climate (1960–2009) and eight future scenarios (high and low CO₂ emission, dry and wet precipitation scenarios, in 2030 and 2050). The results highlighted the substantial spatial variability of frost and heat events across the Australian wheatbelt in current and future climates. As both ‘last frost’ and ‘first heat’ events would occur earlier in the season, the ‘target’ sowing and flowering windows (defined as risk less than 10% for frost (<0 °C) and less than 30% for heat (>35 °C) around flowering) would be shifted earlier by up to 2 and 1 month(s), respectively, in 2050. A short‐season variety would require a shift in target sowing window 2‐fold greater than long‐ and medium‐season varieties by 2050 (8 vs. 4 days on average across locations and scenarios, respectively), but would suffer a lesser decrease in the length of the vegetative period (4 vs. 7 days). Overall, warmer winters would shorten the wheat season by up to 6 weeks, especially during preflowering. This faster crop cycle is associated with a reduced time for resource acquisition, and potential yield loss. As far as favourable rain and modern equipment would allow, early sowing and longer season varieties (i.e. in current climate) would be the best strategies to adapt to future climates.