Are winter-active species vulnerable to climate warming? A case study with the wintergreen terrestrial orchid, Tipularia discolor.

In the eastern United States, winter temperature has been increasing nearly twice as fast as summer temperature, but studies of warming effects on plants have focused on species that are photosynthetically active in summer. The terrestrial orchid Tipularia discolor is leafless in summer and acquires C primarily in winter. The optimum temperature for photosynthesis in T. discolor is higher than the maximum temperature throughout most of its growing season, and therefore growth can be expected to increase with warming. Contrary to this hypothesis, experimental warming negatively affected reproductive fitness (number of flowering stalks, flowers, fruits) and growth (change in leaf area from 2010 to 2012) in T. discolor. Temperature in June-July was critical for flowering, and mean July temperature greater than 29 °C (i.e., 2.5 °C above ambient) eliminated reproduction. Warming of 1.2 °C delayed flowering by an average of 10 days and fruiting by an average of 5 days. Warming of 4.4 °C reduced relative growth rates by about 60%, which may have been partially caused by the direct effects of temperature on photosynthesis and respiration. Warming indirectly increased vapor pressure deficit (VPD) by 0.2-0.5 kPa, and leaf-to-air VPD over 1.3 kPa restricted stomatal conductance of T. discolor to 10-40% of maximum conductance. These results highlight the need to account for changes in VPD when estimating temperature responses of plant species under future warming scenarios. Increasing temperature in the future will likely be an important limiting factor to the distribution of T. discolor, especially along the southern edge of its range.