Different Leaf Strategies Between Lithophytic and Terrestrial Orchids in a Subtropical Karst Forest

The leaf economic spectrum framework explains how plants optimize leaf traits for productivity, distribution, and stress tolerance. Orchids in Southwestern China's karst forests, especially lithophytic species, are challenged by prolonged drought and limited light availability. This study investigated different leaf strategies between lithophytic and terrestrial orchids under the harsh karst environment. We measured key leaf traits, including photosynthesis, structure, biomechanics, nitrogen allocation, and water relations, in twenty-two lithophytic and six terrestrial orchids in a subtropical karst forest. After accounting for phylogenetic influences, we found that lithophytic orchids had a higher leaf mass per area, cuticle thickness, and biomechanical resistance (F_p) but a lower maximum photosynthetic rate (A_max-mass), nitrogen allocation to photosynthesis (N_T), and saturated water content (SWC) than terrestrial orchids. These results suggest that lithophytic orchids prioritize structural investment and stress tolerance over photosynthetic efficiency. Across species, N_T correlated positively with A_max-mass and negatively with F_p, highlighting nitrogen allocation as a key mechanism in leaf cost-benefit strategies. Additionally, SWC emerged as a critical driver of variation in multiple traits, supporting its integration into the leaf economic spectrum for orchids in karst ecosystems. This study offers new insights into orchid adaptation in subtropical karst environments, with implications for plant resilience under changing climates.