From glacial refugia to hydrological microrefugia: Factors and processes driving the persistence of the climate relict tree Zelkova sicula

With only two tiny populations, the climate relict Zelkova sicula (Sicily, Italy) is one of the rarest trees in the world. It also represents the most marginal member of genus Zelkova that was widespread in the broadleaved forests thriving in warm-temperate climates throughout Eurasia until the Last Glacial Age. Occurring at the westernmost range of the genus under typical Mediterranean climate, the micro-topographic settings have always appeared crucial for the survival of this relict. However, the factors and processes actually involved in its persistence in the current refugia, as well as the response of similar relict trees in arid environments, are poorly understood worldwide. In the aim to elucidate these aspects, in the two sites hosting Z. sicula analyses of topographical attributes were combined with investigations on soil moisture dynamics. Additionally, plants' growth and spatial distribution patterns were analyzed to detect fine-scale differences between populations and assess the possible ecological amplitude of the species. Results revealed that convergent topographies are basic determinants of microrefugia in arid environments. Within the investigated sites, underground moisture never decreases below 25%, buffering seasonal rainfall fluctuations. Therefore, hydrological microrefugia play a key role in decoupling from regional climate, supporting the target species in coping with an unsuitable climatic envelope. Additionally, the inter-population variability of biometric attributes showed that individual growth is site-dependent and the species retains a relative ecological plasticity, whereas the strongly clumped spatial patterns confirmed the common clonal growth. On one hand, deeply incised landforms have acted as effective hydrologic microrefugia, on the other clonality coupled with triploidy supposedly improved the resistance of Z. sicula to harsh environments, though entailing inability to reproduce sexually. Most likely, sterility and environmental/physical barriers that have existed for millennia have prevented this relict from leaving the last suitable microrefugia, resulting in the two current rear edge populations.