Stones as Fire Refugia for Ground-Dwelling Macroinvertebrates: Management Implications in Mediterranean Forestry

Abstract

Fire refugia are critical for post-disturbance recovery, yet microhabitats such as stones remain understudied despite their ubiquity and thermal persistence. This study tested whether the depth- and area-dependent refugial capacity of stones previously demonstrated in Mediterranean oak forests also operates in intensively managed plantations and how forest type and management modulate this capacity. Immediate wildfire effects (1–8 days post-fire) on ground-dwelling macroinvertebrates were quantified under 660 stones across burnt and unburnt native maritime pine and exotic eucalypt plantations following a medium- to high-severity wildfire. Stones acted as thermal refugia in both plantation types, with burial depths greater than 5 cm and surface areas greater than 500 cm2 predicting survival. Despite severe impacts (richness declined by 56% in pine and 63% in eucalypt; overall mortality exceeding 50%), diverse taxa persisted under stones, particularly ground spiders, ants, centipedes, rock bristletails, and harvestmen, while plant-associated and moisture-dependent groups suffered the highest losses. Native pine supported a higher abundance and richness per stone than exotic eucalypt in both burnt and unburnt conditions, reflecting management-driven differences in stone size, depth, and availability. These findings show that retaining sufficiently large, deeply buried stones during plantation establishment can enhance post-fire biodiversity recovery in increasingly fire-prone production landscapes.