Compounds of the 20 elements needed by living organisms are relatively soluble in water and therefore vulnerable to being leached and lost from terrestrial ecosystems during mineralization. Intact systems have thus acquired retention mechanisms that sequester nutrients and minimize losses. Such mechanisms should be particularly important where rainfall is very high but direct evidence of retention mechanisms in tropical forests is scarce (Snaddon et al. 2012, Turner et al. 2007).

Bird's nest ferns (Asplenium nidus complex) (Yatabe & Murakami 2003) are common epiphytes of the Old World tropics and house a high abundance of arthropods (Ellwood & Foster 2004). Through interception and retention of leaf litter (Paoletti et al. 1991, Piggott 1996, Richardson 1999), epiphytes interrupt litterfall dynamics (Clark et al. 1998, Nadkarni & Matelson 1991) and delay the return of nutrients to the forest floor (Nadkarni 1984). Precipitation percolating through the canopy as throughfall is enriched as nutrients are leached from plant surfaces (Levia & Frost 2006). Water flowing down the trunk of trees as stemflow is further enriched from prolonged contact and accumulated nutrient deposits on the trunk (Levia & Frost 2003, Liu et al. 2002). Epiphytes can alter stemflow nutrient concentrations by slowing water percolation and by nutrient uptake and release (Awasthi et al. 1995, Strigel et al. 1994).

Leaf defences, leaf nutritional quality and leaf expansion rates may vary with resource availabilities to plants. Such variation could affect rates of leaf loss to herbivores, particularly along the steep resource gradients in disturbed forests. Intraspecific and interspecific variation in leaf damage and leaf expansion rates were measured on dipterocarp seedlings planted into secondary forests 1, 5 and 15 y after logging, and in adjacent primary forest of Sabah, Malaysia. Herbivory rates or amounts of leaf damage were compared across habitats and species for expanding, recently expanded, and mature leaves of Shorea leprosula and Dryobalanops lanceolata (Dipterocarpaceae). In all four habitats, leaves of the faster growing S. leprosula sustained higher rates and amounts of leaf-area loss than did the tougher leaves of slower growing D. lanceolata. Expanding leaves accumulated more leaf-area loss per week than did mature leaves. In all habitats and in both species, more than 25% of expanding leaves disappeared entirely. Rates of leaf-area loss per week differed among habitats for expanding leaves but not for mature leaves. In a relatively open, 1-y-old logged forest, faster leaf expansion reduced the time leaves spent in the most vulnerable stage; however, in S. leprosula a greater rate of leaf area loss countered the shorter expansion time. Thus, leaves accumulated similar total damages across habitats, and herbivory did not produce differences among habitats in seedling growth or mortality. High levels of resources may increase both leaf palatability and leaf expansion rates, with counteracting effects on herbivory.