Canopy seed banks as time capsules of biodiversity in pasture-remnant tree crowns

Tropical pastures present multiple barriers to tree regeneration and restoration. Relict trees serve as "regeneration foci" because they ameliorate the soil microclimate and serve as safe spots for dispersers. Here, we describe another mechanism by which remnant trees may facilitate pasture regeneration: the presence of seed banks in the canopy soil that accumulates from decomposing epiphytes within the crowns of mature remnant trees in tropical cloud forest pastures. We compared seed banks of canopy soils (histosols derived from fallen leaves, fruits, flower, and twigs of host trees and epiphytes, dead bryophytes, bark, detritus, dead animals, and microorganisms, and dust that accumulate on trunks and the upper surfaces of large branches) in pastures, canopy soils in primary forest trees, and soil on the forest floor in Monteverde, Costa Rica. There were 5211 epiphytic and terrestrial plant seeds in the three habitats. All habitats were dominated by seeds in a relatively small number of plant families, most of which were primarily woody, animal pollinated, and animal dispersed. The density of seeds on the forest floor was greater than seed density in either pasture-canopy or forest-canopy soils; the latter two did not differ. Eight species in 44 families and 61 genera from all of the habitats were tallied. There were 37 species in the pasture-canopy soil, 33 in the forest-canopy soil, and 57 on the forest floor. Eleven species were common to all habitats. The mean species richness in the pasture canopy was significantly higher than the forest canopy (F =83.38; p < 0.02). Nonmetric multidimensional scaling ordination revealed that the communities were distinct. Greenhouse experiments verified that many of these seeds were viable, with 29 taxa germinating (23 taxa in pruned mats [mimic of exposed conditions] and 16 taxa in control mats [intact conditions]) within 2 months of observation. Nearly half the species that germinated were characteristic of primary forests (primary forest samples, 19%; pasture samples, 29%). This supports the idea that canopy seed banks of pasture trees can function as time capsules by providing propagules that are removed in both space and time from the primary forest. Their presence may enhance the ability of pastures to regenerate more quickly, reinforcing the importance of trees in agricultural settings.