DIRECTING SPATIAL PATTERNS OF RECRUITMENT DURING AN EXPERIMENTAL URBAN WOODLAND RECLAMATION

Abstract

Studies of biological invasions indicate that natural recruitment of new spe- cies can occur as a nucleation phenomenon, in which scattered colonization foci spread and coalesce. Ecological reclamation of damaged lands might make use of this potential for enhanced natural dispersal, by inoculating sites with multiple small plantings to attract animal dispersers and other mutualists from nearby remnants of natural habitat. We con- ducted an experimental test of this proposition. On a 6-ha section of an abandoned municipal landfill in the New Jersey Meadowlands, we installed 16 clusters of 21 trees and shrubs in an array of fenced plots. Clusters contained seven native species known to: (1) attract bird dispersers to introduce propagules from remnants of off-site habitat; (2) contribute propagules by virtue of high reproductive output and clonal growth; and (3) accelerate woodland succession on open, degraded habitats. Average plant size was varied, with half the plots receiving larger trees and shrubs, to test whether woody plant size would enhance any attractive function. An additional eight empty plots were studied to estimate background rates of recruitment and to test for a fencing effect. Site preparation included the addition of 90 cm of fresh substrate, including organic matter, and a cover crop of annual grasses. Recruitment of woody plants inside and sur- rounding the experimental plots was examined for five years, and results were compared on the basis of treatment and recruitment mode (avian, wind, or clonal dispersal). Woody plant recruitment into experimental plots was rapid and substantial, primarily via dispersal from natural sources. Plots with larger plants attracted significantly more recruits at the outset, but this difference diminished over time. Fall seed rain samples yielded a mean estimate of 426 seeds/M2 within plots. However, size distributions of re- cruiting woody species increasingly shifted toward larger individuals each year. Experi- mental manipulations that opened seed beds for woody plant recruitment had short-lived effects, indicating a narrow window of opportunity for establishment. Spread of the planted species themselves was generally weak, although clonal growth contributed substantially to spread on the margins of plots. Most recruitment outside experimental plots was from external sources. A strong proximity component was found for bird-dispersed recruits, which were highly clustered near planted plots, with the highest densities near source populations on the site margin. Wind-dispersed trees and shrubs, by contrast, were not associated with planted plots and were concentrated near one corner of the site. Discounting plot interiors, total recruitment density for the site after 5 yr was -800 woody stems/ha, 36% via avian dispersal, 10% via clonal spread, and the remainder via wind-borne propagules. New recruits represented 26 woody plant species, all but four from external sources, and only five common species contributed more than a few recruits. We conclude that techniques for manipulating natural seed dispersal hold promise for ecological restoration, provided that background populations are available to supply colonists.