Productivity, nutrient cycling, and succession in single- and mixed-species plantations of Casuarina equisetifolia, Eucalyptus robusta, and Leucaena leucocephala in Puerto Rico

Tree growth, biomass productivity, litterfall mass and nutrient content, changes in soil chemical properties and understory forest succession were evaluated over a 8.5-year period in singleand mixed-species (50 : 50) plantations of two N2-®xing species, Casuarina equisetifolia and Leucaena leucocephala, and a non-®xing species, Eucalyptus robusta. At the optimal harvest age for maximum biomass production (4 years), total aboveground biomass ranged from 63 Mg haÿ1 in the Eucalyptus monoculture to 124 Mg haÿ1 in the Casuarina/Leucaena mixture, and was generally greater in the mixed-species than in single-species treatments due to increased productivity of the N-®xing species in the mixed stands. Total litterfall varied from 5.3 to 10.0 Mg haÿ1 yearÿ1 among treatments, or between 5.9% and 13.2% of net primary production. Litterfall production and rates of nutrient return for N, P, K, Ca and Mg were generally highest for Leucaena, intermediate for Casuarina and lowest for Eucalyptus. These rates were usually higher in the mixed-species than in monospeci®c stands due to differences in biomass productivity, but varied considerably depending on their species composition. Total system carbon and nutrient pools (in biomass plus soils to 40-cm depth) for N, P, K, Ca, Mg, Mn at four years were consistently greater in the plantation treatments than in the unplanted control plots. Relative to the single-species plantations, these system pools were generally larger in the mixed-species plantations for C (ÿ10% to ‡10%), N (‡17% to‡50%), P (ÿ1% to‡63%), K (ÿ19% to‡46%), Ca (ÿ10% to‡48%), Mg (‡5% to‡57%) and Mn (‡19% to‡86%). Whole-tree harvests at four years would result in substantial system carbon and nutrient losses, although these estimated losses would not exceed the estimated gains realized during the four-year period of tree growth at this site. At 7.5 years, soil organic matter and effective cation exchange capacity were reduced in all plantation treatments relative to the control. Changes in soil nutrient content from 0 to 7.5 years were highly variable and not signi®cantly different among treatments, although stands containing Leucaena generally showed higher rates of nitrogen and phosphorus accretion in soils than those with Eucalyptus and/or Casuarina. Natural regeneration of secondary forest tree and shrub species increased over time in all plantation treatments. A total of 24 native or naturalized forest species were recorded in the plantations at 8.5 years. Woody species abundance at this age was signi®cantly greater beneath Casuarina than either Eucalyptus or the Eucalyptus/Leucaena mixed stands. Species richness and diversity, however, were greatest beneath stands containing Eucalyptus and/or Leucaena than in stands with Casuarina. # 1999 Elsevier Science B.V. All rights reserved.