The effect of mixing ground leaf litters to soil on the development of pitch pine ectomycorrhizal and soil arthropod communities in natural soil microcosm systems

The addition of leaf litter to soil influences both the nutrients and polyphenols of soil. It is likely that contrasting nutrient and polyphenolic composition of different plant litters may affect plant growth, mycorrhizal and soil arthropod communities. We report results from a microcosm experiment of effects of incorporation of three single leaf litter species and a mixture of all three on pitch pine seedling growth, their ectomycorrhizal community and soil arthropod community. The three litter species (pine, oak and huckleberry) represent co-dominant species within the New Jersey pine barrens ecosystem. We show that the leaf litters have different composition of nutrients and polyphenols, with rooting matrix containing pine litter having lower inorganic nitrogen content (1.6 microgram g(-1)) than oak (19.9 microgram g(-1)) and huckleberry (4.4 microgram g(-1)), but oak litter having the highest extractable phosphorus (13.3 cf. 0-0.08 microgram g(-1)) and total phenol content and lowest condensed tannin content. These differences were imparted to rooting matrix of homogenized humic (Oa) layer of pine barrens soil to which milled leaf litter was added and used in the microcosms. Pitch pine seedlings grew significantly better in un-amended rooting matrix (0.33+/-0.02 g) than any of the litter treatments (0.15+/-0.02-0.17+/-0.01 g) and tissue P concentrations tracked phosphate concentrations in the rooting matrix. Total P accumulation into plant tissue was higher in oak than control, attributable to a significantly higher (P<0.05) accumulation in roots (3.3+/-0.19 mg g(-1)) compared to other species (1.1+/-0.04-2.3+/-0.08 mg g(-1)). No relationship was seen between tissue N concentration and soil N, but seedlings growing in huckleberry litter amended soil accumulated less N than control. The effect of leaf litters on the ectomycorrhizal community composition were determined by PCA (first two axes accounted for 81% of the variance) and stepwise multiple regression analysis. These analyses showed that huckleberry leaf litter had a significant impact on mycorrhizal community composition with morphotypes Cg and DB being more abundant in the presence of huckleberry litter (178+/-13 cf. 68+/-15-106+/-15 for Cg and 141+/-11 cf. 88+/-23-111+/-18 for DB) and its influence of elevating nitrate nitrogen, organic nitrogen, total phenols and protein precipitation content of the rooting matrix. Mycorrhizal morphotypes BS and SB were significantly more abundant in the community where these soil factors were low in the absence of leaf litter addition. Total ectomycorrhizal abundance was negatively related to hydrolysable tannin concentration in the rooting matrix (r2=0.132, P<0.05). There was no influence of leaf litter type on mite density (dominated by non-burrowing phthiracarids), but collembolan density (dominated by Folsomia spp) showed a greater than threefold reduction in population density in the presence of leaf litter (F=6.47, P<0.05). Collembolan density was positively correlated with mycorrhizal morphotypes GS and SB (P<0.05) and negatively related to morphotypes DB (P<0.05) and soil extractable NH4-N (P<0.05), suggesting a possible selection of fungal species in their diet and a relationship between collembola and nitrification.