Broadleaf Litter Controls Feather Moss Growth in Black Spruce and Birch Forests of Interior Alaska

Abstract

Plant–soil feedbacks can maintain or reinforce alternative states within ecological systems. In Alaskan boreal forests, changes in fire characteristics have stimulated the replacement of needle-leaf black spruce (Picea mariana) by broadleaf deciduous trees. Feather mosses have strong associations with forest type: They dominate black spruce forest understories and are uncommon in broadleaf stands, with consequences for nutrient cycling and carbon storage. Here we test a long-standing hypothesis that broadleaf litter directly excludes mosses with a field experiment in broadleaf paper birch (Betula neoalaskana) and black spruce stands. We established 30 plots (15 each in birch and spruce dominated areas) with three Hylocomium splendens transplants treated with one of three treatments in each plot (ambient leaf litter deposition, birch leaf litter exclusion or addition), and 30 natural H. splendens areas. We measured moss growth and reproductive potential over 3 years. A 1-year experiment assessed leaf leachate and physical structure impacts on moss growth. Moss shoot growth in natural patches was larger in spruce than in birch stands (24.8 vs. 17.3 mg) and H. splendens made large contributions to ecosystem productivity in spruce stands. In both stand types, we observed a 40% reduction in moss biomass between litter addition and exclusion treatments and litter additions decreased sporophyte production. We found no difference in growth for mosses treated for 1 year with leaf leachates or physical litter structures. Leaf litter effects appear strong enough to exclude mosses from broadleaf forests, providing experimental support for hypothesized plant–soil interactions that may stabilize alternate forest types.