Abstract. Over time scales of thousands to millions of years, and in the absence of rejuvenating disturbances that initiate primary or early secondary succession, ecosystem properties such as net primary productivity, decomposition, and rates of nutrient cycling undergo substantial declines termed ecosystem retrogression. Retrogression results from the depletion or reduction in the availability of nutrients, and can only be reversed through rejuvenating disturbance that resets the system; this differs from age-related declines in forest productivity that are driven by shorter-term depression of nutrient availability and plant ecophysiological process rates that occur during succession. Here we review and synthesize the findings from studies of long-term chronosequences that include retrogressive stages for systems spanning the boreal, temperate, and subtropical zones. Ecosystem retrogression has been described by ecologists, biogeochemists, geologists, and pedologists, each of which has developed somewhat independent conceptual frameworks; our review seeks to unify this literature in order to better understand the causes and consequences of retrogression. Studies of retrogression have improved our knowledge of how long-term pedogenic changes drive shorter-term biological processes, as well as the consequences of these changes for ecosystem development. Our synthesis also reveals that similar patterns of retrogression (involving reduced soil fertility, predictable shifts in organismic traits, and ecological processes) occur in systems with vastly different climatic regimes, geologic substrates, and vegetation types, even though the timescales and mechanisms driving retrogression may vary greatly among sites. Studies on retrogression also provide evidence that in many regions, high biomass or "climax" forests are often transient, and do not persist indefinitely in the absence of rejuvenating disturbance. Finally, our review highlights that studies on retrogressive chronosequences in contrasting regions provide unparalleled opportunities for developing general principles about the long-term feedbacks between biological communities and pedogenic processes, and how these control ecosystem development.

Long-term changes in soil phosphorus influence ecosystem development and lead to a decline in the productivity of forests in undisturbed landscapes. Much of the soil phosphorus occurs in a series of organic compounds that differ in their availability to organisms, but changes in the relative abundance of these compounds during pedogenesis remain unknown. We used alkaline extraction and solution phosphorus-31 nuclear magnetic resonance spectroscopy to assess the chemical nature of soil organic phosphorus along a 120,000-year postglacial chronosequence at Franz Josef, New Zealand. Inositol phosphates, DNA, phospholipids, and phosphonates accumulated rapidly during the first 500 years of soil development characterized by nitrogen limitation of biological productivity, but then declined slowly to low concentrations in older soils characterized by intense phosphorus limitation. However, the relative contribution of the various compounds to the total organic phosphorus varied along the sequence in dramatic and surprising ways. The proportion of inositol hexakisphosphate, conventionally considered to be relatively recalcitrant in the environment, declined markedly in older soils, apparently due to a corresponding decline in amorphous metal oxides, which weather to crystalline forms during pedogenesis. In contrast, the proportion of DNA, considered relatively bioavailable in soil, increased continually throughout the sequence, due apparently to incorporation within organic structures that provide protection from biological attack. The changes in soil organic phosphorus coincided with marked shifts in plant and microbial communities, suggesting that differences in the forms and bioavailability of soil organic phosphorus have ecological significance. Overall, the results strengthen our understanding of phosphorus transformations during pedogenesis and provide important insight into factors regulating the composition of soil organic phosphorus.

Understanding the mechanisms that regulate microbial community structure and activity is important if we Increasing the abundance of fungi relative to bacteria should favor are to manage C and N stocks in soils. In general, fungi C accrual, because fungi use C more efficiently, and are composed of more recalcitrant C compounds. We examined changes in soil become more important determinates of decomposition microbial community structure following cessation of tillage-based in soils returned to a more natural state (e.g., Bardgett agriculture and through subsequent succession in a C-accruing tall

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Summary1 Plant community theory often invokes competition to explain why species diversity declines as productivity increases. Competition for all resources might become more intense and lead to greater competitive exclusion or, alternatively, competition for light only could become more intense and exclude poor light competitors. 2 To test these hypotheses, we constructed communities of seven old-field species using combined monocultures. Constructs experienced no interspecific competition, only shoot competition or only root competition, with and without fertilizer. Diversity in these limited interaction communities was compared to diversity in unfertilized and fertilized mixtures of the seven species. 3 Diversity decreased with fertilization in mixtures and in communities with only root competition. Shoot competition had small effects on the community and did not contribute to changes in diversity with fertilization. 4 Root competition may strongly impact plant community structure in unproductive communities where light never becomes limiting, or under non-equilibrium conditions following human disturbances.