Sarah E. Hobbie

Consequences of changing biodiversity

Human alteration of the global environment has triggered the sixth major extinction event in the history of life and caused widespread changes in the global distribution of organisms. These changes in biodiversity alter ecosystem processes and change the resilience of ecosystems to environmental change. This has profound consequences for services that humans derive from ecosystems. The large ecological and societal consequences of changing biodiversity should be minimized to preserve options for future solutions to global environmental problems.

Stoichiometry of soil enzyme activity at global scale

Extracellular enzymes are the proximate agents of organic matter decomposition and measures of these activities can be used as indicators of microbial nutrient demand. We conducted a global-scale meta-analysis of the seven-most widely measured soil enzyme activities, using data from 40 ecosystems. The activities of beta-1,4-glucosidase, cellobiohydrolase, beta-1,4-N-acetylglucosaminidase and phosphatase g(-1) soil increased with organic matter concentration; leucine aminopeptidase, phenol oxidase and peroxidase activities showed no relationship. All activities were significantly related to soil pH. Specific activities, i.e. activity g(-1) soil organic matter, also varied in relation to soil pH for all enzymes. Relationships with mean annual temperature (MAT) and precipitation (MAP) were generally weak. For hydrolases, ratios of specific C, N and P acquisition activities converged on 1 : 1 : 1 but across ecosystems, the ratio of C : P acquisition was inversely related to MAP and MAT while the ratio of C : N acquisition increased with MAP. Oxidative activities were more variable than hydrolytic activities and increased with soil pH. Our analyses indicate that the enzymatic potential for hydrolyzing the labile components of soil organic matter is tied to substrate availability, soil pH and the stoichiometry of microbial nutrient demand. The enzymatic potential for oxidizing the recalcitrant fractions of soil organic material, which is a proximate control on soil organic matter accumulation, is most strongly related to soil pH. These trends provide insight into the biogeochemical processes that create global patterns in ecological stoichiometry and organic matter storage.

Nitrogen limitation constrains sustainability of ecosystem response to CO2

Enhanced plant biomass accumulation in response to elevated atmospheric CO2 concentration could dampen the future rate of increase in CO2 levels and associated climate warming. However, it is unknown whether CO2-induced stimulation of plant growth and biomass accumulation will be sustained or whether limited nitrogen (N) availability constrains greater plant growth in a CO2-enriched world. Here we show, after a six-year field study of perennial grassland species grown under ambient and elevated levels of CO2 and N, that low availability of N progressively suppresses the positive response of plant biomass to elevated CO2. Initially, the stimulation of total plant biomass by elevated CO2 was no greater at enriched than at ambient N supply. After four to six years, however, elevated CO2 stimulated plant biomass much less under ambient than enriched N supply. This response was consistent with the temporally divergent effects of elevated CO2 on soil and plant N dynamics at differing levels of N supply. Our results indicate that variability in availability of soil N and deposition of atmospheric N are both likely to influence the response of plant biomass accumulation to elevated atmospheric CO2. Given that limitations to productivity resulting from the insufficient availability of N are widespread in both unmanaged and managed vegetation, soil N supply is probably an important constraint on global terrestrial responses to elevated CO2.

Plant functional types as predictors of transient responses of arctic vegetation to global change

Abstract. The plant functional types (growth forms) traditionally recognized by arctic ecologists provide a useful framework for predicting vegetation responses to, and effects on, ecosystem processes. These functional types are similar to those objectively defined by cluster analysis based on traits expected to influence ecosystem processes. Principal components analysis showed that two major suites of traits (related to growth rate and woodiness) explain the grouping of species into functional types. These plant functional types are useful because they (1) influence many ecological processes (e.g. productivity, transpiration, and nutrient cycling) in similar ways, (2) predict both responses to and effects on environment, including disturbance regime, and (3) show no strong relationship with traits determining migratory ability (so that no functional type will be eliminated by climatic change simply because it cannot migrate). Circumstantial evidence for the ecological importance of these functional types comes from the distribution of types along environmental gradients and the known ecological effects of traits (e.g., effects of litter quality on decomposition and of plant height on winter albedo) that characterize each functional type. The paleorecord provides independent evidence that some of these functional types have responded predictably to past climatic changes. Field experiments also show that plant functional types respond predictably to changes in soil resources (water and nutrients) but less predictably to temperature. We suggest that evidence for the validity of arctic plant functional types is strong enough to warrant their use in regional models seeking to predict the transient response of arctic ecosystems to global change.

NUTRIENT LIMITATION OF DECOMPOSITION IN HAWAIIAN FORESTS

In Hawaiian montane forests, we assessed whether the same nutrients limit decomposition and aboveground net primary production (ANPP) along a soil chronosequence where nutrients demonstrably and predictably limit ANPP. At three sites that vary in parent material age (300, 20 000, and 4.1 × 106 yr), we used fertilization to assess whether nitrogen (N) and/or phosphorus (P) limit decomposition. Reciprocal transplants using litter bags allowed us to distinguish limitation by externally supplied nutrients vs. limitation by nutrients within litter. Nutrient limitation of decomposition was not predictable from nutrient limitation of ANPP, in that elevated litter and soil N had only small, if any, effects on decomposition, even at the young site where N limits ANPP. At the oldest site where P limits ANPP, both elevated litter P and increased availability of soil N and P increased decomposition rates. Thus, nutrients may limit decomposition more strongly in low-P than in low-N ecosystems. Fertilization affected litter nutrient dynamics more strongly than it did decomposition, and we observed uptake of both N and P by decomposers that was not always accompanied by changes in decomposition rates. Such nutrient incorporation into decomposing litter may retain nutrients within ecosystems, even when nutrients do not limit decomposition rates.

TREE SPECIES EFFECTS ON DECOMPOSITION AND FOREST FLOOR DYNAMICS IN A COMMON GARDEN

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.

Impacts of Biodiversity Loss Escalate Through Time as Redundancy Fades

Give It Time Experimental ecological studies in recent years have provided a great deal of insight into how species diversify and influence ecosystem properties, but in most cases the experiments have been relatively brief (up to ∼5 years). Reich et al. (p. 589; see the Perspective by Cardinale) performed two 13- and 15-year grassland experiments and found that the effects of plant species richness on community-level processes like biomass production tend to be saturating at early stages but that those impacts grow stronger and more linear as experiments run longer. Stronger influences through time were largely driven by increasing amounts of “complementarity” among species, and these trends were correlated with greater expression of functional diversity in multispecies assemblages. Thus, the effects of diversity grow stronger through time as species gain more and more opportunity to vary in their use of the limiting biological resources in their environment, which emphasizes the functional importance of maintaining diversity in ecosystems. Long-term grassland experiments show that high-diversity species combinations become more functionally diverse with time. Plant diversity generally promotes biomass production, but how the shape of the response curve changes with time remains unclear. This is a critical knowledge gap because the shape of this relationship indicates the extent to which loss of the first few species will influence biomass production. Using two long-term (≥13 years) biodiversity experiments, we show that the effects of diversity on biomass productivity increased and became less saturating over time. Our analyses suggest that effects of diversity-dependent ecosystem feedbacks and interspecific complementarity accumulate over time, causing high-diversity species combinations that appeared functionally redundant during early years to become more functionally unique through time. Consequently, simplification of diverse ecosystems will likely have greater negative impacts on ecosystem functioning than has been suggested by short-term experiments.

THE RESPONSE OF TUNDRA PLANT BIOMASS, ABOVEGROUND PRODUCTION, NITROGEN, AND CO2 FLUX TO EXPERIMENTAL WARMING

We manipulated air temperature in tussock tundra near Toolik Lake, Alaska, and determined the consequences for total plant biomass, aboveground net primary production (ANPP), ecosystem nitrogen (N) pools and N uptake, and ecosystem CO2 flux. After 3.5 growing seasons, in situ plastic greenhouses that raised air temperature during the growing season had little effect on total biomass, N content, or growing-season N uptake of the major plant and soil pools. Similarly, vascular ANPP and net ecosystem CO2 exchange did not change with warming, although net primary production of mosses decreased with warming. Such general lack of response supports the hypothesis that productivity in tundra is constrained by the indirect effects of cold temperatures (e.g., low nutrient availability or short growing-season length) rather than by cold growing-season temperatures per se. Despite no effect on net ecosystem CO2 flux, air warming stimulated early-season gross photosynthesis (GP) and ecosystem respiration (ER) througho...

Interactions between litter lignin and soil nitrogen availability during leaf litter decomposition in a Hawaiian Montane Forest.

Previous work in a young Hawaiian forest has shown that nitrogen (N) limits aboveground net primary production (ANPP) more strongly than it does decomposition, despite low soil N availability. In this study, I determined whether (a) poor litter C quality (that is, high litter lignin) poses an overriding constraint on decomposition, preventing decomposers from responding to added N, or (b) high N levels inhibit lignin degradation, lessening the effects of added N on decomposition overall. I obtained leaf litter from one species, Metrosideros polymorpha, which dominates a range of sites in the Hawaiian Islands and whose litter lignin concentration declines with decreasing precipitation. Litter from three dry sites had lignin concentrations of 12% or less, whereas litter from two wet sites, including the study site, had lignin concentrations of more than 18%. This litter was deployed 2.5 years in a common site in control plots (receiving no added nutrients) and in N-fertilized plots. Nitrogen fertilization stimulated decomposition of the low-lignin litter types more than that of the high-lignin litter types. However, in contrast to results from temperate forests, N did not inhibit lignin decomposition. Rather, lignin decay increased with added N, suggesting that the small effect of N on decomposition at this site results from limitation of decomposition by poor C quality rather than from N inhibition of lignin decay. Even though ANPP is limited by N, decomposers are strongly limited by C quality. My results suggest that anthropogenic N deposition may increase leaf litter decomposition more in ecosystems characterized by low-lignin litter than in those characterized by high-lignin litter.

Winter regulation of tundra litter carbon and nitrogen dynamics

Mass and nitrogen (N) dynamics of leaf litter measured in Alaskan tussock tundra differed greatly from measurements of these processes made in temperate ecosystems. Nearly all litter mass and N loss occurred during the winter when soils were mostly frozen. Litter lost mass during the first summer, but during the subsequent two summers when biological activity was presumably higher than it is during winter, litter mass remained constant and litter immobilized N. By contrast, litter lost significant mass and N over both winters of measurement. Mass loss and N dynamics were unaffected by microsite variation in soil temperature and moisture. Whether wintertime mass and N loss resulted from biological activity during winter or from physical processes (e.g., fragmentation or leaching) associated with freeze-thaw is unknown, but has implications for how future climate warming will alter carbon (C) and N cycling in tundra. We hypothesize that spring runoff over permafrost as soils melt results in significant losses of C and N from litter, consistent with the observed influx of terrestrial organic matter to tundra lakes and streams after snow melt and the strong N limitation of terrestrial primary production.