Non-additive Effects Of Litter Mixing Research Articles

Species diversity and chemical properties of litter influence non-additive effects of litter mixtures on soil carbon and nitrogen cycling.

Decomposition of litter mixtures generally cannot be predicted from the component species incubated in isolation. Therefore, such non-additive effects of litter mixing on soil C and N dynamics remain poorly understood in terrestrial ecosystems. In this study, litters of Mongolian pine and three dominant understory species and soil were collected from a Mongolian pine plantation in Northeast China. In order to examine the effects of mixed-species litter on soil microbial biomass N, soil net N mineralization and soil respiration, four single litter species and their mixtures consisting of all possible 2-, 3- and 4-species combinations were added to soils, respectively. In most instances, species mixing produced synergistic non-additive effects on soil microbial biomass N and soil respiration, but antagonistic non-additive effects on net N mineralization. Species composition rather than species richness explained the non-additive effects of species mixing on soil microbial biomass N and net N mineralization, due to the interspecific differences in litter chemical composition. Both litter species composition and richness explained non-additive soil respiration responses to mixed-species litter, while litter chemical diversity and chemical composition did not. Our study indicated that litter mixtures promoted soil microbial biomass N and soil respiration, and inhibited net N mineralization. Soil N related processes rather than soil respiration were partly explained by litter chemical composition and chemical diversity, highlighting the importance of functional diversity of litter on soil N cycling.

Leaf litter mixtures and neighbour effects: Low-nitrogen and high-lignin species increase decomposition rate of high-nitrogen and low-lignin neighbours

In natural ecosystems plant litter is typically a mixture of more than one species and the rate of decomposition can be faster (synergistic) or slower (antagonistic) than the average of its component species (non-additive effects). We analysed the decomposition rates of two-species mixtures to determine if there were consistent non-additive effects of litter mixing on decomposition and how do they compare with the effects of species identity on mixture decomposition. Then we tested if non-additive effects were consistently associated with the presence of particular species in the mixture, to the combination of Fast- or Slow-decomposing species, or to initial litter quality of mixtures. We found: (a) that species identity was the primary determinant of the decomposition rate of mixtures, and (b) we detected significant, but weak, non-additive effects which were consistently synergistic in the most chemically heterogeneous mixtures. However, slower decomposing species appeared to increase the decomposition rate of faster decomposing species (30 times out of 41 after 2 months of incubation, and 17 times out of 24 after 9 months of incubation). During the initial stages of decomposition, low-lignin mixtures showed mostly synergistic effects, whereas high-lignin mixtures showed antagonistic effects. At more advanced stages of decomposition, mixtures containing species with highest difference in initial N content had more synergistic effects, whereas those with similar initial N content showed both synergistic and antagonistic effects. Our results confirm previous findings about the importance of chemical heterogeneity of mixtures as a driver of decomposition rates of litter mixtures. We propose that mechanisms related to carbon priming may be related to synergistic effects in most heterogeneous mixtures, while nitrogen interaction with carbon may be resulting in antagonistic effects in homogeneous and Slow-decomposing species mixtures.

Non-Additive effects on decomposition from mixing litter of the invasive Mikania micrantha H.B.K. with native plants.

A common hypothesis to explain the effect of litter mixing is based on the difference in litter N content between mixed species. Although many studies have shown that litter of invasive non-native plants typically has higher N content than that of native plants in the communities they invade, there has been surprisingly little study of mixing effects during plant invasions. We address this question in south China where Mikania micrantha H.B.K., a non-native vine, with high litter N content, has invaded many forested ecosystems. We were specifically interested in whether this invader accelerated decomposition and how the strength of the litter mixing effect changes with the degree of invasion and over time during litter decomposition. Using litterbags, we evaluated the effect of mixing litter of M. micrantha with the litter of 7 native resident plants, at 3 ratios: M1 (1∶4, = exotic:native litter), M2 (1∶1) and M3 (4∶1, = exotic:native litter) over three incubation periods. We compared mixed litter with unmixed litter of the native species to identify if a non-additive effect of mixing litter existed. We found that there were positive significant non-additive effects of litter mixing on both mass loss and nutrient release. These effects changed with native species identity, mixture ratio and decay times. Overall the greatest accelerations of mixture decay and N release tended to be in the highest degree of invasion (mix ratio M3) and during the middle and final measured stages of decomposition. Contrary to expectations, the initial difference in litter N did not explain species differences in the effect of mixing but overall it appears that invasion by M. micrantha is accelerating the decomposition of native species litter. This effect on a fundamental ecosystem process could contribute to higher rates of nutrient turnover in invaded ecosystems.

Plant identity influences decomposition through more than one mechanism.

Plant litter decomposition is a critical ecosystem process representing a major pathway for carbon flux, but little is known about how it is affected by changes in plant composition and diversity. Single plant functional groups (graminoids, legumes, non-leguminous forbs) were removed from a grassland in northern Canada to examine the impacts of functional group identity on decomposition. Removals were conducted within two different environmental contexts (fertilization and fungicide application) to examine the context-dependency of these identity effects. We examined two different mechanisms by which the loss of plant functional groups may impact decomposition: effects of the living plant community on the decomposition microenvironment, and changes in the species composition of the decomposing litter, as well as the interaction between these mechanisms. We show that the identity of the plant functional group removed affects decomposition through both mechanisms. Removal of both graminoids and forbs slowed decomposition through changes in the decomposition microenvironment. We found non-additive effects of litter mixing, with both the direction and identity of the functional group responsible depending on year; in 2004 graminoids positively influenced decomposition whereas in 2006 forbs negatively influenced decomposition rate. Although these two mechanisms act independently, their effects may be additive if both mechanisms are considered simultaneously. It is essential to understand the variety of mechanisms through which even a single ecosystem property is affected if we are to predict the future consequences of biodiversity loss.

Influences of plant litter diversity on decomposition, nutrient mineralization and soil microbial community structure

Although plant litter identity has long been recognized as one of the important drivers of terrestrial ecosystem processes, our understanding on this issue is limited. We incubated leaf litter of four grassland species showing different functional types (C3, C4, legumes and forbs) with identical soil singly as well as in mixture of multiple species litter (two-species, three-species and all four-species litter) for 60 days under laboratory conditions to address the following questions: (i) how litter mixing (composition and richness) affects ecosystem functions including nitrogen (N) mineralization and mass loss rate; and (ii) how litter mixing impacts soil microbial community structure. Soil microbial community structure was evaluated by phospholipid fatty acid (PLFA) profiling and polymerase chain reaction (PCR)-based denaturing gradient gel electrophoresis (DGGE) analysis following amplification of 16S rDNA and 18S rDNA fragments for bacteria and fungi, respectively. The results revealed an additive effect of litter mixing on N mineralization rate. However, the results also showed that litter mixing had a synergistic effect on mass loss rate, albeit the effect tended to depend on the species identity present in the mixture. Microbial community structure measured by PLFA analysis and PCR-DGGE method showed non-additive effects of litter mixing. Bacterial community composition showed a significant positive correlation with the mass loss rate indicating a feedback between plant and soil microbial communities. Overall, the results of this study indicated the importance of litter identity rather than litter species diversity per se in driving decomposition processes.

Non-additive effects of litter mixtures on decomposition of leaf litters in a Mediterranean maquis

Many studies across a range of ecosystems have shown that decomposition in mixed litter is not predictable from single-species results due to synergistic or antagonistic interactions. Some studies also reveal that species composition and relative abundance may be more important than just richness in driving non-additive effects. Most studies on litter decomposition in Mediterranean maquis, an high-diversity shrubby ecosystem, have dealt exclusively with single species. In this study we investigated, at the individual-litter level, as well as at the litter-mixture level, the effect of litter mixing on decomposition of 3-species litter assemblages with different relative abundance of the component litters; we set up two types of litter assemblages that reflected the heterogeneity of bush cover in the inner maquis and at the edge maquis/gaps, as related to the leaf traits, i.e. sclerophylly vs mesophylly. We measured mass loss, decay of lignin, cellulose and ADSS (acid detergent soluble substances) and fungal mycelium ingrowth. The results show that over a 403-day incubation period, the decomposition of individual litters in mixtures deviated from that of monospecific litters and had different directions. In litter mixtures of the sclerophylls Phillyrea angustifolia and Pistacea lentiscus with the mesophyll Cistus, decomposition was lower than expected (antagonistic effect); in the mixtures of litters with similar physical structure (Ph. angustifolia and P. lentiscus with Quercus ilex) decomposition was faster than expected (synergistic effect). When considering the different decomposition phases, both negative and positive effects occurred in Quercus mixtures depending on the phase of decomposition. In both types of 3-species litter assemblages the greatest effect occurred in uneven mixtures rather than in even mixtures. Our results show that species composition drives the direction whilst the decomposability and the relative abundance drive the magnitude of non-additive effects of litter mixing on decomposition.

Nutrient dynamics in litter mixtures of four Mediterranean maquis species decomposing in situ

In natural conditions, litters shed from different species become mixed with each other, and decompose together. Most studies deal with decomposition of individual species; few studies investigate the influence of litter mixing on decomposition and nutrient dynamics; the results are contradictory as positive, negative, or no effect, of litter mixing have been observed. In this study we test the hypothesis: i) that litter mixing in the Mediterranean maquis, a nutrient poor, high diversity ecosystem, produces non-additive effects on nutrient dynamics; ii) that the effects vary with the composition in species of the mixture and with the relative amount of the species component the mixture. Two types of 3-species mixtures were set up; one contained three sclerophylls, Phillyrea angustifolia, Pistacia lentiscus and Quercus ilex; the other contained the first two species with the mesophyll Cistus. Litterbags, containing monospecific litters and even and uneven mixtures, were incubated under natural condition in situ; even mixtures had the 3 species in equal proportion, whereas uneven mixtures had one of the species as dominant (50%) and the other two species in equal proportion (25%:25%). Litterbags were retrieved after 92, 188 and 403 days; litters from the mixtures were separately analyzed for mass loss and content of nitrogen (N), potassium (K), sodium (Na), calcium (Ca), magnesium (Mg), manganese (Mn), iron (Fe), copper (Cu), and zinc (Zn). Results indicate that mixing influences the dynamics of N, Mn, Ca, Mg, Fe, Cu and Zn, but scarcely affects the dynamics of K and Na. The comparison of observed to expected values for changes of nutrients in litterbags indicates the occurrence of non-additive effects of litter mixing on movements of N, Fe, Cu, and Zn to or from the litterbags containing the mixtures. The effects depend on the composition in species of the mixture, whereas the relative amount of the species component the mixture is not relevant.

Species shift drives decomposition rates following invasion by hemlock woolly adelgid

Insect and disease outbreak is an important cause of selective species removal and accompanying functional change in North American forests. Outbreak of hemlock woolly adelgid, Adelgies tsugae– HWA, is causing selective removal of eastern hemlock Tsuga canadensis at a regional scale. Impacts of outbreak-caused canopy mortality and shifts in dominant species on litter decay were compared across sites that range in HWA-caused canopy damage and subsequent canopy dominance by black birch Betula lenta. Senescent litter from eastern hemlock, black birch, and equal litter mixes were decomposed in the field for 36 months within nine sites in Connecticut and Massachusetts USA. Mass loss and % N accumulation of black birch was 65% and 52% greater compared to eastern hemlock. In contrast, outbreak related canopy damage increased litter mass loss by 11.5% in high mortality stands relative to uninfested stands but canopy damage had no impact on % N dynamics. Non-additive effects of litter mixing influenced chemical dynamics of decaying litter; black birch accumulated less N and eastern hemlock accumulated more N compared to each species decaying alone. However, these changes offset and mixed litter bags overall showed no differences in N dynamics compared to values from each species decaying alone. In eastern hemlock stands invaded by hemlock woolly adelgid, canopy damage influences the rates and dynamics of decay but species differences between hemlock and black birch leaf litter are the dominant mechanisms of decomposition changes and a long-lasting driver of increased N cycling rates. Species shifts may be the dominant driver of altered ecosystem processes for other insect outbreaks, particularly when replacement species have very different characteristics regulating decomposition and N cycling.

Linkages between below and aboveground communities: Decomposer responses to simulated tree species loss are largely additive

Inputs of aboveground plant litter influence the abundance and activities of belowground decomposer biota. Litter-mixing studies have examined whether the diversity and heterogeneity of litter inputs affect decomposer communities in ways that can be predicted from monocultures. They have mainly attempted to detect non-additive effects of litter mixing, although individual species effects (additivity) as well as species interactions (non-additivity) may alter decomposition rates. To determine potential impacts of plant species loss on aboveground-decomposer linkages, we assessed both additive and non-additive effects of litter mixing on decomposer communities. A full-factorial litterbag experiment with leaves from four deciduous tree species was conducted, to assess responses of bacteria, fungi, nematodes, and microarthropods. Data were analyzed using a statistical method that first looked for additive effects based on the presence or absence of species and then any significant species interactions. We observed almost exclusively additive effects of all four litter species on decomposer biota, with each species exerting effects on different aspects of the community. These results imply that the consequences of species loss for the decomposer community will be largely predictable from knowledge of single species litter dynamics. The two species at opposite ends of the quality spectrum exerted the most effects. High-quality Liriodendron tulipifera supported a more diverse arthropod community and drove bottom-up effects on the decomposer food web. Low-quality Rhododendron maximum had negative effects on most groups of biota. Litter of mid-quality species exerted fewer effects. The influence of litter species richness on the Tylenchidae (nematodes) was the only non-additive effect of litter mixing. Together, these data demonstrate an effect of plant community composition on decomposer biomass, abundance, and diversity, confirming a link between above and belowground communities. We were able to identify the species to which the decomposer community is most sensitive, aiding predictions of the consequences of the loss of these dominant species on the decomposer community, with potential feedbacks for organic matter and nutrient turnover.

Nitrogen and Phosphorus Release from Mixed Litter Layers is Lower than Predicted from Single Species Decay

Ecosystem-level nutrient dynamics during decomposition are often estimated from litter monocultures. If species effects are additive, we can statistically predict nutrient dynamics in multi-species systems from monoculture work, and potential consequences of species loss. However, if species effects are dependent on interactions with other litter species (that is, non-additive), predictions based on monoculture data will likely be inaccurate. We conducted a 3-year, full-factorial, mixed-litter decomposition study of four dominant tree species in a temperate forest and measured nitrogen and phosphorus dynamics to explore whether nutrient dynamics in mixtures were additive or non-additive. Following common approaches, we used litterfall data to predict nutrient dynamics at the ecosystem-level. In mixtures, we observed non-additive effects of litter mixing on nutrient dynamics: the presence of nutrient-rich species in mixture facilitated nutrient release, whereas nutrient-poor species facilitated nutrient retention. Fewer nutrients were released from mixtures containing high-quality litter, and more immobilized from mixtures containing low-quality litter, than predicted from monocultures, creating a difference in overall nutrient release between predicted and actual dynamics in litter mixtures. Nutrient release at the ecosystem-level was greatly overestimated when based on monocultures because the effect of species interactions on nutrient immobilization was not accounted for. Our data illustrate that the identity of species in mixtures is key to their role in non-additive interactions, with repercussions for mineral nutrient availability and storage. These results suggest that predictions of ecosystem-level nutrient dynamics using litter monoculture data likely do not accurately represent actual dynamics because the effects of litter species interactions are not incorporated.