Linear and non-linear impacts of a non-native plant invasion on soil microbial community structure and function

Biological invasions can alter ecosystem functions such as litter decomposition and nutrient cycling, but little is known about how invader abundance influences the impact on the ecosystem. It is often assumed that impacts are proportional to invasion density, but this assumption has never been tested and has little justification. We tested the hypothesis that the microbial community structure and function of a mixed hardwood forest soil changed after invasion by Japanese barberry (Berberis thunbergii), an invasive shrub commonly found in eastern hardwood forests, and that changes were proportional to the density of invasion. We constructed microcosms with mixtures of native and invasive leaf litter, and measured microbial community structure (phospholipid fatty acids) and function (litter decomposition). Decomposition was linearly related to the degree of invasion (R ²?=?0.945), but the ratio of bacteria to fungi exhibited a strongly non-linear, threshold response (R ²?=?0.513). These results indicate that impacts of Japanese barberry invasion are not always proportional to invasion density. This finding has implications for the study of biological invasions as well as practical implications for the management of exotic invasive species.     

Rapid nutrient cycling in leaf litter from invasive plants in Hawai’i

Physiological traits that contribute to the establishment and spread of invasive plant species could also have impacts on ecosystem processes. The traits prevalent in many invasive plants, such as high specific leaf areas, rapid growth rates, and elevated leaf nutrient concentrations, improve litter quality and should increase rates of decomposition and nutrient cycling. To test for these ecosystem impacts, we measured initial leaf litter properties, decomposition rates, and nutrient dynamics in 11 understory plants from the Hawaiian islands in control and nitrogen + phosphorus fertilized plots. These included five common native species, four of which were ferns, and six aggressive invasive species, including five angiosperms and one fern. We found a 50-fold variation in leaf litter decay rates, with natives decaying at rates of 0.2–2.3 year^–1 and invaders at 1.4–9.3 year^–1. This difference was driven by very low decomposition rates in native fern litter. Fertilization significantly increased the decay rates of leaf litter from two native and two invasive species. Most invasive litter types lost nitrogen and phosphorus more rapidly and in larger quantities than comparable native litter types. All litter types except three native ferns lost nitrogen after 100 days of decomposition, and all litter types except the most recalcitrant native ferns lost >50% of initial phosphorus by the end of the experiment (204–735 days). If invasive understory plants displace native species, nutrient cycling rates could increase dramatically due to rapid decomposition and nutrient release from invasive litter. Such changes are likely to cause a positive feedback to invasion in Hawaii because many invasive plants thrive on nutrient-rich soils.     

Subordinate plant species impact on soil microbial communities and ecosystem functioning in grasslands: Findings from a removal experiment

Despite their low relative abundance, subordinate plant species may have larger impacts on ecosystem functioning than expected, but their role in plant communities remains poorly understood. The aim of this study was to test how subordinate plant species influence the functioning of a species-rich semi-natural grasslands. A plant removal experiment was set-up in the mountain grasslands of the Jura Mountains (Switzerland) to test the impact of subordinate plant species on soil microbial communities and ecosystem functioning. The experiment included three treatments: removal of all subordinate species, partial biomass removal of dominant species, and a no biomass removal control. After 2 years of treatments, we determined soil microbial community (bacteria and mycorrhizal fungi) by T-RFLP analysis and measured litter decomposition, soil respiration, soil inorganic nitrogen (DIN) availability and throughout above-ground biomass production as measures of ecosystem function. The removal of subordinate plant species strongly affected bacterial and weakly influenced mycorrhizal fungi communities and decreased rates of plant litter decomposition, soil respiration and DIN availability with larger effects than the partial loss of dominant biomass. The removal of subordinate plant species did not modify plant community structure, but it did reduce total above-ground biomass production compared to the control plots. Collectively, our findings indicate that the loss of subordinate species can have significant consequences for soil microbial communities and ecosystem functions, suggesting that subordinate species are important drivers of ecosystem properties.     

Phenotypic responses of invasive species to removals affect ecosystem functioning and restoration

Reducing the abundances of invasive species by removals aims to minimize their ecological impacts and enable ecosystem recovery. Removal methods are usually selective, modifying phenotypic traits in the managed populations. However, there is little empirical evidence of how removal‐driven changes in multiple phenotypic traits of surviving individuals of invasive species can affect ecosystem functioning and recovery. Overcoming this knowledge gap is highly relevant because individuals are the elemental units of ecological processes and so integrating individual‐level responses into the management of biological invasions could improve their efficiency. Here we provide novel demonstration that removals by trapping, angling and biocontrol from lakes of the globally invasive crayfish Procambarus clarkii induced substantial changes in multiple phenotypic traits. A mesocosm experiment then revealed that these changes in phenotypic traits constrain recovery of basic ecosystem functions (decomposition of organic matter, benthic primary production) by acting in the opposite direction than the effects of reduced invader abundance. However, only minor ecological impacts of invader abundance and phenotypic traits variation remained a year after its complete eradication. Our study provides quantitative evidence to an original idea that removal‐driven trait changes can dampen recovery of invaded ecosystems even when the abundance of invasive species is substantially reduced. We suggest that the phenotypic responses of invaders to the removal programme have strong effects on ecosystem recovery and should be considered within the management of biological invasions, particularly when complete eradication is not achievable.     

Effects of Exotic Plant Invasions on Soil Nutrient Cycling Processes

Although it is generally acknowledged that invasions by exotic plant species represent a major threat to biodiversity and ecosystem stability, little attention has been paid to the potential impacts of these invasions on nutrient cycling processes in the soil. The literature on plant–soil interactions strongly suggests that the introduction of a new plant species, such as an invasive exotic, has the potential to change many components of the carbon (C), nitrogen (N), water, and other cycles of an ecosystem. I have reviewed studies that compare pool sizes and flux rates of the major nutrient cycles in invaded and noninvaded systems for invasions of 56 species. The available data suggest that invasive plant species frequently increase biomass and net primary production, increase N availability, alter N fixation rates, and produce litter with higher decomposition rates than co-occurring natives. However, the opposite patterns also occur, and patterns of difference between exotics and native species show no trends in some other components of nutrient cycles (for example, the size of soil pools of C and N). In some cases, a given species has different effects at different sites, suggesting that the composition of the invaded community and/or environmental factors such as soil type may determine the direction and magnitude of ecosystem-level impacts. Exotic plants alter soil nutrient dynamics by differing from native species in biomass and productivity, tissue chemistry, plant morphology, and phenology. Future research is needed to (a) experimentally test the patterns suggested by this data set; (b) examine fluxes and pools for which few data are available, including whole-site budgets; and (c) determine the magnitude of the difference in plant characteristics and in plant dominance within a community that is needed to alter ecosystem processes. Such research should be an integral component of the evaluation of the impacts of invasive species.     

Litter and soil biodiversity jointly drive ecosystem functions

The decomposition of litter and the supply of nutrients into and from the soil are two fundamental processes through which the above- and belowground world interact. Microbial biodiversity, and especially that of decomposers, plays a key role in these processes by helping litter decomposition. Yet the relative contribution of litter diversity and soil biodiversity in supporting multiple ecosystem services remains virtually unknown. Here we conducted a mesocosm experiment where leaf litter and soil biodiversity were manipulated to investigate their influence on plant productivity, litter decomposition, soil respiration, and enzymatic activity in the littersphere. We showed that both leaf litter diversity and soil microbial diversity (richness and community composition) independently contributed to explain multiple ecosystem functions. Fungal saprobes community composition was especially important for supporting ecosystem multifunctionality (EMF), plant production, litter decomposition, and activity of soil phosphatase when compared with bacteria or other fungal functional groups and litter species richness. Moreover, leaf litter diversity and soil microbial diversity exerted previously undescribed and significantly interactive effects on EMF and multiple individual ecosystem functions, such as litter decomposition and plant production. Together, our work provides experimental evidence supporting the independent and interactive roles of litter and belowground soil biodiversity to maintain ecosystem functions and multiple services.     

Broussonetia papyrifera controls nutrient return to soil to facilitate its invasion in a tropical forest of Ghana

Aims Non-native invasive plants can alter soil chemistry through litter production and decomposition to facilitate their invasion. However, the important roles of these underlying processes in plant invasion remain poorly understood, particularly in tropical forest ecosystems. Here, we compared litter production, quality and decomposition of two invasive species (Broussonetia papyriferaandCedrela odorata) and two co-occurring native species (CeltismildbraediiandFuntumia elastica), and soil properties under them to elucidate their roles in the invasion of a tropical forest in Ghana.     

Functional Group Identity Does not Predict Invader Impacts: Differential Effects of Nitrogen-fixing Exotic Plants on Ecosystem Function

The introduction of exotic plants can have large impacts on ecosystem functions such as soil nutrient cycling. Since these impacts result from differences in traits between the exotic and resident species, novel physiological traits such as N cycling may cause large alterations in ecosystem function. It is unclear, however, whether all members of a given functional group will have the same ecosystem effects. Here we look at a within functional group comparison to test whether an annual (Lupinus luteus) and a perennial (Acacia saligna) N-fixing exotic species cause the same effects on soil N cycling in the fynbos vegetation of South Africa. We measured litterfall quantity and quality, and soil total nitrogen and organic matter for each vegetation type as well. Available nitrogen was quantified using ion exchange resin bags monthly for 1 year. We used microcosms to evaluate litter decomposition. Although both exotic species increased the available nitrogen in the soil, only Acacia increased the total soil N and organic matter. This could be explained by the slow decomposition of Acacia litter in the microcosm study, despite the fact that Acacia and Lupinus litter contained equivalent N concentrations. Presumably, low carbon quality of Acacia litter slows its decomposition in soil, resulting in retention of organic nitrogen in Acacia stands after clearing for restoration purposes. The differences in long term impacts of these annual and perennial species highlight the fact that not all N-fixing exotic species exert equivalent impacts. Ecologists should consider multiple traits rather than broadly defined functional groups alone when predicting invader impacts.     

外来植物入侵对陆地生态系统地下碳循环及碳库的影响

生物入侵是当今全球性重大环境问题之一, 是全球变化的主要研究内容.评价外来植物入侵对于生态系统影响的研究多集中在地上部分,对于生态系统地下部分影响的研究相对较少.陆地生态系统地下部分对于生态系统过程的重要性之一体现在它处于生态系统碳分配过程的核心环节.入侵种通过影响群落凋落物的输入数量、质量以及输入时间,影响到对于土壤的碳输入,而入侵种与土著种根系的差异以及入侵种对微生物群落的影响是造成土壤呼吸强度发生变化的主要因素,前者土壤呼吸强度一般比后者高.多数研究表明外来植物入侵对生态系统地下碳循环和碳库产生影响,但由于入侵植物种类较多以及研究地点环境条件的不同,关于外来植物入侵对于土壤碳库和土壤有机碳矿化影响的研究结论并不统一.最后,提出了今后该研究领域应加强的一些建议和方向.     

Winter climate change implications for decomposition in northeastern forests: comparisons of sugar maple litter with herbivore fecal inputs

Forests in northeastern North America are influenced by varying climatic and biotic factors; however, there is concern that rapid changes in these factors may lead to important changes in ecosystem processes such as decomposition. Climate change (especially warming) is predicted to increase rates of decomposition in northern latitudes. Warming in winter may result in complex effects including decreased levels of snow cover and an increased incidence of soil freezing that will effect decomposition. Along with these changes in climate, moose densities have also been increasing in this region, likely affecting nutrient dynamics. We measured decomposition and N release from ¹⁵N‐labeled sugar maple leaf litter and moose feces over 20 months in reference and snow removal treatment (to induce soil freezing) plots in two separate experiments at the Hubbard Brook Experimental Forest in New Hampshire, USA. Snow removal/soil freezing decreased decomposition of maple litter, but stimulated N transfer to soil and microbial biomass. Feces decomposed more rapidly than maple litter, and feces N moved into the mineral soil more than N derived from litter, likely due to the lower C : N ratio of feces. Feces decomposition was not affected by the snow removal treatment. Total microbial biomass (measured as microbial N and C) was not significantly affected by the treatments in either the litter or feces plots. These results suggest that increases in soil freezing and/or large herbivore populations, increase the transfer rate of N from plant detritus or digested plants into the mineral soil. Such changes suggest that altering the spatial and temporal patterns of soil freezing and moose density have important implications for ecosystem N cycling.     

Invasive Grass Alters Litter Decomposition by Influencing Macrodetritivores

The results of nitrogen (N) fertilization experiments have shown inconsistent rates of plant litter decomposition, a phenomenon that may be explained by dispropotionate influence of animal detritivores (macro-detritivores) on litter mass loss versus that of microbial decomposers, whose activity may be dependent on inorganic N. In turn, macrodetritivores may be influenced by plant species composition via their selection of optimal food resources and habitats. In our experiment, fertilizer had no apparent effect on litter decomposition, suggesting that microbial decomposers did not use the additional inorganic N and/or that macrodetritivores had a greater influence on decomposition. Manipulation of macrodetritivores suggested that plant species composition—dominated in this study by Festuca arundinacea, an exotic, invasive grass, and Aster ericoides, a native forb—caused shifts in detrivore communities and/or feeding patterns that tended to increase litter mass loss. Canopy cover of F. arundinacea and A. ericoides ranged from 0% to 11%, suggesting that low-intensity invasion may produce significant changes in ecosystem function, such as decomposition.     

<Emphasis Type="Italic">Hedychium gardnerianum</Emphasis> Invasion into Hawaiian Montane Rainforest: Interactions Among Litter Quality,Decomposition Rate,and Soil Nitrogen Availability

Few studies have examined the invasion of understory species into closed-canopy forests and, despite inter-specific differences in litter quality and quantity between understory and dominant canopy trees, the influence of understory invasions on soil nitrogen (N) cycling remains unknown. This paper examines litter quality and decomposition of kahili ginger (Hedychium gardnerianum), an invasive understory herb, to determine the influence of this species on N cycling in a Hawaiian montane rainforest. To examine the potential feedback between increased soil N availability and litter decomposition, litter from the invasive ginger, a native tree, and native tree fern was collected from unfertilized and fertilized plots and decomposed in a reciprocal transplant design. Hedychium litter decomposed faster than litter from the two native species. Across species, decomposition rates were negatively correlated with litter lignin content. Despite rapid decomposition rates of Hedychium litter, soil nitrogen availability and rates of net mineralization in the soil were similar in invaded and uninvaded plots. Nitrogen cycling at this site may be more strongly influenced by native species, which contribute the most to overall stand biomass. A negative effect of fertilization on the decomposition of Hedychium litter suggests that a negative feedback between litter quality and soil N availability may exist over longer timescales.     

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.     

Litter decomposition of a pine plantation is affected by species evenness and soil nitrogen availability

Background and aims

Litter decomposition is a key process controlling flows of energy and nutrients in ecosystems. Altered biodiversity and nutrient availability may affect litter decomposition. However, little is known about the response of litter decomposition to co-occurring changes in species evenness and soil nutrient availability.

Methods

We used a microcosm experiment to evaluate the simultaneous effects of species evenness (two levels), identity of the dominant species (three species) and soil N availability (control and N addition) on litter decomposition in a Mongolian pine (Pinus sylvestris var. mongolica) plantation in Northeast China. Mongolian pine needles and senesced aboveground materials of two dominant understory species (Setaria viridis and Artemisia scoparia) were used for incubation.

Results

Litter evenness, dominant species identity and N addition significantly affected species interaction and litter decomposition. Higher level of species evenness increased the decomposition rate of litter mixtures and decreased the incidence of antagonistic effects. A. scoparia-dominated litter mixtures decomposed faster than P. sylvestris var. mongolica- and S. viridis-dominated litter mixtures. Notably, N addition increased decomposition rate of both single-species litters and litter mixtures, and meanwhile altered the incidence and direction of non-additive effects during decomposition of litter mixtures. The presence of understory species litters stimulated the decomposition rate of pine litters irrespective of N addition, whereas the presence of pine litters suppressed the mass loss of A. scoparia litters. Moreover, N addition weakened the promoting effects of understory species litters on decomposition of pine litters.

Conclusions

Pine litter retarded the decomposition of understory species litters whereas its own decomposition was accelerated in mixtures. Nitrogen addition and understory species evenness altered species interaction through species-specific responses in litter mixtures and thus affected litter decomposition in Mongolian pine forests, which could produce a potential influence on ecosystem C budget and nutrient cycling.     

Nitrogen addition drives convergence of leaf litter decomposition rates between <Emphasis Type="Italic">Flaveria bidentis</Emphasis> and native plant

Evidence is growing that invasive species can change decomposition rates and associated nutrient cycling within an ecosystem by changing the quality of the litter entering a system. However, the relative contribution of their distinct litter types to carbon turnover is less understood, especially in the context of enhanced N deposition. The objective of this study was to investigate the whole-plant responses of an invasive plant Flaveria bidentis in litter decay to simulated N eutrophication. A 1-year study was conducted to assess if N enhancement influenced decomposition and nutrient dynamics of litters from foliage, fine roots and twigs of F. bidentis compared to co-occurring native species Setaria viridis. N fertilization significantly decreased the decomposition rate of the foliage of the invasive F. bidentis by more than 25% relative to the water control, but had relatively minor effects on decomposition of its twigs and fine root litter or leaf litter from the native species. Collectively, decomposition rates of foliar litters of the invasive and native species become convergent over time in the presence of N addition. Moreover, net N loss was predominately influenced by litter species, followed by the litter type, while N addition had little effect on net N loss. Our study showed that the variation in litter decomposition was much greater between litter types of the invasive F. bidentis than between different plant species under the N addition and that the litter of invasive species with higher inherent decomposability did not always decompose more rapidly than the litter of native species in response to predicted N deposition enhancement.     

Disentangling the Litter Quality and Soil Microbial Contribution to Leaf and Fine Root Litter Decomposition Responses to Reduced Rainfall

Climate change-induced rainfall reductions in Mediterranean forests negatively affect the decomposition of plant litter through decreased soil moisture. However, the indirect effects of reduced precipitation on litter decomposition through changes in litter quality and soil microbial communities are poorly studied. This is especially the case for fine root litter, which contributes importantly to forests plant biomass. Here we analyzed the effects of long-term (11 years) rainfall exclusion (29% reduction) on leaf and fine root litter quality, soil microbial biomass, and microbial community-level physiological profiles in a Mediterranean holm oak forest. Additionally, we reciprocally transplanted soils and litter among the control and reduced rainfall treatments in the laboratory, and analyzed litter decomposition and its responses to a simulated extreme drought event. The decreased soil microbial biomass and altered physiological profiles with reduced rainfall promoted lower fine root—but not leaf—litter decomposition. Both leaf and root litter, from the reduced rainfall treatment, decomposed faster than those from the control treatment. The impact of the extreme drought event on fine root litter decomposition was higher in soils from the control treatment compared to soils subjected to long-term rainfall exclusion. Our results suggest contrasting mechanisms driving drought indirect effects on above-(for example, changes in litter quality) and belowground (for example, shifts in soil microbial community) litter decomposition, even within a single tree species. Quantifying the contribution of these mechanisms relative to the direct soil moisture-effect is critical for an accurate integration of litter decomposition into ecosystem carbon dynamics in Mediterranean forests under climate change.     

Litter evenness influences short-term peatland decomposition processes

There is concern that changes in climate and land use could increase rates of decomposition in peatlands, leading to release of stored C to the atmosphere. Rates of decomposition are driven by abiotic factors such as temperature and moisture, but also by biotic factors such as changes in litter quality resulting from vegetation change. While effects of litter species identity and diversity on decomposition processes are well studied, the impact of changes in relative abundance (evenness) of species has received less attention. In this study we investigated effects of changes in short-term peatland plant species evenness on decomposition in mixed litter assemblages, measured as litter weight loss, respired CO₂ and leachate C and N. We found that over the 307-day incubation period, higher levels of species evenness increased rates of decomposition in mixed litters, measured as weight loss and leachate dissolved organic N. We also found that the identity of the dominant species influenced rates of decomposition, measured as weight loss, CO₂ flux and leachate N. Greatest rates of decomposition were when the dwarf shrub Calluna vulgaris dominated litter mixtures, and lowest rates when the bryophyte Pleurozium schreberi dominated. Interactions between evenness and dominant species identity were also detected for litter weight loss and leachate N. In addition, positive non-additive effects of mixing litter were observed for litter weight loss. Our findings highlight the importance of changes in the evenness of plant community composition for short-term decomposition processes in UK peatlands.     

Long-Term Grazing Accelerated Litter Decomposition in Northern Temperate Grasslands

Livestock grazing affects plant community composition, diversity, and carbon (C) and nutrient cycling in grasslands. Grazing leads to plant communities that have higher relative abundance of grazing-tolerant species, which in turn may alter the chemical composition of biomass and subsequent litter decomposition rates. To better understand the effects of long-term grazing and associated vegetation shifts on biogeochemical cycling in northern temperate grasslands of western Canada, we studied litter decomposition over 18 months at 15 locations, stratified across the Mixed-grass Prairie, Central Parkland, and Foothills Fescue natural subregions. At each location, we examined decomposition in an area exposed to grazing and an area where cattle were excluded. We used litterbags containing leaf litter from seven major grass species representing different grazing tolerances and included a local source of community litter from each study site and cellulose paper as standards. Decomposition was affected by litter types, with litter from grazing-tolerant species such as Poa pratensis and Bouteloua gracilis having faster decomposition rates compared to grazing-intolerant species, supporting the hypothesis that changes in vegetation composition due to grazing influences biogeochemical cycling by modifying litter decomposition in grasslands. Litter decomposition was also overall most rapid in the cool–wet Foothills Fescue, followed by the temperate mesic Central Parkland, and slowest in the warmer–drier Mixed-grass Prairie. Combined with known grazing-induced changes in grassland composition, these findings indicate that livestock grazing may accelerate litter decomposition rates in the more mesic Foothills Fescue and parkland regions, but not the more arid Mixed-grass Prairie. Overall, this study elucidates the role of livestock grazing and its associated effects on litter decomposition and ecosystem processes in northern grassland ecosystems.     

Contrasting effects of hemiparasites on ecosystem processes: can positive litter effects offset the negative effects of parasitism?

Hemiparasites are known to influence community structure and ecosystem functioning, but the underlying mechanisms are not well studied. Variation in the impacts of hemiparasites on diversity and production could be due to the difference in the relative strength of two interacting pathways: direct negative effects of parasitism and positive effects on N availability via litter. Strong effects of parasitism should result in substantial changes in diversity and declines in productivity. Conversely, strong litter effects should result in minor changes in diversity and increased productivity. We conducted field-based surveys to determine the association of Castilleja occidentalis with diversity and productivity in the alpine tundra. To examine litter effects, we compared the decomposition of Castilleja litter with litter of four other abundant plant species, and examined the decomposition of those four species when mixed with Castilleja. Castilleja was associated with minor changes in diversity but almost a twofold increase in productivity and greater foliar N in co-occurring species. Our decomposition trials suggest litter effects are due to both the rapid N loss of Castilleja litter and the effects of mixing Castilleja litter with co-occurring species. Castilleja produces litter that accelerates decomposition in the alpine tundra, which could accelerate the slow N cycle and boost productivity. We speculate that these positive effects of litter outweigh the effects of parasitism in nutrient-poor systems with long-lived hemiparasites. Determining the relative importance of parasitism and litter effects of this functional group is crucial to understand the strong but variable roles hemiparasites play in affecting community structure and ecosystem processes.     

Litter quality and nutrient cycling affected by grazing-induced species replacements along a precipitation gradient

One of the potential mechanisms for the impact of herbivores on nutrient cycling is the effect of selective grazing on litter quality through changes in species composition. However, the scarce evidence collected on this mechanism is controversial and seemingly influenced by site-specific variables. In this paper, we explored the consequences of grazing-induced changes in species composition on litter quality and nitrogen cycling with a regional perspective. Along a 900-mm of mean annual rainfall gradient, we selected species promoted and diminished by grazing from three natural rangelands of Argentina, analyzed their litter quality, and determined their decomposition and nutrient release kinetics under common greenhouse conditions. Litter quality and decomposition rates were strongly associated with plant response to grazing. However, the magnitude and direction of these differences depended on the ecosystem considered. In the wettest site, the species promoted by grazing (forbs) had higher nitrogen and phosphorus contents, faster decomposition rates, and higher release of nitrogen to the soil than species diminished by grazing (C₃ and C₄ grasses). In the intermediate and dry sites, species promoted by grazing had lower nitrogen and phosphorus contents, and slower decomposition rates than those diminished by grazing (C₃ grasses in both cases). Decomposition of the entire group of species was not correlated with mean annual rainfall, but when litter of the species diminished by grazing was analyzed, it was negatively correlated with precipitation. Nitrogen was immobilized more often than mineralized, even after one year of incubation. Immobilization was negatively correlated with precipitation. All these results indicate that grazing may significantly alter nutrient cycling by affecting litter quality through changes in species composition. These effects seem to be larger when species replacements induced by grazing either involve functional groups, as it was the case in our wettest site, or change root to shoot ratios. Therefore, the functional groups involved in the replacement of species as well as shifts between belowground and aboveground allocation should play a key role in grazing-induced changes on nitrogen cycling.