Changes in aboveground primary production and carbon and nitrogen pools accompanying woody plant encroachment in a temperate savanna

When woody plant abundance increases in grasslands and savannas, a phenomenon widely observed worldwide, there is considerable uncertainty as to whether aboveground net primary productivity (ANPP) and ecosystem carbon (C) and nitrogen (N) pools increase, decrease, or remain the same. We estimated ANPP and C and N pools in aboveground vegetation and surface soils on shallow clay and clay loam soils undergoing encroachment by Prosopis glandulosa in the Southern Great Plains of the United States. Aboveground Prosopis C and N mass increased linearly, and ANPP increased logarithmically, with stand age on clay loam soils; on shallow clays, Prosopis C and N mass and ANPP all increased linearly with stand age. We found no evidence of an asymptote in trajectories of C and N accumulation or ANPP on either soil type even following 68 years of stand development. Production and accumulation rates were lower on shallow clay sites relative to clay loam sites, suggesting strong edaphic control of C and N accumulation associated with woody plant encroachment. Response of herbaceous C mass to Prosopis stand development also differed between soil types. Herbaceous C declined with increasing aboveground Prosopis C on clay loams, but increased with increasing Prosopis C on shallow clays. Total ANPP (Prosopis+herbaceous) of sites with the highest Prosopis basal area were 1.2 × and 4.0 × greater than those with the lowest Prosopis basal area on clay loam and shallow clay soils, respectively. Prosopis ANPP more than offset declines in herbaceous ANPP on clay loams and added to increased herbaceous ANPP on shallow clays. Although aboveground C and N pools increased substantially with Prosopis stand development, we found no corresponding change in surface soil C and N pools (0–10 cm). Overall, our findings indicate that Prosopis stand development significantly increases ecosystem C and N storage/cycling, and the magnitude of these impacts varied with stand age, soil type and functional plant traits     

Contrasting above‐ and belowground sensitivity of three Great Plains grasslands to altered rainfall regimes

Intensification of the global hydrological cycle with atmospheric warming is expected to increase interannual variation in precipitation amount and the frequency of extreme precipitation events. Although studies in grasslands have shown sensitivity of aboveground net primary productivity (ANPP) to both precipitation amount and event size, we lack equivalent knowledge for responses of belowground net primary productivity (BNPP) and NPP. We conducted a 2‐year experiment in three US Great Plains grasslands – the C₄‐dominated shortgrass prairie (SGP; low ANPP) and tallgrass prairie (TGP; high ANPP), and the C₃‐dominated northern mixed grass prairie (NMP; intermediate ANPP) – to test three predictions: (i) both ANPP and BNPP responses to increased precipitation amount would vary inversely with mean annual precipitation (MAP) and site productivity; (ii) increased numbers of extreme rainfall events during high‐rainfall years would affect high and low MAP sites differently; and (iii) responses belowground would mirror those aboveground. We increased growing season precipitation by as much as 50% by augmenting natural rainfall via (i) many (11–13) small or (ii) fewer (3–5) large watering events, with the latter coinciding with naturally occurring large storms. Both ANPP and BNPP increased with water addition in the two C₄ grasslands, with greater ANPP sensitivity in TGP, but greater BNPP and NPP sensitivity in SGP. ANPP and BNPP did not respond to any rainfall manipulations in the C₃‐dominated NMP. Consistent with previous studies, fewer larger (extreme) rainfall events increased ANPP relative to many small events in SGP, but event size had no effect in TGP. Neither system responded consistently above‐ and belowground to event size; consequently, total NPP was insensitive to event size. The diversity of responses observed in these three grassland types underscores the challenge of predicting responses relevant to C cycling to forecast changes in precipitation regimes even within relatively homogeneous biomes such as grasslands.     

Litter accumulation alters the abiotic environment and drives community successional changes in two fenced grasslands in Inner Mongolia

Fencing is an effective and practical method for restoring degraded grasslands in northern China. However, little is known about the role of excess litter accumulation due to long‐term fencing in regulating abiotic environment and driving changes in community structure and function. We conducted a three‐year field experiment in two fenced grasslands in Inner Mongolia, and monitored light quantity, soil temperature, and soil moisture continuously, and determined community height, community aboveground net primary productivity (ANPP), and the relative dominance of different plant functional groups. Litter accumulation reduced light quantity and soil temperature but increased soil moisture. The regulating effects of litter accumulation on soil temperature and soil moisture fluctuated temporally and gradually weakened over the growing season. Litter accumulation also altered community vertical structure and function by increasing community height and ANPP. The increase in soil moisture increased the relative dominance of rhizome grasses but suppressed bunch grasses, thereby shifting bunch grass grasslands to rhizome grass grasslands. Our findings provide a potential mechanism for community succession in the context of litter accumulation in fenced grasslands and indicate that the vegetation and ecosystem services of degraded grasslands are improved after appropriate fencing.     

Sawdust Addition Reduces the Productivity of Nitrogen‐Enriched Mountain Grasslands

Anthropogenic nutrient enrichment of mountain grasslands has boosted grasses and fast‐growing unpalatable plants at the expense of slow‐growing species, resulting in a significant loss in biodiversity. A potential tool to reduce nutrient availability and aboveground productivity without destroying the perennial vegetation is carbon (C) addition. However, little is known about its suitability under severe climatic conditions. Here, we report the results of a 3‐year field study assessing the effects of sawdust addition on soil nutrients, aboveground productivity, and vegetational composition of 10 grazed and ungrazed mountain grasslands. Of particular interest was the effect of C addition on grasses and on the tall unpalatable weed Veratrum album. After 3 years, soil pH, ammonium, and plant‐available phosphorus were not altered by sawdust application, and nitrate concentrations were marginally higher in treatment plots. However, the biomass of grasses and forbs (without V. album) was 20–25% lower in sawdust‐amended plots, whereas the biomass of V. album was marginally higher. Sawdust addition reduced the cover of grasses but did not affect evenness, vegetation diversity, or plant species richness, although species richness generally increased with decreasing biomass at our sites. Our results suggest that sawdust addition is a potent tool to reduce within a relatively short time the aboveground productivity and grass cover in both grazed and ungrazed mountain grasslands as long as they are not dominated by tall unpalatable weeds. The technique has the advantage that it preserves the topsoil and the perennial soil seed bank.     

Convergence and contingency in production–precipitation relationships in North American and South African C4 grasslands

Mesic grasslands in North America and South Africa share many structural attributes, but less is known of their functional similarities. We assessed the control of a key ecosystem process, aboveground net primary production (ANPP), by interannual variation in precipitation amount and pattern via analysis of data sets (15- and 24-year periods) from long-term research programs on each continent. Both sites were dominated by C₄ grasses and had similar growing season climates; thus, we expected convergence in precipitation–ANPP relationships. Lack of convergence, however, would support an alternative hypothesis—that differences in evolutionary history and purportedly greater climatic variability in South Africa fundamentally alter the functioning of southern versus northern hemisphere grasslands. Neither mean annual precipitation nor mean ANPP differed between the South African and North American sites (838 vs. 857 mm/year, 423.5 vs. 461.4 g/m² respectively) and growing season precipitation–ANPP relationships were similar. Despite overall convergence, there were differences between sites in how the seasonal timing of precipitation affected ANPP. In particular, interannual variability in precipitation that fell during the first half of the growing season strongly affected annual ANPP in South Africa (P < 0.01), but was not related to ANPP in North America (P = 0.098). Both sites were affected similarly by late season precipitation. Divergence in the seasonal course of available soil moisture (chronically low in the winter and early spring in the South African site vs. high in the North American site) is proposed as a key contingent factor explaining differential sensitivity in ANPP to early season precipitation in these two grasslands. These long-term data sets provided no support for greater rainfall, temperature or ANPP variability in the South African versus the North American site. However, greater sensitivity of ANPP to early season precipitation in the South African grassland suggests that future patterns of productivity may be more responsive to seasonal changes in climate compared with the North American site.     

Shrub encroachment in North American grasslands: shifts in growth form dominance rapidly alters control of ecosystem carbon inputs

Shrub encroachment into grass-dominated biomes is occurring globally due to a variety of anthropogenic activities, but the consequences for carbon (C) inputs, storage and cycling remain unclear. We studied eight North American graminoid-dominated ecosystems invaded by shrubs, from arctic tundra to Atlantic coastal dunes, to quantify patterns and controls of C inputs via aboveground net primary production (ANPP). Across a fourfold range in mean annual precipitation (MAP), a key regulator of ecosystem C input at the continental scale, shrub invasion decreased ANPP in xeric sites, but dramatically increased ANPP (>1000 g m⁻²) at high MAP, where shrub patches maintained extraordinarily high leaf area. Concurrently, the relationship between MAP and ANPP shifted from being nonlinear in grasslands to linear in shrublands. Thus, relatively abrupt (<50 years) shifts in growth form dominance, without changes in resource quantity, can fundamentally alter continental-scale pattern of C inputs and their control by MAP in ways that exceed the direct effects of climate change alone.     

Productivity responses to altered rainfall patterns in a C<Subscript>4</Subscript>-dominated grassland

Rainfall variability is a key driver of ecosystem structure and function in grasslands worldwide. Changes in rainfall patterns predicted by global climate models for the central United States are expected to cause lower and increasingly variable soil water availability, which may impact net primary production and plant species composition in native Great Plains grasslands. We experimentally altered the timing and quantity of growing season rainfall inputs by lengthening inter-rainfall dry intervals by 50%, reducing rainfall quantities by 30%, or both, compared to the ambient rainfall regime in a native tallgrass prairie ecosystem in northeastern Kansas. Over three growing seasons, increased rainfall variability caused by altered rainfall timing with no change in total rainfall quantity led to lower and more variable soil water content (0–30 cm depth), a ~10% reduction in aboveground net primary productivity (ANPP), increased root to shoot ratios, and greater canopy photon flux density at 30 cm above the soil surface. Lower total ANPP primarily resulted from reduced growth, biomass and flowering of subdominant warm-season C₄ grasses while productivity of the dominant C₄ grass Andropogon gerardii was relatively unresponsive. In general, vegetation responses to increased soil water content variability were at least equal to those caused by imposing a 30% reduction in rainfall quantity without altering the timing of rainfall inputs. Reduced ANPP most likely resulted from direct effects of soil moisture deficits on root activity, plant water status, and photosynthesis. Altered rainfall regimes are likely to be an important element of climate change scenarios in this grassland, and the nature of interactions with other climate change elements remains a significant challenge for predicting ecosystem responses to climate change.     

Belowground biomass of alpine shrublands across the northeast Tibetan Plateau

Although belowground biomass (BGB) plays an important role in global cycling, the storage of BGB and climatic effects on it are remaining unclear. With data from 49 sites, we aimed to investigate BGB and its climatic controls in alpine shrublands in the Tibetan Plateau. Our study showed that the BGB (both grass‐layer and shrub‐layer biomass) storage in the alpine shrublands was 67.24 Tg, and the mean BGB density and shrublands area were 1,567.38 g/m² and 4.29 × 10⁴ km², respectively. Shrub layer had a larger BGB stock and accounted for 66% of total BGB this area, while only 34% was accumulated in the grass layer. BGB of the grass layer in the Tibetan Plateau shrublands was larger than that of Tibetan alpine grasslands, indicating that shrubland ecosystem played a critical importance role in carbon cycle on the Tibetan Plateau. The BGB in the grass layer and shrub layer demonstrated different correlations with climatic factors. Specifically, the effects from mean annual temperature on shrub‐layer BGB were not significant, similarly to the relationship between mean annual precipitation and grass‐layer BGB. But shrub‐layer BGB had a significantly positive relationship with mean annual precipitation (p < .05), while grass‐layer BGB showed a trend of decrease with increasing mean annual temperature (p < .05). Consequently, the actual and potential increases of BGB varied due to different increases of mean annual precipitation and temperature among different areas of the Tibetan Plateau. Therefore, in the warmer and wetter scenario, due to contrary relationships from mean annual precipitation and temperature on shrub‐layer BGB and grass‐layer BGB, it is necessary to conduct a long‐term monitoring about dynamic changes to increase the precision of assessment of BGB carbon sequestration in the Tibetan Plateau alpine shrublands.     

Nitrogen controls the net primary production of an alpine Kobresia meadow in the northern Qinghai‐Tibet Plateau

Net primary production (NPP) is a fundamental property of natural ecosystems. Understanding the temporal variations of NPP could provide new insights into the responses of communities to environmental factors. However, few studies based on long‐term field biomass measurements have directly addressed this subject in the unique environment of the Qinghai‐Tibet plateau (QTP). We examined the interannual variations of NPP during 2008–2015 by monitoring both aboveground net primary productivity (ANPP) and belowground net primary productivity (BNPP), and identified their relationships with environmental factors with the general linear model (GLM) and structural equation model (SEM). In addition, the interannual variation of root turnover and its controls were also investigated. The results show that the ANPP and BNPP increased by rates of 15.01 and 143.09 g/m² per year during 2008–2015, respectively. BNPP was mainly affected by growing season air temperature (GST) and growing season precipitation (GSP) rather than mean annual air temperature (MAT) or mean annual precipitation (MAP), while ANPP was only controlled by GST. In addition, available nitrogen (AN) was significantly positively associated with BNPP and ANPP. Root turnover rate averaged 30%/year, increased with soil depth, and was largely controlled by GST. Our results suggest that alpine Kobresia meadow was an N‐limited ecosystem, and the NPP on the QTP might increase further in the future in the context of global warming and nitrogen deposition.     

Feedback from plant species change amplifies CO2 enhancement of grassland productivity

Dynamic global vegetation models simulate feedbacks of vegetation change on ecosystem processes, but direct, experimental evidence for feedbacks that result from atmospheric CO₂ enrichment is rare. We hypothesized that feedbacks from species change would amplify the initial CO₂ stimulation of aboveground net primary productivity (ANPP) of tallgrass prairie communities. Communities of perennial forb and C₄ grass species were grown for 5 years along a field CO₂ gradient (250–500 μL L^?1) in central Texas USA on each of three soil types, including upland and lowland clay soils and a sandy soil. CO₂ enrichment increased community ANPP by 0–117% among years and soils and increased the contribution of the tallgrass species Sorghastrum nutans (Indian grass) to community ANPP on each of the three soil types. CO₂‐induced changes in ANPP and Sorghastrum abundance were linked. The slope of ANPP‐CO₂ regressions increased between initial and final years on the two clay soils because of a positive feedback from the increase in Sorghastrum fraction. This feedback accounted for 30–60% of the CO₂‐mediated increase in ANPP on the upland and lowland clay soils during the final 3 years and 1 year of the experiment, respectively. By contrast, species change had little influence on the ANPP‐CO₂ response on the sandy soil, possibly because Sorghastrum increased largely at the expense of a functionally similar C₄ grass species. By favoring a mesic C₄ tall grass, CO₂ enrichment approximately doubled the initial enhancement of community ANPP on two clay soils. The CO₂‐stimulation of grassland productivity may be significantly underestimated if feedbacks from plant community change are not considered.     

Analysis of spatial and temporal patterns of aboveground net primary productivity in the Eurasian steppe region from 1982 to 2013

To explore the importance of the Eurasian steppe region (EASR) in global carbon cycling, we analyzed the spatiotemporal dynamics of the aboveground net primary productivity (ANPP) of the entire EASR from 1982 to 2013. The ANPP in the EASR was estimated from the Integrated ANPP_NDVI model, which is an empirical model developed based on field‐observed ANPP and long‐term normalized difference vegetation index (NDVI) data. The optimal composite period of NDVI data was identified by considering spatial heterogeneities across the study area in the Integrated ANPP_NDVI model. EASR's ANPP had apparent zonal patterns along hydrothermal gradients, and the mean annual value was 43.78 g C m^?2 yr^?1, which was lower than the global grasslands average. Compared to other important natural grasslands, EASR's ANPP was lower than the North American, South American, and African grasslands. The total aboveground net primary productivity (TANPP) was found to be 378.97 Tg C yr^?1, which accounted for 8.18%–36.03% of the TANPP for all grasslands. In addition, EASR's TANPP was higher than that of the grasslands in North America, South America, and Africa. The EASR's TANPP increased in a fluctuating manner throughout the entire period of 1982–2013. The increasing trend was greater than that for North American and South American and was lower than that for African grasslands over the same period. The years 1995 and 2007 were two turning points at which trends in EASR's TANPP significantly changed. Our analysis demonstrated that the EASR has been playing a substantial and progressively more important role in global carbon sequestration. In addition, in the development of empirical NDVI‐based ANPP models, the early–middle growing season averaged NDVI, the middle–late growing season averaged NDVI and the annual maximum NDVI are recommended for use for semi‐humid regions, semi‐arid regions, and desert vegetation in semi‐arid regions, respectively.     

内蒙古草甸草原与典型草原地下生物量与生产力季节动态及其碳库潜力

地下根系是草原生态系统的重要组成部分,其生物量及其净生产力对地下碳库具有直接与间接作用,分析地下生物量季节动态与周转对深入揭示草原生态系统碳库动态及其固碳速率与潜力具有重要意义。应用钻土芯法对不同利用方式或管理措施下内蒙古草甸草原、典型草原地下生物量动态及其与温度、降水的相关性研究表明:草甸草原和典型草原地上生物量季节动态均为单峰型曲线,与上月降水显著正相关(P0.05),但地下生物量季节动态表现为草甸草原呈"S"型曲线,典型草原则是双峰型曲线,与温度、降水相关性均不显著(P0.05);两种草原根冠比和地下生物量垂直分布均为指数函数曲线,根茎型草原地下生物量集中在土壤0—5 cm,丛生型草原地下生物量集中于土壤5—10 cm,根冠比值在生长旺季(7—8月份)最小。草甸草原地下净生产力及碳储量范围分别为2167—2953 g m-2a-1和975—1329 gC m-2a-1,典型草原为2342—3333 g m-2a-1和1054—1450 gC m-2a-1,地下净生产力及其碳储量约为地上净生产力及其碳储量的10倍,具有较大的年固碳能力,且相对稳定;地下净生产力与地上净生产力呈显著负相关性(P0.05);地下生物量碳库是地上生物量碳库的10倍左右,适度放牧可增加地下生产力,但长期过度放牧显著降低其地下生物量与生产力,并使其垂直分布趋向于浅层化。     

Consequences of shrub expansion in mesic grassland: Resource alterations and graminoid responses

Abstract. In the mesic grasslands of the central United States, the shrub Cornus drummondii has undergone widespread expansion in the absence of recurrent fire. We quantified alterations in light, water and N caused by C. drummondii expansion in tall‐grass prairie and assessed the hypothesis that these alterations are consistent with models of resource enrichment by woody plants. Responses in graminoid species, particularly the dominant C₄ grass Andropogon gerardii, were concurrently evaluated. We also removed established shrub islands to quantify their legacy effect on resource availability and assess the capability of this grassland to recover in sites formerly dominated by woody plants. The primary effect of shrub expansion on resource availability was an 87% reduction in light available to the herbaceous understorey. This reduced C uptake and N use efficiency in A. gerardii and lowered graminoid cover and ANPP at the grass‐shrub ecotone relative to undisturbed grassland. Shrub removal created a pulse in light and N availability, eliciting high C gain in A. gerardii in the first year after removal. By year two, light and N availability within shrub removal areas returned to levels typical of grassland, as had graminoid cover and ANPP were similar to those in open grassland. Recovery within central areas of shrub removal sites lagged behind that at the former grass‐shrub ecotone. These results indicate that the apparent alternative stable state of C. drummondii dominance in tall‐grass prairie is biotically maintained and driven by reductions in light, rather than resource enrichment. Within areas of shrub removal, the legacy effect of C. drummondii dominance is manifest primarily through the loss of rhizomes of the dominant grasses, rather than any long‐term changes in resource availability. C. drummondii removal facilitates grassland recovery, but the effort required to initiate this transition is a significant cost of woody plant expansion in mesic grasslands. Prevention of woody plant expansion in remnant tall‐grass prairies is, therefore, a preferred management option.     

Is the removal of aboveground shrub biomass an effective technique to restore a shrub‐encroached grassland?

Encroachment of woody plants into grasslands is a global phenomenon that has substantial impacts on pastoral productivity and ecosystem services. Over the past half century, pastoralists and land management agencies have explored various options to control woody plants in order to improve ecosystem services in shrub‐encroached grasslands. We examined the effectiveness of controlling the encroachment of the shrub Caragana microphylla into grassland in Inner Mongolia, China. We cut and removed all of the aboveground biomass from 450 shrubs, predicting that the effectiveness of this technique to control shrubs would depend on shrub morphology. Specifically, we expected that larger shrubs with more biomass would be more difficult to kill by cutting than smaller shrubs. A year after treatment, we found that cutting killed only 11% of the 450 treated shrubs, and of these, three‐quarters of the locations that they occupied reverted to grasses and one‐quarter to bare soil. Shrubs that survived the cutting treatment produced more stems and leaf biomass, and therefore had a greater leaf to stem ratio. Shrubs that died after cutting had a lower crown area and basal area, and less stem biomass than shrubs that resprouted within 12 months of cutting. There were no effects of shrub height on the fate of treated shrubs. Cutting had no effect on understory plant cover or richness, but reproductive plants were taller under shrubs that were not cut. Overall, our study showed that removing aboveground shrub biomass by cutting is an ineffective technique for “restoring” the original grassland community unless shrubs are very small. Strategic targeting of small shrubs would be a more effective technique for controlling the spread of C. microphylla in the long term.     

青藏高原两种植被类型净初级生产力与气候因素的关系及周转值分析

基于2008—2016年青海海北站9年净初级生产力及气候因子监测数据,分析了青藏高原高寒小嵩草草甸和高寒金露梅灌丛两种植被净初级生产力年际动态,并探讨了气候因子对其影响及其不同土层深度根系周转值特征。结果表明:(1)年际尺度上,小嵩草草甸地上净初级生产力表现为显著增加趋势,增幅为7.02 g m~(-2) a~(-1),而金露梅灌丛地上净初级生产力相对较为稳定;对于其地下净初级生产力和总生产力,小嵩草草甸和金露梅灌丛均表现为增加趋势(P0.05),9年间小嵩草草甸地上、地下和总净初级生产力平均值分别为(217.55±9.95)、(1882.75±161.33) g m~(-2) a~(-1)和(2100.30±163.38) g m~(-2) a~(-1),金露梅灌丛地上、地下和总净初级生产力9年间平均值分别为(256.27±11.4)、(1614.31±173.03) g m~(-2) a~(-1)和(1870.58±177.93) g m~(-2) a~(-1)。(2)不同植被类型地上净初级生产力对气候因素响应不同,金露梅灌丛地上净初级生产力主要受温度影响,而温度对小嵩草草甸地上净初级生产力无显著影响。此外,降水不是限制高寒生态系统草地地上净初级生产力主要因子,相比于降水影响,高寒生态系统地上净初级生产力更受温度调控。(3)年均温和年降水对金露梅灌丛和小嵩草草甸地下净初级生产力均无显著影响(P0.05),表明高寒生态系统,其地下生产力受外界气候条件变化影响微弱,是一个稳定的碳库。(4)两种植被类型其根系周转值均随着土壤深度的增加呈逐渐增加趋势,且高寒灌丛根系周转值明显高于高寒草甸根系周转值。研究表明,在全球气候变暖背景下将会增加金露梅灌丛地上净初级生产力,而对小嵩草草甸地上净初级生产力无显著影响。     

灌丛化对黄土高原草地植物群落结构和地上生物量的影响

中国北方草地普遍出现灌丛化现象,灌丛化改变植物群落结构、植物多样性和生产力,直接影响着草地生态保护与可持续利用。该研究以黄土高原灌丛化草地为研究对象,通过植被调查,分析比较不同坡向的灌丛斑块与禾草斑块植物群落结构(物种组成、优势种及物种多样性)和地上生物量的差异。结果发现:(1)灌丛化草地不同坡向对物种多样性及地上生物量均无显著影响(P 0.1),但不同斑块植物群落结构(P=0.001)及地上生物量(P0.001)存在显著差异。(2)灌丛化草地共出现植物29种,其中禾草斑块有27种,灌丛斑块有18种;灌丛化显著改变了植物群落的物种组成,优势种由长芒草(Stipa bungeana)更替为矮脚锦鸡儿(Caragana brachypoda),且灌丛化降低了草地物种丰富度,增加了群落均匀度。(3)灌丛化显著改变了草地地上生物量,其中灌丛斑块地上生物量较禾草斑块地上生物量增加251.2 g·m~(-2),灌丛斑块中灌木/半灌木地上生物量提高了452.1 g·m~(-2),多年生丛生禾草减少了176.5 g·m~(-2),其余功能群植物的地上生物量减少了24.4 g·m~(-2)。(4)灌丛化过程(从禾草斑块—灌丛斑块)中,植物种丢失对地上生物量减少的影响较小,新增物种和群落优势种更替促进了灌木斑块地上生物量增加;虽然灌丛化导致草地地上生物量增加,但植物物种丰富度降低和优势种更替很有可能改变草地多样性和稳定性维持机制。     

Multi‐symbiotic systems: functional implications of the coexistence of grass–endophyte and legume–rhizobia symbioses

The coexistence of symbionts with different functional roles in co‐occurring plants is highly probable in terrestrial ecosystems. Analyses of how plants and microbes interact above‐ and belowground in multi‐symbiotic systems are key to understand community structure and ecosystem functioning. We performed an outdoor experiment in mesocosms to investigate the consequences of the interaction of a provider belowground symbiont of legumes (nitrogen‐fixing bacteria) and a protector aerial fungal symbiont of grasses (Epichloё endophyte) on nitrogen dynamics and aboveground net primary productivity. Four plants of Trifolium repens (Trifolium, a perennial legume) either inoculated or not with Rhizobium leguminosarum, grew surrounded by 16 plants of Lolium multiflorum (Lolium, an annual grass), with either low or high levels of the endophyte Neotyphodium occultans. After five months, we quantified the number of nodules in Trifolium roots, shoot biomass of both plant species, and the contribution of atmospheric nitrogen fixation vs. soil nitrogen uptake to above ground nitrogen in each plant species. The endophyte increased grass biomass production (+ 16%), and nitrogen uptake from the soil – the main source for the grass. Further, it reduced the nodulation of neighbour Trifolium plants (?50%). Notably, due to a compensatory increase in nitrogen fixation per nodule, this reduced neither its atmospheric nitrogen fixation – the main source of nitrogen for the legume – nor its biomass production, both of which were doubled by rhizobial inoculation. In consequence, the total amount of nitrogen in aboveground biomass and aboveground productivity were greatest in mesocosms with both symbionts (i.e. high rhizobia + high endophyte). These results show that, in spite of the deleterious effect of the endophyte on the establishment of the rhizobia–legume symbiosis, the coexistence of these symbionts, leading to additive effects on nitrogen capture and aboveground productivity, can generate complementarity on the functioning of multi‐symbiotic systems.     

Effects of Animal Husbandry on Secondary Production and Trophic Efficiency at a Regional Scale

Agricultural systems are expected to have higher net secondary production (NSP) than natural systems as a result of higher trophic efficiency and lower interannual variability. These differences, however, have not been quantified across regional gradients. We compiled a dataset of herbivore biomass, consumption, NSP, annual precipitation, and aboveground net primary production (ANPP) for extensive livestock farms across a wide precipitation gradient in Argentina. We compared these data with worldwide published studies of natural systems. In a double-logarithmic scale, NSP of agricultural systems increased with ANPP from semiarid to subhumid systems and decreased from subhumid to humid systems, a response that contrasted with the linear positive increase of natural systems. Compared to natural systems dominated by homeotherms, E _troph (NSP:ANPP) in agricultural systems in semiarid areas was 8 times higher, due to a 2 times higher E _consump (Consumption:ANPP) and a 4 times higher E _prod (NSP:Consumption). In subhumid areas, E _troph was 46 times higher, due to a 13.7 times higher E _consump and a 3.3 times higher E _prod. In humid areas, E _troph was 5 times higher, due to a 2.5 times higher E _consump and a 2 times higher E _prod. The interannual variation of herbivore biomass, a major determinant of NSP, was 60 % lower in agricultural than in natural systems dominated by homeotherms, and was decoupled from the variability of precipitation. Agricultural systems reach higher NSP by (1) diverting a major proportion of ANPP from the detritus to the grazing chain, (2) converting more efficiently consumption into NSP, and (3) stabilizing herbivore biomass across years.     

Shrub expansion stimulates soil C and N storage along a coastal soil chronosequence

Expansion of woody vegetation in grasslands is a worldwide phenomenon with implications for C and N cycling at local, regional and global scales. Although woody encroachment is often accompanied by increased annual net primary production (ANPP) and increased inputs of litter, mesic ecosystems may become sources for C after woody encroachment because stimulation of soil CO₂ efflux releases stored soil carbon. Our objective was to determine if young, sandy soils on a barrier island became a sink for C after encroachment of the nitrogen‐fixing shrub Morella cerifera, or if associated stimulation of soil CO₂ efflux mitigated increased litterfall. We monitored variations in litterfall in shrub thickets across a chronosequence of shrub expansion and compared those data to previous measurements of ANPP in adjacent grasslands. In the final year, we quantified standing litter C and N pools in shrub thickets and soil organic matter (SOM), soil organic carbon (SOC), soil total nitrogen (TN) and soil CO₂ efflux in shrub thickets and adjacent grasslands. Heavy litterfall resulted in a dense litter layer storing an average of 809 g C m^?2 and 36 g N m^?2. Although soil CO₂ efflux was stimulated by shrub encroachment in younger soils, soil CO₂ efflux did not vary between shrub thickets and grasslands in the oldest soils and increases in CO₂ efflux in shrub thickets did not offset contributions of increased litterfall to SOC. SOC was 3.6–9.8 times higher beneath shrub thickets than in grassland soils and soil TN was 2.5–7.7 times higher under shrub thickets. Accumulation rates of soil and litter C were highest in the youngest thicket at 101 g m^?2 yr^?1 and declined with increasing thicket age. Expansion of shrubs on barrier islands, which have low levels of soil carbon and high potential for ANPP, has the potential to significantly increase ecosystem C sequestration.     

Disproportionate effects of non‐colonial small herbivores on structure and diversity of grassland dominated by large herbivores

The response of semiarid grasslands to small, non‐colonial herbivores has received little attention, focusing primarily on the effects of granivore assemblages on annual plant communities. We studied the long‐term effects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would affect more abundant species and proportions of litter‐bare ground‐vegetation cover through non‐selective herbivory, 2) small herbivores would affect less common species through selective but limited consumption, and 3) herbivore effects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14‐year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large‐plus‐small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores affected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not affect uncommon components of the plant community, but had indirect effects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. There was no relationship between ANPP and the intensity of effects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive effects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our findings show that small mammalian herbivores had disproportionately large effects on plant communities relative to their small consumption of biomass. Even in small‐seeded perennial grasslands with a long history of intensive grazing by large herbivores, non‐colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.