Functional diversity of macroinvertebrates on abandoned cattle enclosures in a semi‐arid African savannah

Zimbabwe is experiencing a shift in land‐use, away from livestock farming and towards wildlife conservation. The abandonment of livestock farming may have unforeseen consequences on biodiversity and ecosystem functioning, as cattle kraals create valuable nutrient‐rich patches across the semi‐arid savannah. It is unclear how macroinvertebrates functionally respond to such nutrient‐rich patches in semi‐arid savannahs. We analysed functional diversity of both aboveground and belowground taxa on abandoned cattle kraals and savannah control plots in Save valley Conservancy (SVC). We used distance‐based multivariate techniques to estimate indices of functional diversity. Our results indicated that after two decades of abandonment, kraals had higher functional richness (FRic), functional divergence (FDiv) and functional dispersion (FDis) of macroinvertebrates when both aboveground and belowground species are combined. When aboveground macroinvertebrates were considered alone, no difference was observed for all the considered functional indices. However, only FRic was higher on kraals when belowground macroinvertebrates were separately considered. Our results suggest that two‐decade‐old abandoned kraals may have recovered enough for aboveground species to match the surrounding savannah plot and even surpassed the savannah control for belowground species functional diversity.     

Biomass responses to elevated CO<Subscript>2</Subscript>, soil heterogeneity and diversity: an experimental assessment with grassland assemblages

While it is well-established that the spatial distribution of soil nutrients (soil heterogeneity) influences the competitive ability and survival of individual plants, as well as the productivity of plant communities, there is a paucity of data on how soil heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. To evaluate the effects of elevated CO₂, soil heterogeneity and diversity (species richness and composition) on productivity, patterns of biomass allocation and root foraging precision, we conducted an experiment with grassland assemblages formed by monocultures, two- and three-species mixtures of Lolium perenne, Plantago lanceolata and Holcus lanatus. The experiment lasted for 90 days, and was conducted on microcosms built out of PVC pipe (length 38 cm, internal diameter 10 cm). When nutrients were heterogeneously supplied (in discrete patches), assemblages exhibited precise root foraging patterns, and had higher total, above- and belowground biomass. Greater aboveground biomass was observed under elevated CO₂. Species composition affected the below:aboveground biomass ratio and interacted with nutrient heterogeneity to determine belowground and total biomass. Species richness had no significant effects, and did not interact with either CO₂ or nutrient heterogeneity. Under elevated CO₂ conditions, the two- and three-species mixtures showed a clear trend towards underyielding. Our results show that differences among composition levels were dependent on soil heterogeneity, highlighting its potential role in modulating diversity–productivity relationships. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible to authorized users.     

Nutrient Limitation to Primary Productivity in a Secondary Savanna in Venezuela1

We examined nutrient limitation to primary productivity in a secondary savanna in the interior branch of the Coastal Range of Venezuela, which was converted from forest to savanna more than 100 years ago. We manipulated soil nutrients by adding nitrogen (+N), phosphorus and potassium (+PK), and nitrogen, phosphorus, and potassium (+NPK) to intact savanna. Eleven months after fertilization, we measured aboveground biomass and belowground biomass as live fine roots in the top 20 cm of soil, and species and functional group composition in response to nutrient additions. Aboveground biomass was highest in the NPK treatment ([mean g/m2]; control = 402, +N = 718, +PK = 490, +NPK = 949). Aboveground production, however, appeared to be limited primarily by N. Aboveground biomass increased 78 percent when N was added alone but did not significantly respond to PK additions when compared to controls. In contrast to aboveground biomass, belowground biomass increased with PK additions but showed no significant increase with N (depth 0–20 cm; [mean g/m2]; control = 685, +N = 443, +PK = 827, +NPK = 832). There was also a 36 percent increase in root length with PK additions when compared to controls. Whole savanna shoot:root ratios were similar for control and +PK (0.6), while those for +N or +NPK fertilization were significantly higher (1.7 and 1.2, respectively). Total biomass response (above + belowground) to nutrient additions showed a strong N and PK co‐limitation ([mean g/m2]; control = 1073, +N = 1111, +PK = 1258, +NPK = 1713). Aboveground biomass of all monocots increased with N additions, whereas dicots showed no response to nutrient additions. Trachypogon spp. (T. plumosus+T. vestitus) and Axonopus canescens, the two dominant grasses, made up more than 89 percent of the total aboveground biomass in these sites. Trachypogon spp. responded to NPK, whereas A. canescens, sedges, and the remaining monocots only responded to N. Even though nutrient additions resulted in higher aboveground biomass in N and NPK fertilized plots, this had little effect on plant community composition. With the exception of sedges, which responded positively to N additions and increased from 4 to 8 percent of die plant community, no changes were observed in plant community composition after 11 months.     

Large herbivores may alter vegetation structure of semi-arid savannas through soil nutrient mediation

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.     

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.     

Spatial heterogeneity in soil nutrient supply modulates nutrient and biomass responses to multiple global change drivers in model grassland communities

Changes in the atmospheric concentration of carbon dioxide ([CO₂]), nutrient availability and biotic diversity are three major drivers of the ongoing global change impacting terrestrial ecosystems worldwide. While it is well established that soil nutrient heterogeneity exerts a strong influence on the development of plant individuals and communities, it is virtually unknown how nutrient heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. We conducted a microcosm experiment to evaluate the effect of simultaneous changes in [CO₂], nutrient heterogeneity (NH), nutrient availability (NA) and species evenness on the biomass and nutrient uptake patterns of assemblages formed by Lolium perenne, Plantago lanceolata and Holcus lanatus. When the nutrients were heterogeneously supplied, assemblages exhibited precise root foraging patterns, and had higher above‐ and belowground biomass (average increases of 32% and 29% for above‐ and belowground biomass, respectively). Nutrient heterogeneity also modulated the effects of NA on biomass production, complementarity in nitrogen uptake and below: aboveground ratio, as well as those of [CO₂] on the nutrient use efficiency at the assemblage level. Our results show that nutrient heterogeneity has the potential to influence the response of plant assemblages to simultaneous changes in [CO₂], nutrient availability and biotic diversity, and suggest that it is an important environmental factor to interpret and assess plant assemblage responses to global change.     

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.     

Spatial heterogeneity in ecosystem structure and productivity in a moist Kenyan savanna

Topographic variations and disturbances are key sources of spatial heterogeneity in the ecosystem and may influence its functioning, productivity, and carbon (C) storage. In water controlled ecosystems, structural and functional heterogeneity become distinct during drought when the ecosystem processes are operating at their limits. We examined spatial heterogeneity arising from grazing, abandoned cropland, presence of Acacia trees, and termite mounds (termitaria). Soil water content (SWC) was significantly (P < 0.05) higher in termitaria and fenced (un-grazed) plots. Higher soil nitrogen (N) content occurred in the Acacia, termitaria, and fenced plots while total biomass was highest in the fenced plot. The termitaria plots showed the highest net ecosystem CO₂ exchange (NEE), ecosystem respiration (R _eco), and gross primary production (GPP) and were the only plots that were net CO₂ sinks. Except in fenced plots, maximum GPP was positively correlated with SWC and green biomass in all the other plots. Green biomass and R _eco were positively correlated with SWC. Shifting cultivation (abandoned farmlands) negatively affected soil quality, ecosystem CO₂ assimilation, and productivity. Removal of grazing (cattle) from the ecosystem negatively influenced GPP, while the presence of termitaria and Acacia trees facilitated soil water and N availability and ecosystem productivity. We concluded that soil water availability was responsible for most of the localized differences in the savanna and has a strong influence on ecosystem C capture and storage. We recommend that future studies on savanna productivity and ecosystem CO₂ fluxes should consider heterogeneity in the ecosystem in order to avoid bias and increase the accuracy of any estimates made.     

Hurricane wrack generates landscape‐level heterogeneity in coastal pine savanna

Wrack (vegetation debris) deposited by storm surges of major hurricanes along the northern Gulf of Mexico produces depressant effects that vary from partial to complete mortality of groundcover vegetation in coastal savannas. As wrack decomposes or is relocated by a subsequent hurricane, patches are opened to colonization. We postulated that patterns of wrack deposition and removal, coupled with differential responses by savanna plant species should produce alternate states of groundcover vegetation. We explored extreme effects of wrack deposited by Hurricane Katrina (2005) in savannas dominated by slash pine Pinus elliottii and cordgrass Spartina patens and located above mean high tide at the Grand Bay National Estuarine Research Reserve, Mississippi, USA. In 2008, we established plots in adjacent areas with and without wrack deposits. Almost no groundcover plant species occurred in wrack deposits compared to adjacent groundcover without wrack. We simulated redistribution of wrack during a new storm surge by removing wrack from replicated plots and depositing it in plots without wrack, recording plant species in treatment and control plots before, then one month and one year after manipulations. One year later, about half the species present before wrack addition (especially dominant graminoids) grew back through redistributed wrack, suggesting that some species were resistant to burial of limited amounts of wrack. Wrack removal resulted in germination and establishment of numerous herbaceous plant species not in undisturbed groundcover, doubling total aboveground numbers of species in the pine savanna and shifting groundcover communities to alternate states not present prior to Katrina. Removal of wrack opens space colonized by resilient species, including those transported in wrack and those surviving intervals between disturbances belowground. Wrack dynamics (deposition and removal) generated alternate states that resulted from resistance‐ and resilience‐driven changes in different patches of groundcover in coastal savannas.     

Litter type and termites regulate root decomposition across contrasting savanna land‐uses

Decomposition is a vital ecosystem process, increasingly modified by human activity. Theoretical frameworks and empirical studies that aim to understand the interplay between human land‐use, macro‐fauna and decomposition processes have primarily focused on leaf and wood litter. For a whole‐plant understanding of how land‐use and macro‐fauna influence decomposition, investigating root litter is required. Using litterbags, we quantified rates of root decomposition across contrasting tropical savanna land‐uses, namely wildlife and fire‐dominated protected areas and livestock pastureland without fire. By scanning litterbags for termite intrusion, we differentiated termite and microbial driven decomposition. Root litter was buried underneath different tree canopies (leguminous and non‐leguminous trees) and outside canopies to account for savanna landscape effects. Additionally, we established a termite cafeteria‐style experiment and common garden to explore termite selectivity of root litter and root trait relationships, respectively. After one year, we found no significant differences in root litter mass loss between wildlife dominated areas and pastureland. Instead, we found consistent species differences in root litter mass loss across land‐uses and additive and non‐additive effects of termites on root decomposition across plant species. Termite selectivity for root litter species occurred for both root and leaf litter buried near termite mounds, but was not explained by root traits measured in the common garden. Termite foraging was greater under leguminous tree canopies than other canopies; however, this did not influence rates of root decomposition. Our study suggests that land‐use has a weak direct effect on belowground processes in savannas. Instead, changes in herbaceous species composition and termite foraging have stronger impacts on belowground decomposition. Moreover, termites were not generalist decomposers of root litter, but their impact varies depending on plant species identity and likely associated root traits. This root litter selectivity by termites is likely to be an important contributor to spatial heterogeneity in savanna nutrient cycling.     

Effects of land‐use changes on ecosystem services: decrease in ant predation in human‐dominated landscapes in central Brazil

Anthropogenic disturbances often affect the abundance and diversity of ants (Hymenoptera: Formicidae) but relatively few studies have explored the implications of such changes on the ecosystem services mediated by these insects. Here, we evaluated how the transformation of Cerrado savanna habitats into crop plantations affects the abundance, diversity, and the predatory activity of ants. A survey of the ant faunas foraging above‐ and belowground was performed in six crop and six non‐crop (i.e., native vegetation) sites. Above‐ and belowground rates of ant predation were estimated at these same sites using mealworms, Tenebrio molitor L. (Coleoptera: Tenebrionidae), as baits, simulating crop herbivores. Belowground predation rates were significantly greater in the non‐crop sites, despite the lack of difference in overall abundance and species richness of ants foraging belowground between the crop vs. non‐crop sites. In contrast, we did not detect any significant difference in aboveground predation rates between crop vs. non‐crop sites even though there were significantly more species of ants foraging aboveground in the non‐crop sites. Army ants (subfamily Dorylinae) were the main predatory species belowground, and their abundance was significantly greater in non‐crop sites. In contrast, the main predators aboveground were omnivore ants of the genera Pheidole and Solenopsis, which had similar abundances in the crop and non‐crop sites. Overall, our results indicate that transformation of native Cerrado habitats into crop plantations reduces the abundance of some important predatory species, notably those that forage belowground, and this may negatively affect the potential for ants to provide pest control services in agroecosystems.     

The variable effects of soil nitrogen availability and insect herbivory on aboveground and belowground plant biomass in an old-field ecosystem

Nutrient availability and herbivory can regulate primary production in ecosystems, but little is known about how, or whether, they may interact with one another. Here, we investigate how nitrogen availability and insect herbivory interact to alter aboveground and belowground plant community biomass in an old-field ecosystem. In 2004, we established 36 experimental plots in which we manipulated soil nitrogen (N) availability and insect abundance in a completely randomized plot design. In 2009, after 6 years of treatments, we measured aboveground biomass and assessed root production at peak growth. Overall, we found a significant effect of reduced soil N availability on aboveground biomass and belowground plant biomass production. Specifically, responses of aboveground and belowground community biomass to nutrients were driven by reductions in soil N, but not additions, indicating that soil N may not be limiting primary production in this ecosystem. Insects reduced the aboveground biomass of subdominant plant species and decreased coarse root production. We found no statistical interactions between N availability and insect herbivory for any response variable. Overall, the results of 6 years of nutrient manipulations and insect removals suggest strong bottom-up influences on total plant community productivity but more subtle effects of insect herbivores on aspects of aboveground and belowground production.     

Carbon dioxide exchange and biomass productivity of the herbaceous layer of a managed tropical humid savanna ecosystem in western Kenya

Aims Humid savannas, as a result of high precipitation amounts, are highly productive. They are also hotspots for land use change and potential sources of carbon dioxide (CO₂) due to the large soil carbon (C) stocks. Understanding how ecosystem CO₂ exchange is influenced by changes arising from agricultural land use is vital in future management of these ecosystems and in responding to the ongoing shifts in management and climate. The aim of this study was to identify how ecosystem CO₂ exchange and biomass productivity of the herbaceous layer of a humid savanna in Kenya respond to current management practices.Methods We used flux chambers to quantify CO₂ fluxes, while monthly harvests were undertaken to determine biomass development of the herbaceous layer of three sites that were (i) fenced to exclude livestock grazing, (ii) subjected to grazing by livestock and (iii) abandoned after being cultivated for maize production and also open to grazing by livestock.Important findings The peak aboveground biomass ranged between 380 and 1449g m ?2 and biomass production was significantly (P < 0.05) lower in the grazed and abandoned plots. The maximum gross primary production (GPP) and net ecosystem CO₂ exchange (NEE) ranged between 21.8±1.3 to 32.5±2.7 and ?9.6±0.7 to^-17.9±4.8 μmol m ?2 s^-1, respectively. Seasonal NEE fluctuations ranged between 10 and 21 μmol m ?2 s^-1, while spatial (among sites) differences ranged between 2 and 10 μmol m ?2 s^-1. Ecosystem respiration (R eco) fluctuated between 5 and 10 μmol m ?2 s^-1 during the growing season. R eco was, however, not significantly different among the sites. Unlike in other similar ecosystems where ecosystem respiration is determined by the ambient temperature, we did not find any relationship between R eco and temperature in this savanna. Instead, soil moisture accounted for 38–88% of the spatial and seasonal fluctuations in ecosystem CO₂ fluxes and aboveground biomass production. Management influenced the maximum GPP and NEE rates through modification of soil moisture, plant species composition and aboveground biomass. We concluded that soil moisture is the key determinant of ecosystem CO₂ exchange and productivity in this tropical savanna. Management, however, significantly modifies C fluxes and productivity through its influence on soil moisture, plant species composition and aboveground green biomass and should be taken into consideration in future similar studies.     

Primary production and carbon allocation in relation to nutrient supply in a tropical experimental forest

Nutrient supply commonly limits aboveground plant productivity in forests, but the effects of an altered nutrient supply on gross primary production (GPP) and patterns of carbon (C) allocation remain poorly characterized. Increased nutrient supply may lead to a higher aboveground net primary production (ANPP), but a lower total belowground carbon allocation (TBCA), with little change in either aboveground plant respiration (APR) or GPP. Alternatively, increases in nutrient supply may increase GPP, with the quantity of GPP allocated aboveground increasing more steeply than the quantity of GPP allocated belowground. To examine the effects of an elevated nutrient supply on the C allocation patterns in forests, we determined whole‐ecosystem C budgets in unfertilized plots of Eucalyptus saligna and in adjacent plots receiving regular additions of 65 kg N ha^?1, 31 kg P ha^?1, 46 kg K ha^?1, and macro‐ and micronutrients. We measured the absolute flux of C allocated to the components of GPP (ANPP, TBCA and APR), as well as the fraction of GPP allocated to these components. Fertilization dramatically increased GPP. Averaged over 3 years, GPP in the fertilized plots was 34% higher than that in the unfertilized controls (3.95 vs. 2.95 kg C m^?2 yr^?1). Fertilization‐related increases in GPP were allocated entirely aboveground – ANPP was 85% higher and APR was 57% higher in the fertilized than in the control plots, while TBCA did not differ significantly between treatments. Carbon use efficiency (NPP/GPP) was slightly higher in the fertilized (0.53) compared with the control plots (0.51). Overall, fertilization increased ANPP and APR, and these increases were related to a greater GPP and an increase in the fraction of GPP allocated aboveground.     

Short‐Term and Long‐Term Effects of Soil Ripping,Seeding, and Fertilization on the Restoration of a Tropical Rangeland

Rangeland degradation is a serious problem in semiarid Africa. Extensive areas of bare, compacted, nutrient‐poor soils limit the productivity and biodiversity of many areas. We conducted a set of restoration experiments in which all eight combinations of soil tilling, fertilization, and seeding with native perennial grasses were carried out in replicated plots. After 6 months, little aboveground biomass was produced in plots without tilling, regardless of seeding or fertilization. Tilling alone tripled plant biomass, mostly of herbaceous forbs and annual grasses. Perennial grasses were essentially limited to plots that were both tilled and seeded. The addition of fertilizer had no significant additional effects. After 7 years, vegetation had declined, but there were still large differences among treatments. After 10 years, one tilled (and seeded) plot had reverted to bare ground, but the other tilled plots still had substantial vegetation. Only one seeded grass (Cenchrus ciliaris) was still a contributor to total cover after 10 years. We suggest that restoration efforts on these soils be directed first to breaking up the surface crust, and second to the addition of desirable seed. A simple ripping trial inspired by this experiment showed considerable promise as a low‐cost restoration technique.     

Effects of nutrient additions on plant biomass and diversity of the herbaceous-subshrub layer of a Brazilian savanna (Cerrado)

The Brazilian Cerrado is a diversity hotspot due to its high level of endemism and rapid loss of habitats. It is estimated that the number of herbaceous species is four times higher than that of woody species. Increasing levels of nitrogen additions to natural ecosystems have been indicated as a determinant of biodiversity loss. We investigated the effects of nutrient additions on the productivity (aboveground and belowground) and on diversity of the herbaceous-subshrub layer of a Brazilian savanna (cerrado stricto sensu). The experiment was carried out in the IBGE Ecological Reserve, near Brasília, Brazil. Between 1998 and 2006, N, P, N plus P, or Ca were applied to sixteen 225 m² plots, arranged in a completely randomized design. Aboveground biomass was compared 1 year after the first fertilization and 10 years later. Floristic diversity was significantly different (P < 0.01) between the treatments. The highest and lowest species richness were presented in control and NP, respectively. The addition of P alone or in combination with N induced invasion by Melinis minutiflora (exotic C₄ grass). The aboveground biomass of this species was higher in NP and P plots. In the N treatment, Echinolaena inflexa (native C₃ grass) presented elevated cover and biomass but M. minutiflora was absent. The invasion by alien species resulted in negative impacts on native grass species. Besides changes in aboveground biomass, addition of N and P also led, although to a lesser extent, to changes in the root morphology and biomass, but these responses were modulated by seasonal variation in soil moisture. The results suggest that environmental changes in nutrient availability can lead to important consequences for diversity and functioning of this savanna where the numerous rare species have more chance to persist under dystrophic conditions as some species that tend to be dominant would be less competitive.     

Linking soil food web structure to above‐ and belowground ecosystem processes: a meta‐analysis

Soil fauna can be an important regulator of community parameters and ecosystem processes, but there have been few quantitative syntheses of the role of soil fauna in terrestrial soil communities and ecosystems. Here, we conducted a meta‐analysis to investigate the impacts of invertebrate soil micro‐ and mesofauna (grazers and predators) on plant productivity and microbial biomass. Overall our results indicate that an increase in the biomass of soil fauna increased aboveground plant productivity across ecosystems by 35% and decreased microbial biomass by 8%. In addition, we found no evidence for trophic cascades in terrestrial soil food webs, but the bacterivorous component of soil fauna influenced plant productivity and microbial biomass more than did the fungivorous component. Furthermore, changes in the biomass of soil fauna differentially affected plant productivity among plant functional groups: a higher biomass of soil fauna increased aboveground productivity by 70% in coniferous systems. However, in ecosystems dominated by legumes, a functional group with lower inorganic nitrogen requirements, there was no response of aboveground productivity to increases in the biomass of soil fauna. In sum, the results of this meta‐analysis indicate that soil fauna help to regulate ecosystem production, especially in nutrient‐limited ecosystems.     

Relationships among soil fertility,native plant diversity and exotic plant abundance inform restoration of forb‐rich eucalypt woodlands

Aim Water and nutrient availability are major limits to productivity in semi‐arid ecosystems; hence, ecological restoration often focuses on conserving or concentrating soil resources. By contrast, nutrient enrichment can promote invasion by exotic annuals, leading to restoration approaches that target reduction of soil nutrients. We aimed to explore potential biodiversity trade‐offs between these approaches by investigating relationships among soil nutrients, exotic annuals and native plant diversity and composition. In particular, we investigated the hypothesis that native plant diversity in semi‐arid to temperate woodlands reflects the productivity–diversity hypothesis, leading to hump‐backed relationships with soil nutrients such that (1) native plant diversity declines with increasing nutrient enrichment and (2) native diversity is limited at the lowest levels of soil fertility. Location Fragmented, long‐ungrazed Eucalyptus loxophleba subsp. loxophleba (York gum)–Acacia acuminata (jam) woodlands in the wheatbelt of South‐Western Australia. Methods We conducted stratified surveys of floristic composition and topsoil nutrient concentrations in 112 woodland patches. We used generalized linear models, structural equation models and ordinations to characterize relationships among soil nutrients, rainfall, exotic annuals and patch‐scale (100 m²) native plant composition and diversity. Results Patch‐scale native plant diversity declined strongly with increasing exotic abundance. This was partly related to elevated soil nutrient concentrations, particularly total nitrogen and available phosphorus. By contrast, there was little evidence for positive correlations between soil nutrients and native diversity, even at very low soil nutrient concentrations. Main conclusions Minimizing weed invasions is crucial for maximizing native plant diversity in E. loxophleba woodlands and could include nutrient‐depleting treatments without substantially compromising the functional capacity of soils to maintain native plant richness and composition. More broadly we emphasize that understanding relationships among ecosystem productivity, plant diversity and exotic invasions in the context of associated theoretical frameworks is fundamental for informing ecological restoration.     

Contribution of aboveground litter,belowground litter,and rhizosphere respiration to total soil CO<Subscript>2</Subscript> efflux in an old growth coniferous forest

In an old growth coniferous forest located in the central Cascade Mountains, Oregon, we added or removed aboveground litter and terminated live root activity by trenching to determine sources of soil respiration. Annual soil efflux from control plots ranged from 727 g C m⁻² year⁻¹ in 2002 to 841 g C m⁻² year⁻¹ in 2003. We used aboveground litter inputs (149.6 g C m⁻² year⁻¹) and differences in soil CO₂ effluxes among treatment plots to calculate contributions to total soil efflux by roots and associated rhizosphere organisms and by heterotrophic decomposition of organic matter derived from aboveground and belowground litter. On average, root and rhizospheric respiration (R_r) contributed 23%, aboveground litter decomposition contributed 19%, and belowground litter decomposition contributed 58% to total soil CO₂ efflux, respectively. These values fall within the range of values reported elsewhere, although our estimate of belowground litter contribution is higher than many published estimates, which we argue is a reflection of the high degree of mycorrhizal association and low nutrient status of this ecosystem. Additionally, we found that measured fluxes from plots with doubled needle litter led to an additional 186 g C m⁻² year⁻¹ beyond that expected based on the amount of additional carbon added; this represents a priming effect of 187%, or a 34% increase in the total carbon flux from the plots. This finding has strong implications for soil C storage, showing that it is inaccurate to assume that increases in net primary productivity will translate simply and directly into additional belowground storage.     

Trickle-down effects of aboveground trophic cascades on the soil food web

Trophic cascades are increasingly being regarded as important features of aboveground and belowground food webs, but the effects of aboveground cascades on soil food webs, and vice versa, remains essentially unexplored. We conducted an experiment consisting of model synthesised communities containing grassland plant and invertebrate species, in which treatments included soil only, soil+plants, soil+plants+aphids, and soil+plants+aphids+predators; predator treatments consisted of the lacewing Micromus tasmaniae and ladybird beetle Coccinella undecimpunctata added either singly or in combination. Addition of Micromus largely reversed the negative effects of aphids on plant biomass, while both of the predator species caused large changes in the relative abundances of dominant plant species. Predators of aphids also affected several components of the belowground subsystem. Micromus had positive indirect effects on the primary consumer of the soil decomposer food web (microflora), probably through promoting greater input of basal resources to the decomposer subsystem. Predator treatments also influenced densities of the tertiary consumers of the soil food web (top predatory nematodes), most likely through inducing changes in plant community composition and therefore the quality of resource input to the soil. The secondary consumers of the soil food web (microbe‐feeding nematodes) were, however, unresponsive. The fact that some trophic levels of the soil food web but not others responded to aboveground manipulations is explicable in terms of top‐down and bottom‐up forces differentially regulating different belowground trophic levels. Addition of aphids also influenced microbial community structure, promoted soil bacteria at the expense of fungi, and enhanced the diversity of herbivorous nematodes; in all cases these effects were at least partially reversed by addition of Micromus. These results in tandem point to upper level consumers in aboveground food webs as a potential driver of the belowground subsystem, and provide evidence that predator‐induced trophic cascades aboveground can have effects that trickle through soil food webs.