Nutrient limitations and their implications on the effects of atmospheric deposition in coastal dunes; lime‐poor and lime‐rich sites in the Netherlands

1 A survey of plant and soil parameters was carried out in dry dune grasslands along the Dutch coast in the lime‐ and iron‐poor Wadden district and initially lime‐ and iron‐rich Renodunaal district, in order to detect differences in nutrient availability related to soil characteristics and potential sensitivity to atmospheric deposition of nitrogen.
2 Plant biomass and phosphorus pools in the shoot were higher in the Wadden district. The low foliar nitrogen concentrations and nitrogen/phosphorus ratios in the Wadden district suggested nitrogen‐limitation, while in the Renodunaal district there appeared to be a balanced supply of both nitrogen and phosphorus.
3 Soil pH, soil organic matter, soil nitrogen and phosphorus concentrations and total amounts were generally higher in the Renodunaal district. In both districts mineral phosphorus decreased with acidification and phosphorus oxalate (iron and aluminium bound) increased.
4 In the Wadden district iron is present primarily in iron–organic matter complexes, which leads to reversible binding of phosphorus. In the Renodunaal district large amounts of iron (hydr)oxides occur and at high pH may contribute to reversible phosphorus‐sorption, but at low pH this probably leads to immobilization of phosphorus.
5 While pools of soil phosphorus are low in the Wadden district, the phosphorus availability may be relatively high due to the comparatively loose nature of phosphorus‐sorption. As a result the area may be nitrogen‐limited and grass‐encroachment may thus have resulted from atmospheric deposition of nitrogen.
6 In the Renodunaal district, atmospheric deposition probably only accelerates grass‐encroachment, because deposition of acid and nitrogen increases the availability of both nitrogen and phosphorus and maintains the 'co‐limitation'.     

The role of vegetation–microclimate feedback in promoting shrub encroachment in the northern Chihuahuan desert

Many arid and semi‐arid landscapes around the world are affected by a shift from grassland to shrubland vegetation, presumably induced by climate warming, increasing atmospheric CO₂ concentrations, and/or changing land use. This major change in vegetation cover is likely sustained by positive feedbacks with the physical environment. Recent research has focused on a feedback with microclimate, whereby cold intolerant shrubs increase the minimum nocturnal temperatures in their surroundings. Despite the rich literature on the impact of land cover change on local climate conditions, changes in microclimate resulting from shrub expansion into desert grasslands have remained poorly investigated. It is unclear to what extent such a feedback can affect the maximum extent of shrub expansion and the configuration of a stable encroachment front. Here, we focus on the case of the northern Chihuahuan desert, where creosotebush (Larrea tridentata) has been replacing grasslands over the past 100–150 years. We use a process‐based coupled atmosphere‐vegetation model to investigate the role of this feedback in sustaining shrub encroachment in the region. Simulations indicate that the feedback allows juvenile shrubs to establish in the grassland during average years and, once established, reduce their vulnerability to freeze‐induced mortality by creating a warmer microclimate. Such a feedback is crucial in extreme cold winters as it may reduce shrub mortality. We identify the existence of a critical zone in the surroundings of the encroachment front, in which vegetation dynamics are bistable: in this zone, vegetation can be stable both as grassland and as shrubland. The existence of these alternative stable states explains why in most cases the shift from grass to shrub cover is found to be abrupt and often difficult to revert.     

Shrub expansion alters forest structure but has little impact on native mammal occurrence

The expansion (or encroachment) of shrubs in forests and woodlands is generally considered a serious threat to biodiversity. The effects of shrub expansion on forest fauna, however, are poorly understood and likely to depend on the availability of key resources in shrub‐encroached forest. Coranderrk Bushland, like many conservation reserves in south‐eastern Australia, is considered threatened by the spread of an indigenous shrub. We investigated the associations between cover of Yarra burgan (Kunzea leptospermoides (Myrtaceae)), vegetation structure and the occurrence of terrestrial native mammals within the reserve, basing our predictions on prior knowledge of burgan growth habits and fauna habitat preferences. We quantified burgan cover and other potentially important habitat attributes using structure surveys, and used motion‐sensing cameras to detect terrestrial mammals. Dense burgan cover was associated with less grass, a sparser understorey, and more cryptogams, dead trees and coarse woody debris. However, there was no evidence that these changes negatively affected native mammals: burgan cover had little influence on the occurrence of any species except swamp wallabies (Wallabia bicolor), which occurred in all areas of the reserve but shifted from sites with high burgan cover during the day to sites with low cover at night. Our findings contrast with those from grassland shrub‐expansion studies, where fauna generally show strong responses to shrub cover. The effects of shrub expansion on forest fauna may be mitigated by the greater pre‐existing structural diversity in forests or the longer time required for structural changes to be fully realized. The large quantities of dead wood in areas with high shrub cover may also provide compensatory resources for small mammals, while the proximity to un‐encroached areas may enable large herbivores to move between dense shelter and forage. Shrub‐encroached forests clearly provide resources for some native fauna, and management strategies need to consider the potential impacts of shrub removal on these taxa.     

Direct and indirect effects of grazing constrain shrub encroachment in semi‐arid Patagonian steppes

Question: What are the long‐term effects of grazing exclusion on the population structure and dynamics of, and interactions among, three dominant shrub species? Location: Grass‐shrub Patagonian steppe, Chubut, Argentina. Methods: Permanent plots were established in grazed paddocks and paddocks excluded from grazing in representative Patagonian rangelands. Shrub abundance, population size‐structure, short‐term (two 3‐yr periods) and long‐term (matrix models) population dynamics, and neighborhood interactions of three native and codominant shrub species (Mulinum spinosum, Senecio filaginoides and Adesmia volckmanni) were measured and analysed using different statistical approaches. Results: The total density of shrubs was 74% higher in paddocks excluded from grazing, owing mainly to increases in Mulinum (80%) and Senecio (68%) species. However, differences in size structure between ungrazed and grazed paddocks were only detected in Mulinum. Demographic rates differed between shrub species, time‐periods and grazing conditions. In particular, recruitment in the short term (especially in wet years) and population growth rate in the long term (λ) were higher in paddocks excluded from grazing only in Mulinum populations. Senecio populations showed a marginal increase in recruitment and mortality independent of the grazing condition in the wet and dry period. Grazing exclusion modified the balance of neighborhood interactions among the three shrub species. In grazing‐exclusion paddocks, there was a balance between positive and negative interspecific interactions, while in grazed paddocks there were more negative intraspecific and interspecific interactions, resulting in a net negative balance of neighborhood interactions. Conclusions: Our understanding of woody encroachment in arid rangelands can be informed through evaluation of direct and indirect effects of grazing exclusion on the abundance and demography of dominant woody species. In Patagonian arid steppes, the occurrence of woody encroachment in rangelands excluded from grazing can be explained by altered responses in plant‐animal and plant‐plant interactions among shrub species.     

Interactions among shrub cover and the soil microclimate may determine future Arctic carbon budgets

Arctic and Boreal terrestrial ecosystems are important components of the climate system because they contain vast amounts of soil carbon (C). Evidence suggests that deciduous shrubs are increasing in abundance, but the implications for ecosystem C budgets remain uncertain. Using midsummer CO₂ flux data from 21 sites spanning 16° of latitude in the Arctic and Boreal biomes, we show that air temperature explains c. one‐half of the variation in ecosystem respiration (ER) and that ER drives the pattern in net ecosystem CO₂ exchange across ecosystems. Woody sites were slightly stronger C sinks compared with herbaceous communities. However, woody sites with warm soils (> 10 °C) were net sources of CO₂, whereas woody sites with cold soils (< 10 °C) were strong sinks. Our results indicate that transition to a shrub‐dominated Arctic will increase the rate of C cycling, and may lead to net C loss if soil temperatures rise.     

Temperature and precipitation controls over leaf‐ and ecosystem‐level CO2 flux along a woody plant encroachment gradient

Conversion of grasslands to woodlands may alter the sensitivity of CO₂ exchange of individual plants and entire ecosystems to air temperature and precipitation. We combined leaf‐level gas exchange and ecosystem‐level eddy covariance measurements to quantify the effects of plant temperature sensitivity and ecosystem temperature responses within a grassland and mesquite woodland across seasonal precipitation periods. In so doing, we were able to estimate the role of moisture availability on ecosystem temperature sensitivity under large‐scale vegetative shifts. Optimum temperatures (T_opt) for net photosynthetic assimilation (A) and net ecosystem productivity (NEP) were estimated from a function fitted to A and NEP plotted against air temperature. The convexities of these temperature responses were quantified by the range of temperatures over which a leaf or an ecosystem assimilated 50% of maximum NEP (Ω₅₀). Under dry pre‐ and postmonsoon conditions, leaf‐level Ω₅₀ in C₃ shrubs were two‐to‐three times that of C₄ grasses, but under moist monsoon conditions, leaf‐level Ω₅₀ was similar between growth forms. At the ecosystems‐scale, grassland NEP was more sensitive to precipitation, as evidenced by a 104% increase in maximum NEP at monsoon onset, compared to a 57% increase in the woodland. Also, woodland NEP was greater across all temperatures experienced by both ecosystems in all seasons. By maintaining physiological function across a wider temperature range during water‐limited periods, woody plants assimilated larger amounts of carbon. This higher carbon‐assimilation capacity may have significant implications for ecosystem responses to projected climate change scenarios of higher temperatures and more variable precipitation, particularly as semiarid regions experience conversions from C₄ grasses to C₃ shrubs. As regional carbon models, CLM 4.0, are now able to incorporate functional type and photosynthetic pathway differences, this work highlights the need for a better integration of the interactive effects of growth form/functional type and photosynthetic pathway on water resource acquisition and temperature sensitivity.     

Cascading biodiversity and functional consequences of a global change–induced biome switch

Aim At a regional scale, across southern Africa, woody thickening of savannas is becoming increasingly widespread. Using coupled vegetation and faunal responses (ants), we explore whether major changes in woody cover in savannas represent an increase in the density of savanna trees (C₄ grass layer remains intact) or a ‘regime shift’ in system state from savanna to thicket (=dry forest) where broad‐leaved, forest‐associated trees shade out C₄ grasses. Location Hluhluwe Game Reserve, South Africa. Methods We sampled paired open (low woody cover) and closed (high cover that have undergone an increase in tree density) sites. Vegetation was sampled using belt transects, and a combination of pitfall trapping and Winkler sampling was used for ants. Results Closed habitats did not simply contain a higher density of woody savanna species, but differed significantly in structure, functional composition (high prevalence of broad‐leaved trees, discontinuous C₄ grasses) and system properties (e.g. low flammability). Ant assemblage composition reflected this difference in habitat. The trophic structure of ant assemblages in the two habitats revealed a functional shift with much higher abundances of predatory species in the closed habitat. Main conclusions The predominance of species with forest‐associated traits and concomitant reduction of C₄ grasses in closed sites indicate that vegetation has undergone a shift in fundamental system state (to thicket), rather than simply savanna thickening. This biome shift has cascading functional consequences and implications for biodiversity conservation. The potential loss of many specialist savanna plant species is especially concerning, given the spatial extent and speed of this vegetation switch. Although it is not clear how easily the habitat switch can be reversed and how stable the thicket habitats are, it is likely in the not‐too‐distant future that conservation managers will be forced to make decisions on whether to actively maintain savannas.     

Shifts in functional traits elevate risk of fire‐driven tree dieback in tropical savanna and forest biomes

Numerous predictions indicate rising CO₂ will accelerate the expansion of forests into savannas. Although encroaching forests can sequester carbon over the short term, increased fires and drought‐fire interactions could offset carbon gains, which may be amplified by the shift toward forest plant communities more susceptible to fire‐driven dieback. We quantify how bark thickness determines the ability of individual tree species to tolerate fire and subsequently determine the fire sensitivity of ecosystem carbon across 180 plots in savannas and forests throughout the 2.2‐million km² Cerrado region in Brazil. We find that not accounting for variation in bark thickness across tree species underestimated carbon losses in forests by ~50%, totaling 0.22 PgC across the Cerrado region. The lower bark thicknesses of plant species in forests decreased fire tolerance to such an extent that a third of carbon gains during forest encroachment may be at risk of dieback if burned. These results illustrate that consideration of trait‐based differences in fire tolerance is critical for determining the climate‐carbon‐fire feedback in tropical savanna and forest biomes.     

Seventy years of continuous encroachment substantially increases ‘blue carbon’ capacity as mangroves replace intertidal salt marshes

Shifts in ecosystem structure have been observed over recent decades as woody plants encroach upon grasslands and wetlands globally. The migration of mangrove forests into salt marsh ecosystems is one such shift which could have important implications for global ‘blue carbon’ stocks. To date, attempts to quantify changes in ecosystem function are essentially constrained to climate‐mediated pulses (30 years or less) of encroachment occurring at the thermal limits of mangroves. In this study, we track the continuous, lateral encroachment of mangroves into two south‐eastern Australian salt marshes over a period of 70 years and quantify corresponding changes in biomass and belowground C stores. Substantial increases in biomass and belowground C stores have resulted as mangroves replaced salt marsh at both marine and estuarine sites. After 30 years, aboveground biomass was significantly higher than salt marsh, with biomass continuing to increase with mangrove age. Biomass increased at the mesohaline river site by 130 ± 18 Mg biomass km^?2 yr^?1 (mean ± SE), a 2.5 times higher rate than the marine embayment site (52 ± 10 Mg biomass km^?2 yr^?1), suggesting local constraints on biomass production. At both sites, and across all vegetation categories, belowground C considerably outweighed aboveground biomass stocks, with belowground C stocks increasing at up to 230 ± 62 Mg C km^?2 yr^?1 (± SE) as mangrove forests developed. Over the past 70 years, we estimate mangrove encroachment may have already enhanced intertidal biomass by up to 283 097 Mg and belowground C stocks by over 500 000 Mg in the state of New South Wales alone. Under changing climatic conditions and rising sea levels, global blue carbon storage may be enhanced as mangrove encroachment becomes more widespread, thereby countering global warming.