Interacting ecological filters influence success and functional composition in restored plant communities over time.

A trait-based community assembly framework has great potential to direct ecological restoration, but uncertainty over how traits and environmental factors interact to influence community composition over time limits the widespread application of this approach. In this study, we examined how the composition of seed mixes and environment (north- vs. south-facing slope aspect) influence functional composition and native plant cover over time in restored grassland and shrubland communities. Variation in native cover over 4 years was primarily driven by species mix, slope aspect, and a species mix by year interaction rather than an interaction between species mix and slope aspect as predicted. Although native cover was higher on wetter, north-facing slopes for most of the study, south-facing slopes achieved a similar cover (65%-70%) by year 4. While community-weighted mean (CWM) values generally became more resource conservative over time, we found shifts in particular traits across community types and habitats. For example, CWM for specific leaf area increased over time in grassland mixes. Belowground, CWM for root mass fraction increased while CWM for specific root length decreased across all seed mixes. Multivariate functional dispersion remained high in shrub-containing mixes throughout the study, which could enhance invasion resistance and recovery following disturbance. Functional diversity and species richness were initially higher in drier, south-facing slopes compared to north-facing slopes, but these metrics were similar across north- and south-facing slopes by the end of the 4-year study. Our finding that different combinations of traits were favored in south- and north-facing slopes and over time demonstrates that trait-based approaches can be used to identify good restoration candidate species and, ultimately, enhance native plant cover across community types and microhabitat. Changing the composition of planting mixes based on traits could be a useful strategy for restoration practitioners to match species to specific environmental conditions and may be more informative than using seed mixes based on growth form, as species within functional groups can vary tremendously in leaf and root traits.

Litter and soil properties under woody and non-woody vegetation types: Implication for ecosystem management in a mountainous semi-arid landscape.

There are contrasting reports about whether and how vegetation types influence litter and soil properties. Accurate and comprehensive assessment of the complex relationship between vegetation types, litter and soil characteristics in semi-arid mountain landscapes is almost unknown. Thus, the purpose of this research was to study the effects of (1) Carpinus orientalis Miller., (2) Crataegus melanocarpa M.B., (3) Rhamnus pallasii Fisch. and C.A.Mey, (4) Agropyron longiaristatum Boiss, (5) Bromus tomentolus Bioss. and (6) Hordeum vulgare L. on litter properties and soil physical, chemical, biochemical and biological features in northern Iran. A sampling of the organic layer (litter) and mineral soil (30 × 30 cm) from a depth of 0-10 cm was done for all characteristics in the summer season and for soil microclimate and biological characteristics in the summer and fall seasons. A total of 90 litter samples, 90 soil samples in summer and 90 soil samples in fall (6 vegetation types × 2 seasons × 15 samples) were taken from the area and transferred to the laboratory. Results showed that the Carpinus improved litter properties, soil organic matter contents, total N and available nutrients (P, K, Ca and Mg) and enzyme activities (urease, acid phosphatase, arylsulfatase and invertase). In addition, the population of earthworm groups (epigeic, anecic, and endogeic), acarina, collembola, nematodes, protozoa (especially in the fall season) and bacteria and fungi (especially in the summer season) under Carpinus significantly increased. Data analysis demonstrated higher soil fertility and biological activities in the woody vegetation, which can be assigned to the higher litter input and nutrients. Overall, the findings of this study showed that woody vegetation, especially Carpinus, can improve soil properties at high altitudes of mountainous, semi-arid sites that are often considered as especially fragile and sensitive ecosystems.

Divergent response to abiotic factor determines the decoupling of water and carbon fluxes over an artificial C4 shrub in desert.

Knowledge on relationship and determinants of water and carbon dioxide (CO2) exchange is crucial to land managers and policy makers especially for the desertified land restoration. However, there remains highly uncertain in terms of water use and carbon sequestration for artificial plantation in desert. Here, continuous water and carbon fluxes were measured using eddy covariance (EC) in conjunction with hydrometeorological measurements over an artificial C4 shrub, Haloxylon ammodendron (C. A. Mey.) Bunge, from July 2020 to 2021 in Tengger Desert, China. Throughout 2021, evapotranspiration (ET) was 189.5 mm, of which 85% (150 mm) occurred during growing season, that was comparable with the summation of precipitation (132.2 mm), dew (33.5 mm) and potential other sources (e.g. deep subsoil water). This ecosystem was a strong carbon sink with net ecosystem production (NEP) up to 446.4 g C m-2 yr-1, much higher than surrounding sites. Gross primary production (GPP, 598.7 g C m-2 yr-1) in this shrubland was comparable with that of other shrublands, whereas ecosystem respiration (Re, 152.3 g C m-2 yr-1) was lower. Random Forest showed that environmental factors can explain 71.56% and 80.07% variation of GPP and ET, respectively. Interestingly, environmental factors have divergent effect on water and carbon exchange, i.e., soil hydrothermic factors (soil moisture content and soil temperature) determine the magnitude and seasonal pattern of ET and Re, while aerodynamics factors (net radiation, atmospheric temperature and wind speed) determine GPP and NEP. As such, divergent response of abiotic factors resulted in the decoupling of water and carbon exchange. Our results suggest that H. ammodendron is a suitable species for large-scale afforestation in dryland given its low water use but high carbon sequestration. Therefore, we infer that artificial planting H. ammodendron in dryland could provide an opportunity for climate change mitigation, and the long-term time series data is needed to confirm its sustainable role of carbon sequestration in the future.

Accumulation in nutrient acquisition strategies of arbuscular mycorrhizal fungi and plant roots in poor and heterogeneous soils of karst shrub ecosystems.

BACKGROUND: Arbuscular mycorrhizal (AM) fungi and roots play important roles in plant nutrient acquisition, especially in nutrient poor and heterogeneous soils. However, whether an accumulation strategy of AM fungi and root exists in such soils of karst shrubland ecosystems remains unclear. Root traits related to nutrient acquisition (root biomass, AM colonisation, root acid phosphatase activity and N2 fixation) were measured in two N2-fixing plants (i.e. Albizia odoratissima (Linn. f.) Benth. and Cajanus cajan (Linn.) Millsp.) that were grown in heterogeneous or homogeneous nutrient (ammonium) soil with and without AM fungi inoculation. RESULTS: Both of these plants had higher AM colonisation, root biomass and relative growth rate (RGR), but lower N2 fixation and root acid phosphatase activity in the rhizosphere in the heterogeneous soil environment, than that in the homogeneous soil environment. Plants grown in the AM fungi-inoculated heterogeneous soil environment had increased root biomass and root acid phosphatase activity compared with those grown in soil without inoculation. AM colonisation was negatively correlated with the N2 fixation rate of A. odoratissima, while it was not significantly correlated with the root phosphatase activity. CONCLUSIONS: Our results indicated that enhanced AM symbiosis and root biomass increased the absorptive surfaces for nutrient acquisition, highlighting the accumulation strategies of AM and root traits for plant nutrient acquisition in nutrient poor and heterogeneous soils of the karst shrubland ecosystem.

Extreme pre-fire drought decreases shrub regeneration on fertile soils.

Extreme drought and increasing temperatures can decrease the resilience of plant communities to fires. Not only may extremely dry conditions during or after fires lead to higher plant mortality and poorer recruitment, but severe pre-fire droughts may reduce the seed production and belowground vigor that are essential to post-fire plant recovery, and may indirectly facilitate invasion. We studied survival, recruitment, and growth of shrubs and herbs in chaparral (shrubland) communities in Northern California after a 2015 fire that immediately followed California's extreme 3-yr drought. We followed the same protocols used to study similar, adjacent communities after a 1999 fire that did not follow a drought, and we compared the two recovery trajectories. Overall, the 2015 fire was not more severe than the 1999 fire, yet it caused higher mortality and lower growth of resprouting shrubs on fertile (sandstone) soils. In contrast, the 2015 fire did not affect the mortality or growth of resprouting shrubs on infertile (serpentine) soils, the density of shrub seedlings, or the richness or cover of native herbs differently than the 1999 fire. However, the 2015 fire facilitated a massive increase in exotic herbaceous cover, especially on fertile soils, possibly portending the early stages of a type conversion to exotic-dominated grassland. Our findings indicate that the consequences of climate change on fire-dependent communities will include effects of pre-fire as well as post-fire climate, and that resprouting shrubs are particularly likely to be sensitive to pre-fire drought.

The role of microtopography and resident species in post-disturbance recovery of arid habitats in Hawai'i.

Habitat-suitability indices (HSI) have been employed in restoration to identify optimal sites for planting native species. Often, HSI are based on abiotic variables and do not include biotic interactions, even though similar abiotic conditions can favor both native and nonnative species. Biotic interactions such as competition may be especially important in invader-dominated habitats because invasive species often have fast growth rates and can exploit resources quickly. In this study, we test the utility of an HSI of microtopography derived from airborne LiDAR to predict post-disturbance recovery and native planting success in native shrub-dominated and nonnative, invasive grass-dominated dryland habitats in Hawai'i. The HSI uses high-resolution digital terrain models to classify sites' microtopography as high, medium, or low suitability, based on wind exposure and topographic position. We used a split-plot before-after-control-impact design to implement a disturbance experiment within native shrub (Dodonaea viscosa) and nonnative, invasive grass (Cenchrus clandestinus)-dominated ecosystems across three microtopography categories. In contrast to previous studies using the same HSI, we found that microtopography was a poor predictor of pre-disturbance conditions for soil nutrients, organic matter content, or foliar C:N, within both Dodonaea and Cenchrus vegetation types. In invader-dominated Cenchrus plots, microtopography helped predict cover, but not as expected (i.e., highest cover would be in high-suitability plots): D. viscosa had the greatest cover in low-suitability and C. clandestinus had the greatest cover in medium-suitability plots. Similarly, in native-dominated Dodonaea plots, microtopography was a poor predictor of D. viscosa, C. clandestinus, and total plant cover. Although we found some evidence that microtopography helped inform post-disturbance plant recovery of D. viscosa and total plant cover, vegetation type was a more important predictor. Important for considering the success of plantings, percent cover of D. viscosa decreased while percent cover of C. clandestinus increased within both vegetation types 20 months after disturbance. Our results are evidence that HSIs based on topographic features may prove most useful for choosing planting sites in harsh habitats or those already dominated by native species. In more productive habitats, competition from resident species may offset any benefits gained from "better" suitability sites.

Does a trade-off between growth plasticity and resource conservatism mediate post-fire shrubland responses to rainfall seasonality?

Growth plasticity may allow fire-prone species to maximize their recovery rates during temporary, sporadic periods of rainfall availability in the post-fire environment. However, moisture-driven growth plasticity could be maladaptive in nutrient-limited environments that require tighter control of growth and resource use. We investigated whether a trade-off between plasticity and conservatism mediates growth responses to altered rainfall seasonality in neighbouring shrubland communities that occupy different soils. We monitored post-fire vegetation regrowth in two structurally similar, Mediterranean-type shrublands for 3 years. We investigated the effects of experimentally altered rainfall seasonality on post-fire species' growth rates. We found that moisture-driven growth plasticity was higher among species occupying the fertile soils of the renosterveld site relative to those occupying the nutrient-poor soils of the fynbos site. This resulted in higher overall responsiveness of post-fire recovery patterns in renosterveld to experimental shifts in rainfall seasonality. In post-fire shrubland communities, the trade-off between moisture-dependent growth plasticity and resource conservatism could be mediated by soil nutrient availability. Therefore, edaphic differences between structurally similar shrublands could lead to differences in their sensitivity to post-fire rainfall seasonality.

Enhancement of ecosystem carbon uptake in a dry shrubland under moderate warming: The role of nitrogen-driven changes in plant morphology.

Net ecosystem CO2 exchange is the result of net carbon uptake by plant photosynthesis and carbon loss by soil and plant respiration. Temperature increases due to climate change can modify the equilibrium between these fluxes and trigger ecosystem-climate feedbacks that can accelerate climate warming. As these dynamics have not been well studied in dry shrublands, we subjected a Mediterranean shrubland to a 10-year night-time temperature manipulation experiment that analyzed ecosystem carbon fluxes associated with dominant shrub species, together with several plant parameters related to leaf photosynthesis, leaf morphology, and canopy structure. Under moderate night-time warming (+0.9°C minimum daily temperature, no significant reduction in soil moisture), Cistus monspeliensis formed shoots with more leaves that were relatively larger and denser canopies that supported higher plant-level photosynthesis rates. Given that ecosystem respiration was not affected, this change in canopy morphology led to a significant enhancement in net ecosystem exchange (+47% at midday). The observed changes in shoot and canopy morphology were attributed to the improved nutritional state of the warmed plants, primarily due to changes in nitrogen cycling and higher nitrogen resorption efficiency in senescent leaves. Our results show that modifications in plant morphology triggered by moderate warming affected ecosystem CO2  fluxes, providing the first evidence for enhanced daytime carbon uptake in a dry shrubland ecosystem under experimental warming.

Plant functional types determine how close postfire seedlings are from their parents in a species-rich shrubland.

BACKGROUND AND AIMS: Fine-scale spatial patterns of the seedlings of co-occurring species reveal the relative success of reproduction and dispersal and may help interpret coexistence patterns of adult plants. To understand whether postfire community dynamics are controlled by mathematical, biological or environmental factors, we documented seedling-adult (putative parent) distances for a range of co-occurring species. We hypothesized that nearest-seedling-to-adult distances should be a function of the distance between the closest conspecific seedlings, closest inter-adult distances and seedling-to-parent ratios, and also that these should scale up in a consistent way from all individuals, to within and between species and finally between functional types (FTs). METHODS: We assessed seedling-adult, seedling-seedling and adult-adult distances for 19 co-occurring shrub species 10 months after fire in a species-rich shrubland in south-western Australia. Species were categorized into 2 × 2 FTs: those that are killed by fire [non-(re)sprouters] vs. those that survive (resprouters) in nine taxonomically matched pairs, and those that disperse their seeds prefire (geosporous) vs. those that disperse their seeds postfire (serotinous). KEY RESULTS: For the total data set and means for all species, seedling-adult distance was essentially a mathematical phenomenon, and correlated positively with seedling-seedling distance and adult-adult distance, and inversely with seedlings per adult. Among the four FTs, seedling-adult distance was shortest for geosporous non-sprouters and widest for serotinous resprouters. Why adults that produce few seedlings (resprouters) should be further away from them defies a simple mathematical or biological explanation at present. Ecologically, however, it is adaptive: the closest seedling was usually under the (now incinerated) parent crown of non-sprouters whereas those of resprouters were on average four times further away. CONCLUSIONS: Our study highlights the value of recognizing four reproductive syndromes within fire-prone vegetation, and shows how these are characterized by marked differences in their seedling-adult spatial relations that serve to enhance biodiversity of the community.

The causes and effects of indigenous C4 grass expansion into a hyper-diverse fynbos shrubland.

The cool season rainfall at our study site should favour C3 rather than C4 grasses. There are, however, several locations where C4 grasses have become dominant, suggesting that rainfall seasonality is not a constraint on distribution. Here, we explored the limitations on C4 grass distribution in a fynbos shrubland. Using δ13C values of SOM, we determined when these grasses had established. We also looked at the role of roads as conduits for establishment and asked what impact these grasses may have on fynbos species richness. We then conducted a field experiment designed to examine the extent to which soil moisture, nutrient availability, and competition with fynbos for resources influence the establishment and growth of the grasses. Finally using aerial photography, we explored the role of changes in land use on distribution. Our results showed that the establishment is recent, that roads may be acting as conduits, and that with establishment, there is a reduction in fynbos species richness and diversity. Disturbance and removing below-ground competition for resources open the system to establishment in wetter areas. This study is the first to look at the potential for C4 grasses expanding into cool season rainfall shrublands such as in Western Australia and South Africa. What is interesting about these results is that C4 grasses can establish and dominate in a cool season rainfall regime. Rather than temperature of the growing season, it is competition for resources from fynbos that prevents these grasses from encroaching.