Examining the direct and indirect mechanisms that shrubs facilitate annual seedbank

Seedbanks are important for the persistence of plant diversity in deserts. Shrub facilitation can be a key component of the storage effect, which proposes plant communities benefit from increased spatial heterogeneity of seedbank densities. Shrubs can facilitate seedbank densities directly by physically trapping seeds and indirectly by increasing beneficiary seed production. Thus, shrubs increased the density and spatial heterogeneity of seedbanks, thereby supporting the storage effect. It is critical to assess the facilitation mechanisms that support seedbank densities in deserts because seedbanks increase species coexistence and confer resilience to climate change.


Shift in community structure in an early-successional Mediterranean shrubland driven by long-term experimental warming and drought and natural extreme droughts.

Abstract: Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long-term (17 years) nocturnal-warming (+0.6 °C) and drought (-40% rainfall) experiment in an early-successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundance. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g. community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter-spring and/or summer, indicating sensitivity to water limitation in this early-successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long-term nocturnal-warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity. This article is protected by copyright. All rights reserved.

Pub.: 18 May '17, Pinned: 15 Jun '17

Effects of foliage litter of a pioneer shrub (Artemisia halodendron) on germination from the soil seedbank in a semi-arid sandy grassland in China.

Abstract: Vegetation recovery during succession is an important process for ecological restoration of the soil, especially in degraded sandy land. However, the driving mechanisms, such as how a pioneer species competes with other species, is uncertain. In China's Horqin Sandy Land, Artemisia halodendron is an important shrub that is common on semi-fixed dunes, where it replaces Agriophyllum squarrosum during succession, and is an important indicator species of the second stage of dune stabilization. However, how it outcompetes other species is still unclear. In this study, we conducted a seed bank germination experiment using soil from the native habitats of A. halodendron on semi-fixed dunes. We covered the soil with foliage litter of A. halodendron at a range of concentrations. Seed germination and seedling growth were strongly affected by the foliage litter. Seed germination and seedling growth were not harmed by a low concentration (≤50 g m(-2)) of the foliage litter but severely inhibited by high concentrations (≥100 g m(-2)). Strong allelopathy, indicated by decreased germination, increased seedling loss, and decreased plant biomass, appeared during the later stages of germination (after about 20 days of incubation). Our results suggest that as a pioneer shrub during the vegetation succession that occurs during dune stabilization, A. halodendron outcompeted other species through the allelopathic effect of its foliage litter. This helps to explain the patchy distribution and heterogeneity of vegetation communities in the Horqin Sandy Land.

Pub.: 24 May '17, Pinned: 15 Jun '17

The influence of vegetation, mesoclimate and meteorology on urban atmospheric microclimates across a coastal to desert climate gradient.

Abstract: Many cities are increasing vegetation in part due to the potential for microclimate cooling. However, the magnitude of vegetation cooling and sensitivity to mesoclimate and meteorology are uncertain. To improve understanding of the variation in vegetation's influence on urban microclimates we asked: how do meso- and regional-scale drivers influence the magnitude and timing of vegetation-based moderation on summertime air temperature (Ta), relative humidity (RH) and heat index (HI) across dryland cities? To answer this question we deployed a network of 180 temperature sensors in summer 2015 over 30 high- and 30 low-vegetated plots in three cities across a coastal to inland to desert climate gradient in southern California, USA. In a followup study, we deployed a network of temperature and humidity sensors in the inland city. We found negative Ta and HI and positive RH correlations with vegetation intensity. Furthermore, vegetation effects were highest in evening hours, increasing across the climate gradient, with reductions in Ta and increases in RH in low-vegetated plots. Vegetation increased temporal variability of Ta, which corresponds with increased nighttime cooling. Increasing mean Ta was associated with higher spatial variation in Ta in coastal cities and lower variation in inland and desert cities, suggesting a climate dependent switch in vegetation sensitivity. These results show that urban vegetation increases spatiotemporal patterns of microclimate with greater cooling in warmer environments and during nighttime hours. Understanding urban microclimate variation will help city planners identify potential risk reductions associated with vegetation and develop effective strategies ameliorating urban microclimate.

Pub.: 07 Jun '17, Pinned: 15 Jun '17

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

Abstract: 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 ANPPNDVI 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 ANPPNDVI 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.

Pub.: 06 Jun '17, Pinned: 15 Jun '17

Coexistence and evolutionary dynamics mediated by seasonal environmental variation in annual plant communities.

Abstract: It is well established theoretically that competing species may coexist by having different responses to variation over time in the physical environment. Whereas previous theory has focused mostly on year-to-year environmental variation, we investigate how within-year variation can be the basis of species coexistence. We ask also the important but often neglected question of whether the species differences that allow coexistence are compatible with evolutionary processes. We seek the simplest circumstances that permit coexistence based on within-year environmental variation, and then evaluate the robustness of coexistence in the face of evolutionary forces. Our focus is on coexistence of annual plant species living in arid regions. We first consider environmental variation of a very simple structure where a single pulse of rain occurs, and different species have different patterns of growth activity following the rain pulse. We show that coexistence of two species is possible based on the storage effect coexistence mechanism in this simplest of varying environments. We find an exact expression for the magnitude of the storage effect that allows the functioning of the coexistence mechanism to be analyzed. However, in these simplest of circumstances, coexistence in our models is not evolutionarily stable. Increasing the complexity of the environment to two rain pulses leads to evolutionarily stable species coexistence, and a route to diversity via evolutionary branching. This demonstration of the compatibility of a coexistence mechanism with evolutionary processes is an important step in assessing the likely importance of a mechanism in nature.

Pub.: 05 Jan '13, Pinned: 15 Jun '17

Variation in moisture duration as a driver of coexistence by the storage effect in desert annual plants.

Abstract: Temporal environmental variation is a leading hypothesis for the coexistence of desert annual plants. Environmental variation is hypothesized to cause species-specific patterns of variation in germination, which then generates the storage effect coexistence mechanism. However, it has never been shown how sufficient species differences in germination patterns for multispecies coexistence can arise from a shared fluctuating environment. Here we show that nonlinear germination responses to a single fluctuating physical environmental factor can lead to sufficient differences between species in germination pattern for the storage effect to yield coexistence of multiple species. We derive these nonlinear germination responses from experimental data on the effects of varying soil moisture duration. Although these nonlinearities lead to strong species asymmetries in germination patterns, the relative nonlinearity coexistence mechanism is minor compared with the storage effect. However, these asymmetries mean that the storage effect can be negative for some species, which then only persist in the face of interspecific competition through average fitness advantages. This work shows how a low dimensional physical environment can nevertheless stabilize multispecies coexistence when the species have different nonlinear responses to common conditions, as supported by our experimental data.

Pub.: 19 Nov '13, Pinned: 15 Jun '17

The relative importance of relative nonlinearity and the storage effect in the lottery model.

Abstract: Although it is likely that many coexistence mechanisms contribute to maintenance of species diversity, most approaches to understanding species coexistence proceed as if only one mechanism would be present. In studies of species coexistence in a temporally fluctuating environment, the storage effect, believed to be the most important coexistence mechanism, has been the focus. Although a different coexistence mechanism--relative nonlinearity--is also predicted to arise frequently with environmental variation, its effect has been overshadowed by the storage effect. The relatively nonlinear growth rates on which the mechanism depends arise simply from differences in life history traits. Many kinds of temporal variation can then interact with these nonlinearity differences to create the relative nonlinearity coexistence mechanism. Much is unknown about when this mechanism is important and its total neglect is not justified. Here, we use the lottery model to provide a much needed quantitative assessment of the relative and combined effects of relative nonlinearity and the storage effect. Our analysis takes advantage of recently developed techniques for quantifying coexistence mechanisms when multiple mechanisms operate in concert. We find that relative nonlinearity is able to contribute substantially to species coexistence in the lottery model when two conditions are satisfied: (1) species must differ greatly in their adult death rates, (2) sensitivity of recruitment to environmental variation must be greater for species with larger adult death rates. In addition, relative nonlinearity has a critical role in compensating for a weakened storage effect when there is high correlation between species in their responses to the varying environment. In some circumstances relative nonlinearity is stronger than the storage effect or is even the sole mechanism of coexistence.

Pub.: 27 Aug '15, Pinned: 15 Jun '17