Phenotypic plasticity: A missing element in the theory of vegetation pattern formation.

Regular spatial patterns of vegetation are a common sight in drylands. Their formation is a population-level response to water stress that increases water availability for the few via partial plant mortality. At the individual level, plants can also adapt to water stress by changing their phenotype. Phenotypic plasticity of individual plants and spatial patterning of plant populations have extensively been studied independently, but the likely interplay between the two robust mechanisms has remained unexplored. In this paper, we incorporate phenotypic plasticity into a multi-level theory of vegetation pattern formation and use a fascinating ecological phenomenon, the Namibian "fairy circles," to demonstrate the need for such a theory. We show that phenotypic changes in the root structure of plants, coupled with pattern-forming feedback within soil layers, can resolve two puzzles that the current theory fails to explain: observations of multi-scale patterns and the absence of theoretically predicted large-scale stripe and spot patterns along the rainfall gradient. Importantly, we find that multi-level responses to stress unveil a wide variety of more effective stress-relaxation pathways, compared to single-level responses, implying a previously underestimated resilience of dryland ecosystems.

Discovery of fairy circles in Australia supports self-organization theory.

Vegetation gap patterns in arid grasslands, such as the "fairy circles" of Namibia, are one of nature's greatest mysteries and subject to a lively debate on their origin. They are characterized by small-scale hexagonal ordering of circular bare-soil gaps that persists uniformly in the landscape scale to form a homogeneous distribution. Pattern-formation theory predicts that such highly ordered gap patterns should be found also in other water-limited systems across the globe, even if the mechanisms of their formation are different. Here we report that so far unknown fairy circles with the same spatial structure exist 10,000 km away from Namibia in the remote outback of Australia. Combining fieldwork, remote sensing, spatial pattern analysis, and process-based mathematical modeling, we demonstrate that these patterns emerge by self-organization, with no correlation with termite activity; the driving mechanism is a positive biomass-water feedback associated with water runoff and biomass-dependent infiltration rates. The remarkable match between the patterns of Australian and Namibian fairy circles and model results indicate that both patterns emerge from a nonuniform stationary instability, supporting a central universality principle of pattern-formation theory. Applied to the context of dryland vegetation, this principle predicts that different systems that go through the same instability type will show similar vegetation patterns even if the feedback mechanisms and resulting soil-water distributions are different, as we indeed found by comparing the Australian and the Namibian fairy-circle ecosystems. These results suggest that biomass-water feedbacks and resultant vegetation gap patterns are likely more common in remote drylands than is currently known.

Emerged or imposed: a theory on the role of physical templates and self-organisation for vegetation patchiness.

In this article, we develop a unifying framework for the understanding of spatial vegetation patterns in heterogeneous landscapes. While much recent research has focused on self-organised vegetation the prevailing view is still that biological patchiness is mostly due to top-down control by the physical landscape template, disturbances or predators. We suggest that vegetation patchiness in real landscapes is controlled both by the physical template and by self-organisation simultaneously, and introduce a conceptual model for the relative roles of the two mechanisms. The model considers four factors that control whether vegetation patchiness is emerged or imposed: soil patch size, plant size, resource input and resource availability. The last three factors determine the plant-patch size, and the plant-to-soil patch size ratio determines the impact of self-organisation, which becomes important when this ratio is sufficiently small. A field study and numerical simulations of a mathematical model support the conceptual model and give further insight by providing examples of self-organised and template-controlled vegetation patterns co-occurring in the same landscape. We conclude that real landscapes are generally mixtures of template-induced and self-organised patchiness. Patchiness variability increases due to source-sink resource relations, and decreases for species of larger patch sizes.

Megaripple mechanics: bimodal transport ingrained in bimodal sands.

Aeolian sand transport is a major process shaping landscapes on Earth and on diverse celestial bodies. Conditions favoring bimodal sand transport, with fine-grain saltation driving coarse-grain reptation, give rise to the evolution of megaripples with a characteristic bimodal sand composition. Here, we derive a unified phase diagram for this special aeolian process and the ensuing nonequilibrium megaripple morphodynamics by means of a conceptually simple quantitative model, grounded in the grain-scale physics. We establish a well-preserved quantitative signature of bimodal aeolian transport in the otherwise highly variable grain size distributions, namely, the log-scale width (Krumbein phi scale) of their coarse-grain peaks. A comprehensive collection of terrestrial and extraterrestrial data, covering a wide range of geographical sources and environmental conditions, supports the accuracy and robustness of this unexpected theoretical finding. It could help to resolve ambiguities in the classification of terrestrial and extraterrestrial sedimentary bedforms.

Mechanisms of vegetation-ring formation in water-limited systems.

A common patch form in dryland landscapes is the vegetation ring. Vegetation patch formation has recently been attributed to self-organization processes that act to increase the availability of water to vegetation patches under conditions of water scarcity. The view of ring formation as a water-limited process, however, has remained largely unexplored. Using laboratory experiments and model studies we identify two distinct mechanisms of ring formation. The first mechanism pertains to conditions of high infiltration contrast between vegetated and bare soil, under which overland water flow is intercepted at the patch periphery. The decreasing amount of water that the patch core receives as the patch expands, leads to central dieback and ring formation. The second mechanism pertains to plants with large lateral root zones, and involves central dieback and ring formation due to increasing water uptake by the newly recruited individuals at the patch periphery. In general the two mechanisms act in concert, but the relative importance of each mechanism depends on environmental conditions. We found that strong seasonal rainfall variability favors ring formation by the overland-flow mechanism, while a uniform rainfall regime favors ring formation by the water-uptake mechanism. Our results explain the formation of rings by fast-growing species with confined root zones in a dry-Mediterranean climate, such as Poa bulbosa. They also explain the formation of rings by slowly growing species with highly extended root zones, such as Larrea tridentata (Creosotebush).

Positive impacts of livestock and wild ungulate routes on functioning of dryland ecosystems.

Livestock grazing is often perceived as being detrimental to the quality and functioning of dryland ecosystems. For example, a study in a semiarid Kenyan savanna proposed that cattle form bare spaces throughout the landscape, which indicate ecosystem degradation. Other studies, conducted in north-eastern Spain, where climatic conditions range between semiarid and Mediterranean subhumid, reported that sheep and goat trails have increased the emergence of rill erosion processes. Sometimes, this negative perception is extended to include wild, large ungulate herbivores as well. Here, we challenge this perception by highlighting the generally nonadverse and even ameliorative impacts of moderate animal rate on geoecosystem functioning of hilly drylands. Specifically, trampling routes (also known as treading paths, livestock terracettes, cattle trails, migration tracks, cowtours, etc.) formed across hillslopes by grazing animals-being either domesticated livestock or native large herbivores-transform the original two-phase vegetation mosaic of shrubby patches and interpatch spaces into a three-phase mosaic. The animal routes increase the complexity of ecosystem, by strengthening the spatial redistribution of water and soil resources at the patch scale and decreasing hydrological connectivity at the hillslope scale. As a consequence, the animal routes improve functioning of hilly drylands and increase their resilience to long-term droughts and climatic change. Therefore, instead of viewing the animal routes as degraded spots, they should be perceived at a wider perspective that allows to properly understand their overall role in sustaining dryland geoecosystems.

Geodiversity impacts plant community structure in a semi-arid region.

Geodiversity refers to the variety of geological and physical elements as well as to geomorphological processes of the earth surface. Heterogeneity of the physical environment has an impact on plant diversity. In recent years, the relations between geodiversity and biodiversity has gained attention in conservation biology, especially in the context of climate change. In this study, we assessed the spatial and temporal change in plant's community structure in a semi-arid region, Sayeret Shaked Long Term Ecosystem Research (LTER) station, Israel. Vegetation surveys were conducted on different hillslopes, either with or without rock covers in order to study the spatial trends of hillslope geodiversity. The surveys were conducted for two consecutive years (2016 and 2017), of which the second year was drier and hotter and therefore permitted to investigate the temporal change of plant's community structure. The results of the spatial trends show that (1) geodiversity increases vegetation biodiversity and promotes perennial plants and those of the temporal change show that (2) the positive effect of geodiversity on plants' community structure and species richness is greater in the drier year than that in a wetter year. The main insight is that in these drylands, hillslopes with higher geodiversity appear to buffer the effect of drier years, and supported a more diverse plant community than lower geodiversity hillslopes.

Periodic versus scale-free patterns in dryland vegetation.

Two major forms of vegetation patterns have been observed in drylands: nearly periodic patterns with characteristic length scales, and amorphous, scale-free patterns with wide patch-size distributions. The emergence of scale-free patterns has been attributed to global competition over a limiting resource, but the physical and ecological origin of this phenomenon is not understood. Using a spatially explicit mathematical model for vegetation dynamics in water-limited systems, we unravel a general mechanism for global competition: fast spatial distribution of the water resource relative to processes that exploit or absorb it. We study two possible realizations of this mechanism and identify physical and ecological conditions for scale-free patterns. We conclude by discussing the implications of this study for interpreting signals of imminent desertification.

A model study of terraced riverbeds as novel ecosystems.

Riverbed terracing has been introduced in ancient times to retain water and soil, to reduce hydrological connectivity and erosion and to increase primary and secondary productivity of agro-ecological systems. These presently abandoned human-made landscapes have become novel ecosystems and a potential source of ecosystem services to humans in drylands. We use a mathematical-modeling approach to study factors that regulate terraced riverbeds and affect community and ecosystem attributes such as productivity, functional diversity and resilience to droughts. We introduce a model that captures the relationships between rainfall pattern, runoff coupling between adjacent terraces, and vegetation growth, taking into account competition for water and light. We found that a large number of weak rainfall events results in lower total biomass and functional diversity across the terraced riverbed compared with a few strong rainfall events. We further analyzed the filtering of species traits from pools of functional groups that make different tradeoffs between investment in above-ground biomass to capture canopy resources and investment in below-ground biomass to capture soil resources. Pools characterized by concave tradeoffs give rise to higher functional diversity, lower biomass production and lower resilience to droughts, as compared with convex pools. New empirical studies are needed to test these model predictions.

Critical transitions in Chinese dunes during the past 12,000 years.

Dune systems can have alternative stable states that coexist under certain environmental conditions: a vegetated, stabilized state and a bare active state. This behavior implies the possibility of abrupt transitions from one state to another in response to gradual environmental change. Here, we synthesize stratigraphic records covering 12,000 years of dynamics of this system at 144 localities across three dune fields in northern China. We find side-by-side coexistence of active and stabilized states, and occasional sharp shifts in time between those contrasting states. Those shifts occur asynchronously despite the fact that the entire landscape has been subject to the same gradual changes in monsoon rainfall and other conditions. At larger scale, the spatial heterogeneity in dune dynamics averages out to produce relatively smooth change. However, our results do show different paths of recovery and collapse of vegetation at system-wide scales, implying that hysteretic behavior occurs in spatially extended systems.

Biogenic crust dynamics on sand dunes.

Sand dunes are often covered by vegetation and biogenic crusts. Despite their significant role in dune stabilization, biogenic crusts have rarely been considered in model studies of dune dynamics. Using a simple model, we study the existence and stability ranges of different dune-cover states along gradients of rainfall and wind power. Two ranges of alternative stable states are identified: fixed crusted dunes and fixed vegetated dunes at low wind power; and fixed vegetated dunes and active dunes at high wind power. These results suggest a crossover between two different forms of desertification.

Regime shifts in models of dryland vegetation.

Drylands are pattern-forming systems showing self-organized vegetation patchiness, multiplicity of stable states and fronts separating domains of alternative stable states. Pattern dynamics, induced by droughts or disturbances, can result in desertification shifts from patterned vegetation to bare soil. Pattern formation theory suggests various scenarios for such dynamics: an abrupt global shift involving a fast collapse to bare soil, a gradual global shift involving the expansion and coalescence of bare-soil domains and an incipient shift to a hybrid state consisting of stationary bare-soil domains in an otherwise periodic pattern. Using models of dryland vegetation, we address the question of which of these scenarios can be realized. We found that the models can be split into two groups: models that exhibit multiplicity of periodic-pattern and bare-soil states, and models that exhibit, in addition, multiplicity of hybrid states. Furthermore, in all models, we could not identify parameter regimes in which bare-soil domains expand into vegetated domains. The significance of these findings is that, while models belonging to the first group can only exhibit abrupt shifts, models belonging to the second group can also exhibit gradual and incipient shifts. A discussion of open problems concludes the paper.

Regime shifts in models of dryland vegetation.

Drylands are pattern-forming systems showing self-organized vegetation patchiness, multiplicity of stable states and fronts separating domains of alternative stable states. Pattern dynamics, induced by droughts or disturbances, can result in desertification shifts from patterned vegetation to bare soil. Pattern formation theory suggests various scenarios for such dynamics: an abrupt global shift involving a fast collapse to bare soil, a gradual global shift involving the expansion and coalescence of bare-soil domains and an incipient shift to a hybrid state consisting of stationary bare-soil domains in an otherwise periodic pattern. Using models of dryland vegetation, we address the question of which of these scenarios can be realized. We found that the models can be split into two groups: models that exhibit multiplicity of periodic-pattern and bare-soil states, and models that exhibit, in addition, multiplicity of hybrid states. Furthermore, in all models, we could not identify parameter regimes in which bare-soil domains expand into vegetated domains. The significance of these findings is that, while models belonging to the first group can only exhibit abrupt shifts, models belonging to the second group can also exhibit gradual and incipient shifts. A discussion of open problems concludes the paper.

Why do active and stabilized dunes coexist under the same climatic conditions?

Sand dunes can be active (mobile) or stable, mainly as a function of vegetation cover and wind power. However, there exists as yet unexplained evidence for the coexistence of bare mobile dunes and vegetated stabilized dunes under the same climatic conditions. We propose a model for dune vegetation cover driven by wind power that exhibits bistabilty and hysteresis with respect to the wind power. For intermediate wind power, mobile and stabilized dunes can coexist, whereas for low (or high) wind power they can be fixed (or mobile). Climatic change or human intervention can turn active dunes into stable ones and vice versa; our model predicts that prolonged droughts with stronger winds can result in dune reactivation.

Localized structures in dryland vegetation: forms and functions.

Vegetation patches in drylands are localized structures of biomass and water. We study these structures using a mathematical modeling approach that captures biomass-water feedbacks. Biomass-water structures are found to differ in their spatial forms and ecological functions, depending on species type, soil conditions, precipitation range, and other environmental factors. Asymptotic spot structures can destabilize to form ring structures, expanding in the radial direction, or crescent structures, migrating uphill. Stable spot structures can differ in their soil-water distributions, forming water-enriched patches or water-deprived patches. The various biomass-water structures are expected to function differently in the context of a plant community, forming landscapes of varying species diversity.