How Long Before a Second Defoliation of Actively Growing Grass Plants in the Desert Grassland?

In the Desert Grassland, second and subsequent defoliations on perennial grasses during the active growing season can have substantial impacts on grass recovery and reproduction following herbivory. Land managers implement tactics to avoid multiple defoliations on plants by way of rotational grazing, reduced stocking rates, and/or reduced time spent within a given pasture. We explored frequency and rate of defoliation by cattle on perennial bunchgrasses within an 11-day grazing period in three pastures including distance to water (300 and 600 m) and plant height to determine their influence on animal diet selection. Results indicate that 32% of all marked plants were defoliated by cattle and only 5% of defoliated plants were defoliated a second time by day 10 of the grazing period. Defoliation patterns in the studied pastures did not differ between two distances from water, or in relation to plant height. However, at the second defoliation cattle grazed plants that were shorter than at the first defoliation suggesting a selection for high quality re-growth over larger forage on offer. The results of this study show that a 10-day grazing period during the growing season of the Desert Grassland is an effective strategy to avoid second defoliations on individual perennial grass plants while maintaining sufficient forage for use during the dormant winter grazing season.

Soil organic carbon in drylands: shrub encroachment and vegetation management effects dwarf those of livestock grazing.

Dryland ecosystems occur worldwide and play a prominent, but potentially shifting, role in global biogeochemical cycling. Widespread woody plant proliferation, often associated with declines in palatable grasses, has jeopardized livestock production in drylands and prompted attempts to reduce woody cover by chemical or mechanical means. Woody encroachment also has the potential to significantly alter terrestrial carbon storage. However, little is known of the long-term biogeochemical consequences of woody encroachment in the broader context of its interaction with common dryland land uses, including "brush management" (woody plant clearing) and livestock grazing. Present assessments exhibit considerable variation in the consequences of these land use/land cover changes, with evidence that brush management may counteract sizeable impacts of shrub encroachment on soil biogeochemical pools. A challenge to assessing the net effects of brush management in shrub-encroached grasslands on soil organic carbon (SOC) and total nitrogen (N) pools is that land management practices are typically considered in isolation, when they are co-occurring phenomena. Furthermore, few studies have assessed spatial patterns in brush management and how these are affected in decades following treatment on sites with contrasting grazing histories. To address these uncertainties and interactions, we quantified the impacts of shrub encroachment and their subsequent mortality resulting from brush management (herbicide application) on SOC and N pools in a Sonoran Desert grassland where long-term grazing manipulations (>100 yr) co-occur with shrub encroachment and brush management. Pools of SOC and N associated with herbicided shrubs declined markedly over ~40 yr, offsetting 66% of the increases from shrub encroachment. However, spatial patterns in SOC induced by shrubs persisted over the decades following brush management. Century-long protection from grazing did little to change SOC and N pools. Accordingly, shrub encroachment and shrub mortality from brush management each far outweighed livestock grazing impacts. Consideration of the patterns of SOC and N through space (e.g., bole-to-dripline gradients), time (e.g., shrub age/size), land use (e.g., livestock grazing and brush management), and their interactions will position us to improve predictions of SOC and N responses to land use/land cover change, inform C-based management decisions, and objectively evaluate trade-offs with other ecosystem services.

Shrub encroachment into grasslands: end of an era?

Shifts in the abundance of grasses and woody plants in drylands have occurred several times during the Holocene. However, our understanding of the rates and dynamics of this state-change in recent decades is limited to scattered studies conducted at disparate spatial and temporal scales; the potential misperceptions of shrub cover change could be remedied using cross spatiotemporal scale analyses that link field observations, repeat ground-level photography and remote sensing perspectives. The study was conducted across a semi-arid landscape in southern Arizona. Local data from long-term transects revealed three distinct chronological phases of shrub cover change: expansion (1961-1991, 0.7% y-1), decline (1992-1997, -2.3% y-1) and stabilization (1998-2012, 22-25% with no net cover change). Twenty-eight years (1984-2011) of broad-scale Landsat Thematic Mapper assessments confirm that shrub cover has been relatively stable in recent decades regardless of grazing regimes and landforms with the exception of the proliferation of succulents at lower elevations (verified by repeat photography acquired in 1987 and 2015) where the physical environment is the harshest, reflecting elevated temperature and winter precipitation deficit. Warmer, drier future climates are predicted to reduce woody plant carrying capacity and promote a shift to xerophytic succulents.

Quantifying long-term plant community dynamics with movement models: implications for ecological resilience.

Quantification of rates and patterns of community dynamics is central for understanding the organization and function of ecosystems. These insights may support a greater empirical understanding of ecological resilience, and the application of resilience concepts toward ecosystem management. Distinct types of dynamics in natural communities can be used to interpret and apply resilience concepts, but quantitative methods that can systematically distinguish among them are needed. We develop a quantitative method to analyze long-term records of plant community dynamics using principles of movement ecology. We analyzed dissimilarity of species composition through time with linear and nonlinear statistical models to assign community change to four classes of movement trajectories. Compositional change in each sampled plot through time was classified into four classes, stability, abrupt nonlinear change, transient reversible change, and gradual linear drift, each representing a different aspect of ecological resilience. These competing models were evaluated based on estimated coefficients, goodness of fit, and parsimony. We tested our method's accuracy and robustness through simulations, or the ability to distinguish among trajectories and classify them correctly. We simulated 16,000 trajectories of four types, of which 94-100% were correctly classified. Next, we analyzed 13 long-term vegetation records from North American grasslands (annual grasslands with warm-season and cool-season communities, shortgrass, mixedgrass, and tallgrass prairies, and sagebrush steppe), and a record of primary succession at Mt. St. Helens volcano. Collectively, we analyzed 14,647 observations from 775 plots, between 1915 and 2012. Dynamics could be reliably assigned for 705 plots (91%), and overall statistical fit was high (goodness of fit, 0.77 ± 0.15 SD). Among the perennial grasslands, stability was most common (44% of all plots), followed by gradual linear (22%), abrupt nonlinear (17%), and reversible (6%) change. Among annual grasslands, abrupt nonlinear shifts (33%) were more common in the warm-season community than in the cool-season (20%). As expected, abrupt nonlinear change was common during primary succession (51%) while reversible change was rare (3%). Generally, reversible dynamics often required 2-3 decades. Analysis of long-term community change, or trajectories, with principles of movement ecology provides a quantitative basis to compare and interpret ecological resilience within and among ecosystems.

Considerations for Achieving Cross-Platform Point Cloud Data Fusion across Different Dryland Ecosystem Structural States.

Remotely sensing recent growth, herbivory, or disturbance of herbaceous and woody vegetation in dryland ecosystems requires high spatial resolution and multi-temporal depth. Three dimensional (3D) remote sensing technologies like lidar, and techniques like structure from motion (SfM) photogrammetry, each have strengths and weaknesses at detecting vegetation volume and extent, given the instrument's ground sample distance and ease of acquisition. Yet, a combination of platforms and techniques might provide solutions that overcome the weakness of a single platform. To explore the potential for combining platforms, we compared detection bias amongst two 3D remote sensing techniques (lidar and SfM) using three different platforms [ground-based, small unmanned aerial systems (sUAS), and manned aircraft]. We found aerial lidar to be more accurate for characterizing the bare earth (ground) in dense herbaceous vegetation than either terrestrial lidar or aerial SfM photogrammetry. Conversely, the manned aerial lidar did not detect grass and fine woody vegetation while the terrestrial lidar and high resolution near-distance (ground and sUAS) SfM photogrammetry detected these and were accurate. UAS SfM photogrammetry at lower spatial resolution under-estimated maximum heights in grass and shrubs. UAS and handheld SfM photogrammetry in near-distance high resolution collections had similar accuracy to terrestrial lidar for vegetation, but difficulty at measuring bare earth elevation beneath dense herbaceous cover. Combining point cloud data and derivatives (i.e., meshes and rasters) from two or more platforms allowed for more accurate measurement of herbaceous and woody vegetation (height and canopy cover) than any single technique alone. Availability and costs of manned aircraft lidar collection preclude high frequency repeatability but this is less limiting for terrestrial lidar, sUAS and handheld SfM. The post-processing of SfM photogrammetry data became the limiting factor at larger spatial scale and temporal repetition. Despite the utility of sUAS and handheld SfM for monitoring vegetation phenology and structure, their spatial extents are small relative to manned aircraft.

Direct effects dominate responses to climate perturbations in grassland plant communities.

Theory predicts that strong indirect effects of environmental change will impact communities when niche differences between competitors are small and variation in the direct effects experienced by competitors is large, but empirical tests are lacking. Here we estimate negative frequency dependence, a proxy for niche differences, and quantify the direct and indirect effects of climate change on each species. Consistent with theory, in four of five communities indirect effects are strongest for species showing weak negative frequency dependence. Indirect effects are also stronger in communities where there is greater variation in direct effects. Overall responses to climate perturbations are driven primarily by direct effects, suggesting that single species models may be adequate for forecasting the impacts of climate change in these communities.

Functional response of U.S. grasslands to the early 21st-century drought.

Grasslands across the United States play a key role in regional livelihood and national food security. Yet, it is still unclear how this important resource will respond to the prolonged warm droughts and more intense rainfall events predicted with climate change. The early 21st-century drought in the southwestern United States resulted in hydroclimatic conditions that are similar to those expected with future climate change. We investigated the impact of the early 21st-century drought on aboveground net primary production (ANPP) of six desert and plains grasslands dominated by C4 (warm season) grasses in terms of significant deviations between observed and expected ANPP. In desert grasslands, drought-induced grass mortality led to shifts in the functional response to annual total precipitation (P(T)), and in some cases, new species assemblages occurred that included invasive species. In contrast, the ANPP in plains grasslands exhibited a strong linear function of the current-year P(T) and the previous-year ANPP, despite prolonged warm drought. We used these results to disentangle the impacts of interannual total precipitation, intra-annual precipitation patterns, and grassland abundance on ANPP, and thus generalize the functional response of C4 grasslands to predicted climate change. This will allow managers to plan for predictable shifts in resources associated with climate change related to fire risk, loss of forage, and ecosystem services.

Ecosystem resilience despite large-scale altered hydroclimatic conditions.

Climate change is predicted to increase both drought frequency and duration, and when coupled with substantial warming, will establish a new hydroclimatological model for many regions. Large-scale, warm droughts have recently occurred in North America, Africa, Europe, Amazonia and Australia, resulting in major effects on terrestrial ecosystems, carbon balance and food security. Here we compare the functional response of above-ground net primary production to contrasting hydroclimatic periods in the late twentieth century (1975-1998), and drier, warmer conditions in the early twenty-first century (2000-2009) in the Northern and Southern Hemispheres. We find a common ecosystem water-use efficiency (WUE(e): above-ground net primary production/evapotranspiration) across biomes ranging from grassland to forest that indicates an intrinsic system sensitivity to water availability across rainfall regimes, regardless of hydroclimatic conditions. We found higher WUE(e) in drier years that increased significantly with drought to a maximum WUE(e) across all biomes; and a minimum native state in wetter years that was common across hydroclimatic periods. This indicates biome-scale resilience to the interannual variability associated with the early twenty-first century drought--that is, the capacity to tolerate low, annual precipitation and to respond to subsequent periods of favourable water balance. These findings provide a conceptual model of ecosystem properties at the decadal scale applicable to the widespread altered hydroclimatic conditions that are predicted for later this century. Understanding the hydroclimatic threshold that will break down ecosystem resilience and alter maximum WUE(e) may allow us to predict land-surface consequences as large regions become more arid, starting with water-limited, low-productivity grasslands.

Empirical assessment of state-and-transition models with a long-term vegetation record from the Sonoran Desert.

Resilience-based frameworks, including state-and-transition models (STM), are being increasingly called upon to inform policy and guide ecosystem management, particularly in rangelands. Yet, multiple challenges impede their effective implementation: (1) paucity of empirical tests of resilience concepts, such as alternative states and thresholds, and (2) heavy reliance on expert models, which are seldom tested against empirical data. We developed an analytical protocol to identify unique plant communities and their transitions, and applied it to a long-term vegetation record from the Sonoran Desert (1953-2009). We assessed whether empirical trends were consistent with resilience concepts, and evaluated how they may inform the construction and interpretation of expert STMs. Seven statistically distinct plant communities were identified based on the cover of 22 plant species in 68 permanent transects. We recorded 253 instances of community transitions, associated with changes in species composition between successive samplings. Expectedly, transitions were more frequent among proximate communities with similar species pools than among distant communities. But unexpectedly, communities and transitions were not strongly constrained by soil type and topography. Only 18 transitions featured disproportionately large compositional turnover (species dissimilarity ranged between 0.54 and 0.68), and these were closely associated with communities that were dominated by the common shrub (burroweed, Haplopappus tenuisecta); indicating that only some, and not all, communities may be prone to large compositional change. Temporal dynamics in individual transects illustrated four general trajectories: stability, nondirectional drift, reversibility, and directional shifts that were not reversed even after 2-3 decades. The frequency of transitions and the accompanying species dissimilarity were both positively correlated with fluctuation in precipitation, indicating that climatic drivers require more attention in STMs. Many features of the expert models, including the number of communities and participant species, were consistent with empirical trends, but expert models underrepresented recent increases in cacti while overemphasizing the introduced Lehmann's lovegrass (Eragrostis lehmanniana). Quantification of communities and transitions within long-term vegetation records presents several quantitative metrics such as transition frequency, magnitude of accompanying compositional change, presence of unidirectional trajectories, and lack of reversibility within various timescales, which can clarify resilience concepts and inform the construction and interpretation of STMs.

Woody plants in grasslands: post-encroachment stand dynamics.

Woody plant abundance is widely recognized to have increased in savannas and grasslands worldwide. The lack of information on the rates, dynamics, and extent of increases in shrub abundance is a major source of uncertainty in assessing how this vegetation change has influenced biogeochemical cycles. Projecting future consequences of woody cover change on ecosystem function will require knowledge of where shrub cover in present-day stands lies relative to the realizable maximum for a given soil type within a bioclimatic region. We used time-series aerial photography (1936, 1966, and 1996) and field studies to quantify cover and biomass of velvet mesquite (Prosopis velutina Woot.) following its proliferation in a semidesert grassland of Arizona. Mapping of individual shrubs indicated an encroachment phase characterized by high rates of bare patch colonization. Upon entering a stabilization phase, shrub cover increases associated with recruitment and canopy expansion were largely offset by contractions in canopy area of other shrub patches. Instances of shrub disappearance coincided with a period of below-average rainfall (1936-1966). Overall, shrub cover (mean +/- SE) on sandy uplands with few and widely scattered shrubs in 1902 was dynamically stable over the 1936-1996 period averaging approximately 35% +/- 5%. Shrub cover on clayey uplands in 1936 was 17% +/- 2% but subsequently increased twofold to levels comparable to those on sandy uplands by 1966 (36% +/- 7%). Cover on both soils then decreased slightly between 1966 and 1996 to 28% +/- 3%. Thus, soil properties influenced the rate at which landscapes reached a dynamic equilibrium, but not the apparent endpoint. Although sandy and clayey landscapes appear to have stabilized at comparable levels of cover, shrub biomass was 1.4 times greater on clayey soils. Declines in shrub cover between 1966 and 1996 were accompanied by a shift to smaller patch sizes on both sandy and clayey landscapes. Dynamics observed during the stabilization phase suggest that density-dependent regulation may be in play. If woody cover has transitioned from directional increases to a dynamic equilibrium, biomass projections will require monitoring and modeling patch dynamics and stand structure rather than simply changes in total cover.

Postfire stand structure in a semiarid savanna: cross-scale challenges estimating biomass.

Algorithms relating remotely sensed woody cover to biomass are often the basis for large-scale inventories of aboveground carbon stocks. However, these algorithms are commonly applied in a generic fashion without consideration of disturbances that might alter vegetation structure. We compared field and remote sensing estimates of woody biomass on savannas with contrasting disturbance (fire) histories and assessed potential errors in estimating woody biomass from cover without considering fire history. Field surveys quantified multilayer cover (MLC) of woody and succulent plants on sites experiencing wildfire in 1989 or 1994 and on nearby unburned (control) sites. Remote sensing estimates of the woody cover fraction (WCF) on burned and control sites were derived from contemporary (2005) dry-season Landsat Thematic Mapper imagery (during a period when herbaceous cover was senescent) using a probabilistic spectral mixture analysis model. Satellite WCF estimates were compared to field MLC assessments and related to aboveground biomass using allometry. Field-based MLC and remotely sensed WCFs both indicated that woody cover was comparable on control areas and areas burned 11-16 years ago. However, biomass was approximately twofold higher on control sites. Canopy cover was a strong predictor of woody biomass on burned and control areas, but fire history significantly altered the linear cover-biomass relationship on control plots to a curvilinear relationship on burned plots. Results suggest predictions of woody biomass from "generic" two-dimensional (2-D) cover algorithms may underestimate biomass in undisturbed stands and overestimate biomass in stands recovering from disturbance. Improving the accuracy of woody-biomass estimates from field and/or remotely sensed cover may therefore require disturbance-specific models or detection of vegetation height and transforming 2-D vegetation cover to 3-D vegetation volume.