The encroachment of woody plants into grasslands is a global phenomenon with implications for biodiversity and ecosystem function. Understanding and predicting the pace of expansion and the underlying processes that control it are key challenges in the study and management of woody encroachment. Theory from spatial population biology predicts that the occurrence and speed of population expansion should depend sensitively on the nature of conspecific density dependence. If fitness is maximized at the low-density encroachment edge then shrub expansion should be "pulled" forward. However, encroaching shrubs have been shown to exhibit positive feedbacks, whereby shrub establishment modifies the environment in ways that facilitate further shrub recruitment and survival. In this case there may be a fitness cost to shrubs at low density causing expansion to be "pushed" from behind the leading edge. We studied the spatial dynamics of creosotebush (Larrea tridentata), which has a history of encroachment into Chihuahuan Desert grasslands over the past century. We used demographic data from observational censuses and seedling transplant experiments to test the strength and direction of density dependence in shrub fitness along a gradient of shrub density at the grass-shrub ecotone. We also used seed-drop experiments and wind data to construct a mechanistic seed dispersal kernel, then connected demography and dispersal data within a spatial integral projection model (SIPM) to predict the dynamics of shrub expansion. The SIPM predicted that, contrary to expectations based on potential for positive feedbacks, the shrub encroachment wave is "pulled" by maximum fitness at the low-density front. However, the predicted pace of expansion was strikingly slow (ca. 8 cm/yr), and this prediction was supported by independent re-surveys of the ecotone showing little to no change in spatial extent of shrub cover over 12 years. Encroachment speed was acutely sensitive to seedling recruitment, suggesting that this population may be primed for pulses of expansion under conditions that are favorable for recruitment. Our integration of observations, experiments, and modeling reveals not only that this ecotone is effectively stalled under current conditions, but also why that is so and how that may change as the environment changes.
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Demography and dispersal at a grass‐shrub ecotone: A spatial integral projection model for woody plant encroachment
The encroachment of woody plants into grasslands is a global phenomenon with implications for biodiversity and ecosystem function. Understanding and predicting the pace of expansion and the underlying processes that control it are key challenges in the study and management of woody encroachment. Theory from spatial population biology predicts that the occurrence and speed of expansion should depend sensitively on the nature of conspecific density dependence. If fitness is maximized at the low‐density encroachment edge, then shrub expansion should be “pulled” forward. However, encroaching shrubs have been shown to exhibit positive feedbacks, whereby shrub establishment modifies the environment in ways that facilitate further shrub recruitment and survival. In this case there may be a fitness cost to shrubs at low density causing expansion to be “pushed” from behind the leading edge. We studied the spatial dynamics of creosotebush (
- NSF-PAR ID:
- 10413280
- Publisher / Repository:
- Wiley Blackwell (John Wiley & Sons)
- Date Published:
- Journal Name:
- Ecological Monographs
- Volume:
- 93
- Issue:
- 3
- ISSN:
- 0012-9615
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract Questions Woody encroachment into grasslands is a worldwide phenomenon partially influenced by climate change, including extreme weather events.
Larrea tridentata is a common shrub throughout the warm deserts of North America that has encroached into grasslands over the past 150 years. Physiological measurements suggest that the northern distribution ofL. tridentata is limited by cold temperatures; thus extreme winter events may slow or reverse shrub expansion. We tested this limitation by measuring the response of individualL. tridentata shrubs to an extreme winter cold (−31°C) event to assess shrub mortality and rate of recovery of surviving shrubs.Location Sevilleta National Wildlife Refuge, Socorro County, New Mexico, USA.
Methods Canopy dieback and recovery following an extreme cold event were measured for 869 permanently marked individual
L. tridentata shrubs in grass–shrub ecotone and shrubland sites. Individual shrubs were monitored for amount of canopy dieback, rate of recovery, and seed set for three growing seasons after the freeze event.Results Shrubs rapidly suffered a nearly complete loss of canopy leaf area across all sites. Although canopy loss was high, mortality was low and 99% of shrubs resprouted during the first growing season after the freeze event. Regrowth rates were similar within ecotone and shrubland sites, even when damage by frost was larger in the latter. After three years of recovery,
L. tridentata canopies had regrown on average 23–83% of the original pre‐freeze canopy sizes across the sites.Conclusions We conclude that isolated extreme cold events may temporarily decrease shrubland biomass but they do not slow or reverse shrub expansion. These events are less likely to occur in the future as regional temperatures increase under climate change.
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Abstract Transitions from grass to woody plant dominance, widely reported in arid systems, are typically attributed to changes in disturbance regimes in combination with abiotic feedbacks, whereas biotic mechanisms such as competition and facilitation are often overlooked. Yet, research in semi‐arid and subhumid savannas indicates that biotic interactions are important drivers in systems at risk for state transition. We sought to bridge this divide by experimentally manipulating grass‐on‐shrub and shrub‐on‐shrub interactions in early and late stages of grassland–shrubland state transition, respectively, and to assess the extent to which these interactions might influence arid land state transition dynamics.
Target
Prosopis glandulosa shrubs had surrounding grasses or conspecific neighbours left intact or killed with foliar herbicide, and metrics of plant performance were monitored over multiple years for shrubs with and without grass or shrub neighbours.Productivity of small shrubs was enhanced by grass removal in years with above‐average precipitation, a result not evident in larger shrubs or during dry years. Proxy evidence based on nearest neighbour metrics suggested shrub–shrub competition was at play, but our experimental manipulations revealed no such influence.
Competition from grasses appears to attenuate the rate at which shrubs achieve the size necessary to modify the physical environment in self‐reinforcing ways, but only during the early stages of shrub encroachment. Our results further suggest that at late stages of grassland‐to‐shrubland state transitions, shrub–shrub competition will not slow the rate of shrub expansion, and suggest that maximum shrub cover is regulated by something other than density‐dependent mechanisms. We conclude that grass effects on shrubs should be included in assessments of desert grassland state transition probabilities and rates, and that desertification models in arid ecosystems that traditionally focus on disturbance and abiotic feedbacks should be broadened to incorporate spatial and temporal variations in competitive effects.
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plain language summary is available for this article. -
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Abstract Successful forest expansion into grassland can be limited by seed dispersal and adverse conditions for tree seedlings in the grassland environment. In the high‐elevation Andes, human‐induced fragmentation has exacerbated the patchy distribution of
Polylepis forests, threatening their unique biological communities and spurring restoration interest. Studies ofPolylepis forest extent in Peru suggest that forest borders have remained stable over the past century despite decreasing anthropogenic disturbance, suggesting that tree seedling recruitment is being limited in the open grassland habitat. We studied natural seedling dispersion patterns ofPolylepis sericea andPolylepis weberbaueri (Rosaceae) at forest–grassland edges across a range of environmental conditions to examine seedling recruitment and colonization of grasslands in Huascaran National Park (Peru). Using data from 2367 seedlings found in 48 forest–grassland edge plots (15 m × 15 m) at forest patches between 3900–4500 masl, we employed generalized mixed modelling to identify the significant associations of seedling densities with environmental covariates. In addition, we compared these associations to patterns of adult presence on the landscape. Seedling densities were associated with a combination of variables varying within (distance to forest edge) and among (elevation and dry season solar irradiation) plots across the landscape. For both species, seedling densities decreased with increasing distance away from the forest in a manner consistent with short‐distance seed dispersal by wind. Our results suggest that such short‐distance dispersal may slow forest expansion, but that there also appear to be substantial post‐dispersal limitations to seedling establishment in the grassland.Polylepis sericea densities decreased with elevation, whileP. weberbaueri increased with elevation and decreased with solar irradiation. Associations of adult presence with elevation and solar irradiation mirrored those of seedling densities. Management of areas with forest patches dominated by these species should consider these differences in their environmental tolerances, particularly during species selection and zonation for reforestation. -
Shrub encroachment is a global phenomenon. Both the causes and consequences of shrub encroachment vary regionally and globally. In the southwestern US a common native C3 shrub species, creosotebush, has invaded millions of hectares of arid and semi-arid C4-dominated grassland. At the Sevilleta LTER site, it appears that the grassland-shrubland ecotone is relatively stable, but infill by creosotebush continues to occur. The consequences of shrub encroachment have been and continue to be carefully documented, but the ecological drivers of shrub encroachment in the southwestern US are not well known. One key factor that may promote shrub encroachment is grazing by domestic livestock. However, multiple environmental drivers have changed over the 150 years during which shrub expansion has occurred through the southwestern US. Temperatures are warmer, atmospheric CO2 has increased, drought and rainy cycles have occurred, and grazing pressure has decreased. From our prior research we know that prolonged drought greatly reduces the abundance of native grasses while having limited impact on the abundance of creosotebush in the grass-shrub ecotone. So once established, creosotebush populations are persistent and resistant to climate cycles. We also know that creosotebush seedlings tend to appear primarily when rainfall during the summer monsoon is well above average. However, high rainfall years also stimulate the growth of the dominant grasses creating a competitive environment that may not favor seedling establishment and survival. The purpose of the Mega-Monsoon Experiment (MegaME) is twofold. First, this experiment will determine if high rainfall years coupled with (simulated) grazing promote the establishment and growth of creosotebush seedlings in the grassland-shrubland ecotone at Sevilleta, thus promoting infill and expansion of creosotebush into native grassland. Second, MegaME will determine if a sequence of wet summer monsoons will promote the establishment and growth of native C4 grasses in areas where creosotebush is now dominant, thus demonstrating that high rainfall and dispersal limitation prevent grassland expansion into creosotebush shrubland.more » « less
