Woody plant encroachment is a globally-prevalent vegetation change phenomenon that has shifted grass-dominated ecosystems to mixed grass and woody plant matrices over the last century. In the Rio Grande Plains of Texas, the introduction of N-fixing woody legumes has increased above- and belowground primary productivity and changed the litter chemistry of the system, accelerating rates of belowground biogeochemical processes. The purpose of this study was to assess the impact of grassland to woodland transition on i) P concentrations in soil physical fractions that differ in their organic matter turnover rates, ii) P availability within the soil over the course of woody encroachment and across the landscape, and iii) microbial community composition and diversity. Soil samples were collected in remnant grasslands and four woody landscape elements (clusters, groves, drainage woodlands, and playas) along a 135-yr chronosequence of woody plant encroachment. P was fractionated by the Hedley method and P concentrations were determined by alkaline oxidation and lithium fusion coupled with ascorbic acid colorimetry. Bacterial and fungal communities were characterized by molecular methods. Whole soil P concentrations were 2-5X greater in woody landscape elements than in grasslands, and nutrient concentrations increased linearly with time following woody plant invasion in all but the slowest-cycling physical fractions. Plant-available P and organic P increased dramatically with time following encroachment. Changes in P availability were more pronounced in drainages and playas than in upland clusters and groves. Analysis of the bacterial and fungal communities demonstrated that microbial communities in grasslands differ at both phylum and genus level from the flora of the wooded landscape elements. This study demonstrates that woody encroachment strongly influences the distribution and availability of soil P and indicates that nutrient cycles in the soil are closely linked and similarly affected by increased woody plant abundance. Microbial communities under woody species differ in composition from those of the grasslands, and are likely contributing to the observed changes in nutrient availability. Since N and P are generally the most limiting nutrients in terrestrial ecosystems, increased stores of P are likely to alter rates of microbial processes, plant-microbe and plant-plant interactions, and successional dynamics in this ecosystem and similar landscapes around the world.

Habitat loss, habitat fragmentation and species invasions have been recognized as three of the leading threats to biodiversity. I examined the effects of habitat loss and fragmentation on native and invasive plants in central Texas. During the last century, the density and abundance of woody plants has been increasing in the savannas of eastern Edwards Plateau. This process, known as woody plant encroachment, not only reduces the amount of open herbaceous habitat but also fragments that habitat creating smaller and more isolated patches. In three studies, I investigated the consequences of this habitat loss and fragmentation for plants which do not occur under the cover of woody plants including native grasses and forbs and the invasive Eurasian bunchgrass, Bothriochloa ischaemum (King Ranch Bluestem). In the first study, I show that woody plant encroachment reduces native herbaceous species richness (the number of species in a given area). Using a collection of historical aerial photographs, I demonstrate that current native herbaceous species richness was most strongly related to recent habitat amount, but to the degree of habitat fragmentation at least 50 years ago. In a second study, I show that the presence of B. ischaemum was negatively related to the degree of fragmentation in the surrounding landscape. Finally, I found that B. ischaemum had higher rates of germination and growth in experimental plots where the species commonly lost with woody plant encroachment were removed than in unmanipulated control plots. Together, this work suggests that woody plant encroachment is directly slowing the spread of an invasive species while indirectly facilitating its establishment. / text

Woody plant encroachment (WPE) into historic grasslands not only alters ecosystem structure but also yields a mosaic of vegetative growth-forms that differ in their inherent physiological capacities and physical attributes. C₃ plants tend to have a relatively broad range of temperature function but at the expensive of a lower optimum rate of photosynthesis. In contrast, C₄ grasses have a greater capacity for maximum uptake but across a relatively narrow range of temperatures. In considering which of these functional groups will outcompete the other within these regions undergoing WPE, one must account not only for these leaf physiological traits, but also the growth form induced differences in rooting depth, and therefore, potential access to deeper subsurface water. Laid upon these competitive interactions is an ever-changing environment, which for the semiarid southwestern US is predicted to become progressively warmer and characterized by highly variable precipitation with longer interstorm periods. In addition to aboveground changes in CO₂ assimilation, WPE influences soil nutrient, water, and carbon cycling. The objectives of this dissertation were to quantify: (1) the influence that temperature and available soil moisture have on regulating soil respiratory efflux within the microhabitats that results from WPE to estimate the influence this vegetative change will have on ecosystem CO₂ efflux; (2) the sensitivity of CO₂ uptake within grassland and woodland ecosystems to temperature and precipitation input in an effort to characterize how WPE might influence regional carbon and water balance; and (3) the role access to stable groundwater has in regulating the temperature sensitivity of ecosystems and their component fluxes. Major findings and contributions of this research include illustrating seasonal patterns of soil respiration within the microhabitats that result from WPE, such that an analysis of the relative contributions of these different components could be made. We found that soil respiration was not only consistently greater under mesquites, but that the relative contributions of these microhabitats varied significantly throughout the year, the duration of soil respiration after each rain was habitat-specific, and that the relationship between soil respiration and temperature followed a hysteretic pattern rather than a linear function (Appendix A). We found that a woodland ecosystem demonstrated a lower temperature sensitivity than a grassland across all seasonal periods of varying soil moisture availability, and that by maintaining physiological function across a wider range of temperatures throughout periods of limited precipitation, C₃ mesquites were acquiring large amounts of carbon while C₄ grasses were limited to functioning within a narrower range of temperatures (Appendix B). Finally, we found that having a connectivity to stable groundwater decoupled leaf and ecosystem scale temperature sensitivities relative to comparable sites lacking such access. Access to groundwater not only resulted in the temperature sensitivity of a riparian shrubland being nearly half that of the upland site throughout all seasonal periods, but also actual rates of net ecosystem productivity and leaf level rates of photosynthesis being dramatically enhanced (Appendix C).

Doctor of Philosophy / Department of Biology / John M. Blair / Juniperus virginiana, an important woody plant invader in the U.S. Central Plains, has increased considerably in density and cover in large areas previously dominated by tallgrass prairie. Change in the phenology and nitrogen use efficiency of the dominant plant communities as J. virginiana replaces native prairies may lead to increased plant productivity and biomass accumulation, but may also alter the microclimate and litter quality that affect soil microbial communities responsible for key soil processes. I have focused my investigations on changes in key soil processes that could lead to differences in soil N availability, as well as changes in ecosystem C and N pools and fluxes as J. virginiana expands into native grasslands. Juniperus virginiana forest soils exhibit greater cumulative annual net N mineralization (11.52 ± 0.38 µg N g¯1 soil y¯1) compared to prairie soils (7.90 ± 0.26 µg N g¯1 soil y¯1) (F = 60.67, P = 0.016), yet slightly reduced potential soil C flux. Examination of internal soil N cycling revealed that both J. virginiana and prairie soils minimize potential soil N losses, by rapid microbial immobilization of inorganic N, and constraining nitrification via substrate limitation or environmental constraints. Leaf-level photosynthetic nitrogen use efficiency (NUE) was over a magnitude higher in the dominant grass, Andropogon gerardii, but high annual ecosystem-level NUE and greater soil N availability may contribute to the higher productivity and rapid accrual of C in newly established J. virginiana forests. Increased plant productivity and elimination of fire in J. virginiana forests have allowed at least 80,000 kg ha-1 increase in ecosystem C storage in about half a century. Soil organic C, an important long-term sink, has also increased significantly in J. virginiana forests, with approximately 34% replacement of C4 grass-derived soil C with new C from trees in the A-horizon. The observed high productivity of J. virginiana and increased N availability necessary to support continued plant biomass accumulation are possible because of substantial (~ 44%) increase in ecosystem N in measured pools, which is a likely a result of reduced volatilization of N from biomass burning, possible increased exogenous N inputs, and/or N translocation from deeper soil horizons. Reduced fire return intervals in prairie provide an opportunity for J. virginiana to establish and facilitate N accrual, which may allow this species to accelerate is own establishment through creating conditions of increased N availability and efficient utilization of N.

The conversion of grasslands to woodlands is an ecosystem transformation that threatens grassland biodiversity, the provision of important ecosystem services, and the sustainability of rural livelihoods. A global phenomenon, woody plant encroachment (WPE) has been particularly problematic in the Southern Great Plains of the United States where the actions of private landowners are integral to sustaining grasslands. Increased diversity in landowners’ motivations for owning land have shifted the social landscape of rural areas necessitating a better understanding of landowners’ perspectives about WPE and their subsequent management actions. Towards this purpose, I employed a mail survey to private landowners in the Edwards Plateau of Texas, Central Great Plains of Oklahoma, and Flint Hills of Kansas to investigate landowner perceptions and management responses to WPE. First, I assessed landowners’ acceptance of WPE as a function of how they relate to their land (i.e., sense of place), their beliefs about the positive and negative consequences of woody plants, and their perceived threat of grassland conversion. Then, I examined the drivers of landowners’ goal intentions to manage woody plants and their current use of five adaptive management practices that prevent WPE. My results demonstrate that landowners vary in their sensitivity to WPE based on how they feel connected to their land. This was true even though most landowners had low acceptance thresholds for WPE, believed it led to numerous negative outcomes, and perceived it as increasingly threatening at greater levels of encroachment. Most landowners wanted to control or remove woody plants and were actively engaged in management practices to do so. These findings address uncertainties about landowners’ acceptance of WPE and grassland conservation actions and provide broad implications for how people perceive and respond to ecosystem transformation. / Master of Science / Around the world, grasslands are converting to tree and shrub woodlands at an unprecedented rate. This transformation profoundly reduces habitat available for grassland plants and animals and diminishes many ecosystem services that people and rural communities rely on. This loss of grasslands has been especially far-reaching throughout the Southern Great Plains of the United States. Because most of this region is privately owned, the management actions of landowners play a crucial role in preventing or allowing this conversion to continue. Recent shifts in land ownership motivations expanding beyond traditional agricultural production have created increased uncertainty about how private landowners view and react to this change. To investigate how landowners perceive and respond to this woody plant encroachment (WPE) phenomenon, I conducted a mail survey of landowners in the Edwards Plateau of Texas, the Central Great Plains of Oklahoma, and the Flint Hills of Kansas. Using sense of place, landowners’ beliefs about the potential positive and negative consequences of woody plants, and their perceptions of how threatening grassland conversion is, I assessed the thresholds at which landowners’ do or do not accept WPE. Then, I examined how acceptance of WPE relates to landowners’ management goals and current use of management practices to control or reduce woody plants. I found that most landowners believed that woody plants had many negative consequences and perceived increasing levels of threat at greater levels of encroachment. This related to low levels of acceptance for woody plants in grasslands. However, landowners’ threat perceptions and acceptance of WPE varied based on their sense of place. Finally, most landowners wanted to control or remove woody plants and were actively engaged in management practices to do so. My results provide critical information regarding how current landowners’ view and respond to grassland conversion and offer broad implications for how people perceive and respond to large-scale environmental change.

In the southern Great Plains, the encroachment of grassland ecosystems by mesquite (Prosopis glandulosa), is widespread, and prescribed fire is commonly used in its control. Despite this, substantial quantitative information concerning their influences on the community composition, functional dynamics, and soil organic carbon (SOC) storage potential of grassland ecosystems is lacking. The objectives of this study were to: a) quantify the effects of seasonal prescribed fire treatments and mesquite encroachment on aboveground net primary productivity (ANPP) and herbaceous community composition; b) characterize SOC pool sizes, turnover, and storage potential relative to vegetation type and fire treatment; c) evaluate the structure and diversity of soil microbial communities relative to vegetation type; and d) characterize the functional diversity of these same microbes using the GeoChip functional gene microarray. Repeated winter and summer fires led to increased ANPP rates (average, 434 and 313 g m-2 y-1, respectively), relative to unburned controls (average, 238 g m-2 y-1), altered herbaceous community composition, and increased the storage of resistant forms of SOC, but did not affect overall SOC storage. Herbaceous ANPP rates did not differ significantly as a result of mesquite encroachment, but herbaceous community composition and SOC storage did. Mesquite soils contained significantly more total, slow-turnover, and resistant forms of SOC than those that occurred beneath C3 or C4 grasses. Similarity among the soil bacterial and fungal communities associated with the major vegetation types in this system was low to moderate. Significant differences were detected among soil fungi, with the mesquite-associated fungi harboring significant differences in community structure relative to the fungal communities associated with each of the other vegetation types examined. Despite this result, few significant differences were detected with respect to the functional diversity of these communities, suggesting either a high degree of functional redundancy, or that the functional differences harbored by these communities are beyond the scope of the GeoChip. The results of this study demonstrate that both fire and mesquite encroachment have the potential to alter ecosystem components and processes significantly, providing new insight regarding the effects of these widespread land use and land cover changes on ecosystem structure and function.

Woody plant encroachment, that is, a substantial increase in the abundance of woody plants in a grassland or savanna, occurs in many parts of the world. It often has large effects on plant and animal populations and communities and on ecosystem properties and processes. However, little is known about the dynamics of woody plant encroachment and how these are affected by soils, by topography, and by the spatial pattern of the vegetation. Encroachment in turn can affect the spatial pattern of the vegetation. Using data from historical aerial photographs, I measured changes in woody plant cover and constructed, parameterized and compared a set of dynamic models of woody plant encroachment in central Texas savannas. These models predicted final woody cover from initial woody cover and the initial spatial configuration of woody plants. Then I incorporated soil and topography into these models to determine their effects. Finally, I examined the effects of encroachment on the spatial pattern of the vegetation. Incorporating negative density dependence in our models improved their fit, demonstrating that encroachment is density-dependent. A function that predicted the formation of new woody patches from a density-independent seed supply also improved the models' performance. The improvement in the models that resulted from incorporating the total length of woody-herbaceous edges confirmed that encroachment in this system occurs in part by the outward expansion of woody patches. The spatial pattern of the vegetation changed during woody plant encroachment. Spatial pattern (measured as degree of fragmentation) often had a non-linear relationship with cover. Furthermore, the spatial heterogeneity in fragmentation, that is, plot-to-plot variation in the degree of fragmentation, also changed during encroachment. Topography and soil type had, in general, little effect the dynamics of woody plant encroachment. Therefore, a relatively simple model of woody plant encroachment provided good predictions of woody cover at the end of the time periods. Other systems experiencing woody plant encroachment, forest succession, or invasion by non-native plants could be modeled using the same approach. / text

abstract: Woody plant encroachment is a worldwide phenomenon linked to water availability in semiarid systems. Nevertheless, the implications of woody plant encroachment on the hydrologic cycle are poorly understood, especially at the catchment scale. This study takes place in a pair of small semiarid rangeland undergoing the encroachment of Prosopis velutina Woot., or velvet mesquite tree. The similarly-sized basins are in close proximity, leading to equivalent meteorological and soil conditions. One basin was treated for mesquite in 1974, while the other represents the encroachment process. A sensor network was installed to measure ecohydrological states and fluxes, including precipitation, runoff, soil moisture and evapotranspiration. Observations from June 1, 2011 through September 30, 2012 are presented to describe the seasonality and spatial variability of ecohydrological conditions during the North American Monsoon (NAM). Runoff observations are linked to historical changes in runoff production in each watershed. Observations indicate that the mesquite-treated basin generates more runoff pulses and greater runoff volume for small rainfall events, while the mesquite-encroached basin generates more runoff volume for large rainfall events. A distributed hydrologic model is applied to both basins to investigate the runoff threshold processes experienced during the NAM. Vegetation in the two basins is classified into grass, mesquite, or bare soil using high-resolution imagery. Model predictions are used to investigate the vegetation controls on soil moisture, evapotranspiration, and runoff generation. The distributed model shows that grass and mesquite sites retain the highest levels of soil moisture. The model also captures the runoff generation differences between the two watersheds that have been observed over the past decade. Generally, grass sites in the mesquite-treated basin have less plant interception and evapotranspiration, leading to higher soil moisture that supports greater runoff for small rainfall events. For large rainfall events, the mesquite-encroached basin produces greater runoff due to its higher fraction of bare soil. The results of this study show that a distributed hydrologic model can be used to explain runoff threshold processes linked to woody plant encroachment at the catchment-scale and provides useful interpretations for rangeland management in semiarid areas. / Dissertation/Thesis / M.S. Civil and Environmental Engineering 2013

Thesis (M.Sc. Agriculture (Soil Science)) -- University of Limpopo, 2021 / Woody plant encroachment results in the degradation of grasslands. It is defined here as the increase in density, cover and biomass of woody plants into formerly open grasslands, reducing grassland productivity. Globally, many arid and semi-arid savanna grasslands are affected by this land cover transformation which changes the vegetation structure by altering the ratio of woody plants relative to grass species and influences soil hydrology. In the existing literature there is limited information on the effects of woody plant encroachment on soil physical and hydrological properties, especially in savanna grasslands. This study quantified and compared the soil physical and hydrological properties in the topsoil and subsoil of open and woody plant encroached grassland sites located on two contrasting soil forms, namely Bainsvlei and Rensburg. To achieve this objective, the two soils were sampled at various depth intervals from dug soil profiles at both sites at Towoomba Research Station in Bela Bela, Limpopo Province, South Africa. Soil physical properties including bulk density, porosity and aggregate stability as well as hydrological properties (water retention and hydraulic conductivity) were determined from collected samples. Compared to open grassland, soil bulk density was 11% and 10% greater in the topsoil and subsoil, while porosity was respectively 6% and 9% lower in the topsoil and subsoil of woody plant encroached grassland for Rensburg soils. In Bainsvlei soil, there was a minimal increase and decrease in the soil bulk density and porosity, respectively. Soil aggregate stability increased by 38% in the subsoil of woody plant encroached grasslands in Rensburg soil, due to increasing clay content with depth. In Bainsvlei soil, the soil aggregate stability was 9% and 13% lower in the topsoil and subsoil of the woody plant encroached grasslands compared to open grassland. Furthermore, the results revealed that in both soils, there was lower soil water retention and hydraulic conductivity in the topsoil and subsoil layers of woody plant encroached grassland than in open grasslands. There were no significant differences observed for soil hydraulic conductivity in the Bainsvlei and Rensburg topsoil. The subsoil hydraulic conductivity decreased by 24% in Bainsvlei and 44% in Rensburg soils in the woody plant encroached grassland. The soil water retention (SWR) decreased with an increase in woody plants. Specifically, there was 25% and 42% decrease in SWR with woody plant encroachment in the topsoil and subsoil of Bainsvlei soil, respectively. The same trend was observed in the Rensburg soils with 50% and 19% decrease in SWR in the topsoil and subsoil, respectively. Overall, the results revealed that soil type and depth influenced soil physical and hydrological properties in the studied woody plant encroached savanna grassland. As such, interventions aimed at controlling woody plant encroachment need to factor in soil type and depth in the development of management practices tailored to improve the soil hydrology of savanna grasslands

Woody plant encroachment is one of the major problems worldwide because it affects negatively the herbaceous layer, which provide forage for livestock production. However, the role of ruminants particularly browsers in the dispersal of woody plant seeds still remains a concern for farmers interested in grass production. Seedpods of various woody plant species constitute a crucial part of the diet of herbivores during the dry season because of their high nutritional quality compared to herbaceous material. The interaction of associated diet quality, seed characteristics and animal species among other factors play a pivotal part in the success of livestock faecal seeds dispersion. Furthermore, dispersed seeds that successfully grow into mature woody plants become an important source of protein for herbivores. The use of woody plants as a source forage is known to be limited by plant secondary metabolites (PSMs) such as condensed tannins. The objectives of this study were to determine 1) the effects of condensed tannins and crude protein of Vachellia tortilis and Dichrostachys cinerea pods in seed recovery and germination fed to goats, and 2) the effects of diet mixing on the feed intake of plant species by goats. In the first experiment, a total of 12 female indigenous goats and 12 female Pedi sheep were utilised in this study, with the average body weights of 29.50 kg ± 1.60 (S.E) and 28.70 kg ± 1.60, respectively. Twelve goats were grouped into two groups of six goats per group, one group was fed D. cinerea pods and the other group was fed V. tortilis pods. The group of 12 sheep were divided similarly, the one group was fed D. cinerea pods and the other group was fed V. tortilis pods. Each animal was given V. tortilis and D. cinerea pods at 2.50% of their body weight. All animals were allowed to consume D. cinerea or V. tortilis pods within 24 h, after which the remaining pods were collected and weighed. Faecal collection commenced immediately after the 24 h pods feeding and was carried on until no seeds were discovered in faeces. All faeces extracted from sheep and goats were collected daily in the morning from the faecal bags. In the second experiment, a total of 24 indigenous goats with average body weight of 26.6 kg ± 0.51 were utilised. Goats were arbitrarily selected and grouped into four groups of six goats per group (goats were placed individually in 2 m2 pens). Each group was fed one of the following diets: diet one - Searsia lancea, diet two - S. pyroides, diet three - Euclea crispa and diet four - was a combination of the three plant species (Searsia lancea, S. pyroides and Euclea crispa). Searsia lancea, S. pyroides and E. crispa branches were collected every morning prior to feeding, and were weighed before offering the animals. Refusals were gathered and weighed, and intake was calculated as distinction between weight in and refusals. Plant species foliage were analysed for crude protein, condensed tannin, acid detergent lignin, acid detergent fibre and neutral detergent fibre. During the first experiment, the cumulative percentage seed recovery of V. tortilis from goats (46.00 % ± 1.90) and sheep (52.00 % ± 2.93) was significantly higher than D. cinerea from goats (13 % ± 1.47) and sheep (24.00 % ± 1.16). Germination percentage of D. cinerea seeds that passed through the gastro-intestinal tract of goats (33.12 % ± 2.94) and sheep (36.00 % ± 2.68) was significantly higher than V. tortilis seeds that passed through the gastro-intestinal tract of goats (28.98 % ± 2.68) and sheep (23.04 % ± 2.81). Average D. cinerea (34.56 % ± 1.99) and V. tortilis (26.02 %± 2.10) seeds that went through the gastro-intestinal of goats and sheep had a significantly higher germination rate than the control (i.e. no passage through the gut; D. cinerea = 2.31 % ± 1.55, V. tortilis = 5.07 % ± 2.68). The high mean cumulative percentage seed recovery of V. tortilis (18.80 %) may be attributed to the relatively higher crude protein than D. cinerea (12.20 %). This may encourage animal seed dispersal and germination of woody plant species with relatively high crude protein content. In the second experiment, Searsia lancea contained 8.50 % CP, 21.46 % acid detergent fibre (ADF), 12.50 % ADL and 39.37 % NDF. Searsia pyroides had 9.03 % CP, 27.07 % ADF, 10.89 % ADL and 40.30 % NDF. Euclea crispa had 6.19 % CP, 26.20 % ADF, 16.63 % ADL and 30.02 % NDF. Mixed diet (combination of the three plant species) had 8.96 % CP, 23.72 % ADF, 11.13 % ADL and 38.28 % NDF. Searsia lancea had 2.70 % of CTs while S. pyroides had 5.20 % CT, E. crispa had 6.44 % CT and mixed diet had 7.20 % CT. The mean dry matter intake varied significantly among dietary groups (P < 0.001). Similarly, goats offered a mixed diet consumed more CTs (P < 0.01) than those offered individual forage species. The high mean cumulative percentage seed recovery of V. tortilis may be attributed to the higher crude protein of V. tortilis (18.80 %) than D. cinerea (12.20 %). Higher passage rate may encourage animal seed dispersal and germination of plant species. The results from experiment two support the postulation that animals foraging in mixed diet systems consume more PSMs and achieve higher dry matter intake than animals confined to monocultures or single species feeding systems. Given that woody plant encroachment is already reducing farm-grazing capacities in African savannas and this problem is predicted to double by 2050, strategies that improve herbivore ability to consume woody plants will increase forage availability and inform bush control programmes and policies. Moreover, the concomitant increase in CTs by goats exposed to diets with diverse species also has positive implications for animal / Agriculture, Animal Health and Human Ecology / M. Sc. (Agriculture)