Doctor of Philosophy / Department of Biology / Brett K. Sandercock / Habitat heterogeneity is a key driver of biodiversity in many ecosystems. In native ecosystems, habitat heterogeneity can arise from multiple drivers including nutrients, topoedaphic conditions, and ecological disturbance. Historically, the prairies of North America existed as a heterogeneous mosaic of habitat conditions created by the interaction of fire and grazing by native ungulates. The focus of many grazing systems has been to minimize disturbance caused by grazing by promoting uniform distributions of grazing animals across management units. Patch-burn grazing is an alternative rangeland management practice that has been proposed to restore historical patch dynamics and biodiversity to rangelands by simulating historical disturbance processes. In my dissertation research, I tested the hypothesis that patch- burn grazing restores habitat heterogeneity to rangelands, and that the resulting habitat heterogeneity can promote biodiversity of native wildlife. I focus on responses of small mammals and coyotes to patch-burn grazing to gain a better understanding of wildlife responses to rangeland management, and because grassland mammals are an ecologically important group. My 3.5-year field study of habitat and small mammal responses to rangeland management showed that: 1) patch-burn grazing created greater heterogeneity in vegetative structure and composition of plant functional groups than in positive and negative controls; 2) habitat heterogeneity created by the interaction of fire and grazing increased small mammal richness and diversity compared to a negative control managed for uniform grazing distributions; 3) the interaction of fire and grazing structured small mammal communities in tallgrass prairie; and 4) population dynamic responses of small mammals to fire and grazing disturbance were species- specific. My 3-year study of coyote survival and resource selection revealed that: 1) rangeland management influences resource selection by coyotes in seasons when they depend on small mammal prey, but not during other seasons; and 2) anthropogenic sources of mortality are important for coyotes at a protected area, even in the absence of harvest. My field results show that restoring the drivers of historical patch dynamics to managed rangelands and publicly held grasslands that are not currently grazed could have profound effects on biodiversity conservation in North America, while continuing to provide ecosystem services to society.

Agriculture has been reported to contribute a significant amount of greenhouse gases to the atmosphere among other anthropogenic activities. With still more than 870 million people in the world suffering from under-nutrition and a growing global food demand, it is relevant to study ways for mitigating the environmental impact of food production. The objective of this work was to identify gaps in the knowledge regarding the main factors affecting greenhouse gas (GHG) emissions from beef farming systems, to reduce the uncertainty on carbon footprint predictions, and to study the relative importance of mitigation options at the system level. A lack of information in the literature was identified regarding the quantification of the relevant animal characteristics of extensive beef systems that can impact on methane (CH4) outputs. In a meta-analysis study, it was observed that the combination of physiological stage and type of diet improved the accuracy of CH4 emission rate predictions. Furthermore, when applied to a system analysis, improved equations to predict CH4 from ruminants under different physiological stages and diet types reduced the uncertainty of whole-farm enteric CH4 predictions by up to 7% over a year. In a modelling study, it was demonstrated that variations in grazing behaviour and grazing choice have a potentially large impact upon CH4 emissions, which are not normally mentioned within carbon budget calculations at either local or national scale. Methane estimations were highly sensitive to changes in quality of the diet, highlighting the importance of considering animal selectivity on carbon budgets of heterogeneous grasslands. Part of the difficulties on collecting reliable information from grazing cattle is due to some limitations of available techniques to perform CH4 emission measurements. Thus, the potential use of a Laser Methane Detector (LMD) for remote sensing of CH4 emissions from ruminants was evaluated. A data analysis method was developed for the LMD outputs. The use of a novel technique to assess CH4 production from ruminants showed very good correlations with independent measurements in respiration chambers. Moreover, the use of this highly sensitive technique demonstrates that there is more variability associated with the pattern of CH4 emissions which cannot be explained by the feed nutritional value. Lastly, previous findings were included in a deterministic model to simulate alternative management options applied to upland beef farming systems. The success of the suggested management technologies to mitigate GHG emissions depends on the characteristics of the farms and management previously adopted. Systems with high proportion of their land unsuitable for cropping but with an efficient use of land had low and more certain GHG emissions, high human-edible returns, and small opportunities to further reduce their carbon footprint per unit of product without affecting food production, potential biodiversity conservation and the livelihood of the region. Altogether, this work helps to reduce the uncertainty of GHG predictions from beef farming systems and highlights the essential role of studies with a holistic approach to issues related to climate change that encompass the analysis of a large range of situations and management alternatives.

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Thesis (MSc)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: I investigated the impacts of the invasive wattle species (Acacia mearnsii, A. dealbata, A. decurrens), on the ecological function and productivity of rangelands in South Africa and their ability to sustain livestock production. More specifically, this study set out to: (1) assess grazing areas at a national scale; (2) identify evidence of progressive impacts of these species on livestock production across a selection of magisterial districts; (3) determine the effects of A. mearnsii density on growth form dominance of indigenous plant species, and highlight how this translates into impacts in forage quality and quantity; (4) determine the effects of A. mearnsii invasion on soil resources and conditions (key determinates of ecological function) required to support grazing production; and finally (5) determine to effects that clearing operations have had on the provision of grazing resources. My results indicate that the level of overlap between wattle invasion and areas with a high grazing potential, at the national scale, is relatively low (2.9%). However, at a magisterial district scale, areas with a high grazing potential have a moderate (approximately 35.1% high grazing potential area under wattle invasion for highest invaded district) level of invasion. Scattered and moderate levels of invasion currently dominate or characterise these species invasion patterns. Both light and dense A. mearnsii invasions reduce grazing capacity. Dense invasion had the most severe effect reducing grazing capacity from 2 ha, required to support one large stock unit, to 8 ha. The clearing of this species was found to improve grazing capacity within 5 years. The reduction in grazing capacity following invasion was largely due to reduced basal cover and herbaceous biomass. The clearing of invaded sites was found to increase both basal cover and herbaceous biomass to pre-invasion levels. Acacia mearnsii invasion was found to affect certain soil properties. Under an A. mearnsii canopy, plant litter, carbon content of the soil and nitrogen concentrations were all found to be elevated. Overall, this study demonstrates that Acacia invasions have a significant effect on grazing resources at a local or site scale, and that for the eastern region of South Africa, these effects are ultimately scaled up to a district level. This is likely to have important financial and human well-being repercussions. However clearing and restoration programs are clearly able to reverse or mitigate these effects. This study therefore lends support to or complements other ecosystem service provision arguments for clearing alien invasive species. / AFRIKAANSE OPSOMMING: Die impak van die indringer- wattelbome spesies ( A. mearnsii , A. dealbata , A. decurrens) , op die ekologiese funksie en produktiwiteit van weivelde in Suid-Afrika en hul vermoë om vee produksie te onderhou was ondersoek. Meer spesifiek, is hierdie studie uiteengesit om: (1) ondersoek in te stel na die mate van oorvleueling tussen indringing deur wattelbome spesies en belangrike weidingsgebiede op 'n nasionale skaal; (2) bewyse te verskaf van die progressiewe impak van hierdie spesies op vee-produksie oor 'n seleksie van politieke distrikte, (3) die gevolge van A. mearnsii digtheid op groei vorm en oorheersing van inheemse plante te wys, en hoe dit voer kwaliteit en kwantiteit affekteer; (4) die gevolge van A. mearnsii indringing op grond hulpbronne sowel as die kondisie (sleutel bepalende faktore van ekologiese funksie) wat benodig word om weiding produksie te ondersteun vas te stel, en uiteindelik (5) om die gevolge van skoonmaak bedrywighede op die voorsiening van weiding hulpbronne te bepaaal. My resultate dui daarop dat die vlak van oorvleueling tussen die wattelbome indringing en gebiede met 'n hoë weiding potensiaal, by die nasionale skaal, relatief laag is (2’9 %). Maar op 'n landdrosdistrik skaal, het gebiede met 'n hoë weiding potensiaal 'n matige (ongeveer 35.1 % weiding potensiaal area onder wattelbome indringing vir die hoogste binnegevalde distrik ) vlak van indringing. Verspreide en matige vlakke van indringing oorheers tans of kenmerk hierdie spesies se indringings patrone. Beide lig en digte A. mearnsii besettings verminder dravermoë. Digte inval het die mees ernstige uitwerking en verminder dravermoë van 2 ha, wat nodig is om een grootvee-eenheid te ondersteun na 8 ha. Bevindinge wys dat die verwydering van hierdie spesies weiding kapasiteit kan verbeter binne 'n tydperk van 5 jaar. Die afname in dravermoë na indringing is grootliks te wyte aan die afname in basale bedekking en kruidagtige biomassa. Na die skoonmaak van indringer persele is bevind dat beide basale bedekking en biomassa weer verhoog tot voor- indringings vlakke. Acacia mearnsii indringing beïnvloed sekere grondeienskappe. Onder A. mearnsii blaredak, is alle blaarafval, koolstof-inhoud van die grond en stikstof konsentrasies verhef. In die algeheel, toon hierdie studie dat die Acacia indringings 'n beduidende uitwerking op weidings hulpbronne by 'n plaaslike of perseel skaal het, en vir die oostelike streek van Suid-Afrika, is hierdie effekte ook uiteindelik vertaalbaar tot 'n distrik vlak. Dit sal waarskynlik belangrike finansiële en menslike welsyn gevolge hê, maar die skoonmaak en herstel programme is duidelik daartoe instaat om die gevolge te keer of te verminder. Hierdie studie leen dus ondersteuning aan of komplimenteer ander ekostelsel dienslewering argumente (soos dié van die Werk vir Water program) vir die verwydering van indringerspesies.

The plant communities of the Long Mynd plateau are the culmination of over 3000 years of human intervention that largely deforested the uplands, and subsequently maintained the generally treeless heath and grassland communities now extant. The capacity of these communities to respond to directional change is well known, indeed the traditional mode of heathland management, burning, depends on the regenerative capacity of the target species, generally heather (Calluna vulgaris), for its success. However, changes in post WW2 stocking practice; the loss of ponies followed by an increase in the numbers of sheep and a change to them being overwintered on the hill, led to excessive grazing and damage to the heath. This coincided with the spread over the hill by bracken (Pteridium aquilinum) and other changes in the distribution and nature of the vegetation. A sequence of vegetation surveys made by various individuals and organisations over the past 75 years or so has been analysed in an attempt to delineate spatial and temporal changes in the vegetation. This demonstrated the need for a standardised survey methodology to allow consistent monitoring. The analysis showed that bracken had been infiltrating most of the communities from its origins outside the lower limits of the Common as well as from some of the valley sides. Within the last decade, this expansion has apparently been contained in line with the current management plan for control. A survey of 730 quadrats in some 30 stands was made to characterise the variation of the vegetation on the plateau, and to relate it to some of the associated environmental factors. Classification, unconstrained ordination and ordination constrained by the abiotic environmental variables, showed that, a) the strongest trend in the vegetation distinguished water-flushed communities, b) non-wetland communities differentiate between heathland and grassland, c) this trend can be only partly be attributed to the measured abiotic environmental variables, d) the amount of pure Pteridietum [U20] is limited, although much of the heathland and grassland has bracken within it. There are indications that invasion by bracken often correlates with a loss of dominance of Calluna in favour of Deschampsia flexuosa and Vaccinium myrtillus. Difficulties in associating these trends with measured abiotic variables suggests, other factors probably management processes, are critical in driving this trend. Distribution of ‘heathland’ bryophytes was found to be associated more with the structure of their ‘host’ vascular communities rather than with abiotic factors. Finally, this investigation considers the practical implications with regard to the future encouragement of heather and the control of bracken. Cutting rather than burning appears to be the ecologically most suitable method for heather regeneration and bracken control.

Cattle grazing patterns on Lehmann lovegrass rangelands often create heavily-grazed (HG) areas surrounded by lightly-grazed (LG) or ungrazed patches. The purpose of this study was to characterize the forage resource and ingestive behavior of cattle grazing Lehmann lovegrass plants in both LG and HG patches on the Santa Rita Experimental Range. The general hypothesis was that residual bunchgrass vegetation resulted in sward characteristics which physically inhibited the efficiency of cattle grazing by increasing the manipulative activity per harvested bite. The overall heights of residual stems and green tillers averaged 70 and 49 cm for LG, 8 and 9 cm for HG patches. Biomass of total standing crop (SC), residual vegetation (RV) and green herbage (GH) averaged 4159, 3395 and 764 kg/ha for LG, 345, 185 and 160 kg/ha for HG patches, respectively. Bulk density of SC, RV and GH of LG and HG patches averaged 58, 48, and 19 [(gm/cm³) X 10⁻⁵] for LG, 38, 23 and 20 [(gm/cm³) X 10⁻⁵] for HG patches, respectively. The ratio of green herbage to residual vegetation averaged 0.22 for LG and 0.86 for HG patches. The different sward structure of LG and HG patches affected the ingestive behavior of the grazing cows. Overall handling time for each grazing bite averaged 1.5 and 1.2 sec/bite in LG and HG patches, respectively. Cows employed different foraging tactics in response to the dynamic changes of the sward conditions. Cows employed top biting extensively to harvest the seed-head and other green herbage at the top of the sward surface. As the height of residual stems increased and dominated the upper strata of the sward, side and low biting were mainly used by the cows as grazing methods to bite the plants to reduce the manipulative activity. Avoidance of LG patches or preference for HG patches was related to the sward structure and largely shaped by the build-up of residual vegetation. Removal of residual vegetation through fire, mowing or heavy utilization for short periods late in the growing season to allow for more accessible green herbage could improve both animal and pasture utilization of the range resource.

The abundance of Lehmann lovegrass (Eragrostis lehmanniana, Nees.) on Arizona ranges prompted this research to evaluate the animal production and winter palatability of this species. Sulfurcoated urea (SCU) was applied to desert grassland range to minimize the risks of fertilization that are brought about by drought and unseasonable rainfall. Tebuthiuron (TEB) was also applied to pasture to document any beneficial effects on forage production and palatability. During the 2 years of this project, two experiments were conducted. A preference study examined forage and animal responses to one-time applications of fertilizer or herbicide treatments and combinations. The second experiment compared the potentials for improved animal production from SCU and SCU + TEB with untreated range. Results indicated that the fertilizer and combination treatments produced more forage than the untreated plots. Tebuthiuron improved forage production by 200 percent in the first year but reduced yields in the second year. The high rate of TEB was detrimental in both years. Forage utilization was improved by all treatments. Nitrogen increased the crude protein content of the grasses proportional to the rate applied. Tebuthiuron also enhanced the protein content of the forage. Total available carbohydrates were lowest in TEB treated forages, but digestibility was increased. Only the 31 lb SCU appeared to be economically profitable. However, excellent carry-over by the 60 SCU treatment suggested profitability after 3 years. Results from the grazing study indicated that production was significantly improved by both treatments. Although forage production was similar for the treatments, the SCU-only pasture provided superior animal gains and greater carrying capacity than the SCU + TEB pasture, even though forage digestibility and apparent disappearance were greater on the combination pasture. The difference in animal performance was attributed to low levels of carbohydrates available from the combination pasture. Treatment of Lehmann lovegrass/blue grama range with 40 SCU/acre appeared profitable for raising growing beef animals, but not for maintenance of a breeding cow herd.

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