Elvis Mubamu Makady

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Elvis Mubamu Makady Large herbivore stocking rate effects on plant palatability, forage preference and soil properties in an Alluvium Fynbos-Renosterveld mosaic By Elvis Mubamu Makady Thesis presented in partial fulfilment of the requirements for the degree of Master of Science in conservation ecology at Stellenbosch University Department of Conservation Ecology and Entomology Faculty of AgriSciences Supervisor: Dr. Cornelia B. Krug Co-supervisor: Professor Karen. J. Esler Date: March 2009 DECLARATION By submitting this thesis electronically, I declare that the entirety of the work contained therein is my own, original work, that I am the owner of the copyright thereof (unless to the extent explicitly otherwise stated) and that I have not previously in its entirety or in part submitted it for obtaining any qualification. Signature:…………………….. Date: 03 March 2009 Copyright © 2009 Stellenbosch University All rights reserved II ABSTRACT An understanding of the interactions of herbivores, plant nutrients and soil properties is crucial for grazing management. Of particular interest are plant palatability and herbivore grazing preferences. These aspects, the focus of this thesis, were studied in a Swartland Shale Renosterveld and Swartland Alluvium Fynbos mosaic in the Western Cape of South Africa. This vegetation is classified as critically endangered due to the transformation of its landscape in favour of agricultural production and urban settlement. Pre-colonial herbivore vertebrates were hunted out by 1700, leading to a sparse knowledge of the early dynamics of the vegetation. However, currently game farmers and landowners are re-introducing selected species into the area. Thus, managers require knowledge on how to implement practical guidelines for best-practice grazing management. This study investigates, firstly, the relationship between plants and animals. This included seasonal assessment of seed germination from dung of bontebok and eland; the effect of stocking rates on plant palatability (crude protein, crude fibre, crude fat, moisture, tannin and mineral) of herbage was studied with the aim to understand how palatability influences herbivore forage decisions. Secondly, this study looked at how stocking rates influence soil nutrients (nitrogen, phosphorus), and others soil properties such as pH and soil moisture. The distribution of alien grasses was facilitated by grazers. About 58 % of grasses germinated from the dung of eland and Bontebok were alien species. Additionally, there was significant variation in palatability related to grazing pressure and plant maturity between sites and over seasons. Grasses on the high grazing site had higher carbohydrate (3%-5% higher) and protein content (1%-5% higher) than at the least grazed site. Mature grasses contained less water (10%-20%), but no obvious variation in tannin concentration than immature grasses. However, species such as Ficinia sp. showed variations with grazing pressure and maturity. Ficinia sp. on the high grazing site had higher tannin concentration (5mg/ml-22mg/ml) and crude protein (4%-9%) than at the least grazed site. When the plant ages, tannin concentration increase (ranging from 15% to19% higher). Two types of plant were recoreded according to their chemical response following grazing pressure. These types are 1) plants that are grazing tolerant and which produce more carbohydrate when grazed and 2) plants that are grazing intolerant under high grazing pressure and which produce chemical defence compounds such as tannin to deter herbivores attacks. The results indicate that in fynbos and renosterveld, the relationship between preference and plant chemical compounds is not consistent since no III patterns emerged to explain what compounds drive preference. A combination of chemical compounds may be the reason for the selection by the grazers; alternatively, other compounds not included in the study may influence the forage selection by an herbivore. The second part of the study showed that stocking rates appeared to have a significant effect on soil properties investigated. Soil moisture was significantly affected by the stocking rate in autumn. The high grazing intensity site had the lowest soil moisture especially in autumn (10% lower than the control site), likely due to heavy trampling and soil compaction. pH was the lowest at the high grazing intensity site in all seasons compared to the control site. Likely reasons were the high deposition of nitrogen through dung and urine deposition, high removal of basic cations and animal exportation through hunting activities. Lastly, grazing maintains inorganic nitrogen at stable levels regardless of seasonal changes and increases the concentration of phosphorus especially under high grazing pressure compare to less grazed sites. The grazing dynamics of Swartland Shale Renosterveld - Swartland Alluvium Fynbos mosaic system are complex and multifaceted. Farmers need to pay attention to the nutritional status of plant species grazed to know whether they meet the nutritional requirements of the game in the area. Overstocking should be avoided in this system as it triggers the production of tanniferous compounds that could decrease the survivorship of herbivores. Moderate grazing 0.09 LAU/ha pressure provides the best stocking rates for effective grazing management. IV OPSOMMING Vir suksesvolle weidingsbestuur is dit noodsaaklik om die interaksie tussen herbivore, plantvoedingstowwe en grondeienskappe te verstaan. Van besondere belang is plantsmaaklikheid en herbivore se weidingsvoorkeure. Hierdie aspekte, wat die fokus van hierdie tesis is, is bestudeer in ‘n Swartland Shale Renosterveld en Swartland Alluvium Fynbos-mosaïek in die Wes-Kaap Provinsie van Suid-Afrika. Hierdie plantegroei word as kritiek bedreigd geklassifiseer weens die transformering van sy landskap ten gunste van landbou-aktiwiteite en stedelike vestiging. Pre-koloniale herbivoor vertebrata is teen 1700 deur jagters uitgeroei, wat gelei het tot min kennis oor die vroeë dinamiek van die plantegroei. Wildsboere en grondeienaars is egter besig om geselekteerde spesies in die area te hervestig. Bestuurders moet dus weet hoe om praktiese riglyne vir beste-praktyk weidingsbestuur te implementeer. Hierdie studie ondersoek eerstens die verhouding tussen plante en diere. Dit sluit seisoenale evaluering van saadontkieming uit die mis van bontebokke en elande in; die effek van veegetalle op plantsmaaklikheid (ruproteïen, ruvesel, ru-vet, vogpeil, tannien en minerale) van die weiveld is bestudeer om vas te stel hoe plantsmaaklikheid herbivore se weidingsbesluite beïnvloed. Tweedens het die studie die invloed van veegetalle op grondvoedingstowwe (stikstof, fosfor) ondersoek, asook ander grondeienskappe soos pH en grondvogpeile. Die verspreiding van uitheemse grasse is deur weidende diere gefassiliteer. Ongeveer 58% van alle grasse wat uit die mis van elande en bontebokke ontkiem het, was uitheemse spesies. Verder was daar beduidende variasie in plantsmaaklikheid verwant aan beweidingsdruk en plantvolwassenheid tussen verskillende persele en oor seisoene. Grasse op die hoogs beweide persele het 3%-5% hoër koolhidraatinhoude en 1%-5% hoër proteïeninhoude gehad as die minder beweide persele. Volwasse grasse het 10%-20% minder water bevat as onvolwasse grasse, maar het nie in terme van tannienkonsentrasie van die onvolwasse grasse verskil nie. Spesies soos Ficinia sp. het variasies getoon met beweidingsdruk en volwassenheid. Ficinia sp. het op die hoog-beweide perseel ‘n hoër tannienkonsentrasie (5mg/ml-22mg/ml) en meer ruproteïen (4%-9%) gehad as op die minste beweide perseel. Wanneer die plant verouder, verhoog die toename in tannienkonsentrasie (met tussen 15% en 19%). Twee tipes plante is aangeteken volgens hul chemiese response ná beweidingsdruk. Hierdie tipes is 1) plante wat beweidingstolerant is en wat meer koolhidrate produseer wanneer hulle as weiding dien en 2) plante wat onder hoë beweidingsdruk beweidingsintolerant is en wat chemiese verbindings V soos tannien produseer om herbivooraanvalle af te weer. Die resultate dui aan dat die verhouding tussen voorkeur en plante se chemiese verbindings nie konstant bly in fynbos en renosterveld nie, aangesien geen patrone na vore getree het wat kon verklaar watter verbindings bepalend is vir voorkeur nie. ‘n Kombinasie van chemiese verbindings mag die rede wees waarom die herbivore sekere plante verkies; alternatiewelik mag ‘n herbivoor se plantseleksie beïnvloed word deur chemiese verbindings anders as dié wat in hierdie studie ondersoek is. Die tweede deel van die studie het aangetoon dat veegetalle blyk ‘n beduidende invloed te hê op die grondeienskappe wat ondersoek is. Grondvogpeile is in die herfs beduidend beïnvloed deur die veegetalle. Die perseel met die hoë beweidingsintensiteit het die laagste grondvogpeil gehad, veral in die winter (10% laer as dié van die kontrole-perseel), waarskynlik weens erge getrappel en grondkompaktering. Vergeleke met die kontrole-perseel was pH die laagste op die perseel met die hoë beweidingsintensiteit, in alle seisoene. Waarskynlike redes hiervoor is die hoë neeerslag van stikstof deur mis- en uriene-neerslae, hoë verwydering van basiese katione en dierevervoer wat deur jagaktiwiteite teweeg gebring word. Laatstens behou weiding anorganiese stikstof teen stabiele vlakke ongeag van seisoenale veranderinge en toenames in die konsentrasie van fosfor, veral onder hoë beweidingsdruk vergeleke met minder beweide persele. Die weidingsdinamiek van die Swartland Shale Renosterveld - Swartland
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