An assessment of the potential for utilisation of soil-stored seed, from on- and off 'conservation islands' (isolated mountains), as an indicator of restoration potential of degraded sites in semi-arid Karoo areas

Thesis (MSc)--University of Stellenbosch, 2000.

ENGLISH ABSTRACT: The composition and state of soil-stored seed banks on- and off the mesa Tafelberg, in the Nama Karoo rangelands of the Eastem Cape, South Africa, were investigated within the context of a broader restoration ecology project "Restoration of degraded Nama Karoo rangelands: the role of conservation islands'". The premise for this seed bank study was that restoration of degraded semi-arid rangelands is possible through applied management programs based on the methodology and practice of ecological restoration. Broadly acknowledged properties of non-equilibrium environments (e.g. unpredictable climates and varying degrees of disturbance) and soilstored seed banks (e.g. spatial and temporal distributions) formed the basis for investigating. the general environment and the properties.. of existing seed, banks, in the Tafelberg locality. The potential role of hills as refugia for palatable plant species was an under-lying element of the investigation. Following a brief investigation, of historical and contemporary research and policy on rangeland degradation in semi-arid regions of the world, the fundamental need for comprehensive and applied seed bank research in the Nama Karoo is emphasised. Within the framework of the umbrella project, the seed bank study examined local environmental criteria commencing with an investigation into seed bank- and vegetation habitats. Chemical and physical properties of soils from twenty two sites on- and off Tafelberg were described. Substantial soil habitat variation, between the top, the north west slopes and plains and the south east slopes and plains of Tafelberg, was identified. Micro-site variation between open-canopy (interplant spaces) and closed-canopy (under plant cover) microhabitats was found to be significant Primary soil habitat differences were linked to soil organic matter content which was found to be low on the plains relative to the top and slopes; and, low in open-canopy micro-sites relative to closed-canopy sites. Soil texture and nutrient composition on- and off Tafelberg was found to be highly variable with significant differences between the top and the plains as well as between the north west and south east plains. The slopes were found to be intermediate (showing some level of gradient) between the top and the plains. Soil nutrient variation was interpreted as a function of textural and parent-material properties of soils. Anthropogenic factors for accelerated erosion, deposition, leaching and salt-crust formation were also considered. It was concluded that while properties of soils on- and off Tafelberg are inherently related to parent soils, changes to textural and nutrient properties may be occurring and these changes may have been exacerbated by high levels of grazing. An investigation (focusing on small shrubs) of plant phenological response, in relation to rainfall and grazing gradients, identified trends of peak budding- and flowering seasons following rainfall during summer and autumn respectively. A continuum of seeding activity, with peaks in late autumn and early winter, was construed from quarterly data. Flush vegetative growth was noted for most small shrubs during spring, autumn and summer surveys. Since some form of activity related to reproductive output (flush growth, budding, flowering or seeding) was apparent at almost all times of the year, it is argued that high intensity disturbance (including grazing) might impact negatively on plant survival, leading to reduced reproductive input (i.e. seeds) into future generations. Plant communities on the top and plains are described noting significant differences between vegetation on top of Tafelberg (comprising primarily high production, palatable grass and shrub species) and that on the plains (comprising mainly spinescent, ephemeral, toxic and low production species). Given the high grazing pressure on the plains (relative to the less utilised slopes and top of Tafelberg), differences in vegetation composition are discussed in relation to studies elsewhere that describe degraded rangelands. It is concluded firstly that the plains surrounding Tafelberg are degraded, secondty that long term over-utilisation has altered vegetation composition and finally that inter-grazing rest periods of three- or six months alone may not restore vegetation diversity nor desirable plant species to the plains. Results of germination trials (investigating soil-stored seed banks) from two sampling episodes (spring and autumn 1998) revealed that seed banks in soils removed from the plains, slopes and top of Tafelberg followed distribution patterns observed in above-ground vegetation. Species-specific data was not finalised for this thesis since not all seedlings matured and flowered within given time constraints. In order to compare seedling emergence data, plant categories were developed that distinguished ephemerals versus persistent (perennial) species and dicotyledonous species versus grasses and other monocotyledonous plants. Samples from the top and the middle to upper slopes showed a high percentage of palatable and persistent grass and shrub species present in soil-stored seed banks while over twothirds of plants germinating from plains' soil samples were ephemeral species and most of these were both tiny «Scm) and short-lived «3 months). Most of the perennial species germinating from plains' samples were seedlings of Pentzia incana, Chrysocoma ciliata and succulents (mostly Mesembryanthemaceae) but few perennial grasses were present. On the other hand, roughly 94% of seedlings germinating from samples from the top and 63% of seedlings germinating from two slopes of Tafelberg respectively were persistent- grass or shrubby species. Roughly 89% of ephemeral species recorded from slopes' samples germinated from the lowest footslope sites. Seed densities were closely linked to micro-habitats with roughly three-fold differences between open- (lower seed density) and closed-canopy (higher seed density) micro-sites. Multivariate analysis of variance (MANOVA) indicated that most of the variation in seed density data was explained at the micro-habitat level. Seasonality played a secondary role with significant twoway interaction between the two effects. It is concluded that both factors must be considered when developing restoration programmes that aim to improve both overall plant cover and improved plant species diversity. While the original or "pristine" state of vegetation composition and seed banks on the plains is unknown, concems were raised regarding the apparently degraded state of total above- and below-ground plant diversity on these plains. A likely consequence of habitat degradation is that species with specific soil-, nutrient ratio-, aspect- and altitudinal requirements from the top and slopes of Tafelberg (as well as from nearby plains' refugia) may not be able to establish in degraded habitats on the plains. The identification of pioneer plant species (tolerant of habitat degradation) that allow increased vegetation cover and safe-sites for seedling germination of desirable plant species is recommended. Restoration programmes will need to be coupled with strict grazing management principles that allow seedling germination. establishment and successful reproductive output of desirable plants for future rangeland regeneration. Common and salient features of the soil habitat, plant pbenological response and germination trial studies are brought together in an examination of habitats and related seed bank diversity on- and off Tafelberg. Acknowledging the brevity of this research study, but utilising case studies from elsewhere and integrating both lines of questioning, the conclusion is again reached that the plains surrounding Tafelberg are degraded through decades and probably centuries of grazing by domestic livestock. It is considered crucial that restoration through improvement (or rehabilitation) of habitats and increased seedling safe-sites be considered. Both climate and grazing management appear to play an irrevocably linked role in shaping vegetation composition in rangelands. While rangelands are intrinsiCalfy adapted to surviving extremes of climatic variability found in non-equilibrium regions it is argued that the impacts of grazing, particularly during times of drought and climate change, are slowly reducing the intrinsic "buffer-capacity" of rangelands to withstand these changes and extremes. It seems apparent from research elsewhere that restoration through passive management is slow and probably not economically achievable within a viable time frame. The instifution and promotion of integrated and strategic programmes that identify and address issues of land degradation and land use change in semi-arid rangelands is recommended. The input and endeavours of different authorities, ministries and a broad public participation incentive are encouraged in these proposed programmes in order to ensure broadly-based input into long term sustainability and conservation of the considerable biological diversity of these regions. Seed bank assessment is considered to be a valuable means of indicating restoration potential and ranqeland condition with potential for the identification of both degraded and conservationworthy areas. Finally, some limitations and challenges of this study are examined through a process of firstly identifying alternative approaches to research methodologies and secondly through proposing recommendations for future research projects. While alternative methods could have been applied for the purposes of accomplishing this study it is concluded that, within the given time- and other constraints, the appropriate methods were applied.

AFRIKAANSE OPSOMMING: Die samestelling entoestand van groncbbewaarde saadbanke.op en van die mesa Tafelberg af, in die Nama Karoo weiveld van die Oos-Kaap, Suid-Afrika, is ondersoek binne die konteks van 'n wyer herstelekologieprojek: "Restourasie van oorbeweide Nama Karoo weiveld: die rol van bewannqseilande'". Die uitqanqspunt van hierdie saadbankondersoek was dat dit moontlik is om beskadigde semidroë weiveld te herstel deur middel van toegepaste bestuursprogramme wat gebaseer is op die metodologie en praktyk van ekologiese herstel. Algemeen erkende eienskappe van nieekwilibrium- omgewings (byvoorbeeld onvoorspelbare klimate en wisselende mates van versteuring) en grond-bewaarde saadbanke (byvoorbeeld ruimtelike en temporale verspreidings), het die basis gevorm vir die ondersoek van die algemene omgewing en die eienskappe van bestaande saadbanke in die Tafelberg omgewing. Die potensiële rol van heuwels as skuilplek vir aanvaarbare plantspesies was 'n onder1iggende element van die ondersoek. Na 'n kort ondersoek van historiese ~n kontemporêre navorsing en beleid oor weiveldbeskadiging in semi-droë streke van die wêreld, is die fundamentele behoefte aan omvattende en toegepaste saadbanknavorsing in die Nama Karoo beklemtoon. Binne die raamwerk van die oorkoepelende projek het die saadbankondersoek plaaslike omgewingskriteria bestudeer, beginnende met 'n ondersoek na saadbank- en plantegroeihabitats. Chemiese en fisiese eienskappe van grond vanaf twee en twintig terreine op en van Tafelberg af is beskryf. Omvattende grondhabitatwisseling tussen die kruin, die noordwestelike hange en die vlaktes en die suidoostelike hange en vlaktes van Tafelberg is geïdentifiseer. Mikroterreinwisseling tussen die oop-dak (tussenplantruimtes) en toe-dak (onderplantdekking) -mikrohabitats is as beduidend bevind. Primêre grondhabitatverskille is gekoppel aan die inhoud van die grond se organiese materiaal, wat op die vlaktes as laag bevind is vergeleke met die kruin en hange; en laag bevind is in oop-dak-mikroterreine vergeleke met toe-dak-terreine, Daar is gevind dat die grondtekstuur en voedingstofsamestelling op en van Tafelberg af aansienlik varieer, met beduidende verskille tussen die kruin en die vlaktes, en ook tussen die noordwestelike en suidoostelike vlaktes. Daar is gevind dat die hange intermediêr is (toon 'n mate van gradiënt) tussen die kruin en die vlaktes. Die wisseling in grondvoedingstowwe is vertolk as 'n funksie van teksturele en ouermateriaaleienskappe van grond. Antropogeniese faktore vir versnelde erosie, neerslag, loging en soutkorsvorming is ook oorweeg. Die gevolgtrekking is gemaak dat terwyl die eienskappe van grond op en van Tafelberg af inherent aan ouergrond verwant is, kan veranderinge aan tekstuur en voedingstofeienskappe voorkom, en hierdie veranderinge kon deur hoë vlakke van weiding vererger gewees het. 'n Ondersoek (wat op klein struike gefokus het) van plantfenologiese reaksie met betrekking tot reënval en weidingsgradiënte het tendense van piekbot- en blomseisoene na reënval gedurende die somer en herfs onderskeidelik geïdentifiseer. 'n Kontinuum van saadskietaktiwiteit, met piektye in laat herfs en vroeë winter, is van kwartaallikse data saamgestel. Groeistuwing is vir die meeste klein struike waargeneem gedurende lente-, herfs- en someropnames. Aangesien die een of ander vorm van aktiwiteit met betrekking tot voortplantingsgroei (groeistuwing, bot, blom of saadskiet) op feitlik al die tye van die jaar sigbaar was, word daar geredeneer dat hoëintensiteitversteuring (ook weiding) 'n negatiewe impak op plantoorlewing kan hê, wat sal lei tot verminderde voortplantingsinset (m.a.w. sade) in toekomstige geslagte. Plantgemeenskappe op die kruin en vlaktes word beskryf met beduidende verskille tussen plantegroei op die kruin van Tafelberg (wat hoofsaaklik bestaan uit hoëproduksie, smaaklike gras- en struikspesies) en dié ap die vlaktes (wat bestaan uit hoofsaaklik doringagtige, efemere, toksiese en laeproduksie-spesies). Gegee die hoë weidingsdruk op die vlaktes (vergeleke met die minder benutte hange en kruin van Tafelberg), word verskille in die samestelling van plantegroei bespreek met betrekking tot studies elders wat beskadigde weiveld beskryf. Die gevolgtrekking word eerstens gemaak dat die vlaktes om Tafelberg beskadig is, tweedens dat langtermyn-oorbenutting die samestelling van die plantegroei verander het, en laastens dat interweiding-rusperiodes van drie tot ses maande alleen dalk nie die diversiteit van plantegroei of die verlangde plantspesies op die vlaktes kan herstel nie. Die uitslae van kiemingstoetse (wat ondersoek ingestel het na grond-bewaarde saadbanke) van twee steekproefepisodes (lente en herfs 1'998)het getoon dat saadbanke in grond wat van die vlaktes, hange en kruin van Tafelberg verwyder is, die verspreidingspatrone volg wat in bogrondse plantegroei waargeneem is. Spesie-spesifieke data is nie vir hierdie tesis gefinaliseer nie, aangesien nie alle saailinge binne die gegewe tydsbeperkinge gegroei en geblom het nie. Ten einde saailing-verskyningsdata te vergelyk, is plantkategorieë ontwikkel wat efemere en langdurige (meerjarige) spesies en dikotiele spesies en grassoorte en ander monokotiele plante onderskei. Steekproewe van die kruin en teen die middelste en boonste hange het 'n hoë persentasie van aanvaarbare en standhoudende gras- en struikspesies getoon wat in grond-bewaarde saadbanke teenwoordig is, terwyl meer as twee derdes van plante wat in die vlaktes se grondmonsters ontkiem het, efemere spesies was, en die meeste daarvan was klein «5cm) en met 'n kort leeftyd «3 maande). Die meeste van die meerjarige spesies wat van die vlaktes se steekproewe kom, was saailinge van Pentzia incana, Chrysocoma ciliata en vetplante (hoofsaaklik Mesembryanthemaceae), maar min meerjarige grasse was teenwoordig. Daarenteen was onderskeidelik ongeveer 94% van saailinge wat van monsters van die kruin en 63% van saailinge wat van twee van die hange van Tafelberg ontkiem het, langdurige gras- of struikspesies. Ongeveer 89% van die efemere spesies wat van die hange se steekproewe aangeteken is, het op die laagste voethang-terreine ontkiem. Saaddigthede toon 'n noue verband met mikrohabitats, met ongeveer drievoudige verskille tussen oop- (laer saaddigtheid) en toe-dak (hoër saaddigtheid) -mikroterreine. 'n Multivariaatontleding van variansie (MANOVA) het aangedui dat die meeste van die variasie in saaddigtheidsdata op die mikrohabitat-vlak verduidelik is. Seisoenaliteit het 'n sekondêre rol gespeel, met beduidende tweerigting-interaksie tussen die twee uitwerkings. Die gevolgtrekking word gemaak dat albei faktore oorweeg moet word wanneer herstelprogramme ontwikkel word wat ten doel het om algehele plantbedekking en die diversiteit van verbeterde plantspesies te verhoog. Terwyl die oorspronklike of "ongerepte" toestand van die plantegroei se samestelling en saadbanke op die vlaktes onbekend is, is kommer uitgespreek oor die klaarblyklik beskadigde toestand van die totale bo- en ondergrondse plantdiversiteit op hierdie vlaktes. 'n Waarskynlike gevolg van habitatbeskadiging is dat spesies met spesifieke grond-, voedingstofverhouding-, aspek- en seevlak-vereistes van die kruin en hange van Tafelberg (asook van nabygeleë vlakteskuilings) nie in staat sal wees om in beskadigde habitats op die vlaktes te vestig nie. Die identifikasie van pionierplantspesies (verdraagsaam vir habitatbeskadiging) wat verhoogde plantegroeidekking en veilige terreine vir saailingontkieming van verlangde plantspesies toelaat, word aanbeveel. Herstelprogramme sal gekoppel moet word aan streng weidingsbeginsels wat saailingontkieming, die vestiging en suksesvolle voortplantingsproduksie van gewenste plante vir toekomstige weiveldherstel moontlik maak. Algemene en belangrike eienskappe van die grondhabitat, plantfenologiereaksie en kiemingsproefondersoeke word saamgevoeg in 'n ondersoek van habitats en verwante saadbankdiversiteit op en van Tafelberg af. Met erkenning van die kortstondigheid van hierdie navorsingsondersoek, maar met benutting van gevallestudies van elders en die integrasie van albei vraaglyne, word daarweertot die slotsom gekom dat die vlaktes om Tafelberg beskadig is deur dekades en waarskynlik eeue se beweiding deur mak lewende hawe. Dit word as uiters belangrik beskou dat herstel deur verbetering (of rehabilitasie) van habitats en 'n groter aantal saailing beveiligingsterreine oorweeg moet word. Dit lyk asof klimaat sowel as weidingsbestuur 'n onherroeplik gekoppelde rol speel in die vorming van die samestelling van plantegroei op weiveld. Terwyl weiveld intrinsiek aangepas is by die oorlewing van uiterste klimaatswisseling wat in nie-ekwilibriese streke aangetref word, word daar geredeneer dat die impak van beweiding, veral gedurende droogtetye en klimaatsverandering, stadigaan die intrinsieke "buffervermoë" van weiveld verminder om hierdie veranderinge en uiterstes te weerstaan. Dit blyk uit navorsing elders dat herstel deur passiewe bestuur stadig en waarskynlik nie ekonomies haalbaar is binne 'n lewensvatbare tydsraamwerk nie. Die instelling en bevordering van geïntegreerde en strategiese programme wat kwessies van grondbeskadiging en verandering van grondgebruik in semi-droë gebiede identifiseer en oplos, word aanbeveel. Die insette en pogings van verskillende owerhede en ministeries en deelname deur die breë publiek word aangemoedig in hierdie voorgestelde programme ten einde insette met 'n breë basis in die langtermyn~volhoubaarheid en bewaring van die aansienlike biologiese diversiteit van hierdie streke te verseker. Saadbankbeoordeling word beskou as 'n waardevolle manier om die herstelpotensiaal en weiveldtoestande met die potensiaal vir die identifisering van beskadigde sowel as bewaringswaardige gebiede aan te dui. Laastens word 'n paar beperkings en uitdagings van hierdie studie ondersoek deur 'n proses van eerstens die identifisering van alternatiewe benaderings tot navorsingsmetodologieë, en tweedens deur die voorstel van aanbevelings vir toekomstige navorsingsprojekte. Terwyl alternatiewe metodes toegepas kon gewees het ten einde hierdie studie af te handel, is die gevolgtrekking dat die toepaslike metodes binne die gegewe tyd5- en ander beperkings toegepas is.