Disturbance factors related to conservation of biodiversity in large-scale ecological networks

Joubert, Lize (2014-04)

Thesis (PhD)--Stellenbosch University, 2014.

Thesis

ENGLISH ABSTRACT: Globally, habitat transformation causes biodiversity loss, with the transformed matrix often affecting the disturbance regime in remnant natural patches. In South Africa, significant parts of the Indian Ocean coastal belt and grassland biomes have been transformed into commercial forestry plantations of alien trees, which are detrimental to local biodiversity. Consequently, large scale ecological networks (ENs) of remnant natural vegetation, maintained areas (e.g. firebreaks) and special landscape features (e.g. rocky outcrops and wetlands) have been implemented among forestry compartments to offset the negative effect of this land use on biodiversity. Different grassland areas, which constitute a major portion of ENs, were managed in different ways, as governed by their primary purpose (e.g. fire protection or conservation). The overall aim of this study was to determine how grassland floral and grasshopper herbivore communities responded to different disturbances (mowing, burning and grazing), and how we can adjust management of the major disturbances to effectively conserve these major components of biodiversity in ENs. Sampling was carried out in the commercial forestry ENs in the lower-elevation Zululand area and adjacent reserve area iSimangaliso Wetland Park, as well as in the forestry ENs in the higher-lying Midlands and adjacent iMpendle Nature Reserve. Both the reserves or protected areas (PAs) acted as reference sites, while other sites were chosen to represent the predominant disturbances in ENs at each locality: mowing, annual vs. longer-rotation burning, time since last fire, and domestic cattle grazing. In the Zululand subtropical grassland (chapter 2), I explored the effect of frequent mowing on firebreaks, and the effect of patch size and isolation on plant communities in non-firebreak natural areas of the EN. Frequent mowing resulted in plant species loss and a shift in species composition of firebreaks. Furthermore, small, isolated patches in the EN far away from the PA border had lower plant species richness and greater species turnover than wide, interconnected corridors near the PA border, which, in turn, was similar to reference sites in the PA. As plant species were lost from frequently-mown firebreaks and small, isolated patches in the EN, I recommend that this management practice should be confined to demarcated areas (e.g. forestry compartment edges and firebreaks) and that creation of wide, interconnected corridors should be prioritized when designing ENs. In higher elevation Afromontane grassland (chapter 3), I investigated the effect of annual burning on plant communities in firebreaks by comparing them to less frequently burned grassland in the EN and PA, respectively. Grazing by domestic cattle was taken as an embedded factor for firebreak and less frequently burned sites in the EN. There were three firebreak types: annually-burned with heavy cattle grazing (plantation firebreaks), annually-burned with light cattle grazing (peripheral firebreaks), and annual burning without cattle grazing (PA firebreaks). Burned reference grassland in the EN and PA hosted plant communities that were similar in species richness, composition and turnover. This was also the case for lightly-grazed peripheral EN firebreaks and PA firebreaks. However, species composition and turnover of plantation EN firebreaks with heavy cattle grazing differed from that in the other two firebreak types. Although not significant (P<0.1), plantation EN firebreaks had less plant species than reference burned grassland in the EN, and all firebreak types had less plant species, lower species turnover and different species composition when compared to reference burned grassland in the PA. Annual burning of firebreaks, with and without cattle grazing, caused a significant shift in plant species composition and a reduction in plant species turnover. When annual burning was combined with heavy cattle grazing, plant species were lost, as was the case in plantation EN firebreaks. Therefore, I recommend that this management practice should be confined to firebreaks, and that cattle access to firebreaks should be strictly controlled. In Chapter 4, I considered the effect of cattle grazing (presence vs. absence, as well as intensity) on Afromontane grassland against the natural backdrop of variation caused by time since last fire in grassland with longer fire-return intervals (excluding all firebreaks). Lowest plant species richness and turnover occurred in unburned (i.e. burned >12 months prior to sampling), ungrazed grassland in the PA. Burning and grazing both caused a change in plant species composition that went hand in hand with an increase in plant species richness and turnover. However, burning (burned vs. unburned) only affected plant communities in ungrazed grassland in the PA. Similarly, the presence of large mammalian grazers (EN vs. PA) only affected plant communities in unburned grassland. Unburned plant communities grazed by domestic cattle in the EN were similar to those in the PA grazed by indigenous black wildebeest, indicating that cattle grazing simulates, at least to some degree, the effect of indigenous ungulate grazing. Nevertheless, heavily-grazed grassland had less plant species than moderately-grazed grassland in the EN. I recommend that burning and grazing should continue in grassland ENs, as these natural disturbances are necessary to maintain diverse and dynamic ecosystems. Nevertheless, managers should instigate cattle grazing with caution, as high intensity grazing can be detrimental to conservation efforts. In Chapter 5, I examined the effect of annual burning, cattle grazing (presence vs. absence) and time since last fire on grasshopper assemblages in Afromontane grassland. In general, grasshoppers benefitted from disturbance, and were remarkably resilient to different disturbance regimes. Grasshopper species richness and their abundance were both greatest in annually-burned firebreaks with light cattle grazing, and lowest in moribund grassland in the PA which had not been burned for several years. Yet, time since last fire only affected grasshopper communities in the absence of large grazers (in the PA). None of the individual disturbances had an effect on the grasshopper assemblage. Rather, these insects responded to the combined effect of annual burning with cattle grazing. Sites were similar in grasshopper species richness, composition and abundance whenever either annual burning or cattle were absent, which suggests that these two disturbances drive changes in the grasshopper assemblage in these grasslands. Although grasshoppers benefited from annual burning with light cattle grazing, I would not recommend this disturbance regime outside firebreaks. Rather, management of other grassland areas in the EN should adapt longer fire-return intervals with a rotational cattle grazing system, so that undisturbed habitat is provided for other sensitive taxa. In conclusion, grassland plants and grasshoppers benefited from some form of disturbance, but were lost from small, isolated patches in the EN, as well as from areas with high disturbance frequency and intensity. Simulation of natural disturbances (moderate levels of fire and grazing) in wide, interconnected corridors is necessary for maintaining diverse and dynamic grassland ecosystem in ENs among commercial forestry plantations.

AFRIKAANSE OPSOMMING: Omskepping van natuurlike habitat na lande of plantasies veroorsaak biodiversiteitsverlies wêreldwyd. Boonop het sulke veranderinge dikwels 'n effek op die versteurings binne-in oorblywende kolle natuurlike plantegroei wat verreikende gevolge kan hê. Groot gedeeltes van die Suid-Afrikaanse grasveldbioom is omskep in bosbou plantasies wat bestaan uit uitheemse bome wat 'n baie groot nadelige effek op plaaslike biodiversiteit het. Daarom is grootskaalse ekologiese netwerke (EN’e), wat bestaan uit oorblywende kolle natuurlike plantegroei, brandbane en spesiale habitattipes in die landskap (bv. rotsriwwe en vleilande), tussen bosbouplantasies geïmplimenteer met die doel om die negatiewe effek van plantasies op plaaslike biodiversiteit te verlig. Bestuur van grasvelde, wat 'n groot gedeel van EN’e uitmaak, wissel dikwels en hang af van hulle primêre doel (bv. beskerming van plantasies teen wegholveldbrande of natuurbewaring). Die doel van hierdie projek was om vas te stel hoe plant- en springkaangemeenskappe in grasvelde reageer op verskillende versteurings (grassny, brand en beweiding), en die optimale bestuur van die versteurings om die biodiversiteit in grasvelde beter te bewaar. Steekproewe is geneem in EN’e tussen bosbouplantasies in die laagliggende Zululand en langsliggende wêrelderfenisgebied, iSimangaliso Wetland Park, asook in die hoërliggende Midlands en langsliggende iMpendle Natuurreservaat (NR). NR’e het as verwysing gedien waarteen die effek van grassny, frekwensie van brande, tydsverloop vanaf die laaste brand, en beweiding deur beeste, wat tipiese versteuringe in EN’e is, gemeet is. In hoofstuk 2 het ek vasgestel wat die effek van grassnyfrekwensie op plantgemeenskappe in brandbane is, en hoe plantgemeenskappe in subtropiese grasveld in die res van die EN reageer op die grootte en strukturele isolasie van oorblywende kolle natuurlike plantegroei. 'n Hoë grassnyfrekwensie het 'n verandering in die spesiesamestelling van plantgemeenskappe in brandbane veroorsaak wat gepaard gegaan het met spesiesverlies. Terselfdertyd was daar minder plant spesies in klein, geïsoleerde kolle natuurlike plantegroei as wat daar in wyer, aaneenskakelende gange nader aan die natuurreservaatgrens was. Laasgenoemde het plantgemeenskappe bevat wat baie soortgelyk aan die in die natuurreservaat was. Daarom stel ek voor dat die skep van wye, aaneengeskakelde natuurlike habitat prioriteit moet geniet wanneer nuwe EN’e ontwerp word, en dat gras slegs gereeld gesny moet word in spesifieke, afgebakende areas (bv. brandbane). Die rede hiervoor is dat hierdie bestuurspraktyk nie bevorderlik was vir die bewaring van plantdiversiteit in EN’e nie. In hoofstuk 3 het ek gekyk hoe die plantgemeenskappe in brandbane daarop reageer om elke jaar gebrand te word deur hulle te vergelyk met Afrikaberg grasveld in die EN en NR wat minder gereeld gebrand word. Beweiding deur beeste is gesien as 'n integrale deel van die EN. Ek het onderskei tussen plantasiebrandbane met swaar beweiding, randbrandbane met ligte beweiding en brandbane in die NR sonder beweiding. Die plantspesiesamestelling van brandbane, met ligte of geen beweiding nie, het verskil van grasvelde wat minder gereeld gebrand word. Tog is die hoeveelheid plantspesies nie geraak nie. Alhoewel die plantgemeenskappe in ligbeweide brandbane soos die in onbeweide brandbane in die NR was, het die plantspesiesamestelling van beide verskille getoon wanneer hulle vergelyk is met plantasiebrandbane wat swaarder deur beeste bewei is. Plantspesierykheid in plantasiebrandbane was boonop heelwat laer as wat in NR grasvelde gevind is, en daar was heelwat meer kaal grond in plantasiebrandbane as in enige van die ander areas. Oor die algemeen het plantspesiesrykheid van brandbane nie daaronder gely om elke jaar gebrand te word nie, maar kwesbare plantgemeenskappe in brandbane het wel daaronder gely om swaar bewei te word. Daarom stel ek voor dat jaarlikse brande tot brandbane beperk word en dat beeste se toegang tot brandbane streng beheer word. In die hoofstuk 4 ondersoek ek die effek van beweiding deur beeste (teenwoordigheid teenoor afwesigheid, sowel as beweidingsintensiteit) op die plantspesiesrykheid en samestelling van gebrande en ongebrande Afrikaberg grasvelde wat minder gereeld gebrand word. Die minste plant spesies is aangeteken in ongebrande, onbeweide grasveld in die NR. Brande en beweiding het albei 'n effek op plantspesiesamestelling gehad wat gepaard gegaan het met 'n toename in plantspesiesrykheid. Plantgemeenskappe in grasvelde wat onlangs (<12 maande voor die steekproef geneem is) gebrand is, het slegs van die in ongebrande grasvelde verskil wanneer nie een van die twee areas bewei is nie. Op 'n soortgelyke trant het die teenwoordigheid van beeste (EN teenoor NR) slegs n effek gehad in ongebrande grasvelde. Ongebrande plantgemeenskappe in die EN wat deur beeste bewei is, was baie soos die in die NR wat deur swartwildebeeste bewei is. Dit dui daarop dat beeste die effek van inheemse wildsoorte tot 'n mate naboots. Des nieteenstaande die bogenoemde, het swaar-beweide grasvelde minder plantspesies gehad as grasvelde wat slegs matig bewei is. Ek stel voor dat brande en beweiding deel moet vorm van die bestuur van grasvelde in EN’e, want hierdie natuurlike versteuringe dra by tot 'n diverse, dinamiese grasveldekosisteem. Tog moet bestuurders versigtig wees wanneer hulle die plaaslike gemeenskap se beeste in EN’e toelaat, want swaar beweiding kan bewaringsinisiatiewe in die wiele ry. In hoofstuk 5 het ek die klem na springkane verskuif, en die effek van jaarlikse brande, beweiding deur beeste (teenwoordigheid teenoor afwesigheid) en tydsverloop sedert laaste brand op hierdie sensitiewe insekte in Afrikaberg grasvelde ondersoek. Alhoewel springkaangemeenskappe baat gevind het by versteuringe, het hulle nie beduidend gereageer op enige van die indiwiduele versteuringe nie. Die digste sprinkaan bevolking met die hoogste spesies diversiteit is aangeteken in brandbane in die EN wat liggies deur beeste bewei is. Darenteen is die laagste bevolking en spesies diversiteit aangeteken in grasvelde in die NR wat groot hoeveelhede dooie plantmateriaal bevat wat aandui dat hierdie grasvelde nie onlangs gebrand het nie. Springkaangemeenskappe in gebrande grasvelde het slegs van ongebrande grasvelde verskil wanneer nie een van die twee bewei is nie. Die sleutelkombinasie van versteuringe wat die rykheid en samestelling van springkaangemeenskape bepaal het, was 'n hoë brandfrekwensie (soos in brandbane) en beweiding deur beeste. Wanneer een van hierdie versteuringe afwesig was, was springkaangemeenskappe tussen verskillende areas dieselfde. Alhoewel springkaangemeenskappe daarby baat gevind het wanneer brandbane elke jaar gebrand en deur beeste bewei is, kan ek nie hierdie bestuurspraktyk vir die res van die EN aanbeveel nie. Grasvelde in die res van die EN behoort eerder minder gereeld (elke 2-4 jaar) gebrand en met 'n rotasiestelsel bewei word. Sodoende sal brandbane voorsien in die behoeftes van springkane, en die res van die EN in die behoeftes van sensitiewe taksa wat onversteurde habitat benodig om te floreer. My slotsom is dat versteuringe nodig is om die volle diversiteit van plante en springkane en die dinamika binne-in grasvelde te bewaar. Tog verdwyn daar plantspesies uit areas met 'n hoë versteuringsintensiteit of frekwensie en klein, geïsoleerde kolle natuurlike plantegroei in die EN. Daarom beveel ek aan dat natuurlike versteuringe (brande en beweiding) matig toegepas moet word in wye, aaneengeskakelde gange in die EN. Hierdie benadering tot natuurbewaring kan biodiversiteit tussen bosbouplantasies beveilig teen verdere verlies.

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