Thesis (MSc)--Stellenbosch University, 2015. / ENGLISH ABSTRACT: Habitat loss and fragmentation are major threats to global biodiversity. A cornerstone of traditional conservation involves setting aside land as formally protected areas (PAs). However, for effective biological conservation in the long term there needs to be connectivity between these PAs. When possible, improved connectivity can be achieved using natural corridors at a landscape scale. Even better is to establish a network of corridors and nodes in the form of ecological networks (ENs). ENs are currently being employed by commercial forestry companies in South Africa. While larger corridors and nodes are considered optimum, factors other than design, such as management and environmental heterogeneity, have also been found to be important for species maintenance. This study aims to explore the role of corridor width in driving the composition of invertebrate assemblages across a transformed landscape in KwaZulu-Natal, South Africa, and to investigate other possible environmental variables significant for species distributions. In Chapter 2, I investigated the contribution of smaller grassland corridors within a timber production matrix to overall biodiversity conservation using two important bioindicator taxa. Ants and dung beetles were sampled in grassland corridors of three size classes, plantation blocks and a nearby PA, iMpendle Nature Reserve. The two taxa showed differential responses to landscape level fragmentation. Dung beetles showed a decrease in species richness and corresponding increase in species turnover with increased fragmentation, while ants were unaffected, although counter intuitively smaller corridors even contained more unique ant species compared to larger corridors. Dung beetle assemblages also showed strong differences between the PA and grassland corridors. While the conservation effectiveness of large corridors undoubtedly exceeds that of smaller corridors, for ants it seems that smaller corridors contribute to their overall conservation within this production landscape. In Chapter 3, I explore the importance of spatial and environmental factors for species distribution across this landscape. Dung beetles were split into functional guilds according to size and nesting behaviour for analyses. Within grassland corridors, tunnelling dung beetle species richness was sensitive to landscape level fragmentation, especially for larger species, while elevation and vegetation type influenced ant species richness. Since rolling dung beetles showed a close association with the PA, the marked difference in dung beetle assemblages between these two land-uses may be due to the presence of pellet producing grazers in the protected area and their replacement by pat producing cattle in the grassland corridors. Other environmental variables that were found to be important for dung beetle species composition were elevation, vegetation type, and soil hardness. For ant species composition, only elevation was found to be important. In conclusion, as large corridors were comparable to the PA in dung beetle and ant species richness, ENs act as extensions of formally PAs, given that they are large enough. Nevertheless, smaller corridors had surprisingly high species richness. Including additional information other than species data improved our knowledge of the underlying factors that drive dung beetle species composition. Even though dung beetle and ant species responded differentially to habitat fragmentation, environmental heterogeneity seemed important for both taxa. Incorporating habitat heterogeneity into the current management scheme may improve the conservation effectiveness within this transformed landscape. / AFRIKAANSE OPSOMMING: Die vermindering en fragmentasie van natuurlike habitat is ‘n groot bedreiging vir globale biodiversiteit. ‘n Belangrike tradisionele benadering tot natuurbewaring behels die afbakening van land vir formele beskermde areas (BAs). Ten einde effektiewe biologiese bewaring oor die langtermyn te verseker moet daar verbinding wees tussen hierdie BAs. Indien moontlik kan verbeterde verbinding verkry word deur die gebruik van natuurlike gange op ʼn landskaps-vlak. Nog beter is om ʼn netwerk van gange en nodes in die vorm van ekologies netwerke (ENe) saam te stel. ENe word tans deur kommersiële bosboumaatskappye in Suid Afrika aangewend. Terwyl groter gange en nodes as optimaal beskou word, is ander faktore behalwe ontwerp, soos bestuur en omgewingsheterogeniteit, ook al gevind as belangrik vir die onderhouding van spesies. Hierdie studie is gemik daarop om die rol van gangwydte as dryfkrag vir die samestelling van invertebraatversamelings oor ʼn getransformeerde landskap in KwaZulu-Natal, Suid-Afrika, te ondersoek, asook ander moontlike omgewingsveranderlikes wat belangrik vir spesiesverpreidings kan wees. In Hoofstuk 2 het ek die bydrae van kleiner gange tot totale biodiversiteit-bewaring ondersoek deur twee belangrike bio-indikator taxa te bestudeer. Miere en miskruiers is versamel in grasland-gange van drie grootte-klasse, plantasie blokke en ‘n naby geleë BA, iMpendle Natuurreservaat. Die twee taxa het verskillende reaksies tot landskaps-vlak fragmentasie getoon. Miskruiers het ‘n verlaging in spesiesrykheid en ‘n gesamentlike verhoging in spesiesomset met verhoogde fragmentasie gewys, terwyl miere nie geaffekteer is nie, alhoewel kleiner gange het trouens meer unieke mierspesies bevat as groter gange. Die miskruierversamelings in die BA het ook opmerklik verskil van dié in die grasland-gange. Alhoewel die bewaringsdoeltreffendheid van groot gange beslis dié van kleiner gange oorskry, kom dit voor dat kleiner gange wel bydra tot die totale bewaring van miere binne hierdie produksielandskap. In Hoofstuk 3 het ek die belangrikheid van ruimtelike en omgewingsfaktore vir spesiesverspreiding oor hierdie landskap ondersoek. Miskruiers is ook in funksionele groepe verdeel volgens grootte en nes-gedrag vir aparte analise. Binne grasland-gange was tonnellende miskruierspesies sensitief vir landskaps-vlak fragmentasie, veral groter spesies, terwyl hoogte bo seevlak en vegetasie tipe mier spesiesrykheid beïnvloed het. Aangesien rollende miskruierspesies ‘n nabye assosiasie met die BA gewys het, mag die opmerklike verskil in miskruier versamelings tussen hierdie twee grondgebruike ʼn gevolg wees van die aanwesigheid van korrel-mis produserend beweiders in die BA en hulle vervanging deur nat-mis produserende beeste in die grasland-gange. Omgewingsveranderlikes uitsluitende ganggrootte wat belangrik gevind is vir miskruier spesiessamestelling was hoogte bo seevlak, vegetasie tipe en grond-hardheid. Vir mier spesiessamestelling was slegs hoogte bo seevlak belangrik. Om af te sluit, aangesien groot gange vergelykbaar was met die BA in miskruier en mier spesiesrykheid, tree ENe op as uitbreidings van BAs, mits hulle groot genoeg is. Desnieteenstaande het kleiner gange ‘n verbasende hoë spesiesrykheid gehad, veral onder miere. Die insluiting van addisionele inligting buiten spesiesdata het ons kennis van die onderliggende faktore wat miskruier spesiessamestelling dryf verbeter. Alhoewel miskruier- en mierspesies verskillend gereageer het op habitat fragmentasie, het dit voorgekom asof omgewingsheterogeniteit belangrik was vir die spesiesverspreiding van beide taxa. Die insluiting van habitatheterogeniteit binne die huidige bestuursplan mag die doeltreffendheid van bewaring binne hierdie getransformeerde landskap verbeter.

Biodiversity can be described as the total variation of life forms, where diversity ranges from gene level up to the ecosystem level. The diversity can be calculated in a number of ways, and this study use one of these methods. In this study empirical food webs have been used and analyzed, where eleven characters are defined and used as parameters to calculate the Euclidean distances between food webs that describe the variation that may exist within classes of terrestrial, marine and freshwater habitats. The class who stood out and showed the greatest diversity at the ecosystem level was the marine food webs, which showed a high value of the average euclidean distance. The other networks were not as distinctive and the average of the euclidean distance in these classes was comparatively low.

Ecological networks are complex systems forming hierarchical structures in which energy and matter is transferred between the network’s compartments. Predicting energy flows in food webs usually involves complex parameter-rich models. In this thesis, the application of the principle of maximum entropy (MaxEnt) to obtain least biased probability distributions based on prior knowledge is proposed as an alternative to predict the most likely energy flows in food webs from the network topology alone. This approach not only simplifies the characterisation of food web flow patterns based on little empirical knowledge but can also be used to investigate the role of bottom-up and top-down controlling forces in ecosystems resulting from the emergent phenomena based on the complex interactions on the level of species and individuals. The integrative measure of “flow extent”, incorporating both bottom- up and top-down controlling forces on ecosystems, is proposed as a principle behind ecosystem evolution and evaluated against empirical data on food web structure. It could be demonstrated that the method of predicting energy flow with the help of MaxEnt is very flexible, applicable to many different setting and types of questions in ecology, and therefore providing a powerful tool for modelling the energy transfer in ecosystems. Further research has to show in how far the most likely flow patterns are realised in real-word ecosystems. The concept of flow extent maximisation as a selection principle during ecosystem evolution can enhance the understanding of emergent phenomena in complex ecosystems and maybe help to draw a link between thermodynamics and ecology.

Les méthodes actuelles utilisées pour définir les enjeux de biodiversité peinent à prendre en compte : les espaces agricoles et les pratiques, à la fois réservoir et pression pour la biodiversité et les dynamiques spatiales et temporelles, modifiant l’occupation du sol et la présence, la qualité et la connectivité des habitats. Ce travail a pour objectif de proposer des solutions pour ces 2 aspects.Nous avons identifié des pratiques agricoles favorables à la biodiversité. Pour comprendre les enjeux liés au développement de ces pratiques, nous avons étudié des leviers pour leur mise en place. Nous avons confirmé la difficulté à mesurer un effet général de certaines pratiques sur la biodiversité. Néanmoins, l’hétérogénéité, dans les pratiques, les types de culture et les éléments semi-naturels environnants est un aspect favorisant la biodiversité. Inciter leur développement nécessite de s’appuyer sur les avantages pour les agriculteurs, au travers des services écosystémiques et d'une multitude de supports de diffusion. Nous avons aussi montré l’importance d'intégrer l’agriculture dans l’identification d’enjeux de conservation.Nous avons présenté les limites des scénarios existants pour l’échelle régionale. Nous proposons une méthode alliant ces deux types de scénarios pour explorer l’impact des changements sur les continuités écologiques. Cette utilisation revêt un intérêt particulier pour les questions d’aménagement, par ex. pour mesurer l’effet, cumulé ou non, de certains projets. Avant le transfert vers des projets appliqués, il convient néanmoins de poursuivre la recherche sur cette méthode, notamment en la complétant avec d’autres mesures de continuités écologiques. / Actual ways to define priorities for biodiversity conservation experience difficulties to take into account: agricultural areas and practices, both reservoir and source of pressure for biodiversity and spatial and temporal dynamics modifying land-cover and land-use and the presence, the quality and the connectivity of habitats. This research work have for objective to propose solutions for these two aspects.We wanted to identify biodiversity-friendly agricultural practices. To understand the issues related to the development of these practices, we also studied the levers to their application. We confirm the difficulty to measure a general effect of some practices on biodiversity. Nevertheless, the heterogeneity, in practices, crops types and semi-natural elements, favor the biodiversity. Encourage their development need to rely on benefits for farmers, through ecosystem services and various substrate for information spread. We show the importance to taking into account agriculture in the identification of conservation priorities.We introduced the limits from existing scenarios to explore changes at regional scale. We proposed a method coupling these two types of scenarios to measure the impact of changes on ecological networks, with an indicator of connectedness, for three species. The application of scenarios to ecological networks’ analyses have a particular interest for land planning questions, for example to measure the, cumulative or not, effect of some projects. Before transfer to applied projects, research on this method have to be continued, in particular completing it with others measures of ecological networks.

Thesis (PhD)--Stellenbosch University, 2014. / 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.

Landscape fragmentation by line barriers in the Czech Republic - development, state of the art and priorities of territorial protection Abstract This paper deals with the problem of landscape fragmentation by linear structures and changes in land cover printed in Europe and the Czech Republic. The development of the fragmentation geometry consisting of transport infrastructure and urban areas is described in detail in the years 1920-2020. In this time boundary the development of measure of landscape fragmentation in the Czech Republic is evaluated. This paper examines also the quality of the current unfragmented landscape. The degree of landscape fragmentation refills the value of ecological integrity (according to Burkhard et al., 2009). The results of gap analysis define the most valuable areas of the Czech countryside, which are not covered by existing special protection area (NP, CHKO, Natura 2000). The resulting areas are also compared to the territorial system of ecological stability and migration important area for the large mammals. Keywords landscape fragmentation - landscape connectivity - ecological integrity - gap analysis of landscape protection

Green Infrastructure of Central Europe Green infrastructure is a strategically planned network that broadens the traditional conservation efforts to encompass the concept of ecosystem services. This study aims to identify green infrastructure network in Central Europe. Method presented in this thesis is an analysis of ecological connectivity based on ecosystem services potential quantified for CORINE land cover classes (Burkhard et al., 2009). Design of ecological corridors between the core areas represented by Natura 2000 sites is dependent on the capacity of ecosystems to provide ecological integrity and regulating services. Analysis was performed in ArcGIS Linkage Mapper extension. Green infrastructure network identifies 17 % of area of Central Europe that provides high values of ecosystem services. Corridors also create linkages between Natura 2000 sites and improve biodiversity conservation as well as they support migration corridors for large mammals. Full implementation of the Birds and Habitats directive and promoting of a European Green Infrastructure are two important targets of The EU Biodiversity strategy to 2020. Powered by TCPDF (www.tcpdf.org)

Ecological interactions such as predation, competition, and mutualism are important forces that influence species evolution. Coevolution is defined as reciprocal evolutionary change in interacting species. The Geographic Mosaic Theory of Coevolution (GMTC) provides a theoretical framework to explain how collections of populations should coevolve across space. Two fundamental aspects of the GMTC are gene flow among populations and the presence of selection mosaics, which are collections of localities with particular selection regimes. Several studies have explored how phenotypic trait matching between species evolves in pairs or small groups of species. However, ecological interactions frequently form large networks that connect dozens of species present in a given community. In networks of mutualisms, for instance, the organization of interactions may affect ecological and evolutionary processes. A next step in understanding the coevolutionary process is to investigate how aspects of the GMTC affect the evolution of species embedded in interaction networks. In this dissertation, we tried to fill this gap using a mathematical model of coevolution, complex networks tools, and information on empirical mutualistic networks. Our numerical simulations of the coevolutionary model allow us to draw three main conclusions. First, gene flow affects trait patterns generated by coevolution and may favor the emergence of trait matching depending on the selection mosaic. Second, the organization of mutualistic networks influences coevolution, but this effect may vanish when gene flow favors trait matching. Intimate mutualisms, such as protection of host plants by ants, form small and compartmentalized networks that generate higher trait matching than large and nested networks typical of mutualisms among free-living species, such as pollination. Third, habitat fragmentation resulting in the disruption of gene flow should reduce the reciprocal adaptations between interacting species and at the same time promote adaptations to the local abiotic environment. In conclusion, we show that a complex interplay between gene flow, the geographic structure of selection, and the organization of ecological networks shapes the evolution of large groups of species. Our results therefore allow predictions of how environmental impacts such as habitat fragmentation will modify the evolution of species interactions / Interações ecológicas como predação, competição e mutualismo são importantes forças que influenciam a evolução de espécies. Chamamos de coevolução a mudança evolutiva recíproca em espécies que interagem. A Teoria do Mosaico Geográfico da Coevolução (TMGC) fornece um arcabouço teórico para entender como conjuntos de populações coevoluem ao longo do espaço. Dois aspectos fundamentais da TMGC são o fluxo gênico entre populações e a presença de mosaicos de seleção, isto é, conjuntos de locais com regimes de seleção particulares. Diversos estudos exploraram como o acoplamento entre fenótipos de diferentes espécies evolui em pares ou pequenos grupos de espécies. Entretanto, interações ecológicas frequentemente formam grandes redes que conectam dezenas de espécies presentes em uma comunidade. Em redes de mutualismos, por exemplo, a organização das interações pode influenciar processos ecológicos e evolutivos. Um próximo passo para a compreensão do processo coevolutivo consiste em investigar como aspectos da TMGC influenciam a evolução de espécies em redes de interações. Nesta dissertação, tentamos preencher esta lacuna usando um modelo matemático de coevolução, ferramentas de redes complexas e informação sobre redes mutualistas empíricas. Nossas simulações numéricas do modelo coevolutivo apontam para três principais conclusões. Primeiro, o fluxo gênico influencia os padrões fenotípicos gerados por coevolução e pode favorecer a emergência de acoplamento fenotípico entre espécies dependendo do mosaico de seleção. Segundo, a organização de redes mutualistas influencia a coevolução, mas este efeito pode desaparecer quando o fluxo gênico favorece acoplamento fenotípico. Mutualismos íntimos, como proteção de plantas hospedeiras por formigas, formam redes pequenas e compartimentalizadas que geram um maior acoplamento fenotípico do que as redes grandes e aninhadas típicas de mutualismos entre espécies de vida livre, como polinização. Por fim, a fragmentação de habitat, ao extinguir o fluxo gênico, pode reduzir as adaptações recíprocas entre espécies e ao mesmo tempo tornar cada espécie mais adaptada ao seu ambiente abiótico local. Em suma, mostramos que interações complexas entre fluxo gênico, estrutura geográfica da seleção e organização de redes ecológicas moldam a evolução de grandes grupos de espécies. Dessa forma, podemos traçar previsões sobre como impactos ambientais como a fragmentação de habitat irão alterar a evolução de interações ecológicas

A restauração da Mata Atlântica tem sido considerada prioridade nas iniciativas de manutenção da biodiversidade. Adicionalmente, há consenso de que os parâmetros para avaliação da restauração ecológica devem mensurar o retorno de funções ecológicas. O estudo de interações planta-visitante floral pode ser um caminho adequado para avaliar a eficiência das práticas de restauração, visto que estas interações desempenham função crítica na dinâmica e diversidade da comunidade. Variações na diversidade de espécies de plantas e de seus visitantes florais podem alterar a freqüência de interação entre as espécies, definir a estrutura das redes de interação, determinando os níveis de generalização e especialização na comunidade. Neste contexto, a tentativa de restaurar florestas tropicais pela adição de espécies arbóreas pode ter efeitos sobre a estrutura, estabelecimento de grupos funcionais e níveis de generalização na rede de interação entre flores e visitantes florais. O objetivo principal deste trabalho é o de comparar redes de interação planta-visitante floral em florestas tropicais restauradas após 5 anos do plantio das arbóreas, florestas regeneradas naturalmente e remanescentes de floresta atlântica em uma área sob domínio da Mata Atlântica no sudeste do Brasil. Para atingir esse objetivo, essas florestas foram comparadas quanto suas diversidades estruturais e funcionais em relação aos seguintes aspectos: 1) Riqueza e atributos de história de vida (formas de vida, sistemas sexuais, modos de polinização e de dispersão); 2) redes de interação plantavisitante floral; 3) Grau de generalização e especialização das redes de interação; 4) robustez quanto à perda de espécie em redes de interação, e 5) Formação de grupos funcionais seguindo características florais e de freqüência de visitas. Para cada aspecto avaliamos a contribuição das espécies plantadas. Florestas restauradas tiveram a maior riqueza de espécies em flor, porém com menor similaridade florística com outras florestas locais. A similaridade em abundâncias relativas de arbustos e lianas com outras categorias de florestas indicou a inclusão de outras formas de vida além de árvores nas florestas restauradas. Porém, a alta abundância relativa de árvores nas florestas regeneradas naturalmente também indicou o potencial de regeneração natural em florestas 15 degradadas. A maior diversidade de modos de polinização biótica e de dispersão de sementes nas florestas restauradas veio das plantas regenerantes espontaneamente. Não houve diferenças significativas quanto às métricas de redes de interação flores e visitantes entre os tratamentos, porém houve uma tendência de maior especialização dessas interações nas florestas nativas e maior robustez à perda de espécies em florestas restauradas. Além disso, plantas regenerantes espontaneamente receberam significantemente mais visitas nas florestas regeneradas naturalmente do que em florestas restauradas, sugerindo que árvores plantadas podem estar reduzindo visitação às flores da vegetação regenerante espontânea, possivelmente competindo por visitantes florais. Em relação à diversidade funcional, 21 grupos funcionais baseados em atributos florais foram estabelecidos entre todas as espécies em flor, onde as espécies da floresta restaurada dominaram três grandes grupos e a floresta nativa apresentou representantes distribuídos equitativamente pelos grupos, sem dominância. Pólen foi a variável que mais contribui para diferenciação dos grupos. As espécies plantadas formaram grupos funcionais exclusivos nas florestas restauradas, contribuindo para uma maior diversificação em atributos funcionais florais em tais comunidades, porém não mais do que a diversificação funcional trazida pelas plantas regenerantes espontaneamente. Redes de interação entre grupos funcionais de plantas e categorias taxonômicas de visitantes reforçaram que os visitantes florais parecem não seguir fielmente grupos funcionais por atributos florais. Considerando que as florestas regeneradas naturalmente apresentaram alta abundância relativa de árvores, não apresentaram diferenças significativas quanto às métricas de redes de interação planta-visitantes florais com as florestas restauradas e que a regeneração natural na região estudada ocorre em grande intensidade, sugerimos que seja dada importância relevante às plantas regenerantes espontaneamente em projetos de restauração. Cabe ressaltar que avaliamos restauração após 5 anos da implantação. Assim, todas as conclusões tiradas deste estudo necessitarão ser acompanhada em estudos futuros. / Restoration of the Brazilian Atlantic Forest has been considered priority in initiatives to maintain biodiversity. Additionally, there is consensus that the parameters to evaluate restoration should address the return of ecological processes. The study of flower-visitor interactions can be a reasonable way to evaluate restoration practice, considering that these interactions have critical role in the dynamics and diversity of communities. Variations in the diversity of plant species and their flower visitors could modify frequency of interactions between species; define the structure of interaction networks, and determine generalization and specialization levels in the community as well. In this context, the attempt to restore tropical forests by planting native trees can affect the structural and functional diversity and generalization level in flower-visitor networks. The main objective of this research is to compare flower-visitor networks in 5-year-old restored forests, naturally regenerated forests and native forests in an Atlantic Forest domain in southeastern Brazil. We compared these forests in relation to: 1) Richness of species and life history traits (growth form, sexual system, biotic pollination modes and dispersal modes); 2) Flower-visitor networks; 3) Generalization and specialization levels in ecological networks; 4) Robustness to species loss in ecological networks; and 5) Functional groups by floral traits and visitation frequencies of flower visitors. We evaluated the contribution of planted species on each of these aspects. Restored forests had the highest floristic richness of species in flower, but little floristic similarity with other native local forests. Similarity in the relative abundance of shrubs and lianas among habitat categories indicated the possibility of annexation of other life forms than trees in restored forests. But the presence of high relative abundance of trees in the naturally regenerated forests also indicated the potential of natural regeneration of the degraded forests. Biotic pollination and dispersal modes tended to be more diverse in restored forests, but it comes as a result of the addition of spontaneously regenerated plants to this forest. There were no significant differences in the metrics of flower-visitor networks between forest categories, although there was a trend towards high specialization of 17 interactions between flower and visitors in native forests and high robustness of species loss in restored forests. In addition, spontaneously regenerated plants received significantly more visits in the naturally regenerated forests than in restored forests, suggesting that the planted trees may reduce the visitation to the spontaneously regenerated vegetation, possibly by competing for flower visitors. With regard to functional diversity, 21 functional groups based on floral traits were recognized when all species in flower was pooled. Species of restored forests were dispersed mainly among three groups, while species from native forests were spread among all groups with almost the same number of species per group. Pollen was the variable that most contributed for grouping species. Planted trees species formed exclusive functional groups, contributing for higher diversification of floral trait to the community. However, this diversification was not higher than provided by spontaneous regenerated plants. Interaction networks between plant functional groups and taxonomic categories of flower visitors ensured that flower visitors do not seem to follow the grouping formed by floral traits. Considering that naturally regenerated forests had high relative abundance of trees, were not different from restored forests in relation to network metrics and that natural regeneration was intense in the region, we suggest paying relevant attention to spontaneous regenerated plants in restoration projects. We would like to point out that we evaluated five-year-old restored forests and there is still need to track these forests in the future.

Bacteria are immensely diverse and hold key-positions in essentially all biogeochemical cycles. In freshwater ecosystems, bacteria degrade and mineralize organic compounds, linking the pool of dissolved organic matter to higher trophic levels. Aware of the global biodiversity loss, ecologists have started identifying the relationship of diversity and ecosystem functioning. Central to this is the question if species can functionally replace other species, hence being functionally redundant. Functional redundancy might allow communities to maintain functioning when diversity is lost. Due to their large numbers and great diversity, bacterial communities have been suspected to harbor large amounts of redundancy. The central aim of this thesis is to investigate the coupling of diversity and ecosystem functioning of bacterial communities and to understand how environmental perturbation affects this relationship. I manipulated the diversity of complex communities by a dilution technique, and measured the performance of bacterioplankton and biofilm-forming communities at different diversities. Reduction of bacterial diversity differently affected different functions, and that the presence or absence of certain species might be causing this pattern. However, for ecosystems to function, the interplay of multiple functions, i.e. multifunctionality, has to be sustained over long periods of time. In bacterial biofilm communities reduced diversity affected multifunctionality, as reflected by extracellular enzyme activities. A continuous cultivation system was used to address the importance of diversity for resistance and resilience upon environmental perturbation. The analysis of co-occurrence of bacterial taxa showed that the communities form a dense network before the perturbation and that these patterns are disturbed by the environmental perturbation. The final chapter of the thesis presents experimental evidence for the positive effects of temporal and spatial refuges for bacterial communities and the functions they provide. Overall, I found several indications for a lower amount of functional redundancy as previously assumed and it becomes apparent from this thesis that a multifunctional perspective and the consideration of environmental heterogeneity is pivotal.