Metacommunity ecology explicitly incorporates processes at multiple spatial scales to explain the assembly and dynamics of a community. In a metacommunity, local communities interact with one another through the dispersal of individuals across a region. As such, metacommunities are molded by two sets of processes: local and regional. Local factors are those that directly impact a single local community, such as environmental conditions, competition, and predation. On the other hand, regional factors affect communities across a landscape and include mechanisms such as, immigration and emigration. The potential interactions between local and regional factors make metacommunity dynamics a unique body of theory when compared to classic community theory. However, while the direct influence of dispersal on metacommunity dynamics continues to be a well-researched topic, how dispersal interacts with local factors to shape metacommunity dynamics is a more open topic. In particular, one continuing gap in my knowledge is how dispersal interacts with biotic effect how it may affect metacommunities. One type of local biotic process that can directly affect communities is a strong interactive species, i.e., a species that affects community structure and diversity, and to the best of my knowledge, the interaction between dispersal and strongly interactive species has not been directly addressed experimentally. In the following study, I investigated the interaction of dispersal and a strong interactive species on metacommunity diversity and assembly. I chose Daphnia magna as my strong interacting species due to its biological and physical traits. Dispersal is known to create predictable patterns of diversity as it increases in a metacommunity. We made logical predictions based off of my knowledge of these patterns, and my inclinations regarding how dispersal would interact with a strong interacting species. The following predictions were made in relation to the control: 1) Alpha diversity would be the highest during low dispersal as new species would be introduced and maintained above the extinction threshold. I also predicted beta diversity would decrease with increased dispersal due to the homogenization of communities. 2) In the presence of D. magna, beta diversity would only increase during low dispersal due to possible rescue effects. 3) Temporal variability would decrease for the low dispersal treatment and increase for the high dispersal treatment in the absence of D. magna. 4) Temporal variability would overall increase across all treatments in the presence of D. magna. To carry out the study, I assembled outdoor mesocosms using a 2x3x3x4 factorial design (Daphnia Treatment: no addition of D. magna, addition of D. magna; Dispersal Treatment: no dispersal, low dispersal, high dispersal; three buckets were equivalent to one metacommunity; 4 replicates). There was a significant interaction between D. magna and dispersal. Over time, beta diversity decreased as communities became homogenized; however, the no dispersal treatment homogenized at a slower rate compared to the other treatments. In addition, D. magna appeared to create local selection for certain taxa resulting in the increase of Bosmina and Simocephalus while other taxa decreased, for example Streblocerus. This trend was likely due to the feeding and grazing habits of D. magna which is known to outcompete other large zooplankton for larger phytoplankton taxa. Lastly, D. magna directly influenced temporal variability of metacommunities in the experiment. In particular, the low dispersal treatment increased in temporal variability in the presence of D. magna. Again, this result could likely be attributed to D. magna effects selecting for certain taxa, or by the re-introduction of new or dying species with each dispersal through rescue effects. Overall, the results in my study supported majority of my predictions. It is clear that D. magna had an effect on communities as taxa abundances increased and beta diversity in the no dispersal treatment did not decrease as quickly. This result suggests that the introduction of D. magna as an invasive to non-local waters could pose a threat to local community dynamics. It is important to understand how a strong interactive species can affect communities across a landscape as they can greatly alter diversity and composition. Future studies should focus on expanding the dispersal gradient and incorporating a local strong interactive species and non-local strong interactive species to understand how they may change community dynamics. / Master of Science

Biogeographic patterns across a landscape are developed by the interplay of environmental processes operating at different spatial and temporal scales. This research investigated dynamics of salt marsh vegetation on the Skallingen salt marsh in Denmark responding to environmental variations at large, medium, and fine scales along both spatial and temporal spectrums. At the broad scale, this research addressed the importance of wind-induced rise of the sea surface in such biogeographic changes. A new hypothetical chain was suggested: recent trends in the North Atlantic Oscillation index toward its positive phase have led to increased storminess and wind tides on the ocean surface, resulting in increased frequency, duration, and magnitude of submergence and, hence, waterlogging of marsh soils and plants, which has retarded ecological succession. At the mid-scale, spatial patterns of vegetation and environmental factors were examined across tidal creeks. Sites closer to tidal creeks, compared to marsh interiors, were characterized by the dominance of later-successional species, higher bulk density, and lower nutrient contents and electrical conductivity. This finding implies that locations near creeks have experienced a better drainage condition than the inner marshes, which eventually facilitated the establishment of later-successional plants that are intolerant to physical stress. At the micro-scale, this research examined how the extent and mode of facilitation and competition vary for different combinations of plant species along physical gradients. Both positive and negative relationships were spatially manifested to a greater degree on the low marsh than on the mid marsh. This insight extends our current knowledge of scale-dependent interactions beyond pioneer zones to higher zones. On the low marsh, different types of bivariate point pattern (i.e., clustered, random, and regular) were observed for different combinations of species even at similar spatial scales. This finding implies that it is difficult to generalize at which scales competition and facilitation occur. To conclude, this research stresses the need for a holistic approach in future investigations of salt marsh biogeography. For example, based on results of this current research, it would be meaningful to develop a comprehensive simulation model that incorporates salt marsh ecology, geomorphology, and hydrology observed across scales.

Invasive species are alien to the ecosystem under consideration and cause economic or environmental damage or harm to human health. Two alien insects that fit this description are the brown marmorated stink bug, Halyomorpha halys and the spotted lanternfly, Lycorma delicatula. Both invaders are polyphagous pests that feed on a myriad of plant species and inflict severe crop losses. As sustainable control methods depend on the accurate monitoring of species’ invasion and involve the use of natural enemies, we addressed these two facets by exploring novel monitoring techniques and deciphering host-parasitoid interactions for improved integrated pest management. Thus, we adopted ‘BugMap’, a citizen science initiative that enables students, farmers and everyday citizens to report sightings of H. halys from Italy, with emphasis on Trentino-Alto Adige. Aside from fostering citizen participation in scientific endeavors and the enhanced literacy that ensues, BugMap helped uncover the invasion dynamics of H. halys and forecast its potential distribution in Trentino, all while coordinating technical monitoring and informing management strategies. The most promising agent currently under study for the classical biological control of H. halys is the Asian egg parasitoid Trissolcus japonicus. To assess the wasp’s potential non-target impacts, we investigated its foraging behavior in response to chemical traces ‘footprints’ deposited by its main host H. halys and by a suboptimal predatory species, the spined soldier bug, Podisus maculiventris. Wasps exhibited a ‘motivated searching’ when in contact with footprints originating from both species. However, T. japonicus arrestment was significantly stronger in response to H. halys footprints, compared with P. maculiventris, implying the presence of underlying chemical cues that shape its natural preferences. A series of GC-MS chemical analyses revealed that n-tridecane and (E)-2-decenal were more abundant in H. halys footprints and are probably the key components utilized by the wasp for short range host location. The function of the aforementioned compounds was studied, n-tridecane acted as an arrestant, prolonging T. japonicus residence time, whereas (E)-2-decenal fulfilled its presumed defensive role and repelled the wasp. These results shed new light on the chemical ecology of T. japonicus and help expand the understanding of parasitoid foraging and its implications for classical biological control. Moving to the other invader L. delicatula, an egg parasitoid Anastatus orientalis was reported attacking it at high rates in its native range in Eastern Asia and may play a key role in reducing its populations there. A series of bioassays revealed that wasps responded to footprints deposited by L. delicatula gravid females by initiating a strong searching behavior. Moreover, A. orientalis preferred to oviposit in egg masses with intact oothecae, suggesting that the host’s egg covering functions as a trigger for A. orientalis probing and oviposition. Thus, A. orientalis not only overcomes, but also reverses an important line of host structural defense for its own fitness gains. This dissertation discusses the benefits of combining citizen science with traditional monitoring, and the usefulness of decoding host-parasitoid interactions to design more efficacious management strategies of invasive insect pests.

In this paper, we develop a model for predation movements of a subsocial spider species, Anelosimus studiosus. We expand on a previous model to include multiple spider interaction on the web as well as a latency period during predation. We then use the model to test different spatial configurations to determine the optimal spacing of spiders within a colony for successful capture during predation. The model simulations indicate that spiders uniformly spacing out along the edge of the web results in the most successful predation strategy. This is similar to the behavior observed by Ross (2013) in which it was determined to be statistically significant that during certain times of the day, spiders were positioned along the edge more than expected under complete spatial randomness.

Environmental change is a major threat to marine ecosystems worldwide. Understanding the key biological processes and environmental factors mediating spatial and temporal species' responses to habitat alterations underpins our ability to forecast impacts on marine ecosystems under any range of scenarios. This is especially important for calcifying species, many of which have both a high climate sensitivity and disproportionately strong ecological impacts in shaping marine communities. Although geographic patterns of calcifiers' sensitivity to environmental changes are defined by interacting multiple abiotic and biotic stressors, local adaptation, and acclimation, knowledge on species' responses to disturbance is derived largely from short- and medium-term laboratory and field experiments. Therefore, little is known about the biological mechanisms and key drivers in natural environments that shape regional differences and long-term variations in species vulnerability to global changes. In this thesis, I examined natural variations in shell characteristics, both morphology and biomineralisation, under heterogeneous environmental conditions i) across large geographical scales, spanning a 30° latitudinal range (3,334 km), and ii) over historical times, using museum collections (archival specimens from 1904 to 2016 at a single location), in mussels of the genus Mytilus. The aim was to observe whether plasticity in calcareous shell morphology, production, and composition mediates spatial and temporal patterns of resistance to climate change in these critical foundation species. For the morphological analyses, the combined use of new statistical methods and multiple study systems at various geographical scales allowed the uncoupling of the contribution of development, genetic status, and environmental factors to shell morphology. I found salinity had the strongest effect on the latitudinal patterns of Mytilus shape. Temperature and food supply, however, were the main predictor of mussel shape heterogeneity. My results suggest the potential of shell shape plasticity in Mytilus as a powerful indicator of rapid environmental changes. I found decreasing shell calcification towards high latitudes. Salinity was the best predictor of regional differences in shell deposition, and its mineral and organic composition. In polar, low-salinity environments, the production of calcite and organic shell layers was increased, while aragonite deposition was enhanced under temperate, higher-salinity regimes. Interacting strong effects of decreasing salinity and increasing food availability on compositional shell plasticity predict the deposition of a thicker external organic layer (periostracum) at high latitudes under forecasted future conditions. This response potential of Mytilus shell suggests an enhanced protection of temperate mussels from predators and a strong capacity for increased resistance of polar and subpolar individuals to dissolving water conditions. Analyses of museum specimens indicated increasing shell calcification during the last century. Deposition of individual shell layers was more closely related to temporal changes in the variability of key environmental drivers than to alterations of mean habitat conditions. Calcitic layer and periostracum showed marked responses to alterations of biotic conditions, suggesting the potential of mussels to trade-off between the deposition of calcareous and organic layers as a compensatory response to strategy-specific predation pressure. These changes in biomineralisation indicated a marked resistance to environmental change over the last century in a species predicted to be vulnerable, and how locally heterogeneous environments and predation levels can have a stronger effect on Mytilus responses than global environmental trends. My work illustrates that biological mechanisms and local conditions, driving plastic responses to the spatial and temporal structure of multiple abiotic and biotic stressors, can define geographic and temporal patterns of unforeseen species resistance to global environmental change.

Previous studies suggest that global warming threatens populations of Arctic char (Salvelinus alpinus L.) and that interspecific competition from brown trout (Salmo trutta L.) can have negative effects when the species occur in sympatry. The aim of this study was to investigate if populations of Arctic char have declined in the last decades in lakes where they in the past have coexisted with brown trout. I examined this by revisiting five lakes previously sampled in the mountain region of Västerbotten (Sweden) during the 1960´s. This study repeated earlier sampling methods and examined the population size, biomasses and individual size and weight of all fish. Results showed increased abundances of brown trout and a trend towards declining Arctic char populations in the five lakes. Both Arctic char and brown trout currently had higher mean lengths than previous sampling years. This study also examined the diet and growth rate of both species. The diet analysis showed that the species to some extent overlapped in diet, while they also showed clear differences, where Arctic charr e.g. consumed more pelagic zooplankton. The growth rate was higher for Arctic char and differed between lakes. The shift towards more brown trout dominated lakes could depend on both that the two species are differently affected by a warmer climate, and that fishing habits have changed since the 1960´s. A warmer climate likely changed the species interaction in favor of trout and could lead to more competition and predation on Arctic char.

Complexity science has come into the limelight in recent years as the scientific community begins to grapple with higher-order natural phenomena that cannot be fully explained via the behaviour of components at lower levels of organization. Network modeling and analysis, being a powerful tool that can capture the interconnections that embody complex behaviour, has therefore been at the forefront of complexity science. In ecology, the network paradigm is relatively young and there remain limitations in many ecological network studies, such as modeling only one type of species interaction at a time, lack of realistic network structure, or non-inclusion of community dynamics and environmental stochasticity. I introduce bioenergetic network models that bring together for the first time many of the fundamental structures and mechanisms of species interactions present in real ecological communities. I then use these models to address some outstanding questions that are relevant to understanding ecological networks at the systems level rather than at the level of subsets of interactions. Firstly, I find that realistic red-shifted environmental noise, and synchrony of species responses to noise, are associated with increased variability in ecosystem properties, with implications for predictive ecological modeling which usually assumes white noise. Next, I look at simultaneous species extinction and invasion, finding that as their individual impacts increase, their combined impact becomes decreasingly additive. In addition, the greater the impact of extinction or invasion, the lesser their reversibility via reintroduction or eradication of the species in question. For modifications of pairwise species interactions by third-party species, a phenomenon that has so far been studied one interaction at a time, I find that the many interaction modifications that occur concurrently in a community can collectively have systematic effects on total biomass and species evenness. Finally, examining a higher level of organization in the form of compartmentalized networks, I find that the relationship between intercompartment connectivity and the impacts of species decline depends considerably on network topology and whether the consumer-resource functional response is prey- or ratio-dependent. Overall, the results vary considerably across model communities with different parameterizations, underscoring the contingency and context dependence of nature that scientists and policy makers alike should no longer ignore. This work hopes to contribute to a growing multidisciplinary understanding, appreciation and management of complex systems that is fundamentally transforming the modern world and giving us insights on how to live more harmoniously within our environment.

La biodiversité favorise un grand nombre de fonctions et services écosystémiques des écosystèmes forestiers tels que la production de bois ou la résistance aux attaques d’insectes et aux maladies. Cependant l’impact de la diversité sur l’acquisition et l’utilisation de l’eau et du carbone reste largement méconnu dans ces écosystèmes. De plus, dans le contexte actuel de changement climatique, l’influence de la diversité sur la réponse des écosystèmes forestiers à des événements climatiques extrêmes tels que la sécheresse reste à étudier. L’objectif de ce travail est donc de déterminer l’impact de la diversité en espèces d’arbre sur d’importantes fonctions du cycle de l’eau et du carbone telles que la transpiration, la composition isotopique du carbone ou la profondeur d’extraction de l’eau à l’échelle de l’arbre et de l’écosystème sous des conditions contrastées dedisponibilité en eau du sol. Ce travail a été réalisé dans le cadre du projet FunDivEUROPE sur un réseau de parcelles forestières ainsi que dans des plantations expérimentales le long d’un gradient Nord-Sud en Europe afin de couvrir une importante gamme de conditions climatiques. Nos travaux ont montré une importante variabilité de la réponse à la diversité à l’échelle de l’arbre et de l’écosystème en termes de relations hydriques et carbonées à travers l’Europe. La diversité en espèces ne semble pas influencer les relations hydriques et carbonées des espèces et des écosystèmes forestiers dans des conditions non limitantes de disponibilité en eau. Cependant, un fort effet de la diversité a été observé en conditions de sécheresse pour certains types forestiers. A partir de ces résultats, je discute des mécanismes d’interaction entre espèces qui peuvent expliquer les effets observés. Nos données ont montré que l’influence de la diversité en espèces est fortement dépendante du contexte et peut êtremodifiée par les conditions environnementales locales et les conditions climatiques. En terme de gestion forestière, je suggère que pour certaines régions en Europe, promouvoir la diversité en espèces ainsi que contrôler la densité des parcelles doit être recommandé afin d’adapter les écosystèmes forestiers aux futures conditions climatiques / Biodiversity is known to support and boost a wide range of forest ecosystem functions and services like productivity and resistance against insect pests and diseases. However, whether tree species diversity also promotes water and carbon acquisition and use in forest ecosystems is still unclear. Furthermore, in the current context of global warming, information on how tree species diversity can influence the response of forest ecosystems to extreme climatic events such as drought are urgently needed. In this framework, the objective of my PhD thesis was to determine how tree species diversity influences important functions of the water and carbon cycle including transpiration, carbon isotope composition and water extraction depth at the tree- and ecosystem-Scale under contrasting soil water conditions. My work was conducted within the FunDivEUROPE project in a network of permanent forest stands and tree plantations across a North-South gradient in Europecovering a wide range of climatic conditions. I found considerable variability among species or forest types in the response of transpiration and carbon isotope composition at the tree- and ecosystem-Scale across Europe. Species diversity did not affect the water and carbon relations of tree species and forest ecosystems under non-Limiting soil water conditions. However, a strong effect of species diversity was observed under drought conditions in some forest types. Based on these data, I discuss the potential mechanisms of species interactions that may explain the observed patterns. I also point out that the influence of species diversity is highly context-Dependent, and changes with local environmental and climatic conditions. In terms of forest management applications, I suggest that, at least in some regions, controlling for tree species diversity along with stand density and total basal area could be recommended to help forests adapt to drier conditions

Bats occupy diverse and unique niches and are regarded as important components in maintaining ecosystem health. They are major consumers of nocturnal insects, serve as pollinators, seed disperser, and provide important economic benefits as consumers of agricultural and forest pest insects. Bats have been proposed as good indicators of the integrity of natural communities because they integrate a number of resource attributes and may show population declines quickly if a resource attribute is missing. Establishing community- and population-level data, and understanding species interactions is especially important in changing landscapes and for species whose populations levels are threatened by outside factors of anthropomorphic disturbance from hibernacular visitation to energy production and fungal pathogens. For these reasons I have set out to establish habitat use patterns, detection probabilities, spatial and temporal occupancy, and investigate species interactions. This thesis is broken down into three distinct chapters each intended to be a stand-alone document. The first establishes the basic ecology from natural history accounts, provides an overview of the various sampling strategies, and gives a comprehensive description of the study area. The seconds sets out to identify the factors influencing detection probabilities and occupancy of six sympatric bats species and provide insight into habitat use patterns. The third examines spatial and temporal activity patterns and investigates species interactions. This study can provide understanding into the secretive and poorly understood patterns of free flying bats across the landscape. It can also deliver useful information to land managers regarding potential changes in landscape practices for the conservation of bat species. / Master of Science

Thesis (PhD)--Stellenbosch University, 2011. / ENGLISH ABSTRACT: Both taxonomic diversity and diversity of species interaction networks are experiencing declines as a result of agricultural intensification at habitat and landscape scales. Reversing this trend is a key conservation issue, particularly for important functional groups such as flower-visiting insects and the networks within which they interact. This is of great concern in regions of high conservation priority such as the Cape Floristic Region (CFR), known for its high level of floral and faunal endemism and exceptional species turnover. Holistic approach to conservation in agricultural landscapes involves both preservation of natural land and wildlife friendly management of the farm land to achieve conservation targets. The value of these extensive management approaches is yet to be fully assessed, especially in perennial systems such as vineyards. I examined here the effects of vineyard management and landscape context on species richness and abundance of flower-visiting insects and their species interaction networks. Possible taxon specific effects were verified. I also investigated whether vineyards under organic and conventional management homogenized insect-flower interaction networks and whether vineyards with different management practices vary in patterns of species turnover. I sampled flower-visiting insects and their interactions in organic and conventional vineyards, and in natural reference sites. Inclusion of natural reference sites enabled me to make management recommendations for patches of natural vegetation in CFR agricultural landscape. Statistical models showed taxon-specific benefit of organic farm management, and of landscape (distance to natural habitat). There was benefit to monkey beetles (Scarabaeidae) but not to bees (Apidae). Organic vineyards had a higher number of insect-flower interactions than conventional ones, but vineyards under the two types of management were similar in terms of other important network indices. However, networks of the vineyards were more nested than the natural sites, indicating that they may be potentially more stable to perturbation and random extinctions. Multivariate dispersion tests revealed insect-flower interaction networks were not homogenized by both organic and conventional vineyards across the landscapes. I also found, through additive partitioning, that organic and conventional vineyards were similar in terms of species turnover of bees and flowering plants. The findings of this study provide heuristic value to current debates on the value of vineyard habitats for insect conservation. Both organic and conventional vineyards that promote sustainable management of the non-crop floral vegetation between vineyard rows are potential solutions for conservation of flower-visiting insects and their interactions. Also, attention has to be paid to the quality and connectivity of the natural habitat patches that are within CFR agricultural landscape. Site specific management and assessment of the value of these landscape elements is important. Management approaches such as carefully controlled burning may be beneficial, as the CFR natural vegetation is a fire-driven community. / AFRIKAANSE OPSOMMING: Taksonomiese diversiteit en diversiteit van spesies-interaksie netwerke ondervind beide afnames as gevolg van landboukundige intensifikasie op habitat en landskap skaal. Om die neiging terug te swaai, is ’n sleutel bewaringsaangeleentheid, veral vir belangrike funksionele groepe soos blom-besoekende insekte en die netwerke waarbinne hulle op mekaar inwerk. Dit is van groot kommer in streke met hoë bewaringsprioriteite soos in die Kaapse Floristiese Streek (KFS), bekend vir sy hoë vlak van plant- en dierendemisme en buitengewone spesies kentering. ’n Holistiese benadering tot bewaring van landboukundige landskappe behels beide die bewaring van natuurlike land en natuurlewe-vriendelike bestuur van die plaasgrond om bewaringsdoelwitte te bereik. Die waarde van hierdie ekstensiewe bestuursbenaderings moet nog volledig bepaal word, veral in meerjarige sisteme soos wingerde. Ek het die uitwerkings van wingerdbestuur en landskapsamehang op spesiesrykheid en volopheid van blombesoekende insekte en hulle spesies interaksie netwerke ondersoek. Moontlike takson-spesifieke uitwerkings is nagegaan. Ek het ook ondersoek ingestel of wingerde onder organiese en gebruiklike bestuur ooreenstemmende insek-blom interaksie netwerke met wingerde met verskillende bestuurspraktyke in patroon van spesies kentering gewissel het. Ek het blom-besoekende insekte en hulle interaksies in organiese en konvensionele wingerde, asook in natuurlike verwysingsgebiede gemonster. Insluiting van natuurlike verwysingsgebiede het my in staat gestel om bestuursvoorstelle vir gebiede van natuurlike plantegroei in KFS landboulandskappe voor te stel. Statistiese modelle toon takson-spesifieke voordeel van organiese plaasbestuur en van die landskap (afstand van natuurlike habitat) self. Daar was voordeel vir bobbejaankewers (Scarabaeidae), maar nie vir bye (Apidae) nie. Organiese wingerde het ’n groter getal insek-blom interaksies as konvensionele wingerde gehad, maar wingerde onder beide tipes van bestuur was soortgelyk in terme van ander belangrike netwerk aanduiders. Netwerke van wingerde was egter meer geklomp dan natuurlike gebiede wat aandui dat hulle potensieel meer stabiel betreffende versteuring en lukrake uitsterwings is. Multivariate verspreidingstoetse het aangetoon dat insek-blom interaksie netwerke by beide organiese en konvensionele wingerde oor landskappe nie eenvormig was nie. Ek het ook bevind deur aanvullende verdeling dat organiese en konvensionele wingerde gelykwaardig was in terme van spesies kentering van bye en blomplante. Die bevindings van hierdie studie verskaf heuristise waarde tot huidige debatte oor die waarde van wingerdhabitatte vir insekbewaring. Beide organies en konvensionele wingerde wat volhoubare bestuur van die nie-gewas plantegroei binne wingerdrye bevorder, is moontlike oplossings vir die bewaring van blom-besoekende insekte en hulle wisselwerkings. Bykomend moet aandag gegee word aan die kwaliteit en verbindings van en tussen natuurlike habitat gebiede binne die KFS landboulandskap. Plekspesifieke (plaaslike) bestuur en bepaling van die waarde van hierdie landskapelemente is belangrik. Bestuursbenaderings, soos noukeurig beheerde brand, mag voordelig wees aangesien die KFS natuurlike plantegroei ’n vuurgedrewe gemeenskap is.