Thesis (M.Sc.) - Carleton University, 2006. / Includes bibliographical references (p. 65-76). Also available in electronic format on the Internet.

Thesis (M. Sc. (Zoology & Entomology))--Rhodes University, 2005.

<p>The introduction of the North American beaver (Castor canadensis) to Tierra del Fuego is a widely known example of a successful biological invasion. Beavers have impacted enormously the biodiversity of the island of Tierra del Fuego, and they are now spreading northward on the continent, prompting the governments of Argentina and Chile to seek methods to control their spread. Beavers first established in forests, where they were initially introduced, but by the 1990s they began to establish in the adjacent steppe. In this dissertation, I study the biology of invasive beavers across the two major habitat types in Patagonia and attempt to develop modeling tools that might be useful to manage their spread.</p><p>In chapter one I studied the history of the beaver introduction in Patagonia and provide evidence that the beaver introduction occurred as a single release event of 20 beavers from northern Manitoba, Canada. This not only clarifies the origin of the invasion, but also suggests that the beaver population of Patagonia descends from a smaller number of individuals than previously assumed.</p><p>In chapter two I studied the demography of invasive North American beavers in the two contrasting habitat types of the island of Tierra del Fuego, forest and steppe. Habitat differences can affect vital rates which may in turn impact the speed of the invasion, but this has been rarely addressed when managing the spread of invaders. I use repeated observations, mark-resight methods, telemetry and camera traps to estimate colony size and vital rates of beavers in the two habitats. Colony size and the number of offspring (“kits”) produced per colony per year were higher in the steppe, contrary to the belief that forest is better habitat. Here I suggest this may be the result of the longer time since invasion in the forests of Tierra del Fuego and that the forest subpopulation is showing density dependent regulation. Beaver survival was high in all age classes and was higher than survival rates recorded in North America. My work shows that beaver plasticity and predator release have likely facilitated the invasion in Patagonia. </p><p>In chapter three, I investigated the more recent invasion of beavers in an area of the Patagonian steppe. I utilized repeated high resolution satellite images to identify beaver ponds, and used them to study changes in beaver abundance and habitat use over time. The number of beaver ponds increased 85 % between 2005 and 2014. During this period, beavers changed their habitat selection pattern, presumably as a response to increased density. Beavers established on small watercourses in canyons first, but as more canyons became occupied over time, beavers moved to less preferred watercourses in plains and U-shaped valleys. Potential new beaver colonies established close to existing beaver ponds, suggesting proximity to a beaver pond is an important determinant of beaver colonization. Identifying habitat preferred by beavers in the steppe could help to increase early detection of the invader at the invasion front. This work highlights the importance of the use of high resolution remote sensing technologies to better understand and monitor biological invasions. </p><p>Finally, in chapter four, I built a spatially explicit individual-based model parameterized with data I collected in the field and use it to make management recommendations. Specifically I assessed the efficacy of a potential management strategy in which a “fire-break” (a zone beyond the current population front in which beavers are removed) perpendicular to the population front is instituted to attempt to prevent further northward spread of the beaver in continental Patagonia. I found that even a 100 km wide firebreak is insufficient to contain the spread of beavers, long dispersal events being the major cause of this failure. Further, I found that increasing the fraction of beavers culled within the firebreak does not decrease either the arrival time or the number of beavers that cross the firebreak. Counterintuitively, my model indicates that moderate levels of culling within the firebreak (rather than high) may be a more effective method to manage the invasion, likely as a result of inversely density dependent dispersal.</p> / Dissertation

O Cerrado é o bioma brasileiro que mais perdeu área para o agronegócio em anos recentes, verificando que a expansão da fronteira agrícola sobre o Cerrado provocou a pulverização dos remanescentes em fragmentos cada vez menores, imersos em matrizes de baixa similaridade, causando uma série de impactos negativos sobre o ecossistema natural. A fragmentação de habitats é considerada uma das mais graves ameaças à conservação da biodiversidade, com consequências ecológicas variáveis, intensificadas na faixa de transição entre matriz e fragmento. Os efeitos de borda são decorrentes de mudanças físicas e bióticas nas faixas marginais das ilhas de vegetação natural, que promovem modificações nas características microclimáticas, as quais conduzem a uma série de alterações importantes nas comunidades que compõem os ecossistemas. O objetivo deste estudo foi investigar a existência de efeito de borda e suas consequências sobre a comunidade vegetal do Cerrado stricto sensu. O estudo foi realizado em fragmento de Cerrado com área de 980,8 ha, localizado no município de Iaras, SP, cujo entorno foi ocupado por pastagem durante cerca de um século. Efetuou-se a caracterização estrutural e florística de diferentes estratos da vegetação em diferentes distâncias da borda (0, 10, 20, 40, 80 m) e coletaram-se dados microclimáticos na matriz (a 10 m de distância da borda) e no interior do fragmento, nas diferentes distâncias da borda e em duas estações do ano (seca e chuvosa). Investigou-se a existência de um possível gradiente de estrutura ou composição da vegetação relacionado com a distância da borda e com as variáveis microclimáticas. Os resultados obtidos mostraram não existir variações de microclima e de estrutura e composição do componente arbóreo e herbáceo-arbustivo em relação à distância da borda do fragmento. Embora independente do microclima e, portanto, diferente dos clássicos efeitos de borda, a invasão pela braquiária (Urochloa decumbens) foi detectada como a única consequência da exposição da periferia do Cerrado stricto sensu às pressões da matriz, causando exclusão das gramíneas nativas e inibindo o desenvolvimento de plantas lenhosas de pequeno porte. A ausência de efeito de borda sobre os demais componentes da vegetação sugere, em um primeiro momento, que esta não é uma ameaça importante para a conservação do cerrado. Porém, a invasão biológica pela gramínea africana é relativamente recente (foi introduzida há cerca de três décadas na propriedade) e suas consequências ainda não podem ser avaliadas adequadamente. É possível que seus impactos sobre os estratos superiores da vegetação venham a ser observados no futuro, quando a inibição da regeneração das espécies arbustivas e arbóreas comprometer a dinâmica da comunidade vegetal e a substituição dos adultos nas populações dessas espécies. Além disso, hoje a invasão só causa impactos significativos sobre a faixa de 0 a 20 m de distância da borda, mas não é possível prever se a gramínea invasora vai se expandir rumo ao núcleo do fragmento no futuro. / From the Brazilian biomes, the Cerrado has been the most widely and rapidly converted to agribusiness in recent years. The expansion of agricultural frontier over the Cerrado domain resulted in thousands of small fragments embedded in matrices of low similarity, causing a series of negative impacts on the natural ecosystem. Habitat fragmentation is considered one of the most serious threats to the conservation of biodiversity and ecological consequences vary, being intensified in the range of transition between matrix and fragment. The edge effects are caused by biotic and physical changes in marginal areas of the islands of natural vegetation, promoting changes in microclimatic characteristics, which leads to a series of important changes in the ecosystems. The aim of this study was to investigate the existence of edge effect and their consequences on the plant community of Cerrado sensu stricto. The study was conducted in Cerrado fragment with an area of 980.8 ha, located in the municipality of Iaras, SP, whose surrounding area was occupied by pasture for about a century. Structure and floristic composition of distinct vegetation layers was assessed at different distances from the edge (0, 10, 20, 40 and 80 m). Climatic data were collected in two seasons (dry and wet), at the same distances withing the fragment and also in the matrix (10 m away from the edge). The existence of a gradient of vegetation composition or structure and the microclimatic variables related to distance from the edge was investigated. No changes in microclimate and structure or composition of the wood layers related to distance from the edge was observed. Although independent of the microclimate and therefore different from classic edge effects, invasion by the African grass Urochloa decumbens (braquiária) was detected as the only consequence of exposure of the periphery of the cerrado sensu stricto to the pressures of the matrix, causing the exclusion of native grasses and inhibiting the development of small woody plants. The absence of edge effect on other components of the vegetation suggests, at first, that this is not a major threat to the conservation of the Cerrado. However, the biological invasion by the African grass is relatively recent (it was locally introduced about three decades ago) and its consequences have not been adequately assessed. It is possible that their impacts on the upper strata of the vegetation may be observed in future, when the inhibition of regeneration of shrub and tree species jeopardize the dynamics of plant community and the replacement of adults in the populations of these species. In addition, at present the invasion has significant impacts only over 0-20 m from the edge, but it is impossible to predict whether the invasive grass will expand or not into the core area of the fragment in the future.

The fire-prone mountain fynbos of the southwestern Cape Province of South Africa has been severely invaded by introduced trees and shrubs. These invasions have transformed fynbos shrublands to dense thickets of trees and shrubs in many parts of the region, thus disrupting various natural ecosystem processes. The ecology of invasions by species of Pinus and Hakea (the most successful genera) was studied using a series of natural experiments in conjunction with autecological studies. The study was divided into main four parts: (i) case studies to elucidate major patterns and processes of invasion; (ii) studies of the life history and population ecology of selected invaders; (iii) studies of the determinants of invasibility; and (iv) assessments of the consequences of invasion and of control programmes. Bibliography: pages 235-272.

Evolution of genotypes during range expansion is driven in part by colonization dynamics. I investigated genetic patterns of colonization and dispersal during initial expansion of an invasive bunchgrass, Brachypodium sylvaticum, into Oregon. Using microsatellite markers, I sampled plants at two different scales: at regular intervals along three parallel roads spanning about 30km, and in populations identified throughout Oregon. I also collected field-generated progeny from a subset of populations and used molecular identification of outcrossing events to estimate selfing rates in both central and peripheral populations. Dispersal patterns were similar at both scales, with non-contiguous dispersal responsible for colonization of new populations. High levels of differentiation were observed at all scales, though newly-colonized populations were more differentiated than older populations. Corvallis populations were responsible for colonization of a majority of populations throughout Oregon, while individuals from Eugene were only occasionally found in new populations. Admixture occurs between Corvallis and Eugene populations, decreasing differentiation, and potentially creating novel phenotypes and increasing evolutionary potential of populations. Selfing rates were high, but two populations in the areas of original introduction had lower rates of selfing, suggesting that selfing rates may decrease as population density and diversity increases with age. The influences of founder effects and bottlenecks on phenotypic evolution during range expansion require further investigation, as inbreeding, lag times, and selection may influence evolutionary trajectories of populations.

No description available.

Invasive species are of particular concern as they have the potential to alter community structure and food web relationships within their invaded habitats. African sharptooth catfish Clarias gariepinus, a generalist predator, was introduced through an inter-basin water transfer scheme into the Great Fish and Sundays Rivers, Eastern Cape, South Africa, where it threatens the native riverine biota. This thesis assessed its impact from a trophic perspective. Patterns in catfish distribution and abundance revealed an upstream to downstream gradient that was associated with spatial distribution of most species within the mainstream, and a mainstream to tributary gradient that was associated with the spatial distribution of native minnows. The catfish was predicted to occur widely within the mainstem habitats and to decrease progressively along the mainstrem to tributary gradient with the physico-chemical environment being a good proxy for predicting both its occurrence and abundance. The results suggest the catfish proliferated within mainstem habitats where invasion resistance was possibly reduced due to alteration of flow. Population dynamics and size structuring of two native cyprinid minnows Pseudobarbus afer and Barbus anoplus, threatened by catfish, were examined within uninvaded headwater streams in relation to their proximate physical habitats. Their habitats were characterised by seasonal variation in physico-chemical conditions and a spatial variation in substrata compositions. No evidence of differences was found between seasons for density and capture probability for either species. The population size and density for P. afer was found to increase with increasing proportion of boulders. In comparison, B. anoplus population size and probability of capture increased with increasing proportion of bedrock and bank vegetation, respectively. Size structuring was explained predominantly by seasonality and habitat variables for P. afer and B. anoplus, respectively. Stable isotope ratios of carbon and nitrogen were used to compare the spatial variation in both the community-wide and catfish-specific niches and to estimate catfish prey sources from different habitats within the invaded systems. Aquatic community and catfish niches were statistically different among localities, suggesting that each locality had a distinct community-wide trophic structure. Dispersion metrics indicated no evidence of differences in the clustering among individuals, but provided evidence of differences in path trajectories for the comparisons of catfish populations that suggested dietary plasticity within different localities. Dietary studies revealed both ontogenetic shift and omnivory that suggested that catfish may exhibit less pronounced top-down effects within its invaded habitats. Manipulative experiments were used to test the response of benthic macroinvertebrates within two rivers that were differentially impacted by catfish as a presstype disturbance. Macroinvertebrates were non-responsive to catfish presence within a system where catfish had previously been established. In contrast, excluding catfish in this system indicated a response that suggested the importance of refuge within invaded habitats and the possible recovery pattern of certain macroinvertebrate taxa. By comparison, introduction of catfish within previously uninvaded localities provided evidence of direct catfish impact through elimination of conspicuous taxa. Acoustic telemetry was used to investigate catfish movement patterns within an invaded lentic habitat and provided evidence that habitat utilisation was non-random. The shallow and structured river mouth habitat, which was most utilised, was probably the most ideal for its breeding and feeding. This inferred potential overlap with native species and suggested the risk of predation and competitive interference. Catfish also exhibited both nocturnal and diurnal activity patterns that were probably related to feeding.

Earth mean surface temperature has increased by 1 °C since the industrial revolution, and this has already had considerable effects on animal and plant species. Ecological responses to the warming climate – often facilitated via phenotypic plasticity – are ubiquitous. However, even though evolution can occur rapidly there are only few examples of genetic adaptation to climate change. In my thesis, I used a near-natural system to study if and how organisms have adapted to 30 years of warming, and how this has affected competitive species interactions. I investigated Baltic Sea populations of the aquatic snails Galba truncatula and Theodoxus fluviatilis, which had been subjected to cooling water discharge from power plants, resulting in water temperatures 4 to 10 °C higher than in the surrounding sea. G. truncatula had high upper thermal limits and large acclimation potential. This plasticity may have helped the species to survive under the new conditions, allowing evolution through natural selection to take place. I found that the populations of the two thermal origins had diverged in SNP markers associated with warmer temperature, whereas divergence in selectively neutral markers was mainly related to geographical distance. Adaptation occurred from standing genetic variation, emphasizing the importance of genetic diversity and population size in enabling the persistence of populations. Changes in thermal sensitivity of growth and survival were subtle yet significant, and complied with theoretical models of thermal adaptation in ectotherms. At the community level, pre-adaptation to warmer conditions aided the native T. fluviatilis when competing with the alien Potamopyrgus antipodarum. However, interspecific competition limited the snails most in those traits favored under warming, highlighting the challenge of adapting to different selecting forces during global change. The persistence of species and populations under climate change depends on several factors - plasticity allowing for initial survival, evolvability in allowing the genetic changes, and species interactions affecting the new ecological niches. The results of my thesis indicate that persistence under climate change is possible when these factors align, but the relative roles of ecology and plasticity may explain why there are so few observed instances of evolution in response to climate change.

Thesis (PhD)--Stellenbosch University, 2012. / ENGLISH ABSTRACT: Large-scale human-mediated movements of organisms promote the establishment of species outside their native ranges and a very small proportion of these species become invasive. Invasive species management typically assumes that introduced species are single, static evolutionary units that are genetically analogous to their native counterparts. However, studies have shown that native and introduced populations of a number of introduced plants differ vastly in their genetic composition. These differences may negatively affect the overall success of control and management programmes, particularly for species that are intra-specifically diverse. The influence of intra-specific diversity on the invasion process was tested in two widely exported tree species that are native to Western Australia, Acacia saligna (three subspecies) and Paraserianthes lophantha (two subspecies). Climate matching between the native and introduced range (using species distribution models, SDM) is widely used to forecast future invasion risks, however, it is unknown if SDMs can detect intra-specific niche differences in invasive plants. The SDMs I developed for the subspecies of A. saligna detected intra-specific differences within the native range, but did not predict the full invasive distribution in South Africa. Unsurprisingly, SDMs agreed with genetic analyses (based on nuclear microsatellites, nuclear DNA, and chloroplast DNA) and did not assign South African populations to any subspecies of A. saligna. South African populations were assigned to a novel genetic entity likely produced by human cultivation practices. A global phylogeny identified this cultivated genotype in introduced populations in eastern Australia and Portugal, while the remaining introduced populations differed markedly in their genetic composition. Overall, A. saligna‘s high intra-specific diversity and complex introduction history generated a variety of genetic patterns across the current global distribution of the taxon. Global populations of P. lophantha were processed using a similar approach to that used for A. saligna, and aimed to determine if the same pathways and modes of introduction produced analogous genetic patterns in a closely related species. Diverse arrays of genotypes were identified in introduced populations of P. lophantha, suggesting inconsistent sampling of a variety of native sources. Further work is however needed to clarify the morphological and genetic differences (if any) between the intra-specific entities, and identify exactly which P. lophantha subspecies were introduced outside of their native range, The variation in the global distribution of genetic diversity observed in A. saligna and P.lophantha demonstrated that intra-specific genetic variation, human usage, and the pathway and manner of introduction interact during several phases of the invasion process and collectively determine the introduced genetic patterns. The dissimilarity in the distribution of genotypes in both species suggests that they might not behave the same way throughout their introduced range. Consequently, management insights might not be transferrable between regions. More generally, my findings provide an important contribution to the debate whether (and how quickly) introduced and native populations should be treated as fundamentally different entities. / AFRIKAANSE OPSOMMING: Grootskaalse menslike verskuiwing van organismes bevorder die vestiging van spesies buite hul natuurlike voorkomsareas en 'n klein hoeveelheid van hierdie spesies word indringers. Tydens die bestuur van indringerspesies word dit tipies aanvaar dat ingevoerde indringerspesies enkele, statiese evolusionêre eenhede is wat analoog is aan hul inhmeemse eweknieë. Studies het egter getoon dat inheemse en uitheemse populasies van 'n aantal ingevoerde plante aansienlik verskil in hul genetiese samestelling. Hierdie verskille kan 'n negatiewe invloed op die algehele sukses van beheer- en bestuursprojekte hê, veral vir die spesies wat intra-spesifiek divers is. Die invloed van intra-spesifieke diversiteit op die indringingsproses is getoets aan twee boomspesies, inheems aan Wes-Australië, wat wyd uitgevoer word: Acacia saligna (drie subspesies) en Paraserianthes lophantha (twee subspesies). Vergelyking van klimaatstoestande tussen n spesie se in- en uitheemse voorkomsareas word wyd gebruik om toekomstige indringingsrisiko te voorspel. Dit was voor hierdie navorsing onduidelik of spesie verspreiding modelle (SVMs) intra-spesifieke nis-verskille in indringerplante kan uitwys. SVMs wat vir die subspesies van A. saligna ontwikkel is, kon intra-spesifieke verskille in Wes-Australië uitwys, maar het nie die volle verspreiding van die spesies in Suid-Afrika voorspel nie. Onverbasend, is geen Suid-Afrikaanse populasies deur genetiese analise (gebaseer op die kern mikrosatelliete, kern-DNS, en chloroplas-DNS) toegewys aan 'n subspesie van A. Saligna nie. Suid-Afrikaanse populasies het 'n nuwe genetiese entiteit wat waarskynlik gekweek is deur menslike verbouingspraktyke. 'n Globale filogenie het hierdie verboude genotipe in addisionele ingevoerde populasies in die ooste van Australië en Portugal geïdentifiseer. Mikrosatelliet genotipes van uitheemse populasies wêreldwyd in Oos-Australië, Israel, Italië, Nieu-Seeland, Portugal, Suid-Afrika, Spanje en die VSA verskil merkbaar in hul genetiese samestelling. A. saligna se hoë intra-spesifieke diversiteit en komplekse geskiedenis van invoer (wat verbouing, wye verspreiding en hoë ―propagule druk betrek), het 'n verskeidenheid van genetiese patrone oor die huidige globale verspreiding van die takson gegenereer. Om te bepaal of 'n globale uiteenlopende genetiese patroon binne nouverwante spesies bestaan, is globale bevolkings van Paraserianthes lophantha verwerk deur gebruik te maak van 'n soortgelyke benadering as wat vir A. saligna gebruik is. Globale populasies van beide studie-spesies bestaan uit 'n diverse verskeidenheid van genotipes. Resultate dui daarop dat P. lophantha van 'n verskeidenheid inheemse bronne ingevoer is. Om te identifiseer watter P. lophantha subspesies buite hul natuurlike voorkomsarea versprei is, word verdere werk benodig om die morfologiese en genetiese verskille (indien enige) tussen die intra-spesifieke entiteite vas te stel. In hierdie tesis het ek gewys dat intra-spesifieke genetiese variasie, menslike gebruik en invoering-geskiedenis saam werk om genetiese patrone in uitheemse populasies te vorm. Verder het ek die waarde van die gebruik van verskillende molekulêre benaderings om indringing geskiedenis te verstaan, gedemonstreer. Die verskil in die verspreiding van die genotipes van A. saligna en P. lophantha dui daarop dat hulle moontlik nie op dieselfde manier dwarsdeur hul uitheemse verspreidingsarea mag optree nie. Bestuursinsigte mag gevolglik nie oordraagbaar wees tussen streke nie. Meer algemeen, bied my bevindings 'n belangrike bydrae tot die debat of (en hoe vinnig) inheemse en ingevoerde populasies behandel moet word as fundamenteel verskillende entiteite.