Plants that can self-fertilise should, on average, be more invasive than plants that can not self-fertilise because they can reproduce regardless of the availability of mates and pollinators. Self-fertilisation should have a strong effect on invasiveness because, to become invasive, introduced plants have to pass through bottlenecks of low plant abundance when mates and pollinators are likely to be scarce. Under these conditions, reproduction of plants that can not self-fertilise is often limited by pollen receipt. Selfing may thus contribute to invasiveness by alleviating pollen limitation Allee effects (pollen limitation caused by low abundance) especially as theoretical work indicates that ability to invade and rate of invasion are highly sensitive to fecundity of small and isolated populations and single individuals. Recently, a correlation between ability to self-fertilise and invasiveness has been observed in several invasive floras, consistent with the hypothesis that species that can self-fertilise should be more invasive. However, it has not yet been demonstrated that this relationship arises from reproductive assurance. To establish the causal basis of a correlation between a plant trait and invasiveness, a mechanism linking that trait to invasiveness must be demonstrated. For this it is necessary to show firstly that the trait actually affects performance in the introduced range and secondly that plant performance affects invasiveness. Self-fertilisation is hypothesised to increase invasiveness by enhancing reproductive performance. The first step in testing this hypothesis is therefore to show that being able to self-fertilise increases fecundity, i.e. that it provides reproductive assurance. However, progeny from self-fertilisation often suffer from inbreeding depression – they perform worse than those from cross-fertilisation – so it is also necessary to show that this cost does not outweigh the reproductive assurance benefit of selfing. So far, reproductive assurance has been assessed in only a few invasive plant species. These studies did not assess inbreeding depression and only one investigated reproductive assurance in relation to abundance, finding no relationship. In this thesis I have sought to understand the importance of self-fertilisation for reproduction of invasive plants in the introduced range through case studies. In particular, I assessed whether reproductive assurance from self-fertilization alleviates Allee effects via pollen limitation. To do this I tested whether pollen limitation and reproductive assurance were greater at low plant abundance. Further, I conducted progeny trials to assess inbreeding depression, as this cost of selfing potentially negates reproductive assurance benefits. I also conducted observations and experiments to identify the principle pollinators of my study species as reproductive assurance and its relationship to plant abundance depend on pollinator visitation, The Australian trees Acacia mearnsii and A. dealbata are highly invasive in the study region of KwaZulu-Natal, South Africa. Through controlled pollination experiments I established that A. dealbata was self-compatible and autonomously self-fertilising, while previous studies reported A. mearnsii as self-incompatible. I identified the native honeybee Apis mellifera scutellata as the principal pollinator of A. mearnsii, A. dealbata and a co-occuring related invasive species, Acacia decurrens, in the study region. I conducted pollen supplementation experiments in two of these species, aiming to indirectly assess reproductive assurance from selfing in the self-compatible A. dealbata by comparing pollen limitation between this species and the self-incompatible A. mearnsii. In both species, I conducted pollen supplementation in single isolated trees and trees in continuous populations, to test whether pollen limitation was more severe in isolation. These pollen supplementation experiments were inconclusive with respect to pollen limitation but indicated that if there was pollen limitation in A. mearnsii, it was not related to isolation. Progeny trials in A. dealbata revealed relatively strong inbreeding depression in progeny growth and survival. This suggests that selfed progeny may not reach reproduction, so even if self-fertilisation provides reproductive assurance, it may not contribute to invasion in this species. As floral morphology of Acacia species prohibits the use of emasculation experiments to directly measure reproductive assurance, I conducted further investigations on Lilium formosanum, a large-flowered, autonomously self-pollinating invasive geophyte native to Taiwan. I identified the long tongued hawkmoth Agrius convolvuli as its primary pollinator in its introduced range in KwaZulu-Natal, South Africa. Trials of progeny from self- and crosspollination in the field (to 31 months) and in a controlled shade-house environment (to 26 months) showed no evidence of inbreeding depression in germination, growth or survival. Flowering was assessed in the shade-house as most plants did not flower in the field. Only one of five populations showed inbreeding depression in probability of flowering in the second year of growth but none showed inbreeding depression in the third year. Inbreeding depression was thus generally undetectable in L. formosanum. I tested for reproductive assurance and pollen limitation in L. formosanum by conducting floral emasculations and pollen supplementations in multiple populations across a range of population size and isolation in three different years. These experiments demonstrated that reproductive assurance was substantial and that pollen limitation was low or absent. Contrary to expectations, reproductive assurance was not greater in smaller populations and was greater for more isolated populations in only one of three years. However, that study did not include many very small populations. To assess reproductive assurance at very low abundance, I created arrays of emasculated and intact plants within and around naturally occurring populations at two sites. Isolated plants had higher reproductive assurance than did plants placed inside the continuous population at one site, supporting the hypothesis that selfing provides reproductive assurance against pollen-limitation Allee effects. However, in these studies, generally inadequate pollinator visitation was the main reason that L. formosanum exhibited reproductive assurance through selfing. The substantial reproductive assurance and minimal inbreeding depression displayed by L. formosanum makes a compelling case for the hypothesis that self-pollination promotes invasion. Nevertheless, demographic modelling will be necessary to assess whether increased fecundity through reproductive assurance results in increased rates of population growth and spread, and hence invasion, in this species. To assess whether reproductive assurance accounts for the relationship between ability to self-fertilise and invasiveness in plants generally, the contribution of self-fertilisation to invasiveness will have to be evaluated for a larger sample of invasive and non-invasive introduced species, using the approaches taken in this thesis, followed up by demographic modelling. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2011.

Like orchids, the complexity of flowers found in asclepiads (Asclepiadoideae, Apocynaceae) and the fact that pollen is presented as pollinaria, offers excellent opportunities to study various aspects of plant-pollinator interactions. In this thesis I investigated two broad themes: ecological aspects of the pollination biology of hymenopteran and fly-pollinated asclepiads as well as the degree of specialization to certain pollinators in these species. Colonizing plants often reproduce through self-pollination, or have highly generalized pollination systems, or both. These characteristics facilitate establishment in small founding populations and generates the prediction that reproductive success should be independent of population size in these species. Chapter one examines the pollination biology of Gomphocarpus physocarpus, an indigenous, weedy species and investigates the relationship between reproductive success and population size. In this species, there is no evidence of an Allee effect and reproductive success is not correlated with population size. In addition G. physocarpus is not capable of self-pollination, suggesting it is completely reliant on pollinators for seed set. The lack of a relationship between pollination success and population size is therefore likely explained by the generalized wasp pollination system of this species. Several milkweeds are invasive outside of their native ranges. Invasive species either need to co-opt native pollinators in order to reproduce or reduce their reliance on pollinators through having the ability to self-pollinate. Co-opting native pollinators is expected to be easier in species that have generalized pollination systems, alternatively species with specialized flower morphologies need to rely on similar functional groups of pollinators to be present within the invaded range. Chapter two investigates the pollination biology and pollination success of the invasive milkweed, Araujia sericifera, and finds that in South Africa, this species is visited mainly by native honeybees and nocturnal moths. Moths however contribute little to pollen removal, and deposition. Based on the apparent morphological mismatch between the flower of A. sericifera and native honeybees, I propose that the native pollinators of this species are likely to be larger Hymenoptera (e.g. Bumblebees). Data from a breeding system study, indicated that this species is not capable of automatic self-pollination, but could set fruit from geitonogamous self-pollinations pointing to the importance of native pollinators for successful reproduction. The pollinaria of milkweeds can accumulate on pollinators to form pollen masses large enough to physically interfere with the foraging behaviour of pollinating insects. In chapter three I describe the pollination biology of Cynanchum ellipticum and find that this species is mainly pollinated by honeybees although this species is visited by several other members of Hymenoptera, Lepidoptera and Diptera. Due to the structure of the pollinaria, these chain together relatively efficiently and frequently form large pollinarium loads on the mouthparts of honeybees. However there is little evidence that these pollinarium loads influence the foraging times of pollinators and only a few individual honeybees exhibited longer foraging times and most honeybees were unaffected by the presence of large pollinarium loads. Within the genus Cynanchum there is large variation in the gynostegium structure that may influence the pattern of pollinarium loading on pollinators as well as pollen reception as shown in chapter three. In Chapter four, the pollination biology of Cynanchum obtusifolium is examined, and like that of C. ellipticum, this species is visited by a wide diversity of pollinators but honeybees appear to be the primary pollinators. More importantly this species is shown to be andromonoecious and produces two morphologically different flower types, that may be distinguished based on differences in the gynostegium structure. These two types of flower could mainly be distinguished by the length of the anther wings. I found that flowers with short anther wings function as male flowers by only exporting- and rarely receiving pollinia. Flowers with longer anther wings function as hermaphrodite flowers and can both export and receive pollinia. The ratio of male to hermaphrodite flowers varied at different times during the flowering season, but preliminary data suggested that this was not related to levels of pollination success. The genera Stapelia and Ceropegia are well known for their intricate floral adaptations that mimic the brood and feeding substrates of pollinating flies. Despite several studies that have documented the various adaptations to fly pollination in different species, there is a lack of natural history studies documenting different flower visitors, pollen loads and long term levels of pollination success in these species. In Chapter six I document the pollination biology of Ceropegia ampliata by documenting different pollinators and quantifying average levels of pollination success and the nectar reward. I also experimentally manipulated the trapping hairs of this species to determine whether trapping hairs influence average levels of pollen export and receipt. I show that Ceropegia ampliata is pollinated by a generalist guild of flies (mainly Tachinidae, Sarcophagidae, Muscidae and Lauxaniidae) and produces minute quantities of relatively dilute nectar as a reward. Pollination success was generally low in this species and increases periodically suggesting that the abundance of pollinators is patchy. I found that flowers with trapping hairs that had already wilted had higher levels of pollinarium removal than flowers with erect hairs, however experimentally removing the hairs had no significant effect on pollen export and receipt. In Chapter seven, I document the pollinators, pollen loads and long term levels of pollination success in Stapelia hirsuta var. bayllissi, a rare sapromyiophilous stapeliad. I find that, in contrast to C. ampliata, this species was specialized to pollination by small flies of the family Anthomyiidae. Similar to the results from Chapter seven, I find that long term levels of pollination success were typically low but could increase periodically, although such increases were generally unpredictable. There are currently very few records documenting pollinator interactions in the Periplocoideae. Many species within this subfamily exhibit open-access flowers suggestive of pollination by short-tongued insects. I investigated the pollination biology of Chlorocyathus lobulata, a rare species with a highly localized distribution. I aimed to determine the pollinators, average levels of pollination success and demography of this species in order to determine whether this rare species is suffering from the collapse of a highly specialized pollinator mutualism. I also quantified the high incidence of flower herbivory caused by larvae of the moth, Bocchoris onychinalis. I find that C. lobulata has a highly generalized fly pollination system and average levels of pollination success suggested that a large proportion of flowers had pollen removed and deposited suggesting that this species is not experiencing pollination failure. The large numbers of juveniles present also indicated that recruitment is taking place.

The Allee effect corresponds to a positive relationship between population size and individual fitness. This positive relationship can cause thresholds, that is, critical population sizes below which the population becomes extinct. For species submitted to the Allee effect, the formation and cohesion of groups is therefore crucial to survival. Animals can achieve this collective behaviour through local interactions. Each individual interacts locally with conspecifics and, at the scale of the group, a unity of behaviour emerges: the animals move together, rest in the same place, or choose the same habitat patch to settle on. <p><p>We use a combination of mathematical modelling and experimental work to study certain mechanisms of collective behaviour. In particular we assess the extent to which different individual interactions can induce collective patterns and thereby influence the dynamics of dispersal and settlement of populations. First, we study the collective settlement induced by the arresting effect of a marker secreted by conspecifics. Then, two potential mechanisms for collective movement are examined: following the conspecifics and following a trail laid by conspecifics. Finally, we integrate explicit mechanisms of dispersal behaviour in a dynamic model involving a set of interconnected populations. This allows the study of the interplay between collective movements and Allee effects at the scale of the metapopulation.<p><p>Our work is inspired by the lifestyle of the two-spotted spider mite, Tetranychus urticae, a phytophagous pest of recognised agricultural importance. These subsocial mites live in aggregates on the leaf surface, protected by a collectively spun silk web. Experimental evidence suggests that its population dynamics are subject to the Allee effect. Moreover, these mites show a tendency to migrate collectively, which makes them an appropriate biological model. / Doctorat en Sciences / info:eu-repo/semantics/nonPublished

A dinâmica de crescimento populacional de uma espécie é permeada pela relação entre as desvantagens da competição intraespecífica e os benefícios da presença de conspecíficos. Para muitas espécies, os benefícios da cooperação podem superar as desvantagens da competição. A correlação positiva entre tamanho populacional e adaptabilidade em populações muito pequenas é conhecida como efeito Allee demográfico. Apesar de haver modelos matemáticos isolados para os diferentes tipos de efeitos Allee, não há um modelo simples que os abranja e os conecte a modelos de crescimento mais gerais (como o de Richards). Propomos unificar modelos de efeitos Allee e o de crescimento de Richards em um modelo que permita um novo ponto de vista sobre o efeito Allee demográfico. Um exemplo do aumento das possibilidades descritivas de tal generalização é a emergência de mais de uma transição cooperação-competição quando considerado um caso particular desse novo modelo (Allee-Gompertz). Apesar da importância do crescimento populacional, a maioria dos modelos básicos de transmissão de doenças infecciosas considera o tamanho populacional constante ou adota simplificações pouco plausíveis. Nesta tese, mostramos as deficiências de um modelo compartimental dinâmico de tuberculose já consagrado e propomos um novo modelo com crescimento populacional logístico. Quando comparados, nosso modelo apresenta previsões mais pessimistas para a erradicação da doença a longo prazo quando testado com parâmetros que definem políticas de controle pouco eficientes. Realizamos tais predições adotando estratégias de controle de países desenvolvidos e subdesenvolvidos. Visto que esses modelos compartimentais desprezam aspectos espaciais, desenvolvemos uma modelagem computacional de agentes, baseada no modelo proposto, com duas estruturas subjacentes: redes aleatórias e redes reais. A súbita emergência de tuberculose resistente a drogas como consequência de tratamentos ineficazes é também um resultado das implementações desses modelos em dois cenários distintos. Esses resultados são comparados com os do modelo compartimental e com os de um modelo de estrutura subjacente mais simples e, como novo resultado, surge nos dois modelos a possibilidade de erradicação da doença em menos de uma década após o início do tratamento. Esse resultado é possível desde que sejam adotadas estratégias eficientes de controle. / The one-species population growth dynamics is permeated by the relationship between the harms from the intraspecific competition and the benefits from the presence of conspecifics. For many species, the benefits from conspecific cooperation may outweigh the harms from competition. The positive correlation between population size and total fitness in very small population known as demographic Allee effect. Although there are isolated mathematical models for different types of Allee effects, there is not a simple model that covers and connects them to more general growth models (like Richards). We propose to unify models of Allee effects and the Richards growth one in a model that allows a new perspective on the demographic Allee effect. An example of the increased descriptive possibilities of such generalization is the emergence of more than one transition cooperation-competition when considering a particular case of this new model (Gompertz-Allee). Despite the importance of population growth, most basic models of infectious diseases transmission considers population size constant or adopts implausible simplifications. In this thesis, we show the shortcomings of a dynamic compartmental model of tuberculosis already established and we propose a new model with population logistic growth. When compared, our model provides more pessimistic forecasts for the eradication of the disease in the long term if it is tested with parameters that define inefficient control policies. We perform such predictions adopting control strategies from developed and underdeveloped countries. Since these compartmental model disregards spatial aspects, we developed a computational agent model, based on the proposed model, with two underlying structures: random networks and real networks. The sudden emergence of drug-resistant tuberculosis as a result of ineffective treatments is also a result from the implementations of these models in two distinct scenarios. These results are compared with the ones from a compartimental model and with the ones from a model with simpler underlying structure and, as a new result, the possibility of eradicating the disease in less than a decade after beginning the treatment appears on the two models. This result is possible adopting effective control strategies.

A dinâmica de crescimento populacional de uma espécie é permeada pela relação entre as desvantagens da competição intraespecífica e os benefícios da presença de conspecíficos. Para muitas espécies, os benefícios da cooperação podem superar as desvantagens da competição. A correlação positiva entre tamanho populacional e adaptabilidade em populações muito pequenas é conhecida como efeito Allee demográfico. Apesar de haver modelos matemáticos isolados para os diferentes tipos de efeitos Allee, não há um modelo simples que os abranja e os conecte a modelos de crescimento mais gerais (como o de Richards). Propomos unificar modelos de efeitos Allee e o de crescimento de Richards em um modelo que permita um novo ponto de vista sobre o efeito Allee demográfico. Um exemplo do aumento das possibilidades descritivas de tal generalização é a emergência de mais de uma transição cooperação-competição quando considerado um caso particular desse novo modelo (Allee-Gompertz). Apesar da importância do crescimento populacional, a maioria dos modelos básicos de transmissão de doenças infecciosas considera o tamanho populacional constante ou adota simplificações pouco plausíveis. Nesta tese, mostramos as deficiências de um modelo compartimental dinâmico de tuberculose já consagrado e propomos um novo modelo com crescimento populacional logístico. Quando comparados, nosso modelo apresenta previsões mais pessimistas para a erradicação da doença a longo prazo quando testado com parâmetros que definem políticas de controle pouco eficientes. Realizamos tais predições adotando estratégias de controle de países desenvolvidos e subdesenvolvidos. Visto que esses modelos compartimentais desprezam aspectos espaciais, desenvolvemos uma modelagem computacional de agentes, baseada no modelo proposto, com duas estruturas subjacentes: redes aleatórias e redes reais. A súbita emergência de tuberculose resistente a drogas como consequência de tratamentos ineficazes é também um resultado das implementações desses modelos em dois cenários distintos. Esses resultados são comparados com os do modelo compartimental e com os de um modelo de estrutura subjacente mais simples e, como novo resultado, surge nos dois modelos a possibilidade de erradicação da doença em menos de uma década após o início do tratamento. Esse resultado é possível desde que sejam adotadas estratégias eficientes de controle. / The one-species population growth dynamics is permeated by the relationship between the harms from the intraspecific competition and the benefits from the presence of conspecifics. For many species, the benefits from conspecific cooperation may outweigh the harms from competition. The positive correlation between population size and total fitness in very small population known as demographic Allee effect. Although there are isolated mathematical models for different types of Allee effects, there is not a simple model that covers and connects them to more general growth models (like Richards). We propose to unify models of Allee effects and the Richards growth one in a model that allows a new perspective on the demographic Allee effect. An example of the increased descriptive possibilities of such generalization is the emergence of more than one transition cooperation-competition when considering a particular case of this new model (Gompertz-Allee). Despite the importance of population growth, most basic models of infectious diseases transmission considers population size constant or adopts implausible simplifications. In this thesis, we show the shortcomings of a dynamic compartmental model of tuberculosis already established and we propose a new model with population logistic growth. When compared, our model provides more pessimistic forecasts for the eradication of the disease in the long term if it is tested with parameters that define inefficient control policies. We perform such predictions adopting control strategies from developed and underdeveloped countries. Since these compartmental model disregards spatial aspects, we developed a computational agent model, based on the proposed model, with two underlying structures: random networks and real networks. The sudden emergence of drug-resistant tuberculosis as a result of ineffective treatments is also a result from the implementations of these models in two distinct scenarios. These results are compared with the ones from a compartimental model and with the ones from a model with simpler underlying structure and, as a new result, the possibility of eradicating the disease in less than a decade after beginning the treatment appears on the two models. This result is possible adopting effective control strategies.

Microbes live in complex environments where competitive and cooperative interactions occur that dictate their success and the status of their environment. By furthering our understanding of the interactions between microbes, questions into the evolution of cooperation, disease virulence and biodiversity can be addressed. This will help develop strategies to overcome problems concerning disease, socioeconomics and conservation. We use an approach that combines evolutionary ecology theory with genetics and molecular biology to establish and develop model microbial ecological systems to examine feeding strategies, in what has been termed synthetic ecology. Using the model fungal plant pathogen system of rice blast disease, we generated less virulent gene deletion mutants to examine the sociality of feeding strategies during infection and test a nascent virulence reduction strategy based on competitive exclusion. We revealed that the success of the pathogen is unexpectedly enhanced in mixed strain infections containing the virulent wild-type strain with a less virulent gene deletion mutant of the metabolic enzyme invertase. Our finding is explained by interference between different social traits that occur during sucrose feeding. To test the generality of our result, gene deletion mutants of putative proteases were generated and characterised. We found that if virulence related genes acted ‘privately’, as predicted by social theory, the associated mutants would not make viable strains to use for this virulence reduction strategy by competitive exclusion. Our study then went on to study the fitness of digesting resources extracellularly, as many microbes do, given that this strategy is exposed to social exploitation by individuals who do not pay the metabolic costs. This was investigated by developing an experimental system with Saccharomyces cerevisiae. Though internalising digestion could suppress cheats, the relative fitness of opposing strategies was dependent upon the environmental and demographic conditions. Using this polymorphic system, the influence of competitors on the stability of cooperation, and the influence of cheats on the maintenance of diversity were assessed. To test the fitness of internal versus external digestion in a more natural setting, we generated an internally digesting strain of the rice blast fungus. In addition to suppressing cheats, the strain had enhanced fitness and virulence over the wild-type. We propose that this is caused by a shift in a trade-off between yield and rate. We show how a synthetic ecology approach can capture details of the biology underlying complex ecological processes, while having control over the factors that drive them, so that the underlying mechanisms can be teased apart.

No description available.

Conservation in the EU and all over the world aims at reducing biodiversity loss which has become a great issue in the last decades. However, despite existing efforts, Earth is assumed to face a sixth mass extinction. One major challenge for conservation is to reconcile the targets with conflicting interests, e.g. for food production in intensively used agricultural landscapes. Agriculture is an example of a coupled human-environment system that is approached in this thesis with the help of mathematical models from two directions. Firstly, the ecological subsystem is considered to find processes relevant for the effect of habitat connectivity on population abundances. Modelling theory predicts that the species-specific growth parameters (intrinsic growth rate and carrying capacity) indicate whether dispersal has a positive or negative effect on the total population size at equilibrium (r-K relationship). We use laboratory experiments in combination with a system of ordinary differential equations and deliver the first empirical evidence for a negative effect of dispersal on the population size in line with this theory. The result is of particular relevance for the design of dispersal corridors or stepping stones which are meant to increase connectivity between habitats. These measures might not be effective for biological conservation. A second population model, consisting of two coupled Ricker maps with a mate-finding Allee effect, is analyzed in order to examine the effect of bistability due to the Allee effect in combination with overcompensation in a spatial system. The interplay can cause complex population dynamics including multiple coexisting attractors, long transients and sudden population collapses. Essential extinction teaches us that not only small populations are prone to extinction but chaotic dynamics can drive a population extinct in a short period of time as well. By a comprehensive model analysis, we find that dispersal can prevent essential extinction of a population. In the context of conservation that is: habitat connectivity can promote rescue effects to save a population that exhibits an Allee effect. The two findings of the first part of this thesis have contrasting implications for conservation which shows that universal recommendations regarding habitat connectivity are impossible without knowledge of the specific system. Secondly, a model for the socio-economic subsystem is presented. Agri-environment schemes (AES) are payments that compensate farmers for forgone profits on the condition that they improve the ecological state of the agricultural system. However, classical economic models that describe the cost-effectiveness of AES often do not take the social network of farmers into account. Numerical simulations of the socio-economic model presented in this thesis suggest that social norms can hinder farmers from scheme participation. Moreover, social norms lead to multistability in farmers’ land-use decision behaviour. Informational campaigns potentially decrease the threshold towards more long-term scheme participation and might be a good tool to complement compensation payments if social norms affect land-use decisions. Finally, a coupled human-environment system is analyzed. An integrated economicecological model is studied to investigate the cost-effectiveness of AES if the species of concern exhibits an Allee effect. A numerical model analysis indicates large trade-offs between agricultural production and persistence probability. Moreover, conservation success strongly depends on the initial population size, meaning that conservation is well advised to start before the species is threatened. Spatial aggregation of habitat can promote rescue effects, suggesting land-sparing solutions for conservation. In that case,agglomeration bonuses may serve to increase the effectiveness of AES. Possible causes for population declines are diverse and can be a combination of human influences, e.g. due to habitat degradation and inherent ecosystem properties. That complicates the task of conservation. The models presented in this thesis simplify complex systems in order to extract processes relevant for biological conservation. The analysis of spatial effects and dynamical model complexity, e.g. due to Allee effects or a nonlinear utility function, allows us improve the understanding of coupled human-environment systems.