Tesis sobre el tema "Propagule pressure"

Introductions of exotic fouling species have severely disrupted marine ecosystems. Theory suggests that high propagule pressure increases the probability of successful establishment. Likewise, empirical studies have shown the importance of ships and boats for transporting exotic species. However, few empirical studies have demonstrated the effect of propagule pressure on invasion success. For marine fouling species, uncertainty about vectors and pathways further impedes our ability to calculate propagule pressure and to understand establishment success. My goal was to better understand the links between propagule pressure from maritime activities and the spatial distribution of exotic fouling species. Using empirical data, I (1) determined the importance of boat characteristics and propagule exposure on boat fouling, (2) demonstrated the roles of commercial shipping and recreational boating in the invasion process, and (3) tested the effect of propagule pressure on population and community diversity. Although boats on the east coast of Canada were less fouled than New Zealand boats, boat-mediated spread may be facilitated in Canada by greater movement of boats among marinas than in New Zealand. Propagule exposure better predicted boat fouling than did boat characteristics. The spatial genetic structure of the colonial tunicate, Botryllus schlosseri, suggests that ships have frequently introduced it to the east coast of Canada and that secondary spread occurs gradually around individual ports, facilitated by recreational boating. The diversity of invasive species within commercial ports is positively related to propagule pressure and invasion success at both community (i.e. inter-specific richness) and population levels (i.e. genetic diversity). Diversity was significantly correlated to the numbers of ship arrivals but not to ballast water discharge events or the volume of ballast water discharged. This suggests that hull biofouling is a more important pathway than ballast water for exotic fouling species. Both boats and ships have influenced the invasion process of fouling species in Canadian coastal areas. The research also underlines the point that predictive models for the spread of biofouling species should be based on regional boating patterns, boating characteristics, and local propagule exposure. Defining relationships between propagule pressure and invasion success is one of the great challenges to understanding the rapid changes occurring in marine ecosystems.
L'introduction des salissures biologiques exotiques a sévèrement perturbé les écosystèmes marins. La théorie suggère qu'un apport de propagules élevé augmente le risque d'envahissement. C'est ainsi que les recherches se sont concentrées à évaluer la probabilité que les navires commerciaux et les petites embarcations transportent des espèces exotiques. Malgré son importance, peu d'études empiriques supportent l'effet de l'apport de propagules sur le succès d'envahissement. Pour les salissures biologiques marines, l'incertitude entre les vecteurs et les voies de transports entrave particulièrement notre capacité à calculer l'apport de propagules entrant par les voies maritimes, limitant ainsi notre compréhension du succès d'envahissement. L'objectif de ma thèse est d'apporter une meilleure compréhension de la relation entre l'apport de propagules provenant du transport maritime et la distribution spatiale des salissures exotiques. À partir de données empiriques, (1) j'ai déterminé l'importance relative des caractéristiques des bateaux de plaisance et l'exposition des propagules environnantes sur la probabilité de la colonisation des coques, (2) j'ai démontré le rôle respectif des navires et des plaisanciers dans le processus d'invasion et (3) j'ai décrit la relation entre l'apport de propagules et la diversité des populations et des communautés. Bien que les bateaux de la côte est du Canada sont moins colonisés que les bateaux de la Nouvelle-Zélande, la propagation des salissures exotiques pourrait être facilitée au Canada par un plus grand trafic entre les marinas. L'exposition aux propagules environnantes prédit davantage l'encrassement biologique des coques que les caractéristiques des bateaux. La génétique des populations de l'ascidie Botryllus schlosseri suggère qu'il y a eu plusieurs introductions dans les ports de la côte est du Canada. Facilité par les plaisanciers, le tunicier s'est ensuite dispersé progressivement autour de ceux-ci. Les indices de diversité soutiennent la relation positive entre l'apport de propagules et le succès d'envahissement et cela autant au niveau des communautés (c.-à-d. la richesse inter-spécifique) qu'au niveau des populations (c.-à-d. diversité génétique). La diversité est significativement corrélée au nombre d'arrivées des navires, mais n'est pas corrélée au nombre de déchargements et au volume de l'eau de lest. Ces relations suggèrent une plus grande importance de l'encrassement des navires que l'eau de lest comme voie d'introduction des salissures exotiques. Cette thèse démontre que les petits bateaux et les navires influencent grandement les processus d'invasions des salissures biologiques des régions côtières du Canada. Cette recherche souligne également que les modèles de propagation des salissures exotiques devraient fonder leurs prémisses sur l'exposition des propagules locales, les caractéristiques des bateaux et l'activité nautique régionale. La nécessité de décrire avec précision les relations entre l'apport de propagules et le succès d'envahissement est l'un des grands défis écologiques afin d'améliorer notre compréhension sur les changements rapides actuellement observés dans nos écosystèmes marins.

Increased trading worldwide has created introduction pathways for aquatic invasive species (AIS), particularly through shipping activities. In our research, we used ballast discharge data and estimates of the wetted surface area of vessels to provide preliminary estimates of the potential propagule pressure ballast and hull fouling organisms on Canadian shipping ports from the commercial shipping sector. We compared total wetted surface area, vessel arrivals and ballast discharge across shipping ports and vessel categories in the Atlantic, Great Lakes-St. Lawrence and Pacific shipping regions. Using these potential propagule pressure estimates for ballast organisms, we developed a model, building on that of MacIsaac et al.’s (2002), to characterize the effective propagule pressure of aquatic non-native species to Canadian shipping ports. Our model includes the effects of environmental similarity between destination and source on mortality, which can be considerable. We parameterized the model using recent nationally and regionally collected databases on ship voyages, and abundance surveys yielding mortality rates of several zooplankton species. These empirically derived parameters were used in our model to predict abundances of live individuals after ballast is discharged, with estimates of uncertainty and sensitivity to key assumptions. Our results indicate that for our three shipping regions, aggregate wetted surface area, vessel arrivals, and total ballast discharge were significantly correlated across shipping ports (Spearman’s ρ ranged from 0.57 to 0.87, p<0.05). Correlations between these measures of propagule pressure were more variable and of varying levels of significance across vessel categories (Spearman’s ρ ranged from 0.43 to 0.98). Our modeling results demonstrate that variation in mortality rates across tanks and voyages resulted in high variation in total effective propagule pressure. The variation between tanks and voyages has important implications for the use of mid-ocean exchange as a ballast management method for different ports and species. To our knowledge, our characterization of potential and effective propagule pressure from the commercial shipping sector is the first to be conducted on a nation-wide scale. Our propagule pressure estimates will contribute to future efforts to determine the relationship between the establishment of aquatic invasive species and the environmental similarity between source and discharge areas.

The invasibility, or susceptibility of an ecosystem to biological invasion is influenced by changes in biotic and abiotic resistance often due to shifts in disturbance regime. The magnitude of invasive propagule pressure interacts with an ecosystem's invasibility to determine the extent of a biological invasion. I examined how propagule pressure, forest community structure and disturbance interact to influence the invasibility of temperate Pacific Northwest forests by the newly-invasive grass, Brachypodium sylvaticum. My goal was to identify which of these factors is most instrumental in enabling the shift from establishment to population growth in B. sylvaticum at the edge of its expanding range. Both observational and experimental studies were employed to identify the many ecological components of this problem. Ecological sampling methods were used to identify trends in B. sylvaticum habitat preference and signs of habitat disturbance. In addition, an experimental study was performed to test the effects of soil and vegetation disturbance on B. sylvaticum seedling propagation. I found that while soil disturbance did not have a significant effect on seedling propagation, vegetation disturbance was implicated in B. sylvaticum spread. Higher propagule pressure and coniferous forest type were also strong predictors of increased B. sylvaticum seedling propagation and survival within established sites. My study demonstrates how propagule pressure and plant community dynamics interact to shift the invasibility of Pacific Northwest forests and facilitate the transition from establishment to spread in the invasion of B. sylvaticum.

The spread of invasive species poses a major ecological and economical threat. Consequently there are ongoing efforts to develop a generalizable mechanism to predict establishment success of non-native species. One proposed mechanism to predict establishment success is propagule pressure, which is defined as the number of individuals introduced at a given time. Although some studies have demonstrated a positive correlation between propagule pressure and establishment success, others have not, and the effect of propagule pressure on establishment success remains unclear. To address this challenge, a strain of bacteria engineered with an Allee effect, a growth dynamic that is often associated with establishing species, was used. The timing between successive introduction events that resulted in establishment success was measured. It was observed that if the time between two introduction events was sufficiently long, growth did not occur. By manipulating the growth rate of the bacteria, it was shown that that the minimal time between the two introduction events that resulted in growth was constrained as growth rate decreased. Moreover, it was concluded that asymmetry in the density of bacteria introduced in the introduction events increased the maximum time between introduction events that resulted in growth. These results help to remedy conflicting data in the literature by identifying conditions where propagule pressure has, and does not have, a positive impact on establishment success. These findings can have major implications in understanding and predicting the unique population dynamics of invasive species.

Estimating the probability of establishment of non-indigenous species is a crucial element in managing their spread. In this thesis, I use two approaches to estimate the probability of establishment of Bythotrephes longimanus, a predatory cladoceran that is invading lakes in Ontario and the surrounding American states. At a watershed level, I develop a vector based model to predict the probability of establishment of B. longimanus over time. I use metrics of propagule pressure from anthropogenic and natural dispersal to estimate spread, and extend the model to incorporate spatial and temporal gaps in knowledge of the invasion status of lakes. I found that recreational boating traffic is the dominant vector of spread and that most risk to lakes is due to static hubs of invasion - the five largest lakes in the watershed. Next, at the scale of a local population introduction, I investigate probability of establishment empirically. I follow B. longimanus populations over their entire life cycle and look for evidence of early invasion dynamics that may affect establishment, including Allee effects, demographic and environmental stochasticity, windows of opportunity and bottlenecks during sexual reproduction. I found that populations introduced at low doses exhibit weak Allee effects during sexual reproduction and that these effects strengthen over the season. Further, probability of establishment is positively related to propagule pressure; however, the relation is highly stochastic. The insights obtained on the characteristics of the relation between propagule pressure and probability of establishment at population and watershed scales can be linked in management plans aimed at slowing the spread of B. longimanus in inland lakes.
L'estimation de la probabilité d'établissement d'espèces invasives est un élément important pour la gestion de leur dispersion. Dans cette thèse, je présente deux approches pour estimer la probabilité d'établissement de Bythotrephes longimanus, un cladocère prédateur qui se disperse à travers des lacs en Ontario et des états Américains avoisinants. D'abord, je développe un modèle basé sur les procédés pour évaluer la probabilité d'établissement de B. longimanus au fil du temps dans un bassin versant. J'emploie des métriques de pression de propagules en provenance de disséminations anthropiques et naturelles afin d'estimer leur dispersion, et j'étends le modèle pour incorporer le manque de certaines informations spatiales et temporelles concernant l'état d'invasion des lacs. Mes résultats indiquent que la circulation de bateaux de plaisance est le vecteur dominant de dispersion et que la majorité du risque d'invasion des lacs est dû à plusieurs points statiques d'invasion - les cinq lacs les plus grands du système. J'étudie ensuite la probabilité d'établissement empiriquement. Je mène une expérience contrôlée afin de suivre des populations récemment introduites de B. longimanus au cours de leur cycle de vie et je cherche des facteurs liées a la dynamique du début d'invasion qui pourraient affecter la façon dont laquelle les pressions de propagules et le succès d'établissement sont reliés. Ceci inclut la stochasticité démographique et environnementale, des opportunités saisonnières où l'établissement est possible, ainsi que la diminution des opportunités et les effets Allee pendant la reproduction sexuelle. Mes résultats suggèrent que l'établissement de B. longimanus est limité aux effets Allee ou aux diminutions d'opportunités pendant reproduction sexuelle pour les populations introduites en doses bases et que les effets Allee se renforcent pendant l'automne. De plus, la probabilité d'établissement est re

Doctor of Philosophy (PhD)
Early recognition of plant invaders is key to their successful management. Yet knowledge of the ecological impacts of species before they become widespread is poor. This thesis examines the ecology of invasive Pinus radiata, a species which is known to spread from introduced plantings in Australia but is currently a low profile invader. Pinus invasions are considered major ecological problems in New Zealand and South Africa where wildlings are beginning to dominate natural areas and suppress native vegetation. Invasion success elsewhere and the large softwood estate in Australia suggest that pines may begin to dominate native eucalypt forests bordering large commercial plantations. This research focused on three components of impact of P. radiata; extent, abundance and effect per individual. The borders of 29 P. radiata plantations in NSW were surveyed in order to quantify the current level of invasion and to identify factors facilitating pine spread. Of particular interest was the role of propagule pressure, vegetation type and fire in the invasion process. The area of land in NSW currently invaded by P. radiata was estimated at almost 4 500 ha, although this is likely to be an underestimate due to an inability to detect wildlings (self-sown pines) at long distances from the plantation. Twenty six of the 29 plantations produced wild pines, however most of the sites are in the very early stages of invasion. Noticeable wildling populations were recorded at nine sites indicating that P. radiata is capable of establishing within native vegetation. Pine spread was most severe in the world heritage listed Blue Mountains region where pine densities reached up to 2000 per hectare in areas adjacent to the plantation and isolated pines were recorded up to 4 km from the source. The presence of isolated pines within intact native vegetation suggests that disturbance is not required for pine establishment in forested environments. Furthermore, high pine emergence and survival rates in eucalypt woodland and evidence of self reproduction by wildlings suggest that in the absence of adequate control measures pines may become established invaders in the Australian landscape. While low levels of current invasion at many sites hindered the ability to examine the factors facilitating invasion some variables that appear to be driving pine success were identified. At the landscape scale plantation size and residence time were significant predictors of the level of invasion at a site. Areas of native vegetation vi adjacent to plantations less than 40 years experienced very low levels of invasion suggesting a lag period between plantation establishment and invasion. However, pines with diameters up to 60 cm were observed growing adjacent to plantations younger than 40 years implying that the first colonisers are capable of establishing soon after plantation trees become reproductive. Propagule pressure was also found to have a strong influence on invasion success on a smaller scale manifesting in a significant positive relationship between the age of a plantation compartment and the likelihood of invasion. A negative relationship between plantation size and level of invasion was a surprising result and was influenced by just two large sites that happened to be located in areas of high rainfall. All sites receiving more than 1300 mm annual rainfall experienced low levels of invasion suggesting that this is a limiting factor for pine spread in NSW. There were significant differences in the level of invasion between vegetation types implying that some communities are more susceptible to invasion. Patterns of spread confirmed ideas regarding the facilitative effect of disturbance in the invasion process and the resistance of wet sclerophyll forest to invasion in Australia. An absence of wildlings in cleared land and areas of remnant bushland was attributed to high levels of grazing pressure. Wind direction did not appear to influence the distribution of pines close to the plantation, but evidence of long distance wind dispersal of pines was provided by an investigation of pine spread from the air at one site where large pines were found growing 10 km downwind from a mature plantation. Fire was found to have both a positive and negative influence on the invasion process. High intensity wildfires are capable of destroying large pines with diameters exceeding 50 cm. However, fire can stimulate seed release from cones resulting in large post-fire recruitment pulses. Seedling densities of up to 3050 per hectare were recorded almost 3 years after wildfire, suggesting that follow up control prior to recruits reaching coning age, i.e. within 5 years, would be beneficial. Surveys of wildling pines exposed to low intensity hazard reduction burns suggest that the majority of pines greater than 3 m in height and with a diameter of more than 10 cm will survive the fire. Low intensity prescribed fires that are carried out after pines have reached this size will fail to control wildling populations. To examine the influence of P. radiata once it has established in the native community this study focused on two mechanisms of impact, the addition of pine litter and increased shade due to an increase in canopy cover. Collection of pine litterfall vii within an invaded eucalypt woodland over a 2 year period recorded rates of up to 1400 kg/ha/year in the most heavily invaded area with a pine basal area of 11.3m2/ha. More than 70 % of pine litter fell directly below the pine canopy suggesting that the most severe litter effects will be limited to these areas. Glasshouse and field experiments were conducted to examine the influence of this increased litter load on the emergence of P. radiata and two native species. Recruitment of native plant species was impeded by litter levels of 6000 kg/ha, the equivalent of approximately 4 years of pine litterfall. Both P. radiata and the two natives responded similarly to pine and eucalypt litter suggesting the two litter types are influencing the recruitment phase equally. However, where pines are added to the system, increased litterfall rates could potentially result in the doubling of the litter load and hence a greater barrier to seedling establishment. Pine invaded eucalypt woodlands are also subjected to three fold increases in canopy cover. Trends in reduced emergence of native species under a pine canopy suggest that the addition of pines to eucalypt forests is likely to have a negative influence on native recruitment and may result in a shift towards a shade tolerant community. However, reversal of trends in emergence below pine canopy between seasons implies that quantifying invasion impacts requires a consideration of temporal variation. Increased levels of disturbance, forest fragmentation and an increasing pine estate are likely to lead to the infestation of new areas. Furthermore the lag phase associated with pine spread means that even if no new plantations are established the number of invasion events will increase. This study has identified a number of risk factors that can be used to guide plantation establishment and the management of invasion events. Minimising disturbance at plantation borders and increasing the ‘no planting’ zone will help to reduce the impacts of pines. Where possible new plantations should be established upwind of cleared land or at least, wet sclerophyll forest. Frequent monitoring of the borders of plantations yet to source invasions, particularly those greater than 40 years of age, will help identify problem areas before control becomes difficult and costly. Maps of the 29 plantations marked with areas of pine infestation will help prioritise sites for control and provide base level knowledge for future monitoring of pine spread. Stringent legislation that binds plantation managers to control wildlings beyond their boundaries is critical for the effective management of pine invasions. With infinite numbers of invaders and limited funds to dedicate to their control, a method of triaging species for management is critical. This is particularly difficult viii when information is typically biased towards invaders that are already widespread. By focussing on the ecological impacts of invaders it becomes possible to rank species on the basis of the threat they pose to native communities. Ecological research is capable of providing the knowledge to quantify invasion impacts and must remain at the centre of policy decisions.

The global movement of aquatic non-indigenous species can have severe ecological, environmental and economic impacts emphasizing the need to identify potential invaders and transport pathways. Initial transport is arguably the most important stage of the invasion process owing to its role in selectively determining potential invasion candidates. This study characterizes a well defined human-mediated dispersal mechanism, ballast water transport, as a vector for the introduction of non-indigenous zooplankton. Ballast water exchange in the open ocean is the most widely adopted practice for reducing the threat of aquatic invasions and is mandatory for most foreign vessels intending to release ballast in Canadian waters. Ships entering Canadian ports are categorized into the following three shipping classes based on current regulations: overseas vessels carrying exchanged ballast water, intra-coastal vessels carrying exchanged ballast water or intra-coastal vessels carrying un-exchanged ballast water. This study characterizes zooplankton communities associated with each of these shipping classes sampled from ports on Canada’s Pacific coast, Atlantic coast and the Great Lakes Basin. Ballast water samples were collected and analyzed from 77 vessels between 2006 - 2007. The ballast water environment was found to be diverse, with over 193 zooplankton taxa, 71 of which were non-indigenous to their receiving environments. Intracoastal vessels containing un-exchanged coastal water transported the greatest density of non-indigenous zooplankton into Canadian ports. Total zooplankton density was found to be negatively correlated with ballast water age The absence of mandatory ballast water exchange and the younger ballast water age of coastal un-exchanged vessels is likely responsible for the higher density of non-indigenous zooplankton in intracoastal un-exchanged vessels. Propagule pressure, invasion history and environmental suitability are all useful in evaluating invasion potential and all suggest that intracoastal un-exchanged vessels pose the greatest invasion threat to Canadian aquatic ecosystems. In conclusion, although the risk of primary introductions from overseas ports may have been reduced through open-ocean exchange of ballast water, secondary introductions from previously invaded ports in North America may be the primary threat to Canadian aquatic ecosystems via this transport vector.

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Here, we quantified the patterns of change in the similarity of the fish fauna of the Upper Paraná ecoregion, caused by the introduction of non-native species and anthropogenic factors, such as the construction of dams and the elimination of a biogeographic barrier. Additionally, we evaluated the importance of changes in rates of turnover and richness species in homogenization/differentiation dynamics. For the study, we mapped the spatial distribution of fish species through occurrence data georeferenced of Brazilian zoological museums. To evaluate temporal changes in the composition of the fish fauna, the data with the distribution of the species were divided into two periods: historical period composed entirely of native species, and the contemporary period, consisting of historical fauna plus non-native species. Changes in similarity patterns between fish assemblages were quantified using the Jaccard dissimilarity index to the historical and contemporary periods. To evaluate the influence of the non-native richness species, number of dam and distance to the Itaipu Dam on changes of similarity, we performed simple and partial Mantel tests. The results showed that the fish fauna has become more dissimilar over time, due to increases in turnover rates and addition of species. Furthermore, factors related to the heterogeneity of the basins and propagules pressure after removal of a biogeographical barrier has determined the patterns of changes in the similarity of the fish fauna
Nesse estudo foram quantificados os padrões de mudança na similaridade da fauna de peixes da ecoregião do alto Paraná, produzidas pela introdução de espécies não-nativas e de fatores antropogênicos, como a construção de barragens e a eliminação de uma barreira biogeográfica. Adicionalmente, foi avaliada a importância de mudanças nas taxas de turnover e de riqueza nas mudanças de similaridade das assembleias de peixes. Para realizar o estudo, a distribuição espacial de espécies de peixes de dez bacias de drenagem do alto Paraná foi mapeada através de dados de ocorrência geo-espacializados tombados em museus zoológicos de instituições brasileiras. Para avaliar as mudanças temporais na composição da fauna de peixes, os dados com as distribuição das espécies foi divido em dois períodos: período histórico, composto somente por espécies nativas, e período contemporâneo, composto por espécies nativas mais não-nativas. Os padrões de similaridade entre as assembleias de peixes para os períodos histórico e contemporâneo foram quantificados através do índice de dissimilaridade de Jaccard. As mudanças na similaridade entre assembleias foi calculada através da diferença entre as similaridades contemporâneas e históricas. Para avaliar os efeitos da riqueza de espécies não-nativas, número de barragens e área das bacias, e a distância das bacias até a barragem de Itaipu foram utilizados testes de Mantel simples e parciais. Os resultados demostraram que a fauna de peixes tem se tornado mais dissimilar ao longo do tempo, em função de aumentos nas taxas de turnover e de adição de espécies. Além disso, fatores relacionados à heterogeneidade das bacias e da pressão de propágulos após a eliminação de uma barreira biogeográfica tem determinado os padrões de mudanças na similaridade da fauna de peixes
2017-05-18

O movimento dos animais desempenha papel importante na aptidão dos indivíduos, no fluxo gênico, na dinâmica de metapopulações e na persistência, a longo prazo, de espécies em um ambiente. Dessa forma, o objetivo deste estudo foi compreender aspectos relacionados à ecologia do movimento da rã-manteiga (Leptodactylus latrans) e da rã-touro (Lithobates catesbeianus). Para isso, indivíduos de ambas as espécies foram marcados com radiotransmissores e rastreados com o objetivo de coletar informações sobre seus movimentos. Ressaltamos os seguintes resultados deste trabalho: (1) a área de vida média de L. latrans é de 2034,2 m2; (2) com o aumento do tamanho corpóreo de anuros há o aumento do tamanho da área de vida; (3) indivíduos de L. latrans se locomovem mais em noites escuras, chuvosas e quentes; e (4) o aumento da pressão de propágulos não reflete uma maior probabilidade no aumento da distribuição geográfica de uma dada população de L. catesbeianus. Entender os padrões de locomoção dos anfíbios e como utilizam o espaço na natureza é fundamental para o entendimento de processos ecológicos e evolutivos relacionados às histórias de vida das espécies desse grupo. Além disso, esse conhecimento serve como uma ferramenta importante no estabelecimento de limites de áreas de conservação com significado biológico para espécies ameaçadas. Ainda, saber como espécies invasoras se locomovem é um dos primeiros passos para que se tomem medidas de controle populacional e para o bloqueio do avanço da distribuição de espécies exóticas
The ability of animals to move plays an important role in individual fitness, gene flow, population regulation, metapopulation dynamics and long-term persistence of species. The aim of this study was to understand aspects related to the movement ecology of the Butter Frog (Leptadactylus latrans) and the Bullfrog (Lithobates catesbeianus). We collected movement data by radio tracking individuals of both species. Some of the most important results obtained were: (1) the mean home range of L. latrans is 2034.2 m2; (2) increasing body size increases home range size in anurans; (3) individuals of L. latrans are more likely to move larger distances in dark, rainy and warm nights; (4) the increase in propagule pressure does not reflect an increase in the probability of geographical spread of a given population of L. catesbeianus. Understanding how amphibians move and use the space in nature is fundamental for the understanding of basic ecological and evolutionary processes related to life history of the species. Moreover, this knowledge is a useful tool to establish biologically meaningful conservation boundary areas to threatened species. Yet, understanding how invasive species move in nature is one of the first steps to population control and to block the spread of geographical distribution of exotic species

Les invasions biologiques sont reconnues comme un facteur évolutif important sur une échelle de temps courte. Elles affectent notamment la structure génétique des populations, les patrons d'évolution phénotypique et la richesse des faunes de parasites associées aux populations envahissantes. Cette étude se propose de quantifier les conséquences d'une invasion biologique suivant ces trois niveaux (génétique, phénotypique et parasitologique) en prenant pour exemple le cas du tilapia du Mozambique Oreochromis mossambicus. Ce cichlidé africain présente un statut remarquable en biologie de la conservation puisqu'il est à la fois (i) l'une des espèces les plus envahissantes au monde car dispersée à l'échelle globale au cours du XXème siècle et (ii) une espèce " quasi-menacée " (UICN) sur son aire native (partie du sud-est de l'Afrique) du fait de son hybridation massive avec d'autres Oreochromis sp. introduits. La démarche générale employée ici est de décrire l'histoire récente des populations à l'aide de marqueurs nucléaires (AFLP) et des séquences de l'ADN mitochondrial (ADNmt), puis de mettre en relation ces résultats génétiques avec la diversité morphologique et la parasitologie des populations. Deux systèmes différents ont été étudiés : Au sein de l'aire native, l'étude se focalise sur le Limpopo inférieur et le sous-bassin de la Changane (Mozambique). Des patrons d'introgression incluant trois espèces en présence sont détectés, mais les hybrides sont peu fréquents et leur expansion limitée. Ces résultats sont de plutôt bonne augure pour la conservation d'O. mossambicus et ils permettent d'identifier deux zones de conservation prioritaires. L'étude des parasites indique une plus grande diversité parasitaire mais de faibles prévalences dans les sites de moindre valeur en conservation, ce qui pourrait favoriser le succès des espèces introduites et de leurs hybrides. Parmi les territoires envahis, les AFLP et l'ADNmt soutiennent une homogénéité générale et une diversité génétique faible, qui sont interprétées comme le résultat d'un fort goulot d'étranglement précédant l'expansion à l'échelle mondiale. Une structure des populations en lien avec la géographie à large échelle (Nouvelle-Calédonie, Guadeloupe, Jamaïque) est cependant détectée. La variation de la forme du corps est également structurée à large échelle géographique, ce en dépit des fortes variations environnementales enregistrées à l'échelle locale. Cela suggère un effet des contraintes génétiques sur la diversification morphologique contemporaine. L'absence de parasites monogènes sur les populations introduites en Nouvelle-Calédonie peut être mise en relation avec un évènement fondateur, et est proposé comme l'un des facteurs ayant pu favoriser le succès de l'espèce. En conclusion, une faible diversité génétique ne contraint vraisemblablement pas un potentiel envahissant élevé et une diversification rapide chez les tilapias.

Invasões biológicas tem um papel cada vez mais importante nas políticas ambientais, visto que espécies invasoras desempenham uma crescente influência sobre novos ambientes onde são introduzidas, podendo gerar grandes impactos naturais e financeiros. Estudos na área da biologia da invasão se fazem extremamente necessários para remediar e evitar novas introduções. Dentre as metodologias aplicadas ao estudo das invasões biológicas, a genética de populações apresenta diversas ferramentas uteis para responder perguntas relevantes nos esforços de controle de espécies invasoras. No presente trabalho usamos recursos moleculares aplicados à genética de populações da rã-touro (Lithobates catesbeianus), o anfíbio invasor mais disseminado no planeta. Através deste estudo foi possível compreender mais sobre a estrutura genética das populações invasoras do Brasil e do mundo, além de seu histórico de invasão e sua população nativa de origem. Entender e contextualizar as características e motivos que levam ao sucesso de uma invasão biológica é importante para esforços de combate a pragas e para evitar que outros invasores se fixem em novos ambientes. Este trabalho levantou novos conhecimentos que podem e devem ser usados em políticas de combates à invasão da rã-touro
Biological invasions play an increasingly important role in environmental policies as invasive species represent a growing impact in new environments where they are introduced, potentially causing large natural and financial problems. Studies in the field of invasion biology are extremely necessary to remedy and prevent new introductions. Among the methodologies applied to the study of biological invasions, population genetics presents several useful tools to answer relevant questions in efforts to control invasive species. In the present work we used molecular resources applied to the genetics of populations of the American Bullfrog (Lithobates catesbeianus), the most widespread invasive amphibian on the planet. Through this study it was possible to understand more about the genetic structure of the invasive populations in Brazil and in the world, and its history of invasion and its native population of origin. Understanding and contextualizing the characteristics and motives that lead to the success of a biological invasion is important for pest control efforts and to prevent other invaders from focusing on new environments. This work has raised new knowledge that can and should be used in policies to combat Bullfrog invasion

This project investigated the fundamental principles of the Theory of Island Biogeography. How biodiversity is influenced by habitat age, size, isolation and quality was studied in two little-known ecosystems; pumice-rafted marine communities that travel through the Pacific Ocean and strand on shorelines, and Queensland's critically endangered Melaleuca irbyana forests. This research found that while habitat age, size and isolation were important for species richness; habitat quality, defined by resource availability and climate, was more influential for predicting biodiversity levels. Small pumice stones and small remnant forests can provide the conditions species need to prosper. Therefore, small and isolated habitats are also important to conserve.

Non-indigenous species have become a global issue of increasing importance in recent years, with many causing significant environmental and economic damage. Identifying locations vulnerable to invasion allows for focus of management efforts towards prevention of invasions at those locations. In order to determine whether constructed water bodies, such as reservoirs, ornamental lakes or retired mines, are more easily invaded environments than natural water bodies, owing to decreased biotic resistance, the distributions of native and non-indigenous freshwater calanoid copepod species in the North Island were examined. Calanoid copepods in ports and other coastal environments were also examined, in order to determine whether ports are more frequently invaded owing to increased propagule supplies from visiting ships and other sources. The distributions of the native freshwater calanoid copepod species Boeckella hamata, B. propinqua, B. delicata and B. tanea are confined in the North Island of New Zealand to specific technostratigraphic terranes when natural waters only are examined, and as such each species can be considered to have a native range. The recently colonised calanoid copepod species Boeckella minuta (6 locations), Skistodiaptomus pallidus (3 locations) and Sinodiaptomus valkanovi (2 locations) are to date confined to constructed water bodies. Boeckella symmetrica (2 locations) may be confined to constructed water bodies, but the status of one location is unclear. Boeckella triarticulata, a species common in the South Island, is known only from a single farm dam in the North Island. The native species Boeckella hamata, B. propinqua and B. delicata were found to occur in constructed waters, but only B. propinqua was found in constructed water bodies outside their natural ranges (9 locations). Calamoecia lucasi is found in lakes throughout most of the North Island, and is not confined to any one terrane. My results indicate that constructed water bodies are more easily invaded by non-indigenous species than natural water bodies, represents a potential pathway for future invaders to establish, and provides locations for species to spread. In order to determine whether recently established freshwater calanoid copepod species have the potential to spread from their present habitats into other water bodies, the prosomal lengths of non-indigenous calanoid copepod species were measured and compared with those for native species. The results suggest that dietary overlap should prevent the non-indigenous species present to date from spreading into any water bodies with established Boeckella populations, although Sinodiaptomus valkanovi and Boeckella triarticulata could potentially spread to lakes containing only Calamoecia lucasi. Data on the co-occurrences of native freshwater calanoid copepod species support the theory of dietary exclusion, as Boeckella species have not been found to coexist. In order to test whether New Zealand marine environments have been invaded by non-indigenous calanoid copepods, and whether ports have been more regularly invaded than non-port areas, calanoid copepods were sampled from various coastal locations around the North Island. With the possible exception of Sulcanus conflictus, no non-indigenous species were found, indicating that non-indigenous marine calanoid copepod species are not establishing in New Zealand despite a history of invasion elsewhere.

Many regional and local factors can influence the distribution of native and exotic species in ecological communities. I examined the regional- and local-scale determinants of native and exotic vascular plant species richness and composition in a highly fragmented oak savanna ecosystem on southeastern Vancouver Island. In sharp contrast to most reported results, I found a negative relationship between native and exotic richness at the regional scale, and no relationship at the local scale. Two extrinsic factors, surrounding road density and climate, best explained the regional-scale relationship by each affecting natives and exotics in opposite ways. Road density and climate were also the dominant predictors of native and exotic composition at the regional scale. Patterns in the patch occupancy of individual species confirmed the importance of these factors but I found that low surrounding road densities and cool, wet conditions predicted the presence of many natives and the absence of many exotics. Environmental factors explained variation in richness and composition at the local scale, but these factors were different for natives and exotics. My results suggest that natives and exotics respond to roads and climate in fundamentally different ways. Roads increase both exotic propagule pressure and disturbance, which may facilitate exotic invasion. In contrast, disturbance from roads may increase the likelihood of local extinction for particular natives. Differing climatic preferences within the native and exotic species pools may also partially explain the observed patterns. There was no evidence that native diversity directly affects exotic diversity (or vice versa). Surprisingly, I found that connectivity was not an important predictor of richness or composition despite the high degree of habitat fragmentation in this ecosystem.

Invasive species represent a major concern; they can result in serious ecological and economic losses and are recognised as one of the most serious threats to global species diversity. Plant invasions are of particular concern in New Zealand, which has high proportions of both naturalised and endemic plant species. In this thesis I focussed on the invasive plant Hieracium lepidulum, an exotic weed introduced from Europe to New Zealand prior to 1941. It is invasive in a variety of habitats in the South Island, where it has steadily increased in distribution and abundance over the last 50 years, and is thought to have detrimental impacts on native plant communities. I investigated factors influencing its invasion success and tested for impacts on native plant communities, making extensive use of existing plots into which H. lepidulum was experimentally introduced in 2003. I examined how community richness, turnover, resource availability and propagule pressure of the invader interacted to determine the invasion success of H. lepidulum. Results differed markedly above and below treeline. Above treeline, plots with higher richness and turnover were more invaded; below treeline, plots with higher available light were more invaded. In both habitats, these findings were modified by the influence of propagule pressure; at low propagule pressure, site characteristics were non-significant in explaining invasion success, while at higher propagule pressure these effects became significant. To test for impacts resulting in altered community composition and structure, I looked for changes in community richness, diversity and evenness subsequent to H. lepidulum introduction. As impacts may be more apparent at fine spatial scales, I made measurements at a 5 x 5 cm cell scale in addition to the established 30 x 30 cm plot scale. Plot species richness increased from 2003 to 2009 and a component of this increase was associated with H. lepidulum density. Other relationships between the plant community and H. lepidulum were generally non-significant. Results showed that H. lepidulum has had no negative effects on community richness, evenness or diversity. Despite being able to opportunistically colonise grassland sites with high turnover, and forest sites subject to canopy disturbance, dependant on propagule pressure, it appears H. lepidulum has not impacted community composition or structure.

The spread of exotic species is considered to be one of the most significant threats to ecosystems and emphasises the need for appropriate management interventions. The majority of marine non-indigenous species (NIS) are believed to have been introduced via ship biofouling and their domestic spread continues to take place via this mechanism. In some countries, biosecurity systems have been developed to prevent the introduction of NIS through biofouling. However, implementing biosecurity strategies is difficult due to the challenges around identifying high-risk vectors. Reliable predictors of risk have remained elusive, in part due to a lack of scientific knowledge. Nonetheless, invasion ecology is an active scientific field that aims to build this knowledge. Propagule pressure is of particular interest in invasion ecology as it describes the quantity and quality of the propagules introduced into a recipient region and is considered to be an important determinant in the successful establishment of NIS. Environmental history affects health and reproductive output of an organism and, therefore, it is beneficial to examine this experimentally in the context of biofouling and propagule pressure. The aim of this thesis was to examine how voyage characteristics influence biofouling recruitment, survivorship, growth, reproduction and offspring performance through the ship invasion pathway. This was to provide fundamental knowledge to assist managers with identifying high-risk vessels that are likely to facilitate the introduction or domestic spread of NIS, and to understand the processes affecting biofouling organisms during long-distance dispersal events. Chapter One provides an introduction to the issues addressed in this thesis. Each data chapter (Chapters Two – Five) then focused on a stage of the invasion process and included field experiments using a model organism, Bugula neritina. Finally, Chapter Six provides a summary of key findings, discussion and the implications to biosecurity management. Throughout this thesis, the effect of donor port residency period on the success of recruits was highlighted. Chapter Two focused on recruitment in the donor region. As expected, recruitment increased with residency period. Importantly, recruitment occurred every day on vulnerable surfaces, therefore, periods as short as only a few days are able to entrain recruits to a vessel hull. The study presented in Chapter Three showed that there was high survivorship of B. neritina recruits during 12 translocation scenarios tested. In particular, the juvenile short-residency recruits (1-8 days) survived voyages of 8 days at a speed of 18 knots; the longest and fastest voyage simulated. Interestingly, variation in voyage speed and voyage duration had no effect on the survivorship of recruits, but did have legacy effects on post-voyage growth. Again, B. neritina which recruited over very short residency periods of 1 day continued to perform well after translocation and had the highest level of reproductive output after the voyage scenarios (Chapter Four). Recruits that were older (32-days) and reproductively mature at the commencement of the scenarios failed to release any propagules. Even though the number of ‘at sea’ and ‘port residency’ days were equal, reproductive output was higher after short and frequent voyages than after long and infrequent voyages. Finally, the study presented in Chapter Five examined transgenerational effects of B. nertina. Results showed that although the environmental history of the parent colony had a carry-over effect on offspring performance, it was the offspring environment that was a stronger determinant of success (measured by reproductive output and growth). Although cross-vector spread is possible (i.e. parent and offspring both fouling an active vessel), offspring released from a hull fouling parent into a recipient environment will perform better. In combination, these studies have provided new insights into NIS transport via vessel biofouling. Although shipping pathways are dynamic and complex, these results suggest that juvenile stages that recruit over short residency periods and are then translocated on short voyages, may pose a higher risk for NIS introduction than originally assumed. This has implications for marine biosecurity management as short residency periods are common and short, frequent voyages are typical of domestic vessel movements which are largely unmanaged.

Many recent studies have demonstrated that propagule pressure is a useful predictor of patterns of invasions by non-native species. However, most of these studies have used only current, not historical, data to estimate propagule pressure. Recognizing the potential importance of propagule pressure over time, I used surrogate variables that represent both past and present propagule pressure, for example, the length of time a surrounding area had been developed. I quantified the relationships between these surrogate variables and the distribution and abundance of non-native woody plant species in central Texas. I constructed statistical models predicting native and non-native species richness and the occurrence of five common species using a set of six ecological and five development-related predictor variables. I compared all models using the corrected Akaike information criterion (AICc). Overall, age of residential development surrounding native woodlands was the best predictor, other than community type, of non-native species richness. As expected, areas near older developments had more non-native species than areas near newer developments. Surprisingly, age of development and average city age, two different measures of the length of time that landscaping (a major source of propagules of non-native woody species in this region) had been present nearby, were much better predictors than distance to source populations. Age of development and average city age (weighted by distance from the site) were also both correlated with distance to source populations; this may be true in other systems as well. This suggests that the reason distance to source population has been a successful predictor of invasion may be because it is a surrogate for an underlying causal variable, length of time of exposure to source populations. Future studies of non-native invasions would benefit from taking into account both past and present propagule pressure: age of residential development and city age could be useful surrogates in other systems.
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碩士
國立臺灣大學
昆蟲學研究所
99
Genetic diversity is one of the key determinants for species’ survival and evolution. From the conservation biology perspective, a small population is prone to extinction by inbreeding depression or accumulation of deleterious mutations. Invasive species, however, are generally successful in introduced ranges despite loss of genetic diversity due to small founder population sizes, which gives rise to a paradox between invasion and conservation biology in term of genetic diversity. Social insects, such as ants, wasps, and termites, represent a highly successful group of invasive species not only because their small founder population sizes often lack immediately recognizable signature but also their great dispersal power, high reproductive rate and the competitive ability of “super individuals”. One of the notorious ant invaders, the red imported fire ant (Solenopsis invicta), originating from South America, has been accidentally introduced into the USA, Australia, Taiwan and China. Invasions by fire ants in these areas provide an excellent framework to investigate the factors contributing to the successful establishment of social insects especially since this species utilizes a single locus complementary sex determination (sl-CSD) system. Theoretically, the genetic load is predicted to be much more profound for S. invicta due to the mass production of sterile diploid males imposed by loss of genetic diversity. This study therefore aimed to determine how fire ants overcome such a disadvantage using population genetic approaches. An assessment of genetic variation at 12 microsatellite loci in 120 fire ant colonies in northern Taiwan (Taoyuan) revealed that Taoyuan was colonized by ants on a minimum of two occasions. Furthermore, estimates of genetic diversity at the CSD locus by calculating the proportion of match matings (queens mated with males harboring identical CSD allele) do not differ significantly between fire ant populations in Taiwan and the putative source, the USA, suggesting that the successful invasion of fire ants is likely attributable to adequate genetic diversity maintained by larger propagule number (e.g, multiple invasions). In summary, this study not only highlights that cryptic invasions might be undetected until the deep genetic structure of focal populations has been revealed but also signifies the importance of genetic diversity in shaping the invasion success of fire ants.

Invasive alien species (IAS) pose a serious threat to ecosystems and societies worldwide. Local ecological knowledge (LEK) is increasingly valued as a means of understanding environmental issues; however, its application in the context of IAS research has been limited. The overall objective of this study was to document the LEK of farmers and Weed Supervisors to gain insight into a recent IAS, Odontites serotina. I conducted semi-structured interviews with farmers and Weed Supervisors with O. serotina management experience. Results indicated that the socio-economic impacts for farmers were severe in affected rural communities. However, participants had developed promising control techniques, including the application of compost mulch. I used this LEK as well as data on species occurrence, environmental variables, and measures of propagule pressure to forecast the potential distribution of O. serotina across Manitoba. The risk map generated will be useful for guiding future monitoring and public outreach efforts.

La germination des spores est une étape essentielle dans le cycle de vie de la majorité des champignons filamenteux. Les champignons mycorhiziens à arbuscules (CMA) forment un certain nombre de propagules infectieuses différentes qui augmentent leur potentiel à coloniser les racines. Parmi elles se trouvent les spores extraracinaires et intraracinaires. La paroi cellulaire des spores joue un rôle majeur dans la survie de ces propagules en étant une barrière physique et osmotique. Puisque une cellule peut faire des ajustements considérables dans la composition et la structure de sa paroi, en réponse aux conditions environnementales, il est possible que les parois des spores intraracinaires et extraracinaires montrent des propriétés mécaniques et osmotiques différentes affectant leur germination et leur survie. Pourtant, contrairement à la connaissance de la génétique moléculaire et de la formation de la paroi cellulaire des CMA, peu d’information est disponible au sujet de ces propriétés mécaniques. Les informations sur la germination des CMA dans des conditions hypertoniques sont aussi rares, et les modèles expérimentaux ne séparent généralement pas les effets directs de la forte pression osmotique externe sur la germination des champignons et les effets attribuables aux plantes. Cette étude avait pour but de répondre à deux importantes séries de questions concernant le comportement des spores mycorhiziennes. Nous avons d'abord déterminé la relation entre la composition de la paroi cellulaire, la structure et les propriétés mécaniques du champignon modèle Glomus irregulare (isolat DAOM 197198). La micro-indentation a été utilisée pour mesurer quantitativement les propriétés mécaniques de la paroi cellulaire. La composition (contenu de chitine et de glomaline) de la paroi cellulaire a été quantifiée par immunofluorescence tandis que la microscopie optique a été utilisée pour mesurer l'épaisseur de la paroi cellulaire. La densité locale en glomaline et l’épaisseur de la paroi étaient significativement plus élevées pour les parois des spores extraracinaires alors que la densité locale en chitine et la rigidité n’ont pas montré de variations entre les spores extraracinaires et intraracinaires. La grande variabilité dans les paramètres étudiés nous a empêchés de cibler un facteur principal responsable de la force totale de la paroi lors de la compression. La diminution des concentrations de chitine et de glomaline a été corrélée à l'évolution de la paroi du champignon au cours de son cycle de vie. On a aussi observé une composition différentielle des couches de la paroi: les polymères de chitine et de glomaline furent localisés principalement dans les couches externes et internes de la paroi, respectivement. Dans la deuxième partie de notre travail, nous avons exploré les effets directs d'engrais, par rapport à leur activité de l'eau (aw), sur la germination des spores et la pression de turgescence cellulaire. Les spores ont été soumises à trois engrais avec des valeurs de aw différentes et la germination ainsi que la cytorrhyse (effondrement de la paroi cellulaire) des spores ont été évaluées après différents temps d'incubation. Les valeurs de aw des engrais ont été utilisées comme indicateurs de leurs pressions osmotiques. L'exposition des spores de Glomus irregulare au choc osmotique causé par les engrais dont les valeurs de aw se situent entre 0,982 et 0,882 a provoqué des changements graduels au niveau de leur cytorrhyse et de leur germination. Avec l'augmentation de la pression de turgescence externe, la cytorrhyse a augmenté, tandis que le taux de germination a diminué. Ces effets ont été plus prononcés à des concentrations élevées en éléments nutritifs. La présente étude, bien qu’elle constitue une étape importante dans la compréhension des propriétés mécaniques et osmotiques des spores de CMA, confirme également que ces propriétés dépendent probablement de plusieurs facteurs, dont certains qui ne sont pas encore identifiés.
Spore germination is an essential developmental stage in the life cycle of many filamentous fungi. Arbuscular mycorrhizal fungi (AMF) form a number of different infectious propagules that increase their potential to colonize roots. Among them are extraradical and intraradical spores. The spore cell wall plays a major role in the survival of these propagules by being a physical and osmotic barrier. Because a cell can make considerable adjustments to the composition and structure of its wall in response to environmental conditions, it is possible that intraradical and extraradical spore walls show different mechanical and osmotic properties affecting their survival and germination. However, in contrast to the knowledge on the genetics and molecular composition of AMF cell wall, little is known about its mechanical properties. Information on the germination of AMF under hypertonic conditions is scarce, and experimental designs and methodologies have generally not allowed the direct effects of high external osmotic pressure on fungal germination to be separated from plant-mediated effects. This study had the goal to address two important sets of questions regarding the behavior of mycorrhizal spores. We first determined the relationship between cell wall composition, structure and mechanical properties of the model fungus Glomus irregulare. Micro-indentation was used to quantitatively measure the cell wall mechanical properties. Cell wall composition (chitin and glomalin content) was studied by immunofluorescence whereas optical microscopy was used to measure the cell wall thickness. Glomalin local density and wall thickness were both significantly higher for extraradical spore walls while chitin local density and rigidity were unaffected by origin of spores. High variability in results prevented us from identifying a primary factor responsible for overall wall strength during compression. Decreases of chitin and glomalin concentrations were correlated to the development of the fungal wall throughout its life-cycle. There was also differential association within the wall layers: The chitin and glomalin polymers were localized mostly in the outer and inner walls, respectively. In the second part of our work, we explored the direct effects of fertilizers, in relation to their water activity (aw), on spore germination and cellular turgor pressure. Spores were exposed to three fertilizers with different aw and spore germination and cytorrhysis of spores were assessed after different times of incubation. Water activities of the fertilizers were used as indicators of their osmotic pressures. Osmotic shock exposure of the Glomus irregulare spores to fertilizers at aw values between 0.982 and 0.882 caused gradual changes in cytorrhysis and germination. With the increase of external turgor pressure, cytorrhysis increased while the rate of germination decreased. These effects were most pronounced at high nutrient concentrations. The present investigation, while likely representing a significant step forward in understanding the mechanical and osmotic properties of AMF spores, also confirms that they might depend on many, as yet unidentified factors. Future research should examine differences in the physiology to discern reasons for such differences in spore properties.