Marine seaweeds constitute one of the most productive plant systems known on Earth and a rich fauna including juvenile fish and crustaceans is dependent on the habitats they form. Human influence on marine costal ecosystems has resulted in large scale changes to the abundance and distribution of species, where species introductions constitute an obvious part. The aims of this thesis were to 1) explore how non-native seaweeds impact on ecosystem functions (primary production and decomposition), and 2) study how interactions between non-native seaweeds and native communities affect invasion success. I used a combination of laboratory assays, outdoor mesocosms and field experiments. Paper I and II revealed that the impact on ecosystem functions were substantially different depending on the identity of the invader. The highly successful non-native red alga Heterosiphonia japonica had a large effect on community productivity. Due to the rapid growth of the invader, the primary production increased by more than four times in mixed species communities with the invader compared to  communities with only native species. In contrast, the morphologically similar and equally successful non-native red alga Bonnemaisonia hamifera grew slowly and had no effect on community production. But B. hamifera produces a potent defense compound that deters native herbivores and reduces the growth of micro-organisms. As a direct or indirect effect of this chemical defense, the litter from B. hamifera decomposed considerably slower compared to native seaweed litter. Rapid growth and defense against predation are likely important in explaining how the two invaders have become successful in the invaded range. These results show that traits related to invasion success may determine impacts on native communities. Paper III shows that the rapidly growing invader H. japonica is avoided as food by native herbivores, which likely enables the invader to survive during colder seasons with sub-optimal growth conditions.  In paper IV I found that competition from the native brown alga Fucus vesiculosus decreased growth of the non-native congener Fucus evanescens. Native herbivores caused more damage to the native competitor but it did not relieve F. evanescens from competitive pressure. Several native brown algae grow in the niche of F. evanescens, which may explain why the species only is growing sparingly in the invaded range. The results indicate that competition with native seaweeds have potential to reduce the success of non-native seaweeds in the new range. In summary, this thesis shows that non-native seaweeds differ strongly in their effect on ecosystem functions. Knowledge of which traits are present among abundant non-native species and how these traits relates to different effects may enable us to gain a better understanding of invasion impacts on native communities. The thesis also highlights that competitive interactions can be of importance for invasion success in seaweed communities. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Manuscript. Paper 3: Manuscript. Paper 4: Manuscript.</p> / Alien-native trophic interactions: consequences for invasion success and ecosystem effects of invasions

No Brasil, varias espécies de gramíneas africanas introduzidas se tornaram importantes invasoras dos cerrados e constituem uma das principais ameaças para a sua biodiversidade. Atributos envolvendo trade-offs evolutivos que influenciam na adaptabilidade das espécies - aquisição, alocação e perda de recursos pelas plantas por meio de competição, facilitação, estresse e distúrbio - irão influenciar o desempenho dos indivíduos e a manutenção das populações nas comunidades vegetais. Deste modo, e de extrema importância entender quais os mecanismos que levam uma espécie invasora a ter um desempenho melhor sobre as espécies nativas. Comparações envolvendo características entre espécies exóticas e nativas podem levar a uma melhor compreensão sobre o processo da invasão. Diante disso, este trabalho procurou identificar algumas das estratégias competitivas adotadas por uma gramínea invasora, Urochloa decmbens, e por uma gramínea nativa, Echinolaena inflexa, em ecossistema de cerrado. Para verificar aspectos relativos às estratégias de colonização das espécies foram analisadas suas fenologias reprodutivas, as taxas de viabilidade e de germinação das sementes. Em um experimento com plantas envasadas, sementes de ambas as espécies foram semeadas em diferentes proporções relativas uma a outra. Três censos foram realizados durante o período de um ano para avaliar a sobrevivência, crescimento e fecundidade das espécies. Medidas da capacidade fotossintética e das taxas de assimilação foram tomadas para ambas as espécies tanto em parcelas puras quanto em parcelas mistas; também foi quantificada a área foliar especifica e total das espécies. No experimento com as plantas envasadas, quando sementes de ambas as espécies foram colocadas juntas, a nativa teve seu crescimento suprimido pela invasora. Nos tratamentos puros, os indivíduos da espécie nativa cresceram em tamanho, mas somente três se tornaram reprodutivos. No entanto, muitos indivíduos da espécie invasora cresceram em tamanho e se reproduziram em todos os tratamentos, mostrando uma alocação de recursos tanto para crescimento quanto para reprodução. A espécie invasora apresentou uma taxa fotossintética maior, porem ambas as espécies tiveram sua capacidade fotossintética e sua taxa fotossintética reduzida nos parcelas mistas, sugerindo que a competição afeta tanto a espécie nativa como a espécie invasora. Uma vez que a espécie nativa possui área foliar especifica menor em relação à espécie invasora, espera-se que ela apresente uma perda menor de nutrientes e, conseqüentemente, uma maior conservação/retenção dos nutrientes, o que favoreceria sua persistência em ambientes como o cerrado. No entanto, com a ocorrência de distúrbios e possíveis alterações na disponibilidade de nutrientes, a gramínea nativa pode ser deslocada pela invasora, que apresenta maior capacidade de colonizar novas áreas por meio de maiores taxas de germinação e estabelecimento; e maior produtividade, apresentando maiores taxas fotossintéticas, sendo mais eficiente no uso do nitrogênio. / Many African grasses have been introduced in Brazil and have invaded areas of Brazilian savannas, thus becoming a major threat to the biological diversity of this biome. Traits involving evolutionary trade-offs that influence the species adaptive strategies, such as differences in nutrient uptake, nutrient loss and biomass allocation in response to plant competition and facilitation, and environmental stress and disturbance, will influence individual performance and population maintenance in plant communities. Therefore, it is extremely important to identify the mechanisms associated with invasiveness that lead to a better performance of exotic species over co-occuring natives. Comparisons involving traits of both native and exotic species may lead to a better understanding concerning the success of invasions. This study sought to recognize possible competitive strategies adopted by an invasive grass, Urochloa decmbens, and a native grass, Echinolaena inflexa, in a Brazilian savanna. In order to identify aspects relative to their colonization strategies, the reproductive fenologies and seed viability and germination rates of both species were analyzed. An experiment with potted plants was performed where seeds of both species were sown in different relative proportions to each other. Three census were carried out during a period of 12 months to evaluate survival, growth and fecundity of both species. Gas exchange and chlorophyll fluorescence were quantified for each species in pure and mixed stands in the field; total and specific leaf area for both species were also measured. In the potted experiment, when seeds of both species were sown together, the growth of the native species was suppressed by the exotic species. In pure treatments, however, individuals of the native species grew in size, but only 2% reproduced. On the other hand, individuals of the exotic species grew in size and reproduced in all treatments, demonstrating that this species allocated enough resources for both growth and reproduction. The photosynthetic rates were greater for the invasive species, however, in mixed stands, both species had their photosynthetic rates and capacities decreased. The smaller specific leaf area of the native species suggests lower rates of nutrient loss than the invasive species, which would favor its persistence in nutrient poor environments, as the Brazilian savannas. Yet, with the occurrence of disturbances and variable nutrient availabilities, the native species might be displaced by the invasive species, since the later possesses traits leading to a higher competitive ability in such conditions.

Der Anteil an nicht-einheimischen Pflanzenarten (Neophyten), die durch menschlichen Einfluss in neue Gebiete eingebracht worden sind, hat in den letzten zwei Jahrhunderten deutlich zugenommen. Weltweit gefährdet die Invasion von Neophyten beträchtlich die einheimische Biodiversität und Ökosystemfunktionen. Verschiedene biologische Eigenschaften (z.B. hohe Zuwachsrate und schnelle Vermehrung) führen zu einer besseren Konkurrenzfähigkeit von invasiven Arten und verursachen Veränderungen in der natürlichen Artzusammensetzung. Eine genauere Erfassung der Eigenschaften, die Neophyten zu starken Konkurrenten machen, könnte dabei helfen pflanzliche Invasionen besser zu verstehen und zukünftig effektiver zu steuern. Topfversuche ermöglichen die Untersuchung von Pflanzeninteraktionen unter kontrollierten Bedingungen ohne den schwer kalkulierbaren Einfluss heterogener Umweltfaktoren. Allerdings führen die Langlebigkeit und die größeren Dimensionen von Baumindividuen zu mehr Problemen in Topfversuchen im Vergleich zur Untersuchung krautiger Pflanzen. Aus diesem Grund wurde im Rahmen eines Reviews Literatur ausgewertet, um einen Überblick über die praktische Durchführung von Topfversuchen, die sich ausschließlich mit Baumarten beschäftigen, zu geben. Es ist offensichtlich, dass der Vorteil von Topfversuchen zugleich auch einen Nachteil darstellt: Aufgrund der kontrollierten Bedingungen sind Topfversuche in ihrer Eignung natürliche Gegebenheiten zu imitieren immer eingeschränkt. Die Zuverlässigkeit von Topfversuchen bei der Vorhersage des Baumwachstums unter natürlichen Bedingungen ist daher problematisch. Eine Möglichkeit um die Übertragbarkeit von Topfversuchen zu verbessern, könnte die Durchführung zusätzlicher Felduntersuchungen sein. In einem Topfversuch wurden die, durch Unterschiede in der Wuchsrate, Biomasseproduktion und Biomasseverteilung bedingten, Konkurrenzmechanismen von zwei einheimischen (Quercus robur L., Carpinus betulus L.) und zwei nicht-einheimischen Baumarten untersucht (Prunus serotina Ehrh., Robinia pseudoacacia L.). Einjährige Jungpflanzen wurden verschiedenen intra- und interspezifischen Konkurrenzbedingungen ausgesetzt, mit oder ohne den Einfluss von Wurzelkonkurrenz. Um die Konkurrenzmechanismen genauer zu bestimmen, wurde zwischen Wurzel- und Sprosskonkurrenz unterschieden, indem entweder ober- oder unterirdische Plastiktrennwände in die Töpfe integriert wurden. Es wurde angenommen, dass die Gesamtbiomasseproduktion der Neophyten im Vergleich zur Biomasseproduktion der einheimischen Baumarten signifikant höher ist und dies zu einer Verringerung der Biomasse von Q. robur und C. betulus führt. Des Weiteren wurde der Einfluss der unterirdischen Konkurrenz auf das Wachstum und die Biomasseverteilung der einheimischen Arten gemäß der ‚balanced-growth hypothesis‘ untersucht. Unsere Ergebnisse bestätigen die Annahmen, dass die Biomasseproduktion der beiden Neophyten P. serotina und R. pseudoacacia signifikant höher ist und dies zu einem großen Konkurrenzvorteil und zu einer Biomassereduktion der beigemischten konkurrenzschwächeren einheimischen Arten führt. Der Konkurrenzdruck auf Q. robur und C. betulus wurde vor allem durch die Wurzelkonkurrenz der nicht-einheimischen Arten verursacht. Die Ausschaltung von unterirdischen Pflanzeninteraktionen durch Trennwände führte somit zu einem Anstieg der Biomasseproduktion der beiden einheimischen Arten. Demzufolge scheint sogar ein begrenztes Wurzelvolumen bessere Wachstumsbedingungen zu bieten als direkter Wurzelkontakt mit invasiven Konkurrenten. In Übereinstimmung mit der ‚balanced-growth hypothesis‘ reagieren Q. robur und C. betulus auf die starke unterirdische Konkurrenz durch die Neophyten, indem sie mehr Biomasse in Richtung der Wurzeln transportieren. Die verstärkte Investition der Pflanzen in die Wurzeln geht vor allem zu Lasten von Blatt- und Astbiomasse. Außerdem hat sich gezeigt, dass Artenmischungen aus einheimischen und nicht-einheimischen Bäumen mehr Biomasse produzieren, als man anhand des Wachstums dieser Arten in Monokulturen erwartet hätte. Im Vergleich zu Monokulturen oder Mischungen beider Neophyten war der Konkurrenzdruck für P. serotina und R. pseudoacacia in Mischungen mit den weniger produktiven einheimischen Baumarten geringer. Bei Betrachtung der beiden nicht-einheimischen Arten wird deutlich, dass P. serotina signifikant mehr Biomasse produziert. Trotzdem hat R. pseudoacacia aufgrund der starken Wurzelkonkurrenz einen negativen Einfluss auf die Biomasseproduktion von P. serotina. Wachsen die beiden konkurrenzstarken Neophyten zusammen in einem Topf, produzieren sie weniger Biomasse als in den entsprechenden Monokulturen. Es gibt Anzeichen dafür, dass die starke Konkurrenzfähigkeit der invasiven Neophyten oftmals zu Lasten ihrer Stresstoleranz geht. Damit einhergehend zeigten die beiden nicht-einheimischen Arten im Topfversuch eine höhere Mortalitätsrate: Vor allem P. serotina scheint zudem empfindlich gegenüber Schatten, Trockenheit und Überflutung zu sein. Möglicherweise könnte diese Schwachstelle der Neophyten genutzt werden, um eine weitere Ausbreitung einzudämmen.

The spread of invasive plant species in natural habitats has become a worldwide problem with negative environmental and economic impacts. An increasing number of invasive organisms are responsible for adverse environmental and economic impacts worldwide, including species extinction, crop failures, reduced water supply, and damage to industrial infrastructures (KERCHER et al., 2007). Phalaris arundinacea L. is widespread throughout the world, except Antarctica and Greenland. Center of diversity of this genus is in the Mediterranean. Members of the genus Phalaris occurs in moist habitats from lower to alpine altitudes (ANDERSON, 1997). Phalaris has a plethora of uses. Its most frequent use is as the root wastewater treatment plants. Phalaris grown as feed for livestock and is also used as an ornamental grass. Phalaris have recently received a lot of attention as a new biomass source for the production of renewable energy in USA. In recent years there has been a massive spread of P. arundinacea across North America (currently occurs in 43 states) and Canada (ZEDLER & KERCHER, 2004). Phalaris represents a significant threat to its original wetland vegetation and is classified as a harmful agens in nine state of U.S. states (LAVERGNE & MOLOFSKY, 2004). It is believed that these aggressive population have European origin.

All three jewelweeds species (Impatiens. L.)contain series of secondary metabolites from category of phenolic substances, e.g. naphthoquinones. This substances belong to allelochemicals, they can to affect germinability and growth of other plants. The extracts from jewelweeds (leaves, roots) and their fractions can to affect negatively germinability and growth of the chosen weed's seeds. Knowledges of allelopathic effects in jewelweeds species on other plants might be of important impact on ecological farming in the implementing of alternative herbicides.

No Brasil, varias espécies de gramíneas africanas introduzidas se tornaram importantes invasoras dos cerrados e constituem uma das principais ameaças para a sua biodiversidade. Atributos envolvendo trade-offs evolutivos que influenciam na adaptabilidade das espécies - aquisição, alocação e perda de recursos pelas plantas por meio de competição, facilitação, estresse e distúrbio - irão influenciar o desempenho dos indivíduos e a manutenção das populações nas comunidades vegetais. Deste modo, e de extrema importância entender quais os mecanismos que levam uma espécie invasora a ter um desempenho melhor sobre as espécies nativas. Comparações envolvendo características entre espécies exóticas e nativas podem levar a uma melhor compreensão sobre o processo da invasão. Diante disso, este trabalho procurou identificar algumas das estratégias competitivas adotadas por uma gramínea invasora, Urochloa decmbens, e por uma gramínea nativa, Echinolaena inflexa, em ecossistema de cerrado. Para verificar aspectos relativos às estratégias de colonização das espécies foram analisadas suas fenologias reprodutivas, as taxas de viabilidade e de germinação das sementes. Em um experimento com plantas envasadas, sementes de ambas as espécies foram semeadas em diferentes proporções relativas uma a outra. Três censos foram realizados durante o período de um ano para avaliar a sobrevivência, crescimento e fecundidade das espécies. Medidas da capacidade fotossintética e das taxas de assimilação foram tomadas para ambas as espécies tanto em parcelas puras quanto em parcelas mistas; também foi quantificada a área foliar especifica e total das espécies. No experimento com as plantas envasadas, quando sementes de ambas as espécies foram colocadas juntas, a nativa teve seu crescimento suprimido pela invasora. Nos tratamentos puros, os indivíduos da espécie nativa cresceram em tamanho, mas somente três se tornaram reprodutivos. No entanto, muitos indivíduos da espécie invasora cresceram em tamanho e se reproduziram em todos os tratamentos, mostrando uma alocação de recursos tanto para crescimento quanto para reprodução. A espécie invasora apresentou uma taxa fotossintética maior, porem ambas as espécies tiveram sua capacidade fotossintética e sua taxa fotossintética reduzida nos parcelas mistas, sugerindo que a competição afeta tanto a espécie nativa como a espécie invasora. Uma vez que a espécie nativa possui área foliar especifica menor em relação à espécie invasora, espera-se que ela apresente uma perda menor de nutrientes e, conseqüentemente, uma maior conservação/retenção dos nutrientes, o que favoreceria sua persistência em ambientes como o cerrado. No entanto, com a ocorrência de distúrbios e possíveis alterações na disponibilidade de nutrientes, a gramínea nativa pode ser deslocada pela invasora, que apresenta maior capacidade de colonizar novas áreas por meio de maiores taxas de germinação e estabelecimento; e maior produtividade, apresentando maiores taxas fotossintéticas, sendo mais eficiente no uso do nitrogênio. / Many African grasses have been introduced in Brazil and have invaded areas of Brazilian savannas, thus becoming a major threat to the biological diversity of this biome. Traits involving evolutionary trade-offs that influence the species adaptive strategies, such as differences in nutrient uptake, nutrient loss and biomass allocation in response to plant competition and facilitation, and environmental stress and disturbance, will influence individual performance and population maintenance in plant communities. Therefore, it is extremely important to identify the mechanisms associated with invasiveness that lead to a better performance of exotic species over co-occuring natives. Comparisons involving traits of both native and exotic species may lead to a better understanding concerning the success of invasions. This study sought to recognize possible competitive strategies adopted by an invasive grass, Urochloa decmbens, and a native grass, Echinolaena inflexa, in a Brazilian savanna. In order to identify aspects relative to their colonization strategies, the reproductive fenologies and seed viability and germination rates of both species were analyzed. An experiment with potted plants was performed where seeds of both species were sown in different relative proportions to each other. Three census were carried out during a period of 12 months to evaluate survival, growth and fecundity of both species. Gas exchange and chlorophyll fluorescence were quantified for each species in pure and mixed stands in the field; total and specific leaf area for both species were also measured. In the potted experiment, when seeds of both species were sown together, the growth of the native species was suppressed by the exotic species. In pure treatments, however, individuals of the native species grew in size, but only 2% reproduced. On the other hand, individuals of the exotic species grew in size and reproduced in all treatments, demonstrating that this species allocated enough resources for both growth and reproduction. The photosynthetic rates were greater for the invasive species, however, in mixed stands, both species had their photosynthetic rates and capacities decreased. The smaller specific leaf area of the native species suggests lower rates of nutrient loss than the invasive species, which would favor its persistence in nutrient poor environments, as the Brazilian savannas. Yet, with the occurrence of disturbances and variable nutrient availabilities, the native species might be displaced by the invasive species, since the later possesses traits leading to a higher competitive ability in such conditions.

This dissertation explores the differential effects of fires and rodent communities on native and invasive desert plant communities. Chapter one examines the impacts of fire and repeat fires on fuel loads in two different Utah desert sites, one in the cool Great Basin Desert and one in the hyper-arid Mojave Desert, over the course of four years. We found that both desert sites responded with varying intensities to a single burn, but the effects of a reburn were not as pronounced. We also found that our Great Basin Desert site had a stronger response to fire than our Mojave Desert site, producing a higher plant fuel loads that could potentially exacerbate the changing fire regimes. These data can be used to help map the effects that climate change may have on fuel loads and the fire potential of these deserts. Chapter two uses a full factorial experimental design to compare the effects of rodent herbivory and fire on native versus invasive seedlings at our Mojave Desert field site. We found that rodent herbivory has a more negative impact on the survival of native seedlings than invasive seedlings. This could be because the invasives are grasses that tend to tolerate herbivory better than the native shrubs and forbs. Chapter three again uses a full factorial experimental design to assess the impacts of rodents and fire on the fate of native and invasive seeds at our field site in the Mojave Desert. We found that rodents removed seeds, and they did show preferences for some species of seed over others, but these preferences were not different between native and invasive seeds or seed mass. We propose that the preferences may be influenced by other seed traits such as water content, handling time or physical and chemical defenses. As with the seedlings in chapter two, fire did not have any impact on rodent seed preferences. These data highlight the importance of considering rodent effects on seeds used in restoration effects following wildfires.

The ecological dynamics of co-flowering communities are largely mediated by pollinators. However, current understanding of pollinator-mediated interactions primarily relies on how co-flowering plants influence attraction of shared pollinators, and much less is known about plant–plant interactions that occur via heterospecific pollen (HP) transfer. Invaded communities in particular can be highly affected by the transfer of alien pollen, but the strength, drivers and fitness consequences of these interactions at a community scale are not well understood. Here we analyse HP transfer networks in nine coastal communities in the Yucatan Mexico that vary in the relative abundance of invasive flowers to evaluate how HP donation and receipt varies between native and alien plants. We further evaluate whether HP donation and receipt are mediated by floral traits (e.g. display, flower size) or pollinator visitation rate. Finally, we evaluated whether post-pollination success (proportion of pollen tubes produced) was affected by alien HP receipt and whether the effect varied between native and alien recipients. HP transfer networks exhibit relatively high connectance (c. 15%), suggesting high HP transfer within the studied communities. Significant network nestedness further suggests the existence of species that predominantly act as HP donors or recipients in the community. Species-level analyses showed that natives receive 80% more HP compared to alien species, and that alien plants donate 40% more HP than natives. HP receipt and donation were mediated by different floral traits and such effects were independent of plant origin (native or alien). The proportion of alien HP received significantly affected conspecific pollen tube success in natives, but not that of alien species. Synthesis. Our results suggest that HP transfer in invaded communities is widespread, and that native and alien species play different roles within HP transfer networks, which are mediated by a different suite of floral traits. Alien species, in particular, play a central role as HP donors and are more tolerant to HP receipt than natives—a finding that points to two overlooked mechanisms facilitating alien plant invasion and success within native co-flowering communities.

The ecological dynamics of co-flowering communities are largely mediated by pollinators. However, current understanding of pollinator-mediated interactions primarily relies on how co-flowering plants influence attraction of shared pollinators, and much less is known about plant–plant interactions that occur via heterospecific pollen (HP) transfer. Invaded communities in particular can be highly affected by the transfer of alien pollen, but the strength, drivers and fitness consequences of these interactions at a community scale are not well understood. Here we analyse HP transfer networks in nine coastal communities in the Yucatan Mexico that vary in the relative abundance of invasive flowers to evaluate how HP donation and receipt varies between native and alien plants. We further evaluate whether HP donation and receipt are mediated by floral traits (e.g. display, flower size) or pollinator visitation rate. Finally, we evaluated whether post-pollination success (proportion of pollen tubes produced) was affected by alien HP receipt and whether the effect varied between native and alien recipients. HP transfer networks exhibit relatively high connectance (c. 15%), suggesting high HP transfer within the studied communities. Significant network nestedness further suggests the existence of species that predominantly act as HP donors or recipients in the community. Species-level analyses showed that natives receive 80% more HP compared to alien species, and that alien plants donate 40% more HP than natives. HP receipt and donation were mediated by different floral traits and such effects were independent of plant origin (native or alien). The proportion of alien HP received significantly affected conspecific pollen tube success in natives, but not that of alien species. Synthesis. Our results suggest that HP transfer in invaded communities is widespread, and that native and alien species play different roles within HP transfer networks, which are mediated by a different suite of floral traits. Alien species, in particular, play a central role as HP donors and are more tolerant to HP receipt than natives—a finding that points to two overlooked mechanisms facilitating alien plant invasion and success within native co-flowering communities.

In this diploma thesis current state of vegetation cover was analyzed after spring flood in 2006 on the lower section of the Sázava river. Immediately after the flood there were 38 sites with alluvial deposits identified in the floodplain. The differences in species composition and species diversity between sites with alluvial deposits and sites without deposits were analyzed. The data were collected by using phytosociological relevés in pairs i.e. - plot with an alluvial deposit and plot without. There were 86 vascular plant species recorded out of which 10 species were non-native but not invasive species and 7 species were non-native invasive species. Results of this thesis revealed that species composition and diversity of invasive species differ between plots with alluvial deposits and plots without alluvial deposits. Moreover, plots without alluvial deposits host more invasive species. Keywords: vegetation succession, river floodplain, extreme floods, species diversity, plant invasion