Thesis (PhD (Botany and Zoology))--University of Stellenbosch, 2009. / ENGLISH ABSTRACT: Although the impacts of biological invasions are widely appreciated, a bias exists in research effort to post‐dispersal processes because of the difficulties of measuring propagule pressure and the detecting of newly established species. Here the Antarctic is used as a model system in which to quantify the initial dispersal of alien species and investigate the factors that contribute to the establishment and range dynamics of alien species once they have arrived in the region. Human movements are known to transport alien species into the Antarctic, some of which have successfully established and had wide ranging consequences in recipient ecosystems. Considering terrestrial flora, this research found that over 700 seeds from 99 taxa, including some species known to be invasive, are transported into the Antarctic each year in association with South African National Antarctic Programme (SANAP) passenger luggage and cargo. The first ever assessment of propagule drop‐off indicated that 30‐50% of these propagules will enter the recipient environment. Further results suggested that the construction of the British Antarctic Survey Halley VI station will facilitate the transport of over 5000 seeds from 34 taxa into the region, making this a significant pathway for introductions. Propagule pressure due to SANAP logistics is also considerable for marine species. Fouling assemblages on the external hull surfaces of the SANAP resupply vessel, the SA Agulhas, form only once the vessel’s anti‐fouling paint has been damaged by travel through sea ice and are characterised by low diversity. Ice scour prevents fouling assemblages from being transported to the Antarctic continent, but assemblages remain largely intact when travelling to sub‐Antarctic Islands. In the sea‐chests of the vessel populations of a known invasive, Mytilus galloprovincialis, were found with some individuals having survived transportation to the Antarctic region on multiple occasions. Once species have overcome initial dispersal barriers, they face further ecological and physiological challenges in order to establish in the recipient region. The parasitoid wasp Aphidius matricariae was first recorded on Marion Island in 2001. Surveys around the island show that adult abundance and the frequency of aphid parasitism are highest adjacent to a common anchor point of the SA Agulhas and decline away from this region. Genetic diversity was low, suggesting that the population was established from a single introduction. This highlights that high propagule pressure is not necessary for successful establishment of introduced invertebrates. Another species that has overcome the dispersal barrier is the terrestrial slug Deroceras panormitanum, which was introduced to Marion Island in the 1970’s and has since spread throughout much of the coastal habitat of the island. For this species range limits are set by intolerance of low temperature and salinity, and abundance structure is characterized by patches and gaps which are associated with this species inability to tolerate dry conditions. To prevent further alien introductions in the region, targeted management of high risk pathways is required. In addition, increased vigilance is needed to detect and manage newly established aliens before their ranges expand. / AFRIKAANSE OPSOMMING: Alhoewel die impak van biologiese indringings in die breë waardeer word, bestaan daar vooroordeel in navorsingspogings ten opsigte van na‐verspreidingsprosesse, weens die moeilikhede om verspreidingseenheid druk te meet en in die opsporing van nuut gevestigde spesies. Hier word die Antarktiese streek as ‘n model sisteem gebruik waarin die aanvanklike verspreiding van uitheemse spesies gekwantifiseer kan word en om die faktore wat bydrae tot die vestiging en grens dinamika van uitheemse spesies te ondersoek, wanneer hulle in ’n streek aangekom het. Menslike bewegings is bekend daarvoor om uitheemse spesies na die Antarktiek te vervoer, sommige waarvan suksesvol gevestig het en omvattende gevolge in die ontvanger ekosisteme gehad het. Aangaande terrestriële flora het hierdie navorsing gevind dat oor die 700 sade van 99 taxa, insluitende sommige spesies wat bekend is om indringend te wees, jaarliks na die Antarktiek vervoer word in assosiasie met die Suid Afrikaanse Nasionale Antarktiese Program (SANAP) se bagasie en vrag. Die eerste waardebepaling van verspreidingseenheid afgooi het aangedui dat 30‐50% van hierdie verspreidingseenhede die ontvanger omgewing sal binnedring. Verdere resultate het voorgestel dat die konstruksie van die Britse Antarktiese Opname Halley VI stasie die vervoer van 5000 sade van 34 taxa sal fasiliteer die streek in, wat dit ’n betekenisvolle weg maak vir indringings. Verspreidingseenheid druk is ook, as gevolg van die SANAP logistiek, aanmerklik vir mariene spesies. Vuilgoed versamelings op die eksterne omhullende oppervlaktes van die SANAP her‐voorsieningsvaartuig, die SA Agulhas, vorm wanneer die vaartuig se anti‐vuilgoed verf beskadig is met reis deur see‐ys en word gekenmerk deur lae diversiteit. Ys skuring voorkom dat vuilgoed versamelings vervoer word na die Antarktiese kontinent, maar versamelings bly grootliks vasgeheg wanneer na sub‐Antarktiese eilande gereis word. Populasies van ’n welbekende indringer, Mytilus galloprovincialis, is gevind in die see‐storingsarea van die vaartuig, met sommige individue wat die vervoering na die Antarktiese streek oorleef het op verskeie geleenthede. Wanneer spesies die aanvanklike verspreidingshindernisse oorkom het, staar hulle verdere ekologies en fisiologiese uitdagings in die gesig ten einde in die ontvanger streek te vestig. Die parasitiese wespe Aphidius matricariae is vir die eerste keer op Marion aangeteken in 2001. Opnames om die eiland toon dat volwasse oorvloed en die frekwensie van plantluis parasitisme die hoogste is aangrensend die algemene ankerpunt van die SA Agulhas en afneem weg van die omgewing af. Genetiese diversiteit was laag, wat voorstel dat die populasie gevestig het van ’n enkele bekendstelling. Dit lig uit dat hoë verspreidingseenheid druk nie nodig is vir die suksesvolle vestiging van bekendgestelde invertebrata nie. ‘n Ander spesie wat die verspreidingshindernisse oorkom het is die landslak Deroceras panormitanum wat in die 1970’s op Marion Eiland bekendgestel is en wat sedertdien versprei het oor meeste van die kushabitat van die eiland. Vir hierdie spesie word grens limiete vasgestel deur onverdraagsaamheid vir lae temperature en saliniteit en die oorvloedstruktuur word gekenmerk deur laslappe en gapings wat geassosieer word met die spesie se onvermoë om droë kondisies te hanteer. Om verdere uitheemse bekendstellings in die streek te voorkom, word geteikende bestuur van hoë risiko weë vereis. Toenemende waaksamheid is bykomend nodig vir die opsporing en bestuur van nuutgevestigde uitheemse spesies voordat hulle grense verbreed.

<p>Semi-natural grasslands, which are a declining and fragmented habitat in Europe, contain a high biodiversity, and are therefore of interest to conservation. This thesis examines how plant diversity is influenced by the landscape context, and how plant and fungal diversity can be targeted by practical conservation using indicator species and congruence between species groups. Reproduction and recruitment of the dioecious herb <i>Antennaria dioica </i>was also investigated, providing a case study on how fragmentation and habitat degradation may affect grassland plants.</p><p>Grassland size and heterogeneity were of greater importance for plant diversity in semi-natural grassland, than present or historical connectivity to other grasslands, or landscape characteristics. Larger grasslands were more heterogeneous than smaller grasslands, being the likely reason for the species-area relationship.</p><p>A detailed study on <i>A. dioica </i>discovered that sexual reproduction and recruitment may be hampered due to skewed sex-ratios. Sex-ratios were more skewed in small populations, suggesting that dioecious plants are likely to be particularly sensitive to reduced grassland size and fragmentation.</p><p>A study on indicators of plant species richness, used in a recent survey of remaining semi-natural grasslands in Sweden, revealed several problems. A high percentage of all indicator species were missed by the survey, removing an otherwise significant correlation between indicator species and plant species richness. Also, a null model showed that the chosen indicator species did not perform significantly better than species chosen at random from the available species pool, questioning the selection of the indicators in the survey. Diversity patterns of the threatened fungal genus <i>Hygrocybe</i> were not congruent with plant species richness or composition. Plants are thus a poor surrogate group for Hygrocybe fungi, and probably also for other grassland fungi. Implications from this thesis are that conservation of semi-natural grasslands should target several species groups, and that an appropriate scale for plant conservation may be local rather than regional.</p>

Introduced species present one of the greatest threats to biodiversity of native species, and knowledge of introduced species ecology is imperative for the development of management plans to ensure conservation of native species populations. We sought to determine the distribution and nesting behavior of an introduced population of red-bellied squirrels (Sciurus aureogaster) on islands of the Florida Keys currently managed as part of Biscayne National Park, and document potential for the species to impact native flora and fauna. Squirrels were difficult to observe in the dense vegetation of the subtropical forest, so we relied on their leaf nests, which were highly visible in the canopy of trees, to determine current presence and distribution on the Park's islands. We found nests throughout the mixed-hardwood forests of Elliott Key and Sands Key, and also documented a single, old nest on Old Rhodes Key, the first ever documentation of the species that far south in the Upper Keys. Nests were located in tall trees with more canopy linkages than random focal trees, and nests were placed in the upper canopy on the north side of the nest tree more often than expected by chance. Squirrels selected West Indies mahogany (Swietenia mahagoni) to place nests more often than available in the forest. Squirrels used areas with greater tree density and canopy cover, but lower recent hurricane damage and fewer woody shrub stems, than areas available at random in the forest. Squirrels built nests only in mixed-hardwood forest. Overall, this introduced species exhibited nest site selection behavior similar to other tree squirrels, and appears capable of continued spread despite the initial site of introduction on an oceanic island. Knowledge obtained from this research is being used by managers and applied to an eradication program to remove this invasive species from Biscayne National Park.

Humans are modifying the global landscape at an unprecedented scale and pace. As a result, species are declining and going extinct at an alarming rate. Here, I investigate two main aspects of species’ declines: what factors are contributing to their declines and how effective our conservation efforts have been. I assessed one of the main mechanisms for protecting species by looking at the Endangered Species Act (ESA) in the United States. I examined three separate indicators of species declines for different groups of species: range contractions in Canadian imperilled species, declines in abundance in global amphibian populations and increases in temporal variability in abundance in North American breeding birds. I found that change in recovery status of ESA listed species was only very weakly related to the number of years listed, number of years with a recovery plan, and funding. These tools combined explained very little of the variation in recovery status among species. Either these tools are not very effective in promoting species’ recovery, or species recovery data are so poor that it is impossible to tell whether the tools are effective or not. I examined patterns of species’ declines in three different groups in relation to a number of anthropogenic variables. I found high losses of Canadian imperiled bird, mammal, amphibian and reptile species in regions with high proportions of agricultural land cover. However, losses of imperiled species are significantly more strongly related to the proportion of the region treated with agricultural pesticides. This is consistent with the hypothesis that agricultural pesticide use, or something strongly collinear with it (perhaps intensive agriculture more generally), has contributed significantly to the decline of imperiled species in Canada. Global increases in UV radiation do not appear to be a major cause of amphibian population declines. At individual sites, temporal changes in amphibian abundance are not predictably related to changes in UV intensity. Variability in species’ abundance of North American breeding birds, after accounting for mean abundance, is not systematically higher in areas of high human-dominated land cover or climate change. Rather, it appears that areas with a high proportion of human-dominated cover come to have a higher proportion of highly abundant, and thus more variable, species.

Den här studien tar upp problematiken med fragmentering och dess påverkan på arters spridning och förekomst. Huvudsyftet med studien var att testa teorin om öbiogeografi genom att undersöka förekomsten av ett antal kärlväxter på olika åkerholmar med olika avstånd till närmaste spridningskälla i form av ett bryn i en närliggande skog. Vi har inventerat 40 stycken par av åkerholmar (öar) och skogsbryn (fastland) i ett jordbrukslandskap i Stockholms län. Inventeringsresultatet har analyserats med hjälp av univeriata linjära modeller-, och multiveriat ordination. Resultatet visar att teorin om öbiogeografi bara delvis kan förklara förekomstmönster för kärlväxter på åkerholmar. Vidare fann vi att spridningen av arter begränsas av åkerholmens storlek. Avstånd hade en effekt på artsammansättningen på åkerholmar. Vårt resultat visar på ett behov av att se till fragmentsstorlek samt grad av isolering som viktiga faktorer som styr möjligheten för att arter i framtiden ska kunna sprida sig till nya habitat. / This study address the problem of fragmentation, and its impact on species distribution and abundance. The main aim of the study was to test the theory of island biogeography using plant species prevalence from field islets at different distances from nearest “mainland”. We have studied 40 pairs of field islets (islands) and forest edges (mainland) in agricultural landscapes in the Stockholm County. The result of the inventory was analyzed using univariate linear models-, and multivariate ordination. Our result show that the theory of island biogeography partly can explain species prevalence patterns, but not all of them. Furthermore, we found that species distributions were limited by the size of the field islets. Distance from mainland had an effect on the species composition in the field islets. Our results indicate a need to include the size of habitat fragments, and the degree of isolation as important factors determining the ability for future species dispersal to newly established habitats.

Unionoid mussels are globally threatened and their conservation requires species-specific knowledge on their ecology and parasite-host interaction. Unio crassus is one of Europe’s most threatened unionoid species and has a temporary obligate parasitic life stage (glochidia) on fish. A lack of suitable hosts is probably a major limitation for mussel recruitment, but host species composition, suitability and availability in time and space have yet to be fully explored. This thesis examines different aspects of the host fish species, including their composition, suitability and ecological importance, in relation to U. crassus, using both field and laboratory studies. The effects of mussel and host density on mussel reproductive potential were considered, as were aspects of evolutionary adaptations between mussels and fish and how climate change may affect their interaction. The results show that U. crassus is a host generalist, parasitizing a variety of fish species. Host suitability and density, which varied among fish species and rivers, affected the level of glochidia encapsulation, hence mussel reproductive potential, more so than the density of mussels taking part in reproduction. Ecologically important hosts included both highly suitable primary hosts, and less suitable hosts that were highly abundant. Whether or not U. crassus has specific adaptations to its hosts to enhance juvenile transformation remains unclear. No distinct pattern of local adaptation was found, nor was there an effect of host fish presence on the timing of glochidia release by adult mussels. Instead, temperature played a major role, with results suggesting that changes in spring water temperature regimes can cause temporal and spatial mismatches in the mussel-host interaction. This thesis indicates that investigations of local mussel-host interactions help in identifying mechanisms important for unionoid conservation management and prioritization. / Många sötvattenmusslor har en komplex livscykel där larverna (glochidier) under sin utveckling till frilevande musslor parasiterar på gälarna hos lämpliga värdfiskar. Flera av våra musslor, såsom den tjockskaliga målarmusslan (Unio crassus), är globalt hotade och för att kunna bevara och förvalta dessa arter på bästa sätt behöver vi lära oss mer om deras ekologi och samspelet mellan musslan och dess värdfiskar. Avsaknaden av värdfiskar innebär förmodligen en stor begränsning för rekryteringen av juvenila musslor, men det finns trots detta en begränsad kunskap om hur artsammansättningen i fisksamhället och dess tillgänglighet påverkar musselpopulationer.  Min avhandling undersöker olika aspekter av interaktioner mellan U. crassus och dess värdfiskar, som hur värdfisksamhällen och fiskarters värdlämplighet påverkar musslans reproduktionspotential. Jag har även studerat hur tätheter av olika fiskarter och vuxna musslor påverkar rekryteringen, eventuella evolutionära anpassningar samt om en förhöjd temperatur skulle kunna påverka interaktionen mellan U. crassus och dess värdfiskar. Resultaten visar att U. crassus är en generalist som parasiterar på en mängd olika fiskarter. Jag fann dock en stor variation i dominerande fiskarter och lämpliga värdar mellan olika åar, vilket påverkade reproduktionspotentialen hos musslorna mer än vad tätheten vuxna musslor som deltog i reproduktionen gjorde. Som ekologiskt viktiga värdar fanns således både särskilt lämpliga, primära värdarter, men också mindre lämpliga arter som förekom i höga tätheter. Ingen tydlig lokal anpassning kunde observeras, och fiskens närvaro påverkade inte tidpunkten för när de vuxna musslorna släppte sina glochidielarver. Däremot fann jag att temperaturen spelade en viktig roll för musslans reproduktion, där ökad temperatur föreslås ha negativa effekter på interaktionen mellan musslan och dess värdfiskar. Avhandlingen visar på vikten av att studera interaktioner mellan den tjockskaliga målarmusslan och dess värdar på lokal skala för att bättre kunna identifiera och prioritera viktiga naturvårdsåtgärder. / UnioCrassusforLIFE (European LIFE+ project: LIFE10 NAT/SE/000046)

New farming needs and innovations have, over time, led to changes in land use. Arable fields have been turned into pastures and semi-natural pastures into arable fields. Due to the ecological value of continually grazed semi-natural pastures in Sweden the aim of this study is to find out how former arable fields that are permanently grazed differ from semi-natural pastures. In this study I selected appropriate semi-natural pastures from a national monitoring program on seminatural- pastures and meadows. Plots continuously grazed were compared with grazed plots on former arable fields. Pair-wise differences in the Ellenberg indicator values Light (L), Soil moisture (F), soil pH (R) and soil nitrogen (N), vertical coverage of trees, bushes and vegetation, species richness and species composition were tested. Except for species composition there were only small differences between former arable fields that are permanently grazed and semi-natural pastures. That means former arable fields can with time and grazing gain many of the values continuously grazed semi-natural pastures have.

A bacia do rio Piracicaba tem recebido atenção especial das autoridades municipais, pois possui grandes problemas de aproveitamento de seus recursos hídricos. A bacia do rio Corumbataí, um dos principais afluentes da margem direita do rio Piracicaba, é regionalmente importante, não só porque ainda possui águas de boa qualidade, mas também por apresentar elementos raros na paisagem do interior do estado. Esta tese teve o objetivo de caracterizar as assembléias de peixes na bacia do rio Corumbataí e fornecer ferramentas para avaliação de seu status ambiental. Foram escolhidos 4 rios principais com 3 pontos de coleta em cada um. Foram realizadas coletas no período de março a junho e setembro a dezembro de 2001, totalizando 24 coletas. Os dados bióticos foram avaliados através de medidas de diversidade. Para testar a hipótese de variação espaço-temporal da assembléia de peixes foi aplicado o modelo linear ANCOVA onde a variável resposta foi a riqueza de espécies; o fator foi ordem do rio; sendo utilizadas 2 covariáveis ambientais: temperatura e número de indivíduos, revelando uma variação espaço-temporal e padrões reconhecidos no meio acadêmico: relação espécie-área e conceito de rio contínuo. Técnicas multivariadas foram aplicadas para determinar a correlação entre a riqueza e a paisagem no entorno do ponto de coleta, revelando que existem mais espécies em locais com maior cobertura vegetal e mata ciliar preservada. Algumas espécies de peixes se mostraram bons indicadores ambientais: Hypostomus strigaticeps, Hoplosternum littorale e Salminus hilarii; por outro lado o Astyanax altiparanae não se mostrou um bom indicador / The Piracicaba river basin has received special attention from local authorities because it has presented a lot of problems about the misutilization of its hydric resources. The Corumbataí river basin, one of the major streams of the right side of the Piracicaba river, is regionally important, not only because it still has good water quality but also because it has unique elements inside the landscape of São Paulo state. This thesis aims to characterize the fish assemblage of the Corumbataí river basin and provide tools to assess the present environmental status of the basin. It has been chosen 4 major streams with 3 unit samples in each one of the streams. The samples were carried out from March to June and from September to December in the year of 2001, making up a total of 24 samples. The biotic data were assessed using diversity measurement. An ANCOVA was used to test the hypothesis of time-space variation of the fish assemblage, where the dependent variable was the species richness; the factor was the stream order and 2 environmental co-variates were used: temperature and number of individuals, thus revealing both time-space variation and some patterns which are recognized academically: species-area relationship and river continuum concept (RCC). Multivariate techniques were used to determine the correlation between richness and the landscape characteristics in the sampling unit, revealing that there is a greater number of species along the sites with a larger riparian zone. Some fish species have showed to be a good indicator: Hypostomus strigaticeps, Hoplosternum littorale and Salminus hilarii; on the other hand Astyanax altiparanae has not showed to be a good indicator species

Small mammals are a keystone guild in arid ecosystems; often exhibiting top-down control of the diversity and structure of plant communities. However, changing climate, shifting fire regimes, and the invasion of exotic plants are modifying the structure of arid systems. Environmental changes in these arid systems are likely altering small mammal communities, and therefore, their ecological role. We examined two aspects of the community composition of small mammals in the Great Basin: changes in community composition since large scale sampling of the region began in 1930, and the current population of a sensitive species of small mammal, the dark kangaroo mouse (Microdipodops megacephalus). In Chapter 1, we compared diversity and composition of present day small mammal communities to communities sampled between the years of 1930 and 1980. We sampled 234 historical locations across the eastern Great Basin region during the summers of 2014 and 2015. Our results indicated that diversity, richness, and evenness of small mammals in the Great Basin have declined significantly over the last century (P=0.002, P=0.03, P=0.002). The relative abundance of generalist species has increased, while specialist species have declined (P<0.001, P<0.001). Also, community composition at each site has changed significantly over the past century. Alterations in the community structure of small mammals may have cascading implications for the future of the Great Basin ecoregion. In Chapter 2, we conducted a region-wide survey for the dark kangaroo mouse in western Utah. Four teams sampled 232 locations across western Utah during the summers of 2014-2015. Of the 232 sites sampled, only 5 sites resulted in dark kangaroo mouse captures, totaling 15 individuals. These results could indicate a state-wide population decline for this species, both compared to historic population levels and to the populations surveyed less than ten years ago. The rapid decline may be a result of habitat degradation associated with invasive plant species and increasing fire frequency, the effects of which are exacerbated by the dark kangaroo mouse's life history as an ecological specialist. Unless large-scale habitat restoration and preservation is conducted for remaining populations, it is likely the dark kangaroo mouse will continue to decline within the state.

Invasive species introductions are associated with negative economic and environmental impacts, including reductions in native species populations. Successful invasive species populations often grow rapidly and a new food web equilibrium is established. Invasive, predatory northern pike (Esox lucius; hereafter pike) were detected in 2010 in Utah Lake, UT, a highly-degraded ecosystem home to the endemic, endangered June sucker (Chasmistes liorus). Here we test whether pike predation could hinder the restoration efforts of June sucker using the number of June sucker consumed by pike at various population densities as our metric. More specifically, we considered pike density at which the population could consume all June sucker stocked a critical threshold. Currently the number of naturally recruited June sucker is drastically lower than the number stocked. Thus, the metric we used to determine whether the pike population could hinder the June sucker restoration efforts is the number of pike that could consume the number of June sucker stocked. We combined pike growth and foraging observations with an energy-budget, bioenergetics consumption model to quantify lake-wide pike predation on June sucker. We also used an age-structured density dependent population model to estimate the pike population growth trajectory under various mitigation scenarios. Of 125 pike, we found an average pike consumes 0.8-1.0% June sucker and 40% sport fish. According to our bioenergetics model simulations, a population of adult pike at a very high density (60 pike per hectare) has the potential to consume nearly 6 million age-0 June sucker per year, which is likely more June sucker consumed than exist in the environment. In addition, our model suggests that an adult pike density greater than 1.5 pike per hectare has the potential to consume all June sucker stocked annually. Our age-structured population model suggests the pike population will reach equilibrium around 2026 at between 8 and 12 adult pike per hectare with the potential to consume between 0.8 and 1.2 million age-0 June sucker per year, respectively. The growing pike population could hamper restoration efforts and threaten endangered June sucker, a population with a mere 2,000 adults, in jeopardy of extinction. Our findings not only inform pike management efforts, but also highlight the importance of allocating resources toward habitat restoration to provide refuge for juvenile June sucker from predation, preventing the spread of aquatic invasive species, and the need for aquatic invasive species education.