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Biodiversity is determined by a myriad of complex processes acting at different scales. Given the current rates of biodiversity loss and change, it is of paramount importance that we improve our understanding of the underlying structure of ecological communities. In this thesis, I focused on Species Abundance Distributions (SAD), as a synthetic measure of biodiversity and community structure, and on Beta (β) diversity patterns, as a description of the spatial variation of species composition. I systematically assessed the effect of scale on both these patterns, analysing a broad range of community data, including different taxa and habitats, from the terrestrial, marine and freshwater realms. Knowledge of the scaling properties of abundance and compositional patterns must be fully integrated in biodiversity research if we are to understand biodiversity and the processes underpinning it, from local to global scales. SADs depict the relative abundance of the species present in a community. Although typically described by unimodal logseries or lognormal distributions, empirical SADs can also exhibit multiple modes. However, the existence of multiple modes in SADs has largely been overlooked, assumed to be due to sampling errors or a rare pattern. Thus, we do not know how prevalent multimodality is, nor do we have an understanding of the factors leading to this pattern. Here, I provided the first global empirical assessment of the prevalence of multimodality across a wide range of taxa, habitats and spatial extents. I employed an improved method combining two model selection tools, and (conservatively) estimated that ~15% of the communities were multimodal with strong support. Furthermore, I showed that the pattern is more common for communities at broader spatial scales and with greater taxonomic diversity (i.e. more phylogenetically diverse communities, since taxonomic diversity was measured as number of families). This suggests a link between multimodality and ecological heterogeneity, broadly defined to incorporate the spatial, environmental, taxonomic and functional variability of ecological systems. Empirical understanding of how spatial scale affects SAD shape is still lacking. Here, I established a gradient in spatial scale spanning several orders of magnitude by decomposing the total extent of several datasets into smaller subsets. I performed an exploratory analysis of how SAD shape is affected by area sampled, species richness, total abundance and taxonomic diversity. Clear shifts in SAD shape can provide information about relevant ecological and spatial mechanisms affecting community structure. There was a clear effect of area, species richness and taxonomic diversity in determining SAD shape, while total abundance did not exhibit any directional effect. The results supported the findings of the previous analysis, with a higher prevalence of multimodal SADs for larger areas and for more taxonomically diverse communities, while also suggesting that species spatial aggregation patterns can be linked to SAD shape. On the other hand, there was a systematic departure from the predictions of two important macroecological theories for SAD across scales, specifically regarding logseries distributions being selected only for smaller scales and when species richness and number of families were proportionally much smaller than the total extent. β diversity quantifies the variation in species composition between sites. Although a fundamental component of biodiversity, its spatial scaling properties are still poorly understood. Here, I tested if two conceptual types of β diversity showed systematic variation with scale, while also explicitly accounting for the two β diversity components, turnover and nestedness (species replacement vs species richness differences). I provided the first empirical analysis of β diversity scaling patterns for different taxa, revealing remarkably consistent scaling curves. Total β diversity and turnover exhibit a power law decay with log area, while nestedness is largely insensitive to scale changes. For the distance decay of similarity analysis, while area sampled affected the overall dissimilarity values, rates of similarity were consistent across large variations in sampled area. Finally, in both these analyses, turnover was the main contributor to compositional change. These results suggest that species are spatially aggregated across spatial scales (from local to regional scales), while also illustrating that substantial change in community structure might occur, despite species richness remaining relatively stable. This systematic and comprehensive analysis of SAD and community similarity patterns highlighted spatial scale, ecological heterogeneity and species spatial aggregation patterns as critical components underlying the results found. This work expanded the range of scales at which both theories deriving SAD and community similarity studies have been developed and tested (from local plots to continents). The results here showed strong departures from two important macroecological theories for SAD at different scales. In addition, the overall findings in this thesis clearly indicate that unified theories of biodiversity (or assuming a set of synthetic minimal assumptions) are unable to accommodate the variability in SADs shape across spatial scales reported here, and cannot fully reproduce community similarity patterns across scales. Incorporating more realistic assumptions, or imposing scale dependent assumptions, may prove to be a fruitful avenue for ecological research regarding the scaling properties of SAD and community similarity patterns. This will allow deriving new predictions and improving the ability of theoretical models to incorporate the variability in abundance and similarity patterns across scales.

Orientador: Itamar Alves Martins / Banca: Luis Cesar Schiesari / Banca: Lilian Casatti / Resumo: Girinos são um importante componente de ecossistemas de água doce e, ao longo dos últimos trinta anos, vêm sendo utilizados como modelos para testar hipóteses em ecologia de comunidades. O conjunto de informações disponível atualmente sugere que a presença de predadores influencia a abundância e o uso de hábitat por girinos e, conseqüentemente, a estrutura da comunidade. O objetivo deste estudo foi determinar o uso de recursos e investigar a influência de predadores na distribuição espacial de espécies e na estrutura de comunidades de girinos. As principais questões deste estudo foram: 1) qual o papel dos fatores abióticos na estruturação de comunidades de girinos?; 2) qual o grau de sobreposição de nicho entre as espécies nos três principais eixos de recursos: tempo (ocorrência sazonal), alimento e espaço (hábitat)?; 3) os girinos apresentam um padrão de distribuição não-aleatório em relação aos predadores? As amostragens tiveram freqüência mensal com coletas simultâneas de girinos e insetos predadores. Na caracterização dos corpos d'água foram utilizados dez descritores ambientais. Para determinar as guildas de espécies com relação ao uso de hábitat e ocorrência mensal foi implementada, respectivamente, uma análise de agrupamento e de ordenação por escalonamentro multidimensional não métrico. Para investigar a associação entre a abundância de girinos e os gradientes ambientais foi usada uma análise de correspondência canônica. A partilha de recursos entre girinos foi analisada utilizando-se uma análise de sobreposição de nicho em conjunto com um modelo nulo, considerando como eixos de recursos a ocorrência mensal, ocorrência nas poças e os itens alimentares. A análise da co-ocorrência pareada de girinos e predadores foi efetuada utilizando-se... (Resumo completo, clicar acesso eletrônico abaixo) / Abstract: Tadpoles are an important component of freshwater ecossystems and in the last thirty years has been used as model organisms to test hypothesis in community ecology. The data currently available suggest that the presence of predators influences the abundance and habitat use by tadpoles and, consequently the community structure. The aims of this study were to establish how tadpoles divide resources and to investigate the influence of predators on species distribution and on community structure of tadpoles. The main questions were: 1) What is the role of abiotic factors in the community structure?; 2) What is the extent of niche overlap among species, considering the three main resource axes: time (seasonal occurrence), diet items and space (macrohabitat)?; 3) Tadpoles exhibit a non-random distribution pattern in relation to predaceous insects? We conducted monthly samplings to collect tadpoles and predaceous insects. We also used 10 environmental descriptors to characterize the water bodies. To recognize species guilds in relation to habitat use and monthly occurrence, we employed respectively a cluster analysis and a n-MDS, respectively. To investigate the association between species abundance and environmental gradients we used a canonical correspondence analysis. The resource partitioning among tadpoles was analyzed using a null model, considering as recource axes monthly occurrence, pond occupancy and diet items. The pair-wise co-occurrence analysis between tadpoles and predaceous insects was carried out using a null model. Our main findings demonstrate that forest canopy cover and pond hydroperiod were the main factors influencing species abundance along water bodies sampled. There was a species sorting along these two environmental gradients, with species restricted to both ends of these continuums. There was a high overlap in seasonal occurence... (Complete abstract click electronic access below) / Mestre

This work applies advanced sampling (mark-release-recapture) and comparative approaches addressing the patterns in composition, structure and variability of larval trematode communities in three species of gastropod molluscs (Lymnaea stagnalis, Planorbarius corneus and Radix auricularia) at two nested scales of community organisation in typical Central European eutrophic environments. Hypothesis-testing with the application of null-model analyses, logistic regression modelling and multivariate randomisation techniques, revealed determinants of transmission rates, levels of infection and community structure in freshwater snail hosts in Central Europe and elucidated the mechanisms linking the spatial and temporal environmental variability with the action of complex community assembly rules in freshwater pulmonate snails.

Some climate models predict that, within the next 30-50 years, sea surface temperatures (SSTs) will frequently exceed the current thermal tolerance of corals (Fitt et al. 2001; Hughes et al. 2003; Hoegh-Guldberg et al. 2007). A potential consequence is that mass coral bleaching may take place (i) during warm El Niño-Southern Oscillation (ENSO) events which are predicted to occur in some regions more frequently than the current 3-7 year periodicity (Hoegh-Guldberg 1999; Sheppard 2003) or (ii) perhaps as often as annually or biannually if corals and their symbionts are unable to acclimate to the higher SSTs (Donner et al. 2005, 2007). Global data also indicate an upward trend toward increasing frequencies, intensities, and durations of tropical hurricanes and cyclones (Emanual 2005; Webster et al. 2005). As coral communities have been shown to require at least 10-30 years to recover after a major disturbance (e.g. Connell 1997; Ninio et al. 2000; Bruno & Selig 2007; Burt et al. 2008), it is possible that future coral communities may be in a constant state of recovery, with regeneration times exceeding the periods between disturbances. Life history traits (e.g. reproduction, recruitment, growth and mortality) vary among species of hard corals; thus, gradients in community structures may have a strong influence on susceptibilities to disturbance and rates of recovery (Connell 1997; Ninio & Meekan 2002). Taxa which are more susceptible to bleaching and mechanical disturbance (e.g. tabular and branching acroporids and pocilloporids) may experience continual changes in population structure due to persistent cycles of regeneration or local extirpation, while the more resistant taxa (e.g. massive poritids and faviids) may display relatively stable population structures (Woodley et al. 1981; Hughes & Connell 1999; Baird & Hughes 2000; Marshall & Baird 2000; Loya et al. 2001; McClanahan & Maina 2003). Determining whether resistant coral taxa have predictable responses to disturbances, with consistent patterns over wide spatial scales, may improve predictions for the future affects of climate change and the composition of reefs (Done 1999; Hoegh-Guldberg 1999; McClanahan et al. 2004). The work presented in this dissertation describes the spatial and temporal patterns in community structures for high latitude coral assemblages that have experienced the types of natural disturbances which are predicted to occur in tropical reef systems with increasing frequency as a result of climate change. The primary area of focus is the southeastern Arabian Gulf, where the coral communities are exposed to natural conditions that exceed threshold limits of corals elsewhere in the world, with annual temperature ranges between 14-36°C (Kinzie 1973; Shinn 1976) and salinities above 40 ppt. Two additional regions are included in this study for comparisons of high latitude coral community structures. The northwestern Gulf of Oman is adjacent to the southeastern Arabian Gulf (i.e. the two bodies of water are connected by the Strait of Hormuz); however, the environmental conditions are milder in the Gulf of Oman such that the number of coral taxa therein is threefold that found in the southeastern Arabian Gulf (i.e. 107 coral species in the Gulf of Oman compared to 34 species in this region of the Arabian Gulf (Riegl 1999; Coles 2003; Rezai et al. 2004)). Broward County, Florida is geographically remote from the Gulfs and, therefore, serves as a benchmark for testing whether consistent patterns in community structures exist despite different climatic and anthropogenic influences. The coral communities within the southeastern Arabian Gulf, the northwestern Gulf of Oman, and Broward County, Florida have been exposed to recurrent elevated sea surface temperature (SST) anomalies, sequential cyclone and red tide disturbances, and frequent hurricanes and tropical storms, respectively. These disturbances and other impacts (e.g. bleaching episodes, disease outbreaks, anthropogenic stresses) have affected the more susceptible acroporids and pocilloporids, resulting in significant losses of coral cover by these families and shifts towards massive corals as the dominant taxa. During the post-disturbance scarcity or absence of branching and tabular corals, the resistant massive taxa have become the crux of the essential hard coral habitat for fish, invertebrates and other marine organisms. Because recovery to pre-disturbance community structures may take decades or may not occur at all, it is vital that scientists and resource managers have a better understanding of the spatial and temporal ecology patterns of the corals that survive and fill in the functional gaps that are created by such disturbances. To aid in this understanding, this dissertation presents spatial and temporal patterns for the coral assemblages which have developed after the respective disturbances. Spatial ecology patterns are analyzed using graphical descriptions (e.g. taxa inventories, area cover, densities, size frequency distributions), univariate techniques (e.g. diversity indices), distributional techniques (e.g. k-dominance curves) and multivariate techniques (e.g. hierarchical clustering, multidimensional scaling). Temporal comparisons at monitoring sites within the southeastern Arabian Gulf and northwestern Gulf of Oman describe the coral population dynamics and are used to create size class transition models that project future population structures of massive corals in the recovering habitats.

Abstract Freshwater ecosystems are severely threatened by a variety of anthropogenic stressors. In order to keep track with at least part of the changes, it is important to efficiently assess and monitor freshwater biological diversity. Biological assessment programs are developed to detect human-induced changes in the ecological state of aquatic systems. These programs typically rely on the assumption that environmental conditions are the sole drivers of biological communities occupying a site and, thus, these local communities would correctly inform about environmental conditions. Recently, this background principle of current bioassessment methods has faced some criticism, stemming from the idea that community structuring is a more complex process than just a mere result of local environmental conditions. In this thesis, I studied the natural and anthropogenic drivers of freshwater biodiversity. I was particularly interested if the various biodiversity metrics studied showed any spatial patterns and if so, for which reasons these patterns might occur. To obtain a comprehensive picture of spatial patterns in biodiversity, I studied multiple spatial contexts, biological groups and indices. I found that environmental conditions were not the only drivers of freshwater biodiversity. Instead, different spatial patterns, likely stemming from dispersal processes, were surprisingly powerful drivers of aquatic communities and index values derived from them. The spatial context (i.e. spatial extent and connectivity) of the aquatic study systems likely played a major role in structuring biodiversity. I also found that the distinct biological groups and indices studied were partly related to different predictor variables. The findings of this thesis are of importance to the development of new bioassessment methods. The results of this thesis also suggest that the spatial context of the study setting should be acknowledged when interpreting results based on current bioassessment methods. / Tiivistelmä Makeanveden ekosysteemit ovat hyvin alttiita ihmistoiminnalle. Ekosysteemissä mahdollisesti tapahtuvien muutosten havaitseminen vaatii tehokkaita vesistöjen ekologisen tilan ja luonnon monimuotoisuuden arviointi- ja seurantamenetelmiä. Näiden menetelmien toimintaperiaatteen yleisenä tausta-ajatuksena on, että biologiset yhteisöt määräytyvät paikallisten ympäristöolojen mukaan. Tietyn paikan yhteisön oletetaan siis heijastavan kyseisen paikan ympäristön tilaa. Viime aikoina tausta-ajatus paikallisten ympäristöolojen merkityksestä ainoana eliöyhteisöjä muovaavana tekijänä on kuitenkin kohdannut kritiikkiä. Kriitikot painottavat, että biologisten yhteisöjen rakenteeseen vaikuttavat monet muutkin asiat kuin paikalliset ympäristöolosuhteet ja niissä tapahtuvat ihmisperäiset muutokset. Väitöskirjassani tutkin sisävesien luonnon monimuotoisuuteen vaikuttavia tekijöitä. Olin erityisen kiinnostunut siitä, näkyykö tutkituissa biologisissa parametreissa maantieteellisessä tilassa ilmeneviä spatiaalisia säännönmukaisuuksia. Saadakseni mahdollisimman laaja-alaisen käsityksen luonnon monimuotoisuudessa esiintyvistä spatiaalisista säännönmukaisuuksista, tutkin useaa spatiaalista kontekstia, eliöryhmää ja indeksiä. Tutkimuksessa selvisi, että paikalliset ympäristöolosuhteet eivät ole ainoita luonnon monimuotoisuuteen vaikuttavia tekijöitä. Erilaiset spatiaaliset säännönmukaisuudet, todennäköisesti eliöiden levittäytymiseen liittyvien seikkojen aiheuttamina, olivat yllättävän yleisiä makeiden vesien eliöyhteisöjen rakenteessa ja niihin perustuvien indeksien arvoissa. Tutkimussysteemien spatiaalinen konteksti (alueen laajuus ja paikkojen väliset spatiaaliset suhteet) selvästi vaikutti luonnon monimuotoisuutta kuvastavien indeksien arvojen vaihteluun. Lisäksi selvisi, että eri eliöryhmät ja indeksit olivat useimmiten liitoksissa hyvin erilaisiin selittäviin muuttujiin, osoittaen, että nämä mittarit kuvastavat eri asioita. Väitöskirjassa esitetyt havainnot on tärkeää huomioida vesistöjen ekologisen tilan ja luonnon monimuotoisuuden arviointi- ja seurantamenetelmiä kehitettäessä. Spatiaalisen kontekstin merkitys olisi hyvä huomioida myös nykyisten arviointi- ja seurantamenetelmien tuottamien tulosten tulkinnassa.

Climate change is expected to increase the precipitation ~30% in higher latitudes during the next century, increasing the land runoff via rivers to aquatic ecosystems. The Baltic Sea will receive higher river discharges, accompanied by larger input of allochthonous dissolved organic matter (DOM) from terrestrial ecosystems. The salinity will decrease due to freshwater dilution. The allochthonous DOM constitute a potential growth substrate for microscopic bacterioplankton and phytoplankton, which together make up the basal trophic level in the sea. The aim of my thesis is to elucidate the bacterial processing of allochthonous DOM and to evaluate possible consequences of increased runoff on the basal level of the food web in the Baltic Sea. I performed field studies, microcosm experiments and a theoretical modeling study. Results from the field studies showed that allochthonous DOM input via river load promotes the heterotrophic bacterial production and influences the bacterial community composition in the northern Baltic Sea. In a northerly estuary ~60% of bacterial production was estimated to be sustained by terrestrial sources, and allochthonous DOM was a strong structuring factor for the bacterial community composition. Network analysis showed that during spring the diversity and the interactions between the bacteria were relatively low, while later during summer other environmental factors regulate the community, allowing a higher diversity and more interactions between different bacterial groups. The influence of the river inflow on the bacterial community allowed “generalists” bacteria to be more abundant than “specialists” bacteria.    Results from a transplantation experiment, where bacteria were transplanted from the northern Baltic Sea to the seawater from the southern Baltic Sea and vice versa, showed that salinity, as well as the DOM composition affect the bacterial community composition and their enzymatic activity. The results showed that α-proteobacteria in general were favoured by high salinity, β-proteobacteria by low salinity and terrestrial DOM compounds and γ-proteobacteria by the enclosure itself. However, effects on the community composition and enzymatic activity were not consistent when the bacterial community was retransplanted, indicating a functional redundancy of the bacterial communities.  Results of ecosystem modeling showed that climate change is likely to have quite different effect on the north and the south of the Baltic Sea. In the south, higher temperature and internal nutrient load will increase the cyanobacterial blooms and expand the anoxic or suboxic areas. In the north, climate induced increase in riverine inputs of allochthonous DOM is likely to promote bacterioplankton production, while phytoplankton primary production will be hampered due to increased light attenuation in the water. This, in turn, can decrease the production at higher trophic levels, since bacteria-based food webs in general are less efficient than food webs based on phytoplankton. However, complex environmental influences on the bacterial community structure and the large redundancy of metabolic functions limit the possibility of predicting how the bacterial community composition will change under climate change disturbances.