Molluscan radiations and landscape evolution in Miocene Amazonia

This PhD study aims to exploit the rich archive provided by the Miocene mollusc fauna of the Pebas Formation and other inland Miocene Amazonian formations to reconstruct landscape evolution and biotic development in lowland Amazonia during the Neogene. Over 160 samples from more than 70 Pebas Formation outcrops mostly collected by the author were processed for this study. Additional samples were collected in Andean areas of Colombia and Venezuela and further material from other northwestern South American basins was studied in museums. Pebas Formation samples and well log data made available by Occidental Peru from three wells in the Marañon Basin in Peru were also investigated. During this study four genera and 74 species from the Pebas Formation have been described and a further 13 species have been introduced in open nomenclature, and several species were reported for the first time. The number of mollusc species attributed to the Pebas fauna has increased from around 50 to 156. The Pebas fauna is characterised as aquatic, endemic and extinct, and is a typical representative of a long-lived lake fauna. Fluvial taxa are not common, (marginal) marine taxa are rare. An additional molluscan fauna from the Miocene Solimões Formation of Brazil, containing 13 fresh water species was also described. The newly documented fauna was used to improve biostratigraphic framework of Miocene Amazonian deposits. Twelve mollusc zones were introduced, the upper eleven of which cover a time interval of approximately seven million years covered previously by only three pollen zones. An age model calculated for the borehole data indicates that the Pebas Formation was deposited between c. 24 and 11 Ma. The areal distribution of the outcropping mollusc zones uncovered a broad dome structure, termed here the Iquitos-Araracuara anteclise in the study area. The structure appears to have influenced river courses and also contributed to edaphic heterogeneity that may have been in part responsible for the current high biodiversity in the study area. The Pebas system was a huge system (> one million km2) dominated by relatively shallow lakes, but also containing swamps and rivers. The system was fed by rivers draining the emergent Andes in the west and lowlands and cratons to the east. The Pebas system was located at sea level and was open to marine settings through a northern portal running through the Llanos Basin and East Venezuela Basin towards the Caribbean. Cyclical baselevel changes possibly related to Mylankhovitch cycles, have been documented in depositional sequences of the Pebas Formation. The composition of the Pebasian mollusc fauna implies that the system was mostly a fresh water system. Such an interpretation is matched by strontium isotope ratios as well as very negative δ18O ratios found in the shells, but is at odds with oligohaline and mesohaline ichnofacies found in the same strata. The mollusc fauna of the Pebas Formation diversified through most of the existence of the lake system. The diversification was mostly the result of in-situ cladogenesis. The success of some of the Pebasian endemic clades is explained by adaptation to fresh water, low oxygen, common unconsolidated lake bottoms (soup grounds) as well as high predation intensity. Maximum diversity was reached at the base of the late Middle to early Late Miocene Grimsdalea pollen zone, some 13 Ma. At the time some 85 species co-occurred, 67 of which are considered as Pebasian endemics. A subsequent drop in species richness coincides with indications of elevated salinities, although a causal relation still needs to be established. Apparently the Pebas fauna went (almost) entirely extinct with the replacement of the lake system into a fluvio-tidal system during the Early Late Miocene, some 11 Ma.

Past, present and future - assessing changes in aquatic environments using subfossil diatoms

The deterioration of surface waters is one of the most important issues in the environmental management of the European Union. Thus, the EU Water Framework Directive 2000/60/EC (WFD) requires “good ecological and chemical status” of surface waters by 2015 allowing only a slight departure from ecological reference conditions characterized by the biological communities typical for the conditions of minimal anthropogenic impact. The WFD requires the determination of ecological reference conditions and the present ecological status of surface waters. To meet this legislative demand, sedimentary diatom assemblages were used in these studies with various methods 1) to assess natural and human activity induced environmental changes, 2) to characterize background conditions 3) to evaluate the present ecological status and 4) to predict the future of the water bodies in the light of palaeolimnological data. As the WFD refers to all surface waters, both coastal and inland sites were included. Two long and two short sediment cores from the Archipelago Sea in the northern Baltic Sea were examined for their siliceous microfossils in order to assess (1) the Holocene palaeoenvironmental history and (2) the recent eutrophication of the area. The diatom record was divided into local diatom assemblage zones (LDAZ, long cores) and diatom assemblage zones (DAZ, short cores). Locally weighted weighted averaging regression and calibration (LWWA) was applied for the quantitative reconstruction of past TN concentrations (short cores). An age model for the long cores was constructed by using independent palaeomagnetic and AMS-14C methods. The short cores were dated using radiometric (210Pb, 226Ra and 137Cs) methods. The long cores date back to the early history of the Archipelago Sea, which was freshwater – no salinity increase referable to the brackish phase of the Yoldia Sea is recognized. The nutrient status of the lacustrine phase was slightly higher in the Archipelago Sea than in the Baltic Proper. Initial brackish-water influence is observed at 8 150 ±80 cal. BP (LDAZ4), but fully brackish conditions were established at 7 700 ±80 cal. BP (LDAZ5). The diatom assemblages indicate increasing salinity, warming climate and possible eutrophic conditions during the lacustrine to brackish-water transition. The decreasing abundance of Pseudosolenia calcar-avis (Schultze) Sundström and the increasing abundance of the ice-cover indicator species Pauliella taeniata (Grunow) Round and Basson indicate decreasing salinity and climatic cooling after ca. 5 000 cal. BP. Signs of eutrophication are visible in the most recent diatom assemblage zones of both short cores. Diatom-inferred total nitrogen (DI-TN) reconstructions partially fail to trace the actual measured total nitrogen concentrations especially from the late 1980s to the mid 1990s. This is most likely due to the dominating diatom species Pauliella taeniata, Thalassiosira levanderi Van Goor and Fragilariopsis cylindrus (Grunow) W. Krieger being more influenced by factors such as the length of the ice-season rather than nutrient concentrations. It is concluded that the diatom assemblages of the study sites are principally governed by climate fluctuations, with a slight influence of eutrophication visible in the most recent sediments. There are indications that global warming, with reduced ice cover, could impact the spring blooming diatom species composition in the Archipelago Sea. In addition, increased sediment accumulation in the early 90s coincides with the short ice-seasons suggesting that warming climate with decreasing ice-cover may increase sedimentation in the study area. The diverse diatom assemblages dominated by benthic species (54 %) in DAZ1 in the Käldö Fjärd core can be taken as background diatom assemblages for the Archipelago Sea. Since then turbidity has increased and the diatom assemblages have been dominated by planktonic diatoms from around the mid 1800s onwards. The reconstructed reference conditions for the total nitrogen concentrations fluctuate around 400 μg l-1. Altogether two short sediment cores and eight short cores for top-bottom analysis were retrieved from Lake Orijärvi and Lake Määrjärvi to assess the impact of the acid mine drainage (AMD) derived metals from the Orijärvi mine tailings on the diatom communities of the lakes. The Cu (Pb, Zn) mine of Orijärvi (1757 – 1956) was the first one in Finland where flotation techniques (1911 – 1955) were used to enrich ore and large quantities of tailings were produced. The AMD derived metal impact to the lakes was found to be among the heaviest thus far recorded in Finland. Concentrations of Cu, Pb and Zn in Lake Orijärvi sediments are two to three orders of magnitude higher than background values. The metal inputs have affected Lake Orijärvi and Lake Määrjärvi diatom communities at the community levels through shifts in dominant taxa (both lakes) and at the individual level through alteration in frustule morphology (Lake Orijärvi). At present, lake water still has elevated heavy metal levels, indicating that the impact from the tailings area continues to affect both lakes. Lake Orijärvi diatom assemblages are completely dominated by benthic species and are lacking planktonic diatoms. In Lake Määrjärvi the proportion of benthic and tychoplanktonic diatoms has increased and the planktonic taxa have decreased in abundance. Achnanthidium minutissimum Kützing and Brachysira vitrea (Grun.) R. Ross in Hartley were the most tolerant species to increased metal concentrations. Planktonic diatoms are more sensitive to metal contamination than benthic taxa, especially species in the genus Cyclotella (Kützing) Brébisson. The ecological reference conditions assessed in this study for Lake Orijärvi and Lake Määrjärvi comprise diverse planktonic and benthic communitites typical of circumneutral oligotrophic lakes, where the planktonic diatoms belonging to genera Cyclotella , Aulacoseira Thwaites, Tabellaria Ehrenberg and Asterionella Hassall dominate in relative abundances up to ca. 70%. The benthic communities are more diverse than the planktonic consisting of diatoms belonging to the genera Achnanthes Bory, Fragilaria Lyngbye and Navicula St. Vincent. This study clearly demonstrates that palaeolimnological methods, especially diatom analysis, provide a powerful tool for the EU Water Frame Work Directive for defining reference conditions, natural variability and current status of surface waters. The top/bottom approach is a very useful tool in larger-scale studies needed for management purposes. This “before and after” type of sediment sampling method can provide a very time and cost effective assessment of ecological reference conditions of surface waters.

Climate driven changes in temperature, pH and food quality : effects on copepod reproduction

Increased emissions of greenhouse gases into the atmosphere are causing an anthropogenic climate change. The resulting global warming challenges the ability of organisms to adapt to the new temperature conditions. However, warming is not the only major threat. In marine environments, dissolution of carbon dioxide from the atmosphere causes a decrease in surface water pH, the so called ocean acidification. The temperature and acidification effects can interact, and create even larger problems for the marine flora and fauna than either of the effects would cause alone. I have used Baltic calanoid copepods (crustacean zooplankton) as my research object and studied their growth and stress responses using climate predictions projected for the next century. I have studied both direct temperature and pH effects on copepods, and indirect effects via their food: the changing phytoplankton spring bloom composition and toxic cyanobacterium. The main aims of my thesis were: 1) to find out how warming and acidification combined with a toxic cyanobacterium affect copepod reproductive success (egg production, egg viability, egg hatching success, offspring development) and oxidative balance (antioxidant capacity, oxidative damage), and 2) to reveal the possible food quality effects of spring phytoplankton bloom composition dominated by diatoms or dinoflagellates on reproducing copepods (egg production, egg hatching, RNA:DNA ratio). The two copepod genera used, Acartia sp. and Eurytemora affinis are the dominating mesozooplankton taxa (0.2 – 2 mm) in my study area the Gulf of Finland. The 20°C temperature seems to be within the tolerance limits of Acartia spp., because copepods can adapt to the temperature phenotypically by adjusting their body size. Copepods are also able to tolerate a pH decrease of 0.4 from present values, but the combination of warm water and decreased pH causes problems for them. In my studies, the copepod oxidative balance was negatively influenced by the interaction of these two environmental factors, and egg and nauplii production were lower at 20°C and lower pH, than at 20°C and ambient pH. However, presence of toxic cyanobacterium Nodularia spumigena improved the copepod oxidative balance and helped to resist the environmental stress, in question. In addition, adaptive maternal effects seem to be an important adaptation mechanism in a changing environment, but it depends on the condition of the female copepod and her diet how much she can invest in her offspring. I did not find systematic food quality difference between diatoms and dinoflagellates. There are both good and bad diatom and dinoflagellate species. Instead, the dominating species in the phytoplankton bloom composition has a central role in determining the food quality, although copepods aim at obtaining as a balanced diet as possible by foraging on several species. If the dominating species is of poor quality it can cause stress when ingested, or lead to non-optimal foraging if rejected. My thesis demonstrates that climate change induced water temperature and pH changes can cause problems to Baltic Sea copepod communities. However, their resilience depends substantially on their diet, and therefore the response of phytoplankton to the environmental changes. As copepods are an important link in pelagic food webs, their future success can have far reaching consequences, for example on fish stocks.

Dispersal and metacommunity dynamics in a soft-sediment benthic system : how well is the seafloor connected?

Connectivity depends on rates of dispersal between communities. For marine soft-sediment communities continued small-scale dispersal as post-larvae and as adults can be equally important in maintaining community composition, as initial recruitment of substrate by pelagic larvae. In this thesis post-larval dispersal strategies of benthic invertebrates, as well as mechanisms by which communities are connected were investigated. Such knowledge on dispersal is scarce, due to the difficulties in actually measuring dispersal directly in nature, and dispersal has not previously been quantified in the Baltic Sea. Different trap-types were used underwater to capture dispersing invertebrates at different sites, while in parallel measuring waves and currents. Local community composition was found to change predictably under varying rates of dispersal and physical connectivity (waves and currents). This response was, however, dependent on dispersal-related traits of taxa. Actively dispersing taxa will be relatively better at maintaining their position, as they are not as dependent on hydrodynamic conditions for dispersal and will be less prone to be passively transported by currents. Taxa also dispersed in relative proportions that were distinctly different from resident community composition and a significant proportion (40 %) of taxa were found to lack a planktonic larval life-stage. Community assembly was re-started in a large-scale manipulative field experiment over one year across several sites, which revealed how patterns of community composition (α-, β- and λ-diversity) change depending on rates of dispersal. Results also demonstrated that in response to small-scale disturbance, initial recruitment was by nearby-dominant species after which other species arrived from successively further away. At later assembly time, the number of coexisting species increased beyond what was expected purely by local niche requirements (species sorting), transferring regional differences in community composition (β-diversity) to the local scale (α-diversity, mass effect). Findings of this thesis complement more theoretical studies in metacommunity ecology by demonstrating that understanding how and when individuals disperse relative to underlying environmental heterogeneity is key to interpreting how patterns of diversity change across different spatial scales. Such information from nature is critical when predicting responses to, for example, different types of disturbances or management actions in conservation.

Taxonomy and Species Richness of Amazonian Pimplinae and Rhyssinae (hymenoptera: ichneumonidae)

The Amazonian region, the biggest rain forest of our planet, is known for its extraordinary biodiversity. Most of this diversity is still unexplored and new species of different taxa are regularly found there. In this region, as in most areas of the world, insects are some of the most abundant organisms. Therefore, studying this group is important to promote the conservation of these highly biodiverse ecosystems of the planet. Among insects, parasitoid wasps are especially interesting because they have potential for use as biodiversity indicators and biological control agents in agriculture and forestry. The parasitoid wasp family Ichneumonidae is one of the most species rich groups among the kingdom Animalia. This group is still poorly known in many areas of the world; the Amazonian region is a clear example of this situation. Ichneumonids have been thought to be species poor in Amazonia and other tropical areas. However, recent studies are suggesting that parasitoid wasps may be quite abundant in Amazonia and possibly in most tropical areas of the world. The aim of my doctoral thesis is to study the species richness and taxonomy of two of the best known ichneumonid subfamilies in the Neotropical region, Pimplinae and Rhyssinae. To do this I conducted two extensive sampling programs in the Peruvian Amazonia. I examined also a large number of Neotropical ichneumonids deposited to different natural history museums. According to the results of my thesis, the species richness of these parasitoids in the Amazonian region is considerably higher than previously reported. In my research, I firstly further develop the taxonomy of these parasitoids by describing many new species and reporting several new faunistic records (I, II, III). In this first part I focus on two genera (Xanthopimpla and Epirhyssa) which were thought to be rather species poor. My thesis demonstrates that these groups are actually rather species rich in the Amazonian region. Secondly, I concentrate on the species richness of these parasitoids in a global comparison showing that the Neotropical region and especially the Peruvian Amazonia is one of the most species rich areas of Pimpliformes ichneumonids (V). Furthermore, I demonstrate that with the data available to date no clear latitudinal gradient in species richness is visible. Thirdly, increasing the macroecological knowledge of these parasitoids I show that some previously unreported ichneumonid subfamilies are present in the Amazonian region (IV). These new insights and the results of the global comparison of ichneumonid inventories suggest that the previous belief of low diversity in the tropics is most likely related to a lack of sampling effort in the region. Overall, my research increases the knowledge of Neotropical ichneumonids highlighting the importance of Peruvian Amazonia as one of the diversity hotspots of parasitoid wasps.