Evolutionary Ecology of Mountain Birch in Subarctic Stress Gradients: Interplay of Biotic and Abiotic Factors in Plant-Plant Interactions and Evolutionary Processes

In nature, variation for example in herbivory, wind exposure, moisture and pollution impact often creates variation in physiological stress and plant productivity. This variation is seldom clear-cut, but rather results in clines of decreasing growth and productivity towards the high-stress end. These clines of unidirectionally changing stress are generally known as ‘stress gradients’. Through its effect on plant performance, stress has the capacity to fundamentally alter the ecological relationships between individuals, and through variation in survival and reproduction it also causes evolutionary change, i.e. local adaptations to stress and eventually speciation. In certain conditions local adaptations to environmental stress have been documented in a matter of just a few generations. In plant-plant interactions, intensities of both negative interactions (competition) and positive ones (facilitation) are expected to vary along stress gradients. The stress-gradient hypothesis (SGH) suggests that net facilitation will be strongest in conditions of high biotic and abiotic stress, while a more recent ‘humpback’ model predicts strongest net facilitation at intermediate levels of stress. Plant interactions on stress gradients, however, are affected by a multitude of confounding factors, making studies of facilitation-related theories challenging. Among these factors are plant ontogeny, spatial scale, and local adaptation to stress. The last of these has very rarely been included in facilitation studies, despite the potential co-occurrence of local adaptations and changes in net facilitation in stress gradients. Current theory would predict both competitive effects and facilitative responses to be weakest in populations locally adapted to withstand high abiotic stress. This thesis is based on six experiments, conducted both in greenhouses and in the field in Russia, Norway and Finland, with mountain birch (Betula pubescens subsp. czerepanovii) as the model species. The aims were to study potential local adaptations in multiple stress gradients (both natural and anthropogenic), changes in plant-plant interactions under conditions of varying stress (as predicted by SGH), potential mechanisms behind intraspecific facilitation, and factors confounding plant-plant facilitation, such as spatiotemporal, ontogenetic, and genetic differences. I found rapid evolutionary adaptations (occurring within a time-span of 60 to 70 years) towards heavy-metal resistance around two copper-nickel smelters, a phenomenon that has resulted in a trade-off of decreased performance in pristine conditions. Heavy-metal-adapted individuals had lowered nickel uptake, indicating a possible mechanism behind the detected resistance. Seedlings adapted to heavy-metal toxicity were not co-resistant to others forms of abiotic stress, but showed co-resistance to biotic stress by being consumed to a lesser extent by insect herbivores. Conversely, populations from conditions of high natural stress (wind, drought etc.) showed no local adaptations, despite much longer evolutionary time scales. Due to decreasing emissions, I was unable to test SGH in the pollution gradients. In natural stress gradients, however, plant performance was in accordance with SGH, with the strongest host-seedling facilitation found at the high-stress sites in two different stress gradients. Factors confounding this pattern included (1) plant size / ontogenetic status, with seedling-seedling interactions being competition dominated and host-seedling interactions potentially switching towards competition with seedling growth, and (2) spatial distance, with competition dominating at very short planting distances, and facilitation being strongest at a distance of circa ¼ benefactor height. I found no evidence for changes in facilitation with respect to the evolutionary histories of plant populations. Despite the support for SGH, it may be that the ‘humpback’ model is more relevant when the main stressor is resource-related, while what I studied were the effects of ‘non-resource’ stressors (i.e. heavy-metal pollution and wind). The results have potential practical applications: the utilisation of locally adapted seedlings and plant facilitation may increase the success of future restoration efforts in industrial barrens as well as in other wind-exposed sites. The findings also have implications with regard to the effects of global change in subarctic environments: the documented potential by mountain birch for rapid evolutionary change, together with the general lack of evolutionary ‘dead ends’, due to not (over)specialising to current natural conditions, increase the chances of this crucial forest-forming tree persisting even under the anticipated climate change.

Swan Lake ballet

Äänitetty: 14.-15.12.1954, University of Minnesota, Northrop Memorial Auditorium.

Functional diversity of macrophyte communities within and between Pyrenean lakes

Submersed vegetation is a common feature in about 70% Pyrene an high mountain (>1500 m a.s.l.) lakes. Isoetids and soft-water elodeids are common elements of this underw ater flora and can form distinct vegetation units (i.e. patches of vegetation dominated by different species) within complex mosaics of vegetation in shallow waters (<7 m). Since is oetids exert a strong influence on sediment biogeochemistry due to high radial oxygen loss, we examined the small scale characteristics of the lake environment (water and sediment) associated to vegetation patches in order to ascertain potential functional differences among them. To do so, we characterised the species composition and biomass of the main vegetation units from 11 lakes, defined plant communities based on biomass data, and then related each community with sediment properties (redox and dissolved nutrient concentration in the pore water) and water nutrient concentration within plant canopy. We also characterised lake water and sediment in areas without vegetation as a reference. A total of twenty-one vegetation units were identified, ranging from one to five per lake. A cluster analysis on biomass species composition suggested seven different macrophyte communities that were named after the most dominant species:Nitella sp.,Potamogeton praelongus, Myriophyllum alterniflorum, Sparganium angustifolium , Isoetes echinospora,Isoetes lacustris and Carex rostrata . Coupling between macrophyte communities and their immediate environment (overlying water and sediment) was manifested mainly as variation in sediment redox conditions and the dominant form of inorganic nitrogen in pore-water. These effects depended on the specific compositi on of the community, and on the allocation between above- and belowground biomass, and could be predicted with a model relating the average and standard deviation of sediment redox potential from 0 down to -20 cm, across macrophyte communities. Differences in pore-water total dissolved phosphorus were related to the trophic state of the lakes. There was no correlation between sediment and water column dissolved nutrients. However, nitrate concentrations tended to be lower in the water overlaying isoetid communities, in apparent contradiction to the patterns of dissolved nitrates in the pore-water. These tendencies were robust even when comparing the water over laying communities within the same lake, thus pointing towards a potential effect of isoetids in reducing dissolved nitrogen in the lakes.

Feeding ecology and trophic position of three sympatric demersal chondrichthyans in the Northwestern Mediterranean.

Understanding how marine predators interact is a scientific challenge. In marine ecosystems, segregation in feeding habits has been largely described as a common mechanism to allow the coexistence of several competing marine predators. However, little is known about the feeding ecology of most species of chondrichthyans, which play a pivotal role in the structure of marine food webs worldwide. In this study, we examined the trophic ecology of 3 relatively abundant chondrichthyans coexisting in the Mediterranean Sea: the blackmouth catshark Galeus melastomus , the velvet belly lanternshark Etmopterus spinax and the rabbit fish Chimaera monstrosa. To examine their trophic ecology and interspecific differences in food habits, we combined the analysis of stomach content and stable isotopes. Our results highlighted a trophic segregation between C. monstrosa and the other 2 species. G. melastomus showed a diet composed mainly of cephalopods, while E. spinax preyed mainly on shrimps and C. monstrosa on crabs. Interspecific differences in the trophic niche were likely due to different feeding capabilities and body size. Each species showed different isotopic niche space and trophic level. Specifically, C. monstrosa showed a higher trophic level than E. spinax and G. melastomus. The high trophic levels of the 3 species highlighted their important role as predators in the marine food web. Our results illustrate the utility of using complementary approaches that provide information about the feeding behaviour at short (stomach content) and long-term scales (stable isotopes), which could allow more efficient monitoring of marine food-web changes in the study area.

Population structure in a highly pelagic seabird, the Cory's shearwater (Calonectris diomedea): an examination of genetics, morphology and ecology

Increasing evidence suggests oceanic traits may play a key role in the genetic structuring of marine organisms. Whereas genetic breaks in the open ocean are well known in fishes and marine invertebrates, the importance of marine habitat characteristics in seabirds remains less certain. We investigated the role of oceanic transitions versus population genetic processes in driving population differentiation in a highly vagile seabird, the Cory"s shearwater, combining molecular, morphological and ecological data from 27 breeding colonies distributed across the Mediterranean (Calonectris diomedea diomedea) and the Atlantic (C. d. borealis). Genetic and biometric analyses showed a clear differentiation between Atlantic and Mediterranean Cory"s shearwaters. Ringing-recovery data indicated high site fidelity of the species, but we found some cases of dispersal among neighbouring breeding sites (<300 km) and a few long distance movements (>1000 km) within and between each basin. In agreement with this, comparison of phenotypic and genetic data revealed both current and historical dispersal events. Within each region, we did not detect any genetic substructure among archipelagos in the Atlantic, but we found a slight genetic differentiation between western and eastern breeding colonies in the Mediterranean. Accordingly, gene flow estimates suggested substantial dispersal among colonies within basins. Overall, genetic structure of the Cory"s shearwater matches main oceanographic breaks (Almería-Oran Oceanic Front and Siculo-Tunisian Strait), but spatial analyses suggest that patterns of genetic differentiation are better explained by geographic rather than oceanographic distances. In line with previous studies, genetic, phenotypic and ecological evidence supported the separation of Atlantic and Mediterranean forms, suggesting the 2 taxa should be regarded as different species.

Holocene Varved Lake Sediments as Records of Palaeoenvironmental Change and Geomagnetic Palaeosecular Variation in Eastern Finland

This dissertation discusses Holocene palaeoenvironmental and palaeomagnetic secular variation (PSV) records reconstructed from sediments preserved in Lake Lehmilampi (63º37´N, 29º06´E) and Lake Kortejärvi (63º37´N, 28º56´E) in eastern Finland. Several piston and freeze cores were obtained from both lakes for varve and magnetic analyses. Sediment samples were impregnated in low-viscosity epoxy and physical parameters of varves, including varve thickness and relative grey-scale values, were recorded using x-ray densitometry combined with semiautomatic digital image analysis. On average, varve records of Lehmilampi and Kortejärvi cover 5122 and 3902 years, respectively. Past solar activity, as estimated by residual ₁₄C data, compares favourably with varve thicknesses from Lehmilampi during the last 2000 years. This indicates the potential of clastic-organic varves to record sensitively climatic variations. Bulk magnetic parameters, including magnetic susceptibility together with natural, anhysteretic and isothermal remanent magnetizations, were measured to describe mineral magnetic properties and geomagnetic palaeosecular variation recorded in the sediments. Main stages in the development of the investigated lakes are reflected in the variations in the mineral magnetic records, sediment lithology and composition. Similar variations in magnetic parameters and sediment organic matter suggest contribution of bacterial magnetite in the magnetic assemblages of Lehmilampi. Inclination and relative declination records yielded largely consistent results, attesting to the great potential of these sediments to preserve directional palaeosecular variation in high resolution. The PSV data from Lehmilampi and Kortejärvi were stacked into North Karelian PSV stack, which may be used for dating homogenous lake sediments in the same regional context. Reconstructed millennial variations in relative palaeointensity results are approximately in agreement with those seen in the absolute palaeointensity data from Europe. Centennial variations in the relative palaeointensity, however, are influenced by environmental changes. Caution is recommended when using varved lake sediments in reconstructing relative palaeointensity.

Reproductive ecology of the exotic tree Muntingia calabura L. (Muntingiaceae) in southeastern Brazil

The exotic tree Muntingia calabura L. (Muntingiaceae), a species native to Central America, is used as fish feed and fiber and cellulose production in Brazil. This study was carried out in urban areas and verified the reproductive biology of this plant species. Flower and fruit morphology, compatibility system, reproductive phenology, pollination and frugivore animals, and germination of disseminated seeds were recorded by standard field and laboratory procedures. This tree is self-compatible and autonomously self-pollinated, with its flowers being mainly visited by bees and its fruits consumed by birds and bats. Germination of its dispersed seeds is fast and occurs at a high rate. The results of this work suggest that M. calabura is very adaptable to cultivated areas,thus being an excellent choice for urban reforestation. However, its reproductive characteristics place this plant as an invasive species with significant potential in southeastern Brazil.

Effects of anthropogenic and natural hydrological changes on the behavior of the acidic metal discharge from acid sulfate soils in a river- and lake system in western Finland

Larsmo-Öjasjön i Österbotten skapades genom invallningar på 1960-talet pga. industrins behov av sötvatten. Sedan dess har vattenområdet drabbats av återkommande försurning och fiskdöd, och invallningen har ofta beskyllts för problemen. Avhandlingen undersöker syrabelastningen i området; bl.a. hur markanvändning, hydrologi och klimatförändringen påverkar belastningen. Konsekvenserna undersöks med fiskyngel som bioindikator, och olika miljömetoder testas och diskuteras. Ökad kunskap om försurningen hjälper oss att tillämpa effektiva miljömetoder och få förbättrad vattenkvalitet i framtiden. Den primära orsaken till den försämrade vattenkvaliteten under de senaste 40 åren är intensiv dikning av svavelrika sediment. Detta leder till oxidering av svavlet till svavelsyra och uppkomst av sura sulfatjordar. Syran löser upp mängder med toxiska metaller som spolas ut i vattendragen. Undersökningen visar att tiotusentals ton svavelsyra tillsammans med stora mängder metaller rinner till Larsmo-Öjasjön per år från sura sulfatjordar. Åarna bidrar med mest belastning, men den sammanlagda belastningen från de otaliga dikena och bäckarna är oväntat stor. Andra potentiella källor till försurningen, t.ex. muddringar och humussyror, beräknas vara obetydliga. Syra- och metallbelastningen varierar kraftigt med hydrologin, dvs. störst belastning sker under vår- och höstflöden. En eventuell klimatförändring kan ändra på avrinningsmönstret och orsaka mera belastning vintertid. Den årligt återkommande syra- och metallbelastningen kan ofta hindra lakens förökning, vilket kan ha större långtgående konsekvenser för fiskpopulationerna än de relativt sällsynta stora surchockerna med synlig fiskdöd. För att förebygga skador på vattendragen bör man undvika att dränera svavelrika sedimenten. På redan existerande sura sulfatjordar visade sig kontroll av grundvattennivån kunna möjliggöra en effektiverad markanvändning utan märkbart ökade miljökonsekvenser.

Long-term nutrient load management and lake restoration: Case of Säkylän Pyhäjärvi (SW Finland)

Eutrophication caused by anthropogenic nutrient pollution has become one of the most severe threats to water bodies. Nutrients enter water bodies from atmospheric precipitation, industrial and domestic wastewaters and surface runoff from agricultural and forest areas. As point pollution has been significantly reduced in developed countries in recent decades, agricultural non-point sources have been increasingly identified as the largest source of nutrient loading in water bodies. In this study, Lake Säkylän Pyhäjärvi and its catchment are studied as an example of a long-term, voluntary-based, co-operative model of lake and catchment management. Lake Pyhäjärvi is located in the centre of an intensive agricultural area in southwestern Finland. More than 20 professional fishermen operate in the lake area, and the lake is used as a drinking water source and for various recreational activities. Lake Pyhäjärvi is a good example of a large and shallow lake that suffers from eutrophication and is subject to measures to improve this undesired state under changing conditions. Climate change is one of the most important challenges faced by Lake Pyhäjärvi and other water bodies. The results show that climatic variation affects the amounts of runoff and nutrient loading and their timing during the year. The findings from the study area concerning warm winters and their influences on nutrient loading are in accordance with the IPCC scenarios of future climate change. In addition to nutrient reduction measures, the restoration of food chains (biomanipulation) is a key method in water quality management. The food-web structure in Lake Pyhäjärvi has, however, become disturbed due to mild winters, short ice cover and low fish catch. Ice cover that enables winter seining is extremely important to the water quality and ecosystem of Lake Pyhäjärvi, as the vendace stock is one of the key factors affecting the food web and the state of the lake. New methods for the reduction of nutrient loading and the treatment of runoff waters from agriculture, such as sand filters, were tested in field conditions. The results confirm that the filter technique is an applicable method for nutrient reduction, but further development is needed. The ability of sand filters to absorb nutrients can be improved with nutrient binding compounds, such as lime. Long-term hydrological, chemical and biological research and monitoring data on Lake Pyhäjärvi and its catchment provide a basis for water protection measures and improve our understanding of the complicated physical, chemical and biological interactions between the terrestrial and aquatic realms. In addition to measurements carried out in field conditions, Lake Pyhäjärvi and its catchment were studied using various modelling methods. In the calibration and validation of models, long-term and wide-ranging time series data proved to be valuable. Collaboration between researchers, modellers and local water managers further improves the reliability and usefulness of models. Lake Pyhäjärvi and its catchment can also be regarded as a good research laboratory from the point of view of the Baltic Sea. The main problem in both of them is eutrophication caused by excess nutrients, and nutrient loading has to be reduced – especially from agriculture. Mitigation measures are also similar in both cases.

Germination ecology of Rhynchosia capitata: an emerging summer weed in Asia

Rhynchosia capitata is becoming an increasing problem in summer crops, such as cotton, soybean, pearl millet and mungbean in many Asian countries. Laboratory and greenhouse studies have been conducted to determine the effects of several environmental factors on seed germination patterns and seedling emergence of R. capitata. We investigated whether the diverse ecological factors such as temperature, light, salinity, moisture stress, pH, and soil depth affected germination and seedling emergence of R. capitata. Germination increased as temperature increased from 25ºC and significantly reduced at 45ºC. Presence or absence of light did not influence germination. Germination of R. capitata was sensitive to increased salt and moisture stress, as well as to seed burial depth. Only 48% of seeds germinated at 150 mM salt concentration compared to 100% in control (distilled water). Similarly, 15% of seeds germinated at an osmotic potential of ‑0.8 MPa compared to 88% at ‑0.2 MPa. The optimum pH for seed germination of R. capitata was 7 (98% germination), but the seeds also germinated at lower level of pH 5 (85%) and at higher level of pH 10 (75%). In seed burial trial, maximum seedling emergence of 93% occurred at 2 cm depth, and seedling did not emerge from a depth of 12 cm. The high germination ability of R. capitata under a wide range of ecological factors suggests that this species is likely to be the one to cause more problems in a near future, if not managed appropriately.

Germination ecology of wild onion: a rainfed crop weed

Asphodelus tenuifolius is becoming a more common weed in rain-fed area in Pakistan. Laboratory and greenhouse experiments were conducted to determine the effect of different environmental factors on germination and emergence of A.tenuifolius. Results showed that A.tenuifolius can tolerate a wide range of varying environmental factors. Greatest percentage of germination (80%) was recorded at 15 ºC constant temperature; however, considerable germination occurred at 20 and 25 ºC. Light for 10 h photoperiod stimulate germination of Asphodelus tenuifolius compared with complete darkness. Germination was totally inhibited at osmotic stress higher than -0.8 MPa. There was no significant difference in germination at pH 6 to 8; however, there was a slightly decrease at pH 9, compared with distilled water. Asphodelus tenuifolius was very sensitive to salinity; however, a few seeds of A.tenuifolius were able to germinate even at 150 mM NaCl concentration. Greatest emergence occurred with seed placed at soil surface and emergence decreased with increase in burial depth. No emergence occurred from 4 cm or greater. This information may aid in developing tools and strategies for management.

Population ecology of Paepalanthus polyanthus (Bong.) Kunth: demography and life history of a sand dune monocarpic plant

Aspects of population dynamics and life history of Paepalanthus polyanthus (Bong.) Kunth, a sand dune monocarpic plant, were evaluated. A five year study was carried out on three permanent plots (5 m x 5 m) in a sand dune slack at Joaquina beach, Santa Catarina State, Brazil. From December 1986 to June 1989, the population decreased due to the death of the post reproductive plants and a low emergence of seedlings. In June 1989, a great recruitment occurred, but no plants survived. The population re-established itself by 1990-1991. The emergence and high survival of seedlings depended on periods of high pluviosity. Nevertheless, the summer flooding and episodes of drought represented key factors in mortality. The birth and mortality rates varied among the areas. It is suggested that these differences are related with depth of the ground water and with vegetation cover at each site. Paepalanthus polyanthus can reproduce in the second year of life, but few plants do this. The chances of survival and reproduction increase with the size of the basal leaf rosette. Although the production of seeds increases with size, the risk of unexpected flooding, for instance, suggest that a great delay in reproduction might not be the most favorable strategy.

Pollination ecology of Tabebuia aurea (Manso) Benth. & Hook. and T. ochracea (Cham.) Standl. (Bignoniaceae) in Central Brazil cerrado vegetation

The pollination ecology and breeding systems of Tabebuia aurea (Manso) Benth. & Hook., and T. ochracea (Cham.) Standl. were investigated in an area of cerrado vegetation in the Federal District of Brazil. These species occur sympatrically, flower massively and synchronously for a month, during the dry season (July to September). Both have diurnal anthesis, with similar floral structures, a yellow tubular corolla and produce nectar. Fourteen species of bees visited both Tabebuia species, but, only three Centris species and Bombus morio, were considered potential pollinators, because of their high frequency on the flowers and their efficiency in carrying pollen. Tests on the breeding systems of T. aurea and T. ochracea demonstrated that boths species are self-incompatible, with late-acting self-incompatibility. The proportion of fruit set from cross pollination (T. aurea 17.2% and T. ochracea 12.3%) in both species was low considering the great number of flowers displayed. This suggests a lack of maternal resources for fruit-set. The great amount of seeds per fruit (about 92 in T. aurea and 285 in T. ochracea) may represent an investment of maternal resources allocated on higher quality of fertilized ovules.