Dung beetle radiations in Madagascar

Speciation on islands is affected by island size and the range of habitats and resources available and often also by limited interactions with other taxa. An ancestral population may evolve into a large number of species via an adaptive radiation. In Madagascar, most groups of animals and plants have radiated on the island, having arrived via oceanic dispersal during the long isolation of Madagascar. Characteristic features of Malagasy biota are exceptionally high level of endemism, high species richness as well as lack of many higher taxa that are dominant on the African mainland. Malagasy dung beetles are dominated by two tribes, Canthonini and Helictopleurina, with more than 250 endemic species. In this thesis I have reconstructed molecular phylogenies for the two tribes using several gene regions and different phylogenetic methods. Evolution of closely related species and among populations of the same species was examined with haplotype networks. The Malagasy Canthonini consists of three large lineages, while Helictopleurina forms a monophyletic group. The ancestors of each of the four clades colonised Madagascar at different times during Cenozoic. The subsequent radiations differ in terms of the number of extant species (from 37 to more than 100) and the level of ecological differentiation. In addition, Onthophagini (6 species) and Scarabaeini (3) have colonised Madagascar several times, but they have not radiated and the few species have not entered forests where Canthonini and Helictopleurina mostly occur. Among the three Canthonini radiations, speciation appears to have been mostly allopatric in the oldest and the youngest clades, while in the Epactoides clade sister species have diverged in their ecologies but have similar geographical distributions, indicating that speciation may have occurred in regional sympatry. The most likely isolating mechanisms have been rivers and forest refugia during dry and cool geological periods. Most species are generalists feeding on both carrion and dung, and competition among ecologically similar species may prevent their coexistence in the same communities. Some species have evolved to forage in the canopy and a few species have shifted to use cattle dung, a new resource in the open habitats following the introduction of cattle 1500 years ago. The latter shift has allowed species to expand their geographical ranges.

Ozone-induced signaling in Arabidopsis thaliana

Tropospheric ozone (O3) is one of the most common air pollutants in industrialized countries, and an increasing problem in rapidly industrialising and developing countries in Asia, Africa and South America. Elevated concentrations of tropospheric O3 can lead to decrease in photosynthesis rate and therefore affect the normal metabolism, growth and seed production. Acute and high O3 episodes can lead to extensive damage leading to dead tissue in plants. Thus, O3 derived growth defects can lead to reduction in crop yield thereby leading to economical losses. Despite the extensive research on this area, many questions remain open on how these processes are controlled. In this study, the stress-induced signaling routes and the components involved were elucidated in more detail starting from visual damage to changes in gene expression, signaling routes and plant hormone interactions that are involved in O3-induced cell death. In order to elucidate O3-induced responses in Arabidopsis, mitogen-activated protein kinase (MAPK) signaling was studied using different hormonal signaling mutants. MAPKs were activated at the beginning of the O3 exposure. The activity of MAPKs, which were identified as AtMPK3 and AtMPK6, reached the maximum at 1 and 2 hours after the start of the exposure, respectively. The activity decreased back to clean air levels at 8 hours after the start of the exposure. Both AtMPK3 and AtMPK6 were translocated to nucleus at the beginning of the O3 exposure where they most likely affect gene expression. Differences were seen between different hormonal signaling mutants. Functional SA signaling was shown to be needed for the full protein levels and activation of AtMPK3. In addition, AtMPK3 and AtMPK6 activation was not dependent on ethylene signaling. Finally, jasmonic acid was also shown to have an impact on AtMPK3 protein levels and AtMPK3 activity. To further study O3-induced cell death, an earlier isolated O3 sensitive Arabidopsis mutant rcd1 was mapped, cloned and further characterized. RCD1 was shown to encode a gene with WWE and ADP-ribosylation domains known to be involved in protein-protein interactions and cell signaling. rcd1 was shown to be involved in many processes including hormonal signaling and regulation of stress-responsive genes. rcd1 is sensitive against O3 and apoplastic superoxide, but tolerant against paraquat that produces superoxide in chloroplast. rcd1 is also partially insensitive to glucose and has alterations in hormone responses. These alterations are seen as ABA insensitivity, reduced jasmonic acid sensitivity and reduced ethylene sensitivity. All these features suggest that RCD1 acts as an integrative node in hormonal signaling and it is involved in the hormonal regulation of several specific stress-responsive genes. Further studies with the rcd1 mutant showed that it exhibits the classical features of programmed cell death, PCD, in response to O3. These include nuclear shrinkage, chromatin condensation, nuclear DNA degradation, cytosol vesiculation and accumulation of phenolic compounds and eventually patches of HR-like lesions. rcd1 was found to produce extensive amount of salicylic acid and jasmonic acid in response to O3. Double mutant studies showed that SA independent and dependent processes were involved in the O3-induced PCD in rcd1 and that increased sensitivity against JA led to increased sensitivity against O3. Furthermore, rcd1 had alterations in MAPK signature that resembled changes that were previously seen in mutants defective in SA and JA signaling. Nitric oxide accumulation and its impact on O3-induced cell death were also studied. Transient accumulation of NO was seen at the beginning of the O3 exposure, and during late time points, NO accumulation coincided with the HR-like lesions. NO was shown to modify defense gene expression, such as, SA and ethylene biosynthetic genes. Furthermore, rcd1 was shown to produce more NO in control conditions. In conclusion, NO was shown to be involved in O3-induced signaling leading to attenuation of SA biosynthesis and other defense related genes.

Photobiological studies of Baltic Sea phytoplankton

Phytoplankton ecology and productivity is one of the main branches of contemporary oceanographic research. Research groups in this branch have increasingly started to utilise bio-optical applications. My main research objective was to critically investigate the advantages and deficiencies of the fast repetition rate (FRR) fluorometry for studies of productivity of phytoplankton, and the responses of phytoplankton towards varying environmental stress. Second, I aimed to clarify the applicability of the FRR system to the optical environment of the Baltic Sea. The FRR system offers a highly dynamic tool for studies of phytoplankton photophysiology and productivity both in the field and in a controlled environment. The FRR metrics obtain high-frequency in situ determinations of the light-acclimative and photosynthetic parameters of intact phytoplankton communities. The measurement protocol is relatively easy to use without phases requiring analytical determinations. The most notable application of the FRR system lies in its potential for making primary productivity (PP) estimations. However, the realisation of this scheme is not straightforward. The FRR-PP, based on the photosynthetic electron flow (PEF) rate, are linearly related to the photosynthetic gas exchange (fixation of 14C) PP only in environments where the photosynthesis is light-limited. If the light limitation is not present, as is usually the case in the near-surface layers of the water column, the two PP approaches will deviate. The prompt response of the PEF rate to the short-term variability in the natural light field makes the field comparisons between the PEF-PP and the 14C-PP difficult to interpret, because this variability is averaged out in the 14C-incubations. Furthermore, the FRR based PP models are tuned to closely follow the vertical pattern of the underwater irradiance. Due to the photoacclimational plasticity of phytoplankton, this easily leads to overestimates of water column PP, if precautionary measures are not taken. Natural phytoplankton is subject to broad-waveband light. Active non-spectral bio-optical instruments, like the FRR fluorometer, emit light in a relatively narrow waveband, which by its nature does not represent the in situ light field. Thus, the spectrally-dependent parameters provided by the FRR system need to be spectrally scaled to the natural light field of the Baltic Sea. In general, the requirement of spectral scaling in the water bodies under terrestrial impact concerns all light-adaptive parameters provided by any active non-spectral bio-optical technique. The FRR system can be adopted to studies of all phytoplankton that possess efficient light harvesting in the waveband matching the bluish FRR excitation. Although these taxa cover the large bulk of all the phytoplankton taxa, one exception with a pronounced ecological significance is found in the Baltic Sea. The FRR system cannot be used to monitor the photophysiology of the cyanobacterial taxa harvesting light in the yellow-red waveband. These taxa include the ecologically-significant bloom-forming cyanobacterial taxa in the Baltic Sea.

Carbon balance and component CO2 fluxes in boreal Scots pine stands

The forest vegetation takes up atmospheric carbon dioxide (CO2) in photosynthesis. Part of the fixed carbon is released back into the atmosphere during plant respiration but a substantial part is stored as plant biomass, especially in the stems of trees. Carbon also accumulates in the soil as litter and via the roots. CO2 is released into the atmosphere from these carbon stocks in the decomposition of dead biomass. Carbon balance of a forest stand is the difference between the CO2 uptake and CO2 efflux. This study quantifies and analyses the dynamics of carbon balance and component CO2 fluxes in four Southern Finnish Scots pine stands that covered the typical economic rotation time of 80 years. The study was based on direct flux measurements with chambers and eddy covariance (EC), and modelling of component CO2 fluxes. The net CO2 exchange of the stand was partitioned into component fluxes: photosynthesis of trees and ground vegetation, respiration of tree foliage and stems, and CO2 efflux from the soil. The relationships between the component fluxes and the environmental factors (light, temperature, atmospheric CO2, air humidity and soil moisture) were studied with mathematical modelling. The annual CO2 balance varied from a source of about 400 g C/m2 at a recently clearcut site to net CO2 uptake of 200 300 g C/m2 in a middle-aged (40-year-old) and a mature (75-year-old) stand. A 12-year-old sapling site was at the turning point from source to a sink of CO2. In the middle-aged stand, photosynthetic production was dominated by trees. Under closed pine canopies, ground vegetation accounted for 10 20% of stand photosynthesis whereas at the open sites the proportion and also the absolute photosynthesis of ground vegetation was much higher. The aboveground respiration was dominated by tree foliage which accounted for one third of the ecosystem respiration. Rate of wood respiration was in the order of 10% of total ecosystem respiration. CO2 efflux from the soil dominated the ecosystem respiratory fluxes in all phases of stand development. Instantaneous and delayed responses to the environmental driving factors could predict well within-year variability in photosynthetic production: In the short term and during the growing season photosynthesis follows primarily light while the seasonal variation is more strongly connected to temperature. The temperature relationship of the annual cycle of photosynthesis was found to be almost equal in the southern boreal zone and at the timberline in the northern boreal zone. The respiratory fluxes showed instantaneous and seasonal temperature relationships but they could also be connected to photosynthesis at an annual timescale.

Dark Mess

In a world consumed by quests for happiness and personal growth, Grant Stevens��� new exhibition, Dark Mess, delves into the psychic troubles that sometimes lie below the canopy. Working predominantly with video, photography and installation, Stevens��� practice explores how the verbal and non-verbal languages of popular screen culture interface with contemporary subjectivity. This exhibition continues Stevens��� interest in the natural environment as a catalyst and proxy for introspection and self-discovery. Pushing sound and image to distortion, Dark Mess offers a disquieting journey through the undergrowth.

Assessment of crop insect damage using unmanned aerial systems: A machine learning approach

Agricultural pests are responsible for millions of dollars in crop losses and management costs every year. In order to implement optimal site-specific treatments and reduce control costs, new methods to accurately monitor and assess pest damage need to be investigated. In this paper we explore the combination of unmanned aerial vehicles (UAV), remote sensing and machine learning techniques as a promising technology to address this challenge. The deployment of UAVs as a sensor platform is a rapidly growing field of study for biosecurity and precision agriculture applications. In this experiment, a data collection campaign is performed over a sorghum crop severely damaged by white grubs (Coleoptera: Scarabaeidae). The larvae of these scarab beetles feed on the roots of plants, which in turn impairs root exploration of the soil profile. In the field, crop health status could be classified according to three levels: bare soil where plants were decimated, transition zones of reduced plant density and healthy canopy areas. In this study, we describe the UAV platform deployed to collect high-resolution RGB imagery as well as the image processing pipeline implemented to create an orthoimage. An unsupervised machine learning approach is formulated in order to create a meaningful partition of the image into each of the crop levels. The aim of the approach is to simplify the image analysis step by minimizing user input requirements and avoiding the manual data labeling necessary in supervised learning approaches. The implemented algorithm is based on the K-means clustering algorithm. In order to control high-frequency components present in the feature space, a neighbourhood-oriented parameter is introduced by applying Gaussian convolution kernels prior to K-means. The outcome of this approach is a soft K-means algorithm similar to the EM algorithm for Gaussian mixture models. The results show the algorithm delivers decision boundaries that consistently classify the field into three clusters, one for each crop health level. The methodology presented in this paper represents a venue for further research towards automated crop damage assessments and biosecurity surveillance.

Measurements of Volatile Organic Compounds - from Biogenic Emissions to Concentrations in Ambient Air

Volatile organic compounds (VOCs) affect atmospheric chemistry and thereafter also participate in the climate change in many ways. The long-lived greenhouse gases and tropospheric ozone are the most important radiative forcing components warming the climate, while aerosols are the most important cooling component. VOCs can have warming effects on the climate: they participate in tropospheric ozone formation and compete for oxidants with the greenhouse gases thus, for example, lengthening the atmospheric lifetime of methane. Some VOCs, on the other hand, cool the atmosphere by taking part in the formation of aerosol particles. Some VOCs, in addition, have direct health effects, such as carcinogenic benzene. VOCs are emitted into the atmosphere in various processes. Primary emissions of VOC include biogenic emissions from vegetation, biomass burning and human activities. VOCs are also produced in secondary emissions from the reactions of other organic compounds. Globally, forests are the largest source of VOC entering the atmosphere. This thesis focuses on the measurement results of emissions and concentrations of VOCs in one of the largest vegetation zones in the world, the boreal zone. An automated sampling system was designed and built for continuous VOC concentration and emission measurements with a proton transfer reaction - mass spectrometer (PTR-MS). The system measured one hour at a time in three-hourly cycles: 1) ambient volume mixing-ratios of VOCs in the Scots-pine-dominated boreal forest, 2) VOC fluxes above the canopy, and 3) VOC emissions from Scots pine shoots. In addition to the online PTR-MS measurements, we determined the composition and seasonality of the VOC emissions from a Siberian larch with adsorbent samples and GC-MS analysis. The VOC emissions from Siberian larch were reported for the fist time in the literature. The VOC emissions were 90% monoterpenes (mainly sabinene) and the rest sesquiterpenes (mainly a-farnesene). The normalized monoterpene emission potentials were highest in late summer, rising again in late autumn. The normalized sesquiterpene emission potentials were also highest in late summer, but decreased towards the autumn. The emissions of mono- and sesquiterpenes from the deciduous Siberian larch, as well as the emissions of monoterpenes measured from the evergreen Scots pine, were well described by the temperature-dependent algorithm. In the Scots-pine-dominated forest, canopy-scale emissions of monoterpenes and oxygenated VOCs (OVOCs) were of the same magnitude. Methanol and acetone were the most abundant OVOCs emitted from the forest and also in the ambient air. Annually, methanol and mixing ratios were of the order of 1 ppbv. The monoterpene and sum of isoprene 2-methyl-3-buten-2-ol (MBO) volume mixing-ratios were an order of magnitude lower. The majority of the monoterpene and methanol emissions from the Scots-pinedominated forest were explained by emissions from Scots pine shoots. The VOCs were divided into three classes based on the dynamics of the summer-time concentrations: 1) reactive compounds with local biological, anthropogenic or chemical sources (methanol, acetone, butanol and hexanal), 2) compounds whose emissions are only temperaturedependent (monoterpenes), 3) long-lived compounds (benzene, acetaldehyde). Biogenic VOC (methanol, acetone, isoprene MBO and monoterpene) volume mixing-ratios had clear diurnal patterns during summer. The ambient mixing ratios of other VOCs did not show this behaviour. During winter we did not observe systematical diurnal cycles for any of the VOCs. Different sources, removal processes and turbulent mixing explained the dynamics of the measured mixing-ratios qualitatively. However, quantitative understanding will require longterm emission measurements of the OVOCs and the use of comprehensive chemistry models. Keywords: Hydrocarbons, VOC, fluxes, volume mixing-ratio, boreal forest

A generic individual-based spatially explicit model as a novel tool for investigating insect-plant interactions: A case study of the behavioural ecology of frugivorous Tephritidae

Computational modelling of mechanisms underlying processes in the real world can be of great value in understanding complex biological behaviours. Uptake in general biology and ecology has been rapid. However, it often requires specific data sets that are overly costly in time and resources to collect. The aim of the current study was to test whether a generic behavioural ecology model constructed using published data could give realistic outputs for individual species. An individual-based model was developed using the Pattern-Oriented Modelling (POM) strategy and protocol, based on behavioural rules associated with insect movement choices. Frugivorous Tephritidae (fruit flies) were chosen because of economic significance in global agriculture and the multiple published data sets available for a range of species. The Queensland fruit fly (Qfly), Bactrocera tryoni, was identified as a suitable individual species for testing. Plant canopies with modified architecture were used to run predictive simulations. A field study was then conducted to validate our model predictions on how plant architecture affects fruit flies��� behaviours. Characteristics of plant architecture such as different shapes, e.g., closed-canopy and vase-shaped, affected fly movement patterns and time spent on host fruit. The number of visits to host fruit also differed between the edge and centre in closed-canopy plants. Compared to plant architecture, host fruit has less contribution to effects on flies��� movement patterns. The results from this model, combined with our field study and published empirical data suggest that placing fly traps in the upper canopy at the edge should work best. Such a modelling approach allows rapid testing of ideas about organismal interactions with environmental substrates in silico rather than in vivo, to generate new perspectives. Using published data provides a saving in time and resources. Adjustments for specific questions can be achieved by refinement of parameters based on targeted experiments.

Metsik��n rakenteen vaikutus s��teilyn sammumiskertoimeen

Kasvuston G-funktio kuvaa s��teilyn v��henemist�� tai ���sammumista��� auringon eri korkeuskulmissa varjostavan lehtialan suhteen. Sen vuoksi sit�� kutsutaan sammumiskertoimeksi. Tutkimuksen tavoitteena oli selvitt���� vaihteleeko G-funktion muoto eri puulajien metsik��iss��, ja voidaanko puulajit erottaa toisistaan G-funktion muodon avulla. Arvioin lis��ksi G-funktioiden muodossa kasvukaudenaikana tapahtuvia muutoksia. Muita tutkimuksen kannalta mielenkiintoisia tutkimuskysymyksi�� olivat voidaanko ryhmittymist�� tai latvusmuotoa arvioida G-funktion muodon avulla. Maastomittaukset suoritettiin 3.5.2010 -30.9.2010 v��lisen�� aikana. Tutkimusalueena oli Hyyti��l��n mets��aseman (68?59`N, 35?72`E) ymp��rist��, jonka mets��t edustavat tyypillist�� boreaalista havu- ja lehtimets����. Tutkimusta varten perustettiin kuusi yhden puulajin koealaa, kaksi kutakin puulajia kohden. Tutkittavat puulajit olivat m��nty (Pinus sylvestris), kuusi (Picea abies) ja rauduskoivu (Betula pendula). Koealaparit valittiin niin, ett�� kullekin puulajille muodostui varttuneesta mets��st�� ja taimikosta muodostuva pari tai vaihtoehtoisesti tihe��mpi ja harvempi koeala. Koealoille perustettiin 81 mittauspistett�� sis��lt��v�� mittaushila, jossa kunkin pisteen aukkoisuustiedot mitattiin kahden viikon v��lein. Mittaukset suoritettiin kahdella LAI-2000 Plant Canopy Analyzer -laitteella. Laitteiden tulosteista koealoille saatiin aukkoisuustiedot T(?) ja LAI, joiden avulla saatiin laskettua tarkasteltavat G-funktiot. Saman puulajin G-funktiota vertailtiin toisiinsa puulajityypillisten trendien havaitsemiseksi. Keskikes��ll�� eri puulajien G-funktioita verrattiin toisiinsa. Teoreettisten simulointien avulla tutkittiin latvuksen dimensioiden (latvuksen pituus ja s��de) ja sis��isen ryhmitt��isyyden vaikutusta puulajikohtaiseen G-funktioon. Simuloinneissa k��ytettiin hyv��ksi tietoa koealojen puustotunnuksista, lehtialasta sek�� runkoluvusta. Puulajikohtaiset G-funktiot ovat erotettavissa toisistaan funktion minimi- ja maksimiarvojen sijoittumisen sek�� suhteellisen vaihteluv��lin perusteella. Havupuualojen G-funktiot eiv��t juuri muuttuneet kasvukauden aikana. Koivualoilla G-funktion kasvukaudenaikaiset muutokset (pelk��t oksat, hiirenkorvat ja t��ysikasvuiset lehdet) olivat sit�� vastoin helposti havaittavissa. G-funktion muodon avulla voidaan my��s arvioida latvusmuotoa ja ryhmittymist��. M��ntyjen latvusmuoto on approksimoitavissa parhaiten ympyr��kartion avulla. Kuusien ja koivujen latvusmuodon approksimointiin parhaana vaihtoehtona voidaan pit���� ellipsoidia. Teoreettisten simulointien perusteella nuori kuusikko on muita havupuualoja ryhmittyneempi. Tutkimuksen mukaan s��teily sammuu satelliittien yleisimm��ss�� kuvaussuunnassa tehokkaammin kuin kaikkien suuntien yli laskettu keskiarvo (0,5) antaa olettaa. Puulajikohtaisten G-funktioiden avulla voidaan ep��suorasti arvioida sek�� mets��st�� tapahtuvaa heijastusta ett�� metsik��n sis��isi�� s��teilyolosuhteita, sill�� puulaji yhdess�� metsik��n rakenteen kanssa vaikuttaa mets��st�� heijastuvaan s��teilyyn. Puulajikohtainen G-funktio on parametri, jonka avulla voidaan kalibroida malleja, joissa tarvitaan tietoa s��teilyn kulusta erilaisissa kasvustoissa.

Climatic adaptation of Norway spruce (Picea abies (L.) Karsten) in Finland based on male flowering phenology.

Anthesis was studied at the canopy level in 10 Norway spruce stands from 9 localities in Finland from 1963 to 1974. Distributions of pollen catches were compared to the normal Gaussian distribution. The basis for the timing studies was the 50 per cent point of the anthesis-fitted normal distribution. Development up to this point was given in calendar days, in degree days (>5 ��C) and in period units. The count of each parameter began on March 19 (included). Male flowering in Norway spruce stands was found to have more annual variation in quantity than in Scots pine stands studied earlier. Anthesis in spruce in northern Finland occurred at a later date than in the south. The heat sums needed for anthesis varied latitudinally less in spruce than in pine. The variation of pollen catches in spruce increased towards north-west as in the case of Scots pine. In the unprocessed data, calendar days were found to be the most accurate forecast of anthesis in Norway spruce both for a single year and for the majority of cases of stand averages over several years. Locally, the period unit could be a more accurate parameter for the stand average. However, on a calendar day basis, when annual deviations between expected and measured heat sums were converted to days, period units were narrowly superior to days. The geographical correlations respect to timing of flowering, calculated against distances measured along simulated post-glacial migration routes, were stronger than purely latitudinal correlations. Effects of the reinvasion of Norway spruce into Finland are thus still visible in spruce populations just as they were in Scots pine populations. The proportion of the average annual heat sum needed for spruce anthesis grew rapidly north of a latitude of ca. 63�� and the heat sum needed for anthesis decreased only slighty towards the timberline. In light of flowering phenology, it seems probable that the northwesterly third of Finnish Norway spruce populations are incompletely adapted to the prevailing cold climate. A moderate warming of the climate would therefore be beneficial for Norway spruce. This accords roughly with the adaptive situation in Scots pine.

Differences in the climatic adaptation of silver birch (Betula pendula) and downy birch (B. pubescens) in Finland based on male flowering phenology.

Male flowering was studied at the canopy level in 10 silver birch (Betula pendula Roth) stands from 8 localities and in 14 downy birch (B. pubescens Ehrh.) stands from 10 localities in Finland from 1963 to 1973. Distributions of cumulative pollen catches were compared to the normal Gaussian distribution. The basis for the timing of flowering was the 50 per cent point of the anthesis-fitted normal distribution. To eliminate effects of background pollen, only the central, normally distributed part of the cumulative distribution was used. Development up to the median point of the distribution was measured and tested in calendar days, in degree days (> 5 ��C) and in period units. The count of each parameter began on and included March 19. Male flowering in silver birch occurred from late April to late June depending on latitude, and flowering in downy birch took place from early May to early July. The heat sums needed for male flowering varied in downy birch stands latitudinally but there was practically no latitudinal variation in heat sums needed for silver birch flowering. The amount of male flowering in stands of both birch species were found to have a large annual variation but without any clear periodicity. The between years pollen catch variation in stands of either birch species did not show any significant latitudinal correlation in contrast to Norway spruce stands. The period unit heat sum gave the most accurate forecast of the timing of flowering for 60 per cent of the silver birch stands and for 78.6 per cent of the for downy birch stands. Calendar days, however, gave the best forecast for silver birch in 25 per cent of the cases, while degree days gave the best forecast for downy birch in 21.4 per cent of the cases. Silver birch seems to have a local inclination for a more fixed flowering date compared to downy birch, which could mean a considerable photoperiodic influence on flowering time of silver birch. Silver birch and downy birch had different geographical correlations. Frequent hybridization of birch species occurs more often in northern Finland in than in more southern latitudes. The different timing in flowering caused increasing scatter in flowering times in the north, especially in the case of downy birch. The chance of simultaneous flowering of silver birch and downy birch so increased northwards due to a more variable climate and also higher altitudinal variations. Compared with conifers, the reproduction cycles of both birch species were found to be well protected from damage by frost.

Alikasvoksen mittaus ja kartoitus laserkeilauksella

Tasaik��isen mets��n alle muodostuvilla alikasvoksilla on merkityst�� puunkorjuun, mets��nuudistamisen, n��kem��-ja maisema-analyysien sek�� biodiversiteetin ja hiilitaseen arvioinnin kannalta. Ilma-aluksista teht��v�� laserkeilaus on osoittautunut tehokkaaksi kaukokartoitusmenetelm��ksi varttuneiden puustojen mittauksessa. Laserkeilauksen k��ytt����notto operatiivisessa mets��suunnittelussa mahdollistaa aiempaa tarkemman tiedon tuottamisen alikasvoksista, mik��li alikasvoksen ominaisuuksia voidaan tulkita laseraineistoista. T��ss�� ty��ss�� k��ytettiin tarkasti mitattuja maastokoealoja ja kaikulaserkeilausaineistoja (discrete return LiDAR) usealta vuodelta (1���2 km lentokorkeus, 0,9���9,7 pulssia m-2). Laserkeilausaineistot oli hankittu Optech ALTM3100 ja Leica ALS50-II sensoreilla. Koealat edustavat suomalaisia tasaik��isi�� m��nnik��it�� eri kehitysvaiheissa. Tutkimuskysymykset olivat: 1) Mink��lainen on alikasvoksesta saatu lasersignaali yksitt��isen pulssin tasolla ja mitk�� tekij��t signaaliin vaikuttavat? 2) Mik�� on k��yt��nn��n sovelluksissa hy��dynnett��vien aluepohjaisten laserpiirteiden selitysvoima alikasvospuuston ominaisuuksien ennustamisessa? Erityisesti haluttiin selvitt����, miten laserpulssin energiah��vi��t ylempiin latvuskerroksiin vaikuttavat saatuun signaaliin, ja voidaanko laserkaikujen intensiteetille tehd�� energiah��vi��iden korjaus. Puulajien v��liset erot laserkaiun intensiteetiss�� olivat pieni�� ja vaihtelivat keilauksesta toiseen. Intensiteetin k��ytt��mahdollisuudet alikasvoksen puulajin tulkinnassa ovat siten hyvin rajoittuneet. Energiah��vi��t ylempiin latvuskerroksiin aiheuttivat alikasvoksesta saatuun lasersignaaliin kohinaa. Energiah��vi��iden korjaus tehtiin alikasvoksesta saaduille laserpulssin 2. ja 3. kaiuille. Korjauksen avulla pystyttiin pienent��m����n kohteen sis��ist�� intensiteetin hajontaa ja parantamaan kohteiden luokittelutarkkuutta alikasvoskerroksessa. K��ytett��ess�� 2. kaikuja oikeinluokitusprosentti luokituksessa maan ja yleisimm��n puulajin v��lill�� oli ennen korjausta 49,2���54,9 % ja korjauksen j��lkeen 57,3���62,0 %. Vastaavat kappa-arvot olivat 0,03���0,13 ja 0,10���0,22. T��rkein energiah��vi��it�� selitt��v�� tekij�� oli pulssista saatujen aikaisempien kaikujen intensiteetti, mutta hieman merkityst�� oli my��s pulssin leikkausgeometrialla ylemm��n latvuskerroksen puiden kanssa. My��s 3. kaiuilla luokitustarkkuus parani. Puulajien v��lill�� havaittiin eroja siin��, kuinka herk��sti ne tuottavat kaiun laserpulssin osuessa puuhun. Kuusi tuotti kaiun suuremmalla todenn��k��isyydell�� kuin lehtipuut. Erityisen selv�� t��m�� ero oli pulsseilla, joissa oli energiah��vi��it��. Laserkaikujen korkeusjakaumapiirteet voivat siten olla riippuvaisia puulajista. Sensorien v��lill�� havaittiin selvi�� eroja intensiteettijakaumissa, mik�� vaikeuttaa eri sensoreilla hankittujen aineistojen yhdist��mist��. My��s kaiun todenn��k��isyydet erosivat jonkin verran sensorien v��lill��, mik�� aiheutti pieni�� eroavaisuuksia kaikujen korkeusjakaumiin. Aluepohjaisista laserpiirteist�� l��ydettiin alikasvoksen runkolukua ja keskipituutta hyvin selitt��vi�� piirteit��, kun rajoitettiin tarkastelu yli 1 m pituisiin puihin. Piirteiden selitysvoima oli parempi runkoluvulle kuin keskipituudelle. Selitysvoima ei merkitt��v��sti alentunut pulssitiheyden pienentyess��, mik�� on hyv�� asia k��yt��nn��n sovelluksia ajatellen. Lehtipuun osuutta ei pystytty selitt��m����n. Tulosten perusteella kaikulaserkeilausta voi olla mahdollista hy��dynt���� esimerkiksi ennakkoraivaustarpeen arvioinnissa. Sen sijaan alikasvoksen tarkempi luokittelu (esim. puulajitulkinta) voi olla vaikeaa. Kaikkein pienimpi�� alikasvospuita ei pystyt�� havaitsemaan. Lis���� tutkimuksia tarvitaan tulosten yleist��miseksi erilaisiin metsik��ihin.

Tree mortality, fire scar formation and regeneration 8 years after low-intensity fire in managed Pinus sylvestris stands

Fire is an important driver of the boreal forest ecosystem, and a useful tool for the restoration of degraded forests. However, we lack knowledge on the ecological processes initiated by prescribed fires, and whether they bring about the desired restoration effects. The purpose of this study was to investigate the impacts of low-intensity experimental prescribed fires on four ecological processes in young commercial Scots pine (Pinus sylvestris) stands eight years after the burning. The processes of interest were tree mortality, dead wood creation, regeneration and fire scar formation. These were inventoried in twelve study plots, which were 30 m x 30 m in size. The plots belonged to two different stand age classes: 30-35 years or 45 years old at the time of burning. The study was partly a follow-up of study plots researched by Sidoroff et al. (2007) one year after burning in 2003. Tree mortality increased from 183 stems ha-1 in 2003 to 259 stems ha-1 in 2010, corresponding to 15 % and 21 % of stem number respectively. Most mortality was experienced in the stands of the younger age class, in smaller diameter classes and among species other than Scots pine. By 2010, the average mortality of Scots pine per plot was 18%, but varied greatly ranging from 0% to 63% of stem number. Delayed mortality, i.e. mortality that occurred between 2 and 8 years after fire, seemed to become more important with increasing diameter. The input of dead wood also varied greatly between plots, from none to 72 m3 ha-1, averaging at 12 m3 ha-1. The amount of fire scarred trees per plot ranged from none to 20 %. Four out of twelve plots (43 %) did not have any fire scars. Scars were on average small: 95% of scars were less than 4 cm in width, and 75% less than 40 cm in length. Owing to the light nature of the fire, the remaining overstorey and thick organic layer, regeneration was poor overall. The abundance of pine and other seedlings indicated a viable seed source existed, but the seedlings failed to establish under dense canopy. The number of saplings ranged from 0 to 12 333 stems ha-1. The results of this study indicate that a low intensity fire does not necessarily initiate the ecological processes of tree mortality, dead wood creation and regeneration in the desired scale. Fire scars, which form the basis of fire dating in fire history studies, did not form in all cases.

Does the age of fine root carbon indicate the age of fine roots in boreal forests?

To test the reliability of the radiocarbon method for determining root age, we analyzed fine roots (originating from the years 1985 to 1993) from ingrowth cores with known maximum root age (1 to 6 years old). For this purpose, three Scots pine (Pinus sylvestris L.) stands were selected from boreal forests in Finland. We analyzed root 14C age by the radiocarbon method and compared it with the above-mentioned known maximum fine root age. In general, ages determined by the two methods (root 14C age and ingrowth core root maximum age) were in agreement with each other for roots of small diameter (<0.5mm). By contrast, in most of the samples of fine roots of larger diameter (1.5-2mm), the 14C age of root samples of 1987-89 exceeded the ingrowth core root maximum age by 1-10 years. This shows that these roots had received a large amount of older stored carbon from unknown sources in addition to atmospheric CO2 directly from photosynthesis. We conclude that the 14C signature of fine roots, especially those of larger diameter, may not always be indicative of root age, and that further studies are needed concerning the extent of possible root uptake of older carbon and its residence time in roots. Keywords: fine root age, Pinus sylvestris, radiocarbon, root carbon, ingrowth cores, tree ring

Monitoring Indigenous Tropical Montane Forests in the Taita Hills Using Airborne Digital Camera Imagery

The loss and degradation of forest cover is currently a globally recognised problem. The fragmentation of forests is further affecting the biodiversity and well-being of the ecosystems also in Kenya. This study focuses on two indigenous tropical montane forests in the Taita Hills in southeastern Kenya. The study is a part of the TAITA-project within the Department of Geography in the University of Helsinki. The study forests, Ngangao and Chawia, are studied by remote sensing and GIS methods. The main data includes black and white aerial photography from 1955 and true colour digital camera data from 2004. This data is used to produce aerial mosaics from the study areas. The land cover of these study areas is studied by visual interpretation, pixel-based supervised classification and object-oriented supervised classification. The change of the forest cover is studied with GIS methods using the visual interpretations from 1955 and 2004. Furthermore, the present state of the study forests is assessed with leaf area index and canopy closure parameters retrieved from hemispherical photographs as well as with additional, previously collected forest health monitoring data. The canopy parameters are also compared with textural parameters from digital aerial mosaics. This study concludes that the classification of forest areas by using true colour data is not an easy task although the digital aerial mosaics are proved to be very accurate. The best classifications are still achieved with visual interpretation methods as the accuracies of the pixel-based and object-oriented supervised classification methods are not satisfying. According to the change detection of the land cover in the study areas, the area of indigenous woodland in both forests has decreased in 1955 2004. However in Ngangao, the overall woodland area has grown mainly because of plantations of exotic species. In general, the land cover of both study areas is more fragmented in 2004 than in 1955. Although the forest area has decreased, forests seem to have a more optimistic future than before. This is due to the increasing appreciation of the forest areas.