Olli-Pekka Tikkanen

Phenological variation as protection against defoliating insects: the case of Quercus robur and Operophtera brumata.

Phenological synchrony between budburst and emergence of larvae is critical for the fitness of many spring-feeding insect herbivores. Therefore, large intraspecific variation in timing of budburst of the host may have a negative effect on the herbivore. We studied how asynchrony between emergence of larvae and budburst affects the fitness of Operophtera brumata (Lepidoptera: Geometridae), a major defoliator of Quercus robur, which can adapt to the phenology of a single tree. It is known that, in maturing leaves of Q. robur, accumulation of condensed tannins has a negative effect on growth of O. brumata. However, there is no information about the effect of hydrolysable tannins and other phenolics that are potential antifeedants. In this study, we also analysed changes in secondary chemistry of the foliage of Q. robur and how different compounds are correlated with growth and survival of O. brumata. The effect of asynchrony on O. brumata was studied in rearing experiments. The neonate larvae were incubated without food for different periods of time. The decline in nutritional quality of foliage was estimated by rearing cohorts of larvae with manipulated hatching times on the leaves of ten individual Q. robur trees. For the chemical analysis, the foliage of these trees was sampled at regular intervals. In the absence of foliage, mortality of neonate larvae started to increase exponentially soon after the larvae emerged. If the larvae missed budburst, the decline in nutritional quality of the foliage led to increased mortality and lower body mass (= fecundity). Hydrolysable tannins were not significantly correlated with performance of the larvae. Only condensed tannins were found to correlate negatively with the growth and survival of O. brumata. Certain individual trees were unsuitable hosts for O. brumata because the decline in quality of the foliage was very rapid. Based on regression equations for increasing rate of mortality and decreasing fecundity, we calculated that a relatively small mismatch of ±30 degree days between budburst and hatching of larvae leads to a 50% decrease in the fitness of O. brumata. Thus, large phenological variation within a Q. robur stand can limit the colonisation of neighbouring trees by dispersing larvae. Furthermore, the hybridisation of moths adapted to phenologically different trees may lead to maladapted phenology of their offspring.

Spatially dynamic forest management to sustain biodiversity and economic returns

Production of marketed commodities and protection of biodiversity in natural systems often conflict and thus the continuously expanding human needs for more goods and benefits from global ecosystems urgently calls for strategies to resolve this conflict. In this paper, we addressed what is the potential of a forest landscape to simultaneously produce habitats for species and economic returns, and how the conflict between habitat availability and timber production varies among taxa. Secondly, we aimed at revealing an optimal combination of management regimes that maximizes habitat availability for given levels of economic returns. We used multi-objective optimization tools to analyze data from a boreal forest landscape consisting of about 30,000 forest stands simulated 50 years into future. We included seven alternative management regimes, spanning from the recommended intensive forest management regime to complete set-aside of stands (protection), and ten different taxa representing a wide variety of habitat associations and social values. Our results demonstrate it is possible to achieve large improvements in habitat availability with little loss in economic returns. In general, providing dead-wood associated species with more habitats tended to be more expensive than providing requirements for other species. No management regime alone maximized habitat availability for the species, and systematic use of any single management regime resulted in considerable reductions in economic returns. Compared with an optimal combination of management regimes, a consistent application of the recommended management regime would result in 5% reduction in economic returns and up to 270% reduction in habitat availability. Thus, for all taxa a combination of management regimes was required to achieve the optimum. Refraining from silvicultural thinnings on a proportion of stands should be considered as a cost-effective management in commercial forests to reconcile the conflict between economic returns and habitat required by species associated with dead-wood. In general, a viable strategy to maintain biodiversity in production landscapes would be to diversify management regimes. Our results emphasize the importance of careful landscape level forest management planning because optimal combinations of management regimes were taxon-specific. For cost-efficiency, the results call for balanced and correctly targeted strategies among habitat types.

Growth and development of a generalist insect herbivore, Operophtera brumata, on original and alternative host plants

Abstract A generalist feeding strategy is common among eruptive insect herbivores but the ultimate reasons for a generalist strategy are not clear. Although generalist insect herbivores are able to complete their life cycle on several species of host plants, there is wide variation in the performance of individuals grown on different hosts. We examined whether different populations of Operophtera brumata are adapted to use the host species which is locally most abundant, and how the host plant affects growth and development of the insect. We reared two allopatric populations (eastern Finland, Prunus padus; south-west Finland, Quercus robur) on four species of host plants (Pr. padus, Populus tremula, Q. robur, Salix phylicifolia) from neonate larvae to the adult stage and measured the growth and development of individuals and the timing of adult hatching. The performance of both populations was best on Pr. padus, and the south-western population, originally on Q. robur, was well adapted to this host. The host affected the growth of females more than that of males. The host plant had an unexpected effect on hatching times of the adults. Individuals grown on the original host hatched in normal synchrony, i.e. males 6–7 days before females; but on alternative hosts this synchrony was disturbed. As is common in eruptive, capital-breeding generalist moths where female fecundity is linked to weight, host quality is critical for the flightless females of O. brumata. We suggest that in a heterogeneous environment the disturbing effect of alternative host plants on adult emergence may decrease the population density and growth rate compared to the potential maximum in a homogeneous environment.

Cost-effective strategies to conserve boreal forest biodiversity and long-term landscape-level maintenance of habitats

Setting aside parcels of land is the main conservation strategy to reduce the rate of biodiversity loss worldwide. Because funding for biological conservation is limited, it is important to distinguish the most efficient ways to use it. Here, we assess implications of alternative measures to conserve biodiversity in managed boreal forest landscapes. We calculated four alternative spatio-temporal scenarios and compared these to the current management regime over 100-year time period. In the alternative scenarios, a fixed amount of funding was invested in (1) permanent large reserves (each tens of ha in size), (2) permanent small reserves (each a few ha in size), (3) temporary small reserves (based on 10-year contracts with private land owners), and (4) green-tree retention (small groups of trees retained on clear-cuts). To assess biodiversity implications, we used habitat suitability indices to calculate overall habitat availability for five groups of red-listed and habitat-specific species associated with decaying spruce logs. The possibilities for timber harvests did not differ among the scenarios, but biodiversity performance was different. The scenarios with permanent reserves tended to outperform other scenarios, suggesting that conservation policies based on permanent reserves are the most cost-efficient in the long term. Results, however, varied among time scales and species groups. In the short term, a strategy of investment in temporary small reserves was the most efficient. Habitat for species associated with old spruce dead-wood and preferring shade was rare throughout all simulations, and therefore, it is likely that these species cannot be sustained in managed forests. Species that live on fresh dead-wood and are associated with forest edges coped well in all scenarios suggesting that such species will persist in managed landscapes without additional conservation efforts. Explicit definition of conservation objectives and time frames for conservation action are thus prerequisites for successful conservation planning.

Bark salicylates and condensed tannins reduce vole browsing amongst cultivated dark-leaved willows (Salix myrsinifolia)

Summary.Vole feeding amongst herbal willows that have a high concentration of salicylates in their bark and leaves, and may therefore be cultivated for use as raw material for herbal medicine was tested in the field and in laboratory conditions. Eight clones of dark-leaved willow (Salix myrsinifolia Salisb.) were cultivated for two years with six different methods combining three fertilisation levels (none, low and high), black plastic mulch applied for suppressing weed competition and unmulched control. Samples for the laboratory feeding trial were taken from the unfertilised plants during willow winter dormancy and twigs were fed to 16 voles as a multi-choice experiment. The bark area removed was calculated from image analysis of the material left by the voles. The diameter and the bark thickness of the twigs were measured. Concentrations of salicin, salicortin, HCH-salicortin, picein, triandrin, triandrin derivative, gallocatechin, (+)-catechin, luteolin-7-glucoside, hyperin, total condensed tannins and total nitrogen were measured from the twigs fed to voles in the laboratory. Browsing by a natural population of voles amongst winter-dormant willows was measured in the field. In the laboratory, voles browsed on 80% of the twigs and in the field voles browsed on 33% of the twigs. Vole feeding followed similar patterns in the field and in the laboratory experiment; feeding was clearly higher amongst the plants grown in unmulched control compared to those in plastic mulch. The same clones, 1, 2 and 6 were preferred in both experiments. Voles preferred thin twigs to thick ones. Feeding correlated negatively with concentrations of salicylates and tannins. As vole feeding seems to be highly affected by willow cultivation method and plant genotype, careful selection of cultivated clones and cultivation methods can enhance the reliability of herbal willow cultivation.

Use of host plants by Operopthera brumata L. (Lep., Geometridae) during the first recorded outbreak in the subcontinental boreal zone of Fennoscandia.

Abstract: The first known outbreak of Operopthera brumata in the subcontinental boreal zone was found in 1994. In the summer of 1995 use of the host plants by O. brumata was studied during the outbreak. Eight transects were studied from outbreak epicentres to the undefoliated perimeter. Along the transects the defoliation level of each tree and shrub species was estimated. The epicentres were in deciduous tree stands in early secondary succession on abandoned pasture land. The defoliation of different species of deciduous trees growing the area differed significantly. Of the dominant canopy species of the epicentres, Prunus padus and Sorbus aucuparia were the most defoliated, while Alnus incana and Betula pubescens had a medium level of defoliation. The average level of defoliation was 90% for P. padus and S. aucupara, and about 25% for A. incana and B. pubescens. Shrub species such as Rhamnus frangula, Ribes spp. and Sambucus racemosa had very low levels of feeding. In the study area, Quercus robur, normally the main hostplant of O. brumata, was rare; but Prunus padus, which has a similar growing strategy, was very abundant in the outbreak epicentres but absent around the perimeters. In general, P. padus was the most defoliated tree species in the study area. We suggest that P. padus is the main host plant of O. brumata in SE Fennoscandia, and only in those areas where it is abundant are O. brumata populations able to increase to the outbreak level.

No negative correlation between growth and resistance to multiple herbivory in a deciduous tree, Betula pendula

Abstract Plants are assumed to have a trade-off between growth and resistance. This trade-off has been demonstrated in herbaceous plants, but the information on woody plants is conflicting. In contrast to simple annual plant–herbivore systems, trees face highly variable herbivory. In this note, we measured clonal variation in the growth of Betula pendula and in its resistance to nine major herbivore species belonging to five different feeding guilds. There were significant differences in both the growth and the resistance of clones to all the herbivores studied. However, we found no negative correlation between growth and resistance. No clone was generally resistant or susceptible to most of the herbivores. This suggests that, if there is an ultimate trade-off in resource allocation of B. pendula between growth and resistance, it is masked by other factors, for example, by several opposite abiotic and biotic selection pressures.

Spatial pattern of outbreaks of Operophtera brumata in eastern Fennoscandia and their effects on radial growth of trees

Abstract During the mid 1990s, Operophtera brumata outbreaks were reported from different locations in eastern Fennoscandia. In this paper, we analysed the outbreak history of 13 separate 24–47-year-old stands of Prunus padus , the main host of this herbivore in eastern Fennoscandia, based on width of annual growth rings. The stands were located on both sides of the Finnish and Russian border. We also studied the effect of an insect outbreak on radial growth of several dominant tree species in mixed forests. It has been suggested that less-preferred tree species in the mixed stands may benefit from an insect outbreak. Heavily infested tree species sustained a large reduction in radial growth. However, we did not find that less-preferred tree species could directly benefit from an the outbreak, to a degree that is measurable in radial growth. There were no signs of previous outbreaks of O. brumata , and the recent epidemic was almost totally confined to the Russian side. We suggest that a decline in the area of farmland after the 1950s led to development of an extraordinarily large amount of high-quality habitats on the Russian side which, together with a period of mild winters in the early 1990s, created the conditions for this novel O. brumata epidemic.

Factors controlling the evaporation of secondary organic aerosol from α‐pinene ozonolysis

Abstract Secondary organic aerosols (SOA) forms a major fraction of organic aerosols in the atmosphere. Knowledge of SOA properties that affect their dynamics in the atmosphere is needed for improving climate models. By combining experimental and modeling techniques, we investigated the factors controlling SOA evaporation under different humidity conditions. Our experiments support the conclusion of particle phase diffusivity limiting the evaporation under dry conditions. Viscosity of particles at dry conditions was estimated to increase several orders of magnitude during evaporation, up to 109 Pa s. However, at atmospherically relevant relative humidity and time scales, our results show that diffusion limitations may have a minor effect on evaporation of the studied α‐pinene SOA particles. Based on previous studies and our model simulations, we suggest that, in warm environments dominated by biogenic emissions, the major uncertainty in models describing the SOA particle evaporation is related to the volatility of SOA constituents.

Applying a framework for landscape planning under climate change for the conservation of biodiversity in the Finnish boreal forest

Conservation strategies are often established without consideration of the impact of climate change. However, this impact is expected to threaten species and ecosystem persistence and to have dramatic effects towards the end of the 21st century. Landscape suitability for species under climate change is determined by several interacting factors including dispersal and human land use. Designing effective conservation strategies at regional scales to improve landscape suitability requires measuring the vulnerabilities of specific regions to climate change and determining their conservation capacities. Although methods for defining vulnerability categories are available, methods for doing this in a systematic, cost-effective way have not been identified. Here, we use an ecosystem model to define the potential resilience of the Finnish forest landscape by relating its current conservation capacity to its vulnerability to climate change. In applying this framework, we take into account the responses to climate change of a broad range of red-listed species with different niche requirements. This framework allowed us to identify four categories in which representation in the landscape varies among three IPCC emission scenarios (B1, low; A1B, intermediate; A2, high emissions): (i) susceptible (B1 = 24.7%, A1B = 26.4%, A2 = 26.2%), the most intact forest landscapes vulnerable to climate change, requiring management for heterogeneity and resilience; (ii) resilient (B1 = 2.2%, A1B = 0.5%, A2 = 0.6%), intact areas with low vulnerability that represent potential climate refugia and require conservation capacity maintenance; (iii) resistant (B1 = 6.7%, A1B = 0.8%, A2 = 1.1%), landscapes with low current conservation capacity and low vulnerability that are suitable for restoration projects; (iv) sensitive (B1 = 66.4%, A1B = 72.3%, A2 = 72.0%), low conservation capacity landscapes that are vulnerable and for which alternative conservation measures are required depending on the intensity of climate change. Our results indicate that the Finnish landscape is likely to be dominated by a very high proportion of sensitive and susceptible forest patches, thereby increasing uncertainty for landscape managers in the choice of conservation strategies.