Increased summer temperature is associated with reduced calf mass of a circumpolar large mammal through direct thermoregulatory and indirect, food quality, pathways.

Climate change represents a growing ecological challenge. The (sub) arctic and boreal regions of the world experience the most rapid warming, presenting an excellent model system for studying how climate change affects mammals. Moose (Alces alces) are a particularly relevant model species with their circumpolar range. Population declines across the southern edge of this range are linked to rising temperatures. Using a long-term dataset (1988-1997, 2017-2019), we examine the relative strength of direct (thermoregulatory costs) and indirect (food quality) pathways linking temperature, precipitation, and the quality of two important food items (birch and fireweed) to variation in moose calf mass in northern Sweden. The direct effects of temperature consistently showed stronger relationships to moose calf mass than did the indirect effects. The proportion of growing season days where the temperature exceeded a 20 °C threshold showed stronger direct negative relationships to moose calf mass than did mean temperature values. Finally, while annual forb (fireweed) quality was more strongly influenced by temperature and precipitation than were perennial (birch) leaves, this did not translate into a stronger relationship to moose calf weight. The only indirect path with supporting evidence suggested that mean growing season temperatures were positively associated with neutral detergent fiber, which was, in turn, negatively associated with calf mass. While indirect impacts of climate change deserve further investigation, it is important to recognize the large direct impacts of temperature on cold-adapted species.

Simulated moose (Alces alces L.) browsing increases accumulation of secondary metabolites in bilberry (Vaccinium myrtillus L.) along gradients of habitat productivity and solar radiation.

We have addressed the impact of moose (Alces alces L.) on accumulation of secondary metabolites, lignin, and nitrogen in bilberry (Vaccinium myrtillus L.) along gradients of habitat productivity and solar radiation. The study was conducted within a long-term research project on direct and indirect impacts of moose on the ecosystem. In the experiment, browsing, defecation, and urination corresponding to four different moose densities were simulated for eight years before bilberry tissue was collected and analyzed. Some quantitatively dominant flavonoids were affected by the simulated moose browsing and by habitat productivity and light. The content of flavonoids increased with increasing moose density and light, and decreased with increasing habitat productivity. The higher concentration of secondary metabolites in bilberry from nutrient-poor sites may have resulted from the increased photosynthesis relative to growth, which facilitated secondary metabolism. The higher concentration of secondary metabolites in plants subjected to simulated moose- herbivory might have been caused in part by loss of biomass. In addition, in areas with high biomass loss, i.e., high moose density, a more open canopy was created and more solar radiation could have induced secondary metabolism.

Micro and macro-habitat associations in saproxylic beetles: implications for biodiversity management.

Restoration of habitats is critically important in preventing full realization of the extinction debt owed as a result of anthropogenic habitat destruction. Although much emphasis has been placed on macrohabitats, suitable microhabitats are also vital for the survival of most species. The aim of this large-scale field experiment was to evaluate the relative importance of manipulated microhabitats, i.e., dead wood substrates of spruce (snags, and logs that were burned, inoculated with wood fungi or shaded) and macrohabitats, i.e., stand types (clear-cuts, mature managed forests, and forest reserves) for species richness, abundance and assemblage composition of all saproxylic and red-listed saproxylic beetles. Beetles were collected in emergence traps in 30 forest stands in 2001, 2003, 2004 and 2006. More individuals emerged from snags and untreated logs than from burned and shaded logs, but species richness did not differ among substrates. Assemblage composition differed among substrates for both all saproxylics and red-listed saproxylic species, mainly attributed to different assemblage composition on snags. This suggests that the practise of leaving snags for conservation purposes should be complemented with log supplementation. Clear-cuts supported fewer species and different assemblages from mature managed forests and reserves. Neither abundance, nor species richness or assemblage composition differed between reserves and mature managed forests. This suggests that managed stands subjected to selective cutting, not clear-felling, maintain sufficient old growth characteristics and continuity to maintain more or less intact assemblages of saproxylic beetles. Thus, alternative management methods, e.g., continuity forestry should be considered for some of these stands to maintain continuity and conservation values. Furthermore, the significantly higher estimated abundance per ha of red-listed beetles in reserves underlines the importance of reserves for maintaining viable populations of rare red-listed species and as source areas for saproxylic species in boreal forest landscapes.

Browsed twig environmental DNA: diagnostic PCR to identify ungulate species.

Ungulate browsing can have a strong effect on ecological processes by affecting plant community structure and composition, with cascading effects on nutrient cycling and animal communities. However, in the absence of direct observations of foraging, species-specific foraging behaviours are difficult to quantify. We therefore know relatively little about foraging competition and species-specific browsing patterns in systems with several browsers. However, during browsing, a small amount of saliva containing buccal cells is deposited at the bite site, providing a source of environmental DNA (eDNA) that can be used for species identification. Here, we describe extraction and PCR protocols for a browser species diagnostic kit. Species-specific primers for mitochondrial DNA were optimized and validated using twigs browsed by captive animals. A time series showed that about 50% of the samples will amplify up to 12 weeks after the browsing event and that some samples amplify up to 24 weeks after browsing (12.5%). Applied to samples of natural browsing from an area where moose (Alces alces), roe deer (Capreolus capreolus), fallow deer (Cervus dama) and red deer (Cervus elaphus) are sympatric, amplification success reached 75%. This method promises to greatly improve our understanding of multispecies browsing systems without the need for direct observations.

Vole preference of bilberry along gradients of simulated moose density and site productivity.

Browsing by large herbivores might either increase or decrease preference for the plant by other herbivores, depending on the plant response. Using a cafeteria test, we studied the preference by root voles (Microtus oeconomus [Pallas, 1776]) for bilberry (Vaccinium myrtillus L.) previously subjected to 4 levels of simulated moose (Alces alces [Linnaeus, 1758]) density. The different levels of moose density were simulated at population densities relevant for Fennoscandian conditions, in exclosures situated along a site productivity gradient. We expected: (i) voles to prefer bilberry from high productivity sites over low productivity sites; (ii) voles to prefer browsed bilberry, if plants allocate resources to compensatory growth or to avoid browsed bilberry if plants allocate resources to defense; (iii) these effects to increase with increasing simulated moose density; and (iv) the concentration of plant chemicals and the plant morphology to explain vole preference. Specifically, we predicted that voles would prefer: (i) plants with high nitrogen content; (ii) plants with low content of defensive substances; and (iii) tall plants with long shoots. Voles preferred bilberry from the high productivity sites compared to the low productivity sites. We also found an interaction between site productivity and simulated moose density, where voles preferred unbrowsed plants at low productivity sites and intermediate levels of browsing at high productivity sites. There was no effect of plant chemistry or morphology on vole preference. We conclude that moose browsing impacts the food preference of voles. With the current high densities of moose in Fennoscandia, this could potentially influence vole food selection and population dynamics over large geographical areas.

Reconnecting to the biosphere.

Humanity has emerged as a major force in the operation of the biosphere, with a significant imprint on the Earth System, challenging social-ecological resilience. This new situation calls for a fundamental shift in perspectives, world views, and institutions. Human development and progress must be reconnected to the capacity of the biosphere and essential ecosystem services to be sustained. Governance challenges include a highly interconnected and faster world, cascading social-ecological interactions and planetary boundaries that create vulnerabilities but also opportunities for social-ecological change and transformation. Tipping points and thresholds highlight the importance of understanding and managing resilience. New modes of flexible governance are emerging. A central challenge is to reconnect these efforts to the changing preconditions for societal development as active stewards of the Earth System. We suggest that the Millennium Development Goals need to be reframed in such a planetary stewardship context combined with a call for a new social contract on global sustainability. The ongoing mind shift in human relations with Earth and its boundaries provides exciting opportunities for societal development in collaboration with the biosphere--a global sustainability agenda for humanity.

Depression of belowground respiration rates at simulated high moose population densities in boreal forests.

Large herbivores can affect the carbon cycle in boreal forests by changing productivity and plant species composition, which in turn could ultimately alter litter production, nutrient cycling, and the partitioning between aboveground and belowground allocation of carbon. Here we experimentally tested how moose (Alces alces) at different simulated population densities affected belowground respiration rates (estimated as CO2 flux) in young boreal forest stands situated along a site productivity gradient. At high simulated population density, moose browsing considerably depressed belowground respiration rates (24-56% below that of no-moose controls) except during June, where the difference only was 10%. Moose browsing depressed belowground respiration the most on low-productivity sites. Soil moisture and temperature did not affect respiration rates. Impact of moose on belowground respiration was closely linked to litter production and followed Michaelis-Menten dynamics. The main mechanism by which moose decrease belowground respiration rates is likely their effect on photosynthetic biomass (especially decreased productivity of deciduous trees) and total litter production. An increased productivity of deciduous trees along the site productivity gradient causes an unequal effect of moose along the same gradient. The rapid growth of deciduous trees may offer higher resilience against negative effects of moose browsing on litter production and photosynthate allocation to roots.

Managing climate change impacts to enhance the resilience and sustainability of Fennoscandian forests.

Projected warming in Sweden and other Fennoscandian countries will probably increase growth rates of forest trees near their northern limits, increase the probability of new pest outbreaks, and foster northerly migration of both native and exotic species. The greatest challenges for sustainable forestry are to restore and enhance the ecological and socioeconomic diversity of intensively managed forested landscapes. With appropriate management, climate warming may facilitate the regeneration of this diversity. Experimental transplant gardens along latitudinal or altitudinal gradients and high-resolution maps of expected future climate could provide a scientific basis for predicting the climate response of potential migrant species. Management of corridors and assisted migration could speed the movement of appropriate species.

Building resilience and adaptation to manage Arctic change.

Unprecedented global changes caused by human actions challenge society's ability to sustain the desirable features of our planet. This requires proactive management of change to foster both resilience (sustaining those attributes that are important to society in the face of change) and adaptation (developing new socioecological configurations that function effectively under new conditions). The Arctic may be one of the last remaining opportunities to plan for change in a spatially extensive region where many of the ancestral ecological and social processes and feedbacks are still intact. If the feasibility of this strategy can be demonstrated in the Arctic, our improved understanding of the dynamics of change can be applied to regions with greater human modification. Conditions may now be ideal to implement policies to manage Arctic change because recent studies provide the essential scientific understanding, appropriate international institutions are in place, and Arctic nations have the wealth to institute necessary changes, if they choose to do so.

Geographic variations in anthropogenic drivers that influence the vulnerability and resilience of social-ecological systems.

Across the circumpolar North large disparities in the distribution of renewable and nonrenewable resources, human population density, capital investments, and basic residential and transportation infrastructure combine to create recognizable hotspots of recent and foreseeable change. Northern Fennoscandia exemplifies a relatively benign situation due to its current economic and political stability. Northern Russia is experiencing rapid, mostly negative changes reflecting the general state of crisis since the collapse of the Soviet Union. North America enjoys a relatively stable regulatory structure to mitigate environmental degradation associated with industry, but is on the verge of approving massive new development schemes that would significantly expand the spatial extent of potentially affected social-ecological systems. Institutional or regulatory context influences the extent to which ecosystem services are buffered against environmental change. With or without a warming climate, certain geographic areas appear especially vulnerable to damages that may threaten their ability to supply goods and services in the near future. Climate change may exacerbate this situation in some places but may offer opportunities to enhance resilience in the long term.

Cycles and synchrony in the Collared Lemming (Dicrostonyx groenlandicus) in Arctic North America.

Lemming populations are generally characterised by their cyclic nature, yet empirical data to support this are lacking for most species, largely because of the time and expense necessary to collect long-term population data. In this study we use the relative frequency of yearly willow scarring by lemmings as an index of lemming abundance, allowing us to plot population changes over a 34-year period. Scars were collected from 18 sites in Arctic North America separated by 2-1,647 km to investigate local synchrony among separate populations. Over the period studied, populations at all 18 sites showed large fluctuations but there was no regular periodicity to the patterns of population change. Over all possible combinations of pairs of sites, only sites that were geographically connected and close (<6 km) showed significant synchrony in fluctuations. The populations studied may not even be cyclic, at least for the time period 1960 to 1994, and although fluctuating, randomisation tests could not reject the null hypothesis of random fluctuations. These data have implications for the testing of hypotheses regarding lemming cycles and highlight the need for long-term trapping data to characterise the lemming cycle.

Asynchronous population dynamics of Siberian lemmings across the Palaearctic tundra.

The synchrony of Siberian lemming (Lemmus sibiricus L.) population dynamics was investigated during a ship-borne expedition along the Palaearctic tundra coast in the summer of 1994. On 12 sites along the coast from the Kola Peninsula to Wrangel Island, relative densities of lemmings were recorded using a standardised snap-trapping programme. The phase position of the lemming cycle in each of the studied populations was determined based on current density estimates, signs of previous density and the age profile of each population (ageing based on eye lens mass). In addition, dendrochronological methods were used to determine when the last peak in the density of microtine populations occurred at each site. The examined lemming populations were in different phases of the lemming cycle. Some populations were in the peak phase, as indicated by high current densities, an age profile in which older individuals were well represented, and signs of high previous density (abundant old lemming faeces). Other populations were in the decline phase, as reflected in a moderate current density, a predominance of older individuals and signs of high previous density. Populations in the low phase had an extremely low current density and showed signs of high previous density, while populations in the increase phase had a moderate current density, a predominance of younger individuals and showed signs of low previous density. The results of phase determinations based on dendrochronological methods support the findings based on lemming demography. Recent Russian studies carried out on some of the sites also agreed with our phase determination results. Thus, on a regional scale (across the whole Palaearctic tundra), the population dynamics of Siberian lemmings can be considered asynchronous. However, sites situated adjacent to each other were often phase synchronous, suggesting a more fine-grained pattern of dynamics with synchrony over distances as long as 1000 km or so, e.g. the Yamal and Taymyr Peninsulas.

Geographical and latitudinal variation in growth patterns and adult body size of Swedish moose (Alces alces).

We examined the geographical pattern in growth and adult body size among 14 populations of Swedish moose (Alces alces) using data from 4,294 moose (≥1.5 years old) killed during the hunting season in 1989-1992. In both sexes, adult body mass was significantly positively correlated with latitude. Moose in northern populations had a 15-20% larger adult body mass than moose in the south. Juvenile body mass was correlated with neither latitude nor adult body mass. Thus, variation in time (years) and rate of body growth after the juvenile stage were responsible for most of the variation in adult body mass among populations. Moose in northern populations grew for approximately 2 more years of life than southern moose. In contrast to adult body mass, skeletal size (measured as jawbone length) was not correlated with latitude, suggesting that variation in adult body mass was primarily due to differences in fat reserves. Discrimination between population characteristics, such as moose density, climate, and the amount of browse available to moose, showed climatic harshness to be the most important variable explaining geographical variation in body mass among populations. The results support the notion that in mammals body size increases with latitude in accordance with Bergmann's rule. We conclude that (1) variation in patterns of growth after the juvenile stage is the main cause of the latitudinal trend in adult body size in moose, and (2) climatic conditions are a more important factor than population density and availability of food in explaining geographical variation in growth patterns and adult body mass between populations of Swedish moose.

Flowering, Shoot Production, and Vole Bark Herbivory in a Boreal Willow.

Different intensities of vole bark herbivory were simulated on the boreal willow Salix myrsinifolia-phylicifolia in field experiments, and plant responses were followed during 4 yr. Intermediate damage (every second stem barked) resulted the 1st yr in a fourfold increase in the production of basal shoots per individual, and in the following years, a twofold increase in the number of catkins per intact stem compared with control willows. However, 1 yr after damage, the average number of seeds per individual was 21% lower than in control willows and seed germination was reduced 40%. Severe damage (all stems barked) caused a nearly 10-fold increase in the production of basal shoots the 1st yr and a reduction and delay of sexual reproduction. Mortality of damaged stems was high (100%), while genet mortality was < 5% in the group in which all stems were barked. Flowering in female willows was reduced more than in male willows, and males produced catkins on younger basal shoots than females. We suggest that in this system, where the principal herbivore fluctuates between years, compensatory reactions by damaged plants might be of particular benefit, as the period of growth following damage is more or less free from herbivores.

Reduction of aquatic vegetation following the colonization of a Northern Swedish Lake by the muskrat, Ondatra zibethica.

The muskrat (Ondatra zibethica (L.)) utilizes emergent hydrophytes both for housebuilding and for food. The impact of these activities on stands of Equisetum fluviatile L. in a northern Swedish lake was investigated. The areas of open water created by the muskrats were estimated from infrared air photographs, taken in July, on three occasions during a five-year period. During the first three years, with low muskrat densities, the areas of open water were estimated to represent removal of about one percent of the Equisetum stands. In the summer following a population peak the total area of open water had increased to about four percent. In that summer there were, on average, about 160 such areas per hectare and their combined perimeter length was about 1.5 km. The many small areas created by the muskrats were mostly very irregular in shape, each with a long perimeter in relation to its area. If not too numerous, the muskrats have a positive effect on the waterfowl population in well-vegetated lakes, by increasing the feeding areas available for both adults and young birds.