Long-term changes in herbivore community and vegetation impact of wild and domestic herbivores across Iceland.

Changes in wild and domestic herbivore populations significantly impact extensive grazing systems, particularly in low productive environments, where increasing wild herbivore populations are perceived as a threat to farming. To assess the magnitude of these changes in Iceland, we compiled time series on herbivore populations from 1986 to 2020 and estimated changes in species densities, metabolic biomass, and consumption of plant biomass in improved lands and unimproved rangelands. We compared estimates of consumption rates to past and present net primary production. Overall, the herbivore community composition shifted from livestock to wildlife dominated. However, wild herbivores only contributed a small fraction (14%) of the total herbivore metabolic biomass and consumption (4-7%), and livestock dominated the overall herbivore biomass. These insights highlight the necessity of developing improved local integrated management for both wild and domestic herbivores where they coexist.

A regionally coherent ecological fingerprint of climate change, evidenced from natural history collections.

Climate change has dramatic impacts on ecological systems, affecting a range of ecological factors including phenology, species abundance, diversity, and distribution. The breadth of climate change impacts on ecological systems leads to the occurrence of fingerprints of climate change. However, climate fingerprints are usually identified across broad geographical scales and are potentially influenced by publication biases. In this study, we used natural history collections spanning over 250 years, to quantify a range of ecological responses to climate change, including phenology, abundance, diversity, and distributions, across a range of taxa, including vertebrates, invertebrates, plants, and fungi, within a single region, Central Norway. We tested the hypotheses that ecological responses to climate change are apparent and coherent at a regional scale, that longer time series show stronger trends over time and in relation to temperature, and that ecological responses change in trajectory at the same time as shifts in temperature. We identified a clear regional coherence in climate signal, with decreasing abundances of limnic zooplankton (on average by 7691 individuals m-3 °C-1) and boreal forest breeding birds (on average by 1.94 territories km-2 °C-1), and earlier plant flowering phenology (on average 2 days °C-1) for every degree of temperature increase. In contrast, regional-scale species distributions and species diversity were largely stable. Surprisingly, the effect size of ecological response did not increase with study duration, and shifts in responses did not occur at the same time as shifts in temperature. This may be as the long-term studies include both periods of warming and temperature stability, and that ecological responses lag behind warming. Our findings demonstrate a regional climate fingerprint across a long timescale. We contend that natural history collections provide a unique window on a broad spectrum of ecological responses at timescales beyond most ecological monitoring programs. Natural history collections are thus an essential source for long-term ecological research.

Legacy effects of herbivory on treeline dynamics along an elevational gradient.

Treelines are expected to expand into alpine ecosystems with global warming, but herbivory may delay this expansion. This study quantifies long-term effects of temporally varying sheep densities on birch recruitment and growth in the treeline ecotone. We examined treeline ecotone successional trajectories and legacy effects in a replicated experimental setup, where enclosures were present for 14 years with three different sheep densities (0, 25, 80 sheep km-2). Before and after the enclosures were present, the site had an ambient sheep density of 20-25 km-2. We sampled field data 4 years after enclosure removal and compared these to data sampled 8 and 9 years after enclosure erection. We sampled data on birch browsing pressure, birch distribution across life-stages (recruits, saplings, and mature trees), and birch annual radial growth. Fourteen years of increased or decreased sheep density had observable legacy effects depending on birch life-stage. Birch recruit prevalence decreased in areas, where sheep were reintroduced after being absent for 14 years. For the same areas, sapling and mature tree prevalence increased, indicating that these areas have entered alternative successional trajectories compared to areas, where sheep were present the whole time. Birch annual radial growth showed a lag effect of 2 years after enclosure removal, with growth decreasing in areas where sheep had been absent for 14 years and increasing where sheep densities were high. Thus, decadal-scale absences of herbivores can leave legacy effects due to increased numbers of trees that have high resistance to later-introduced herbivore browsing.

Impacts of roads on bird species richness: A meta-analysis considering road types, habitats and feeding guilds.

Roadsides can harbour remarkable biodiversity; thus, they are increasingly considered as habitats with potential for conservation value. To improve construction and management of roadside habitats with positive effects on biodiversity, we require a quantitative understanding of important influential factors that drive both positive and negative effects of roads. We conducted meta-analyses to assess road effects on bird communities. We specifically tested how the relationship between roads and bird richness varies when considering road type, habitat characteristics and feeding guild association. Overall, bird richness was similar in road habitats compared to non-road habitats, however, the two apparently differ in species composition. Bird richness was lowered by road presence in areas with denser tree cover but did not differ according to road type. Richness differences between habitats with and without roads further depended on primary diet of species, and richness of omnivores was positively affected by road presence. We conclude that impacts of roads on bird richness are highly context-dependent, and planners should carefully evaluate road habitats on a case by case basis. This emphasizes the need for further studies that explicitly test for differences in species composition and abundance, to disentangle contexts where a road will negatively affect bird communities, and where it will not.

Will borealization of Arctic tundra herbivore communities be driven by climate warming or vegetation change?

Poleward shifts in species distributions are expected and frequently observed with a warming climate. In Arctic ecosystems, the strong warming trends are associated with increasing greenness and shrubification. Vertebrate herbivores have the potential to limit greening and shrub advance and expansion on the tundra, posing the question of whether changes in herbivore communities could partly mediate the impacts of climate warming on Arctic tundra. Therefore, future changes in the herbivore community in the Arctic tundra will depend on whether the community tracks the changing climates directly (i.e. occurs in response to temperature) or indirectly, in response to vegetation changes (which can be modified by trophic interactions). In this study, we used biogeographic and remotely sensed data to quantify spatial variation in vertebrate herbivore communities across the boreal forest and Arctic tundra biomes. We then tested whether present-day herbivore community structure is determined primarily by temperature or vegetation. We demonstrate that vertebrate herbivore communities are significantly more diverse in the boreal forest than in the Arctic tundra in terms of species richness, phylogenetic diversity and functional diversity. A clear shift in community structure was observed at the biome boundary, with stronger northward declines in diversity in the Arctic tundra. Interestingly, important functional traits characterizing the role of herbivores in limiting tundra vegetation change, such as body mass and woody plant feeding, did not show threshold changes across the biome boundary. Temperature was a more important determinant of herbivore community structure across these biomes than vegetation productivity or woody plant cover. Thus, our study does not support the premise that herbivore-driven limitation of Arctic tundra shrubification or greening would limit herbivore community change in the tundra. Instead, borealization of tundra herbivore communities is likely to result from the direct effect of climate warming.

Urban aliens and threatened near-naturals: Land-cover affects the species richness of alien- and threatened species in an urban-rural setting.

Urbanisation has strong effects on biodiversity patterns, but impacts vary among species groups and across spatial scales. From a local biodiversity management perspective, a more general understanding of species richness across taxonomic groups is required. This study aims to investigate how fine-scale land-cover variables influence species richness patterns of locally threatened and alien species. The study was performed in Trondheim, Norway, covering a steep urbanisation gradient. Spatially correlated Generalised Linear Mixed Effects Models predicting the number of all-, threatened-and alien species by taxon, habitat, habitat heterogeneity and mean aspect within 500 m×500 m grid cells were constructed. The habitat categories were based on detailed land-cover maps. The highest number of threatened species was found in habitats relatively less affected by humans, whereas the number of alien species were only dependent on taxonomic group and spatial correlation. It is shown that land-cover variables within an administrative border can be used to make predictions on species richness within overarching species groups. Recommendations to biodiversity management agencies are to ensure protection of natural habitats to favour locally threatened species, and closely monitor urban areas to mitigate the introduction and spread of alien species.

Herbivory and climate as drivers of woody plant growth: Do deer decrease the impacts of warming?

Vegetation at ecotone transitions between open and forested areas is often heavily affected by two key processes: climate change and management of large herbivore densities. These both drive woody plant state shifts, determining the location and the nature of the limit between open and tree or shrub-dominated landscapes. In order to adapt management to prevailing and future climate, we need to understand how browsing and climatic factors together affect the growth of plants at biome borders. To disentangle herbivory and climate effects, we combined long-term tree growth monitoring and dendroecology to investigate woody plant growth under different temperatures and red deer (Cervus elaphus) herbivory pressures at forest-moorland ecotones in the Scottish highlands. Reforestation and deer densities are core and conflicting management concerns in the area, and there is an urgent need for additional knowledge. We found that deer herbivory and climate had significant and interactive effects on tree growth: in the presence of red deer, pine (Pinus sylvestris) growth responded more strongly to annual temperature than in the absence of deer, possibly reflecting differing plant-plant competition and facilitation conditions. As expected, pine growth was negatively related to deer density and positively to temperature. However, at the tree population level, warming decreased growth when more than 60% of shoots were browsed. Heather (Calluna vulgaris) growth was negatively related to temperature and the direction of the response to deer switched from negative to positive when mean annual temperatures fell below 6.0°C. In addition, our models allow estimates to be made of how woody plant growth responds under specific combinations of temperature and herbivory, and show how deer management can be adapted to predicted climatic changes in order to more effectively achieve reforestation goals. Our results support the hypothesis that temperature and herbivory have interactive effects on woody plant growth, and thus accounting for just one of these two factors is insufficient for understanding plant growth mechanics at biome transitions. Furthermore, we show that climate-driven woody plant growth increases can be negated by herbivory.

Cool as a moose: How can browsing counteract climate warming effects across boreal forest ecosystems?

Herbivory has potential to modify vegetation responses to climatic changes. However, climate and herbivory also affect each other, and rarely work in isolation from other ecological factors, such as plant-plant competition. Thus, it is challenging to predict the extent to which herbivory can counteract, amplify, or interact with climate impacts on ecosystems. Here, we investigate how moose modify climatic responses of boreal trees by using experimental exclosures on two continents and modeling complex causal pathways including several climatic factors, multiple tree species, competition, tree height, time, food availability, and herbivore presence, density, and browsing intensity. We show that moose can counteract, that is, "cool down" positive temperature responses of trees, but that this effect varies between species depending on moose foraging preferences. Growth of preferred deciduous trees was strongly affected by moose, whereas growth of less preferred conifers was mostly driven by climate and tree height. In addition, moose changed temperature responses of rowan in Norway and balsam fir in Canada, by making fir more responsive to temperature but decreasing the strength of the temperature response of rowan. Snow protected trees from browsing, and therefore moose "cooling power" might increase should a warming climate result in decreased snow cover. Furthermore, we found evidence of indirect effects of moose via plant-plant competition: By constraining growth of competing trees, moose can contribute positively to the growth of other trees. Our study shows that in boreal forests, herbivory cooling power is highly context dependent, and in order to understand its potential to prevent changes induced by warming climate, species differences, snow, competition, and climate effects on browsing need to be considered.

Long-term changes in northern large-herbivore communities reveal differential rewilding rates in space and time.

Herbivores have important impacts on ecological and ecosystem dynamics. Population density and species composition are both important determinants of these impacts. Large herbivore communities are shifting in many parts of the world driven by changes in livestock management and exploitation of wild populations. In this study, we analyse changes in large herbivore community structure over 66 years in Norway, with a focus on the contribution of wildlife and livestock. We calculate metabolic biomass of all large-herbivore species across the whole region between 1949 and 2015. Temporal and spatial patterns in herbivore community change are investigated and we test hypotheses that changes in wildlife biomass are driven by competition with livestock. We find that total herbivore biomass decreased from 1949 to a minimum in 1969 due to decreases in livestock biomass. Increasing wild herbivore populations lead to an increase in total herbivore biomass by 2009. Herbivore communities have thus reverted from a livestock dominated state in 1949 (2% of large herbivore metabolic biomass comprised of wildlife species) to a state with roughly equal wildlife and livestock (48% of metabolic biomass comprised of wildlife species). Declines in livestock biomass were a modest predictor of wildlife increases, suggesting that competition with livestock has not been a major limiting factor of wild herbivore populations over the past decades. Instead there was strong geographic variation in herbivore community change, with milder lowland regions becoming more dominated by wild species, but colder mountain and northern regions remaining dominated by livestock. Our findings indicate that there has been notable rewilding of herbivore communities and herbivore-ecosystem interactions in Norway, particularly in milder lowland regions. However, Norwegian herbivores remain mostly regulated by management, and our findings call for integrated management of wild and domestic herbivores.

Can mowing restore boreal rich-fen vegetation in the face of climate change?

Low-frequency mowing has been proposed to be an effective strategy for the restoration and management of boreal fens after abandonment of traditional haymaking. This study investigates how mowing affects long-term vegetation change in both oceanic and continental boreal rich-fen vegetation. This will allow evaluation of the effectiveness of mowing as a management and restoration tool in this ecosystem in the face of climate change. At two nature reserves in Central Norway (Tågdalen, 63° 03' N, 9° 05 E, oceanic climate and Sølendet, 62° 40' N, 11° 50' E, continental climate), we used permanent plot data from the two sites to compare plant species composition from the late 1960s to the early 1980s with that recorded in 2012-2015 in abandoned and mown fens. Changes in species composition and frequency were analysed by multivariate and univariate methods in relation to environmental variables and modelled climate and groundwater data. Mowing resulted in a decline in shrub and Molinia caerulea cover at the continental and oceanic sites respectively, and the total cover of specialist fen species had increased to a significantly greater extent in the mown plots than the unmown at the continental site. However, mowing did not have an effect on the cover of specialist bryophyte species, and some specialist species declined regardless of mowing treatment. Temperature sums had increased at both sites, but precipitation had not changed significantly. Mowing was shown to be the most important determinant of plant community composition at both sites, with local environmental conditions being of secondary importance. In conclusion, the abandonment of traditional management practices results in the loss of characteristic fen species. In order to encourage the restoration of typical rich-fen vegetation, particularly in oceanic areas, additional management measures, such as more intensive mowing, may be required.

No genetic erosion after five generations for Impatiens glandulifera populations across the invaded range in Europe.

BACKGROUND: The observation that many alien species become invasive despite low genetic diversity has long been considered the 'genetic paradox' in invasion biology. This paradox is often resolved through the temporal buildup genetic diversity through multiple introduction events. These temporal dynamics in genetic diversity are especially important for annual invasive plants that lack a persistent seed bank, for which population persistence is strongly dependent on consecutive seed 're-establishment' in each growing season. Theory predicts that the number of seeds during re-establishment, and the levels of among-population gene flow can strongly affect recolonization dynamics, resulting in either an erosion or build-up of population genetic diversity through time. This study focuses on temporal changes in the population genetic structure of the annual invasive plant Impatiens glandulifera across Europe. We resampled 13 populations in 6 regions along a 1600 km long latitudinal gradient from northern France to central Norway after 5 years, and assessed population genetic diversity with 9 microsatellite markers. RESULTS: Our study suggests sufficiently high numbers of genetically diverse founders during population re-establishment, which prevent the erosion of local genetic diversity. We furthermore observe that I. glandulifera experiences significant among-population gene flow, gradually resulting in higher genetic diversity and lower overall genetic differentiation through time. Nonetheless, moderate founder effects concerning population genetic composition (allele frequencies) were evident, especially for smaller populations. Despite the initially low genetic diversity, this species seems to be successful at persisting across its invaded range, and will likely continue to build up higher genetic diversity at the local scale.

Contrasting spatial, temporal and environmental patterns in observation and specimen based species occurrence data.

Species occurrence data records the location and time of an encounter with a species, and is valuable for many aspects of ecological and evolutionary analyses. A key distinction within species occurrence data is between (1) collected and preserved specimens that can be taxonomically validated (i.e., natural history collections), and (2) observations, which are more error prone but richer in terms of number and spread of observations. In this study we analyse the distribution in temporal, spatial, taxonomic and environmental coverage of specimen- and observation based species occurrence data for land plants in Norway, a region with strong climatic and human population density gradients. Of 4.8 million species occurrence records, the majority (78%) were observations. However, there was a greater species richness in the specimen record (N = 4691) than in the observation record (N = 3193) and most species were recorded more as specimens than observations. Specimen data was on average older, and collected later during the year. Both record types were highly influenced by a small number of prolific contributors. The species most highly represented in the observation data set were widespread or invasive, while in the specimen records, taxonomically challenging species were overrepresented. Species occurrence records were unevenly spatially distributed. Both specimen and observation records were concentrated in regions of Norway with high human population density and with high temperatures and precipitation, but in different regions within Norway. Observation and specimen records thus differ in taxonomic, temporal, spatial and environmental coverage for a well-sampled group and study region, potentially influencing the ecological inferences made from studies utilizing species occurrence data. The distribution of observation data dominates the dataset, so inferences of species diversity and distributions do not correspond to the evolutionary or physiological knowledge of species, which is based on specimen data. We make recommendations for users of biodiversity data, and collectors to better exploit the complementary strengths of these distinct biodiversity data types.

Sheep grazing in the North Atlantic region: A long-term perspective on environmental sustainability.

Sheep grazing is an important part of agriculture in the North Atlantic region, defined here as the Faroe Islands, Greenland, Iceland, Norway and Scotland. This process has played a key role in shaping the landscape and biodiversity of the region, sometimes with major environmental consequences, and has also been instrumental in the development of its rural economy and culture. In this review, we present results of the first interdisciplinary study taking a long-term perspective on sheep management, resource economy and the ecological impacts of sheep grazing, showing that sustainability boundaries are most likely to be exceeded in fragile environments where financial support is linked to the number of sheep produced. The sustainability of sheep grazing can be enhanced by a management regime that promotes grazing densities appropriate to the site and supported by area-based subsidy systems, thus minimizing environmental degradation, encouraging biodiversity and preserving the integrity of ecosystem processes.

Case reports to suggest an algorithm for management of total fertilisation failure prior to use of donor gametes.

PURPOSE: Total fertilisation failure (TFF), even with intracytoplasmic sperm injection (ICSI), occurs in approximately 3 % of cycles, can be recurrent and the exact cause is difficult to elucidate. Differentiation between oocyte and sperm-related cause of TFF is possible using mouse oocyte-activation techniques, but is not an option within most clinical settings. Therefore, the management of these couples is clinically driven, and the endpoint, if recurrent, is often the use of donor gametes. However, with the invariable lack of a definitive cause of TFF, any decision between the use of donor sperm or oocytes remains an emotive one. We present two case reports demonstrating the importance of appropriate investigation, activation techniques (mechanical and chemical) and clinical management options to develop a clinical algorithm prior to the use of donor gametes. METHODS: This study is composed of two case reports of assisted reproduction investigation and treatment within an assisted conception unit for couples with recurrent total fertilisation failure. RESULTS: Using appropriate investigation (endocrine, urological and embryological) and treatments (ICSI, IMSI, oocyte-activation techniques), a fertilisation rate of 48 % was achieved in two cycles in couples following a total of nine previous cycles (and 200 previously collected eggs) with TFF. CONCLUSIONS: Oocyte activation requires the triggering of intracellular calcium oscillations by the release of a sperm-specific factor (phospholipase C zeta (PLCζ)) into the oocyte cytoplasm. Although, PLCζ deficiencies have been demonstrated as putative causes of failed activation, impaired oocyte responsiveness may also be a factor. The use of donor gametes is often recommended and is often the required endpoint of treatment. However, these reports outline a clinical algorithm that potentially offers success without donation, and also offers a systematic approach to help decide whether donor oocytes or sperm should be recommended.

Treatment and preservation at the extremes of reproductive age: a case report outlining the ethical dilemmas.

PURPOSE: The purpose of the study was to report a livebirth from a cryopreserved embryo created from autologous oocytes collected at 47 years and 9 months that outlines the ethical difficulties of decision-making at the extreme of reproductive age. METHODS: The method used was IVF and embryo cryopreservation within an assisted conception unit prior to adjuvant cancer treatment in a nulliparous patient diagnosed with breast carcinoma (47 years and 9 months at oocyte collection). RESULTS: A 47-year-old nulliparous woman was diagnosed with breast malignancy during work-up for fertility treatment. Ovarian stimulation yielded one embryo from four oocytes that was cryopreserved to allow completion of adjuvant treatment. Subsequent embryo transfer cycle led to a live birth of a healthy baby girl at term, weighing 3.37 kg. CONCLUSION: This paper demonstrates the oldest reported age of autologous oocyte collection to have achieved a livebirth. In women where most would consider treatment futile, we highlight the difficulties in decision-making in this group of patients.

Poor response cycles: when should we cancel? Comparison of outcome between egg collection, intrauterine insemination conversion, and follow-up cycles after abandonment.

OBJECTIVE: To determine optimal management with one or two mature follicles after stimulation. DESIGN: Retrospective analysis. SETTING: Lister fertility clinic. PATIENT(S): A total of 1,350 IVF/intracytoplasmic sperm injection cycles (7.3% of total) during 1998-2009 were found to have one or two mature follicles. INTERVENTION(S): Group 1 (n = 807) comprised those who proceeded to vaginal egg collection (VEC) (59.8%; outcome per egg collection), group 2 (n=248) those who converted to IUI (18.4%; outcome per insemination) and group 3 (n=259) those who abandoned the current cycle (21.9%; outcome per abandoned cycle in first subsequent cycle). MAIN OUTCOME MEASURE(S): Live birth rate, clinical pregnancy rate, and biochemical pregnancy rate. RESULT(S): Biochemical pregnancy rates of 13.1%, 4.9%, and 9.7%, clinical pregnancy rates of 8.1%, 3.6%, and 7.2%, and ongoing pregnancy rates of 6.8%, 2.0%, and 5.5% were achieved in groups 1, 2, and 3, respectively. All pregnancy outcomes were significantly higher after VEC (group 1) than for those converted to IUI (group 2), and all pregnancy outcomes were higher with borderline significance in group 3 vs. group 2. There was no significant difference in outcome between groups 1 and 3. CONCLUSION(S): Our data suggest that for such poor responders, proceeding to VEC may represent their best chance of successful outcome. Conversion to IUI offers the poorest outcome, and despite the potential for improvements in cycle protocol, abandoning and a further attempt does not improve outcome (using abandoned cycle as the denominator).

Experimental evidence for herbivore limitation of the treeline.

The treeline ecotone divides forest from open alpine or arctic vegetation states. Treelines are generally perceived to be temperature limited. The role of herbivores in limiting the treeline is more controversial, as experimental evidence from relevant large scales is lacking. Here we quantify the impact of different experimentally controlled herbivore densities on the recruitment and survival of birch Betula pubescens tortuosa along an altitudinal gradient in the mountains of southern Norway. After eight years of summer grazing in large-scale enclosures at densities of 0, 25, and 80 sheep/km2, birch recruited within the whole altitudinal range of ungrazed enclosures, but recruitment was rarer in enclosures with low-density sheep and was largely limited to within the treeline in enclosures with high-density sheep. In contrast, the distribution of saplings (birch older than the experiment) did not differ between grazing treatments, suggesting that grazing sheep primarily limit the establishment of new tree recruits rather than decrease the survival of existing individuals. This study provides direct experimental evidence that herbivores can limit the treeline below its potential at the landscape scale and even at low herbivore densities in this climatic zone. Land use changes should thus be considered in addition to climatic changes as potential drivers of ecotone shifts.

A decade of the sperm-washing programme: where are we now?

Since 1999, we have treated HIV-positive men with sperm washing as part of a risk-reduction programme with a year-on-year increase in total infectious cycles performed to over 200 in 2008. Four hundred and thirty nine cycles of IUI, 114 cycles of IVF and 117 cycles of ICSI have been performed in HIV positive men over the decade and of the 259 couples treated, a pregnancy rate and ongoing pregnancy rate per couple of 45.4% and 36.3% have been achieved with over 100 children born with no seroconversions. We outline the continued importance of such risk-reduction measures with 9.7% of samples from men with 'stable' disease on anti-retroviral treatment and undetectable viral load demonstrating detectable viral particles in seminal fluid and discuss measures to improve outcome in this patient group.