Among population differentiation at nuclear genes in native Scots pine (Pinus sylvestris L.) in Scotland

In the Scottish Highlands, Scots pine is at the north-western extreme of its wide natural distribution. Here, the remaining native populations are patchily distributed in highly variable environments, from the more continental, drier eastern Highlands to the milder, wetter Atlantic Ocean coast. As these pinewoods are the remnants of a naturally established forest, they form a valuable system for analysis of genetic and adaptive variation in heterogeneous environments. Using samples from across the Scottish population, we analysed data from nuclear and mitochondrial genes to assess patterns of within and between population genetic variation. Within population diversity levels were high, and significant genetic differentiation among pairs of Scottish populations at relatively small spatial scales was present at several nuclear loci. At these loci, no differentiation had been found among continental populations, even those separated by large geographic distances. Overall, no clear clustering of Scottish samples was found in population structure analysis suggesting that geographically distant populations with high intra-population nucleotide diversity are not strongly isolated and diverged from each other. Scottish populations lacked a mitotype that is widespread in eastern and north-eastern Europe, indicating that pines from that area may not have participated in the most recent colonisation of the British Isles

Genotypic Variation in a Foundation Tree (\u3ci\u3ePopulus tremula\u3c/i\u3e L.) Explains Community Structure of Associated Epiphytes

Community genetics hypothesizes that within a foundation species, the genotype of an individual significantly influences the assemblage of dependent organisms. To assess whether these intra-specific genetic effects are ecologically important, it is required to compare their impact on dependent organisms with that attributable to environmental variation experienced over relevant spatial scales. We assessed bark epiphytes on 27 aspen (Populus tremula L.) genotypes grown in a randomized experimental array at two contrasting sites spanning the environmental conditions from which the aspen genotypes were collected. We found that variation in aspen genotype significantly influenced bark epiphyte community composition, and to the same degree as environmental variation between the test sites. We conclude that maintaining genotypic diversity of foundation species may be crucial for conservation of associated biodiversity

Selection of trees for rubbing by red and roe deer in forest plantations

Antler rubbing is a form of behaviour by which deer may damage and ultimately induce mortality of trees. Understanding factors affecting selection of trees for rubbing may contribute to mitigation of negative effects of such behaviour in plantations or woodlands. We analysed characteristics of trees rubbed by red and roe deer along transects established in plantations of Pinus pinaster (Aiton), Pseudotsuga menziesii (Mirbel) Franco, Betula alba L. and Quercus robur L. in Northeast Portugal. Transects were walked during five sampling periods covering mating seasons of red and roe deer. Red deer preferentially rubbed trees adjacent to the edge of plantations and large clearings whilst roe deer selected those inside plantations within small clearings. There was seasonal segregation in the number of trees rubbed by each deer species with red deer rubbing trees mainly between September and February and roe deer mainly between December and June. Both red and roe deer selected trees with smaller diameter than those of available trees although trees selected by red deer had larger diameters than those selected by roe deer. Roe, but not red deer, tended to avoid trees protected by shrubs. Overall, the selection of trees for rubbing was site-dependent suggesting that generalizations across sites should be made with caution. Mitigating measures, such as deer control, tree protection or provision of alternative rubbing posts should target stands of particular tree species, location of trees in relation to stand clearings and tree size classes.http://www.sciencedirect.com/science/article/B6T6X-4HGM78R-2/1/29fe58190c40581f0716e977b7847d3

Progeny selection for enhanced forest growth alters soil communities and processes

Genetic enhancement of tree species is integral to global forest management practices with mass propagation of enhanced plant material being used to reforest whole landscapes. It is unclear, however, how genetic enhancement of basic traits such as tree growth may influence the function of life supporting soil ecosystems. We studied the potential cascading effects of genetic increases in growth of Norway spruce (Picea abies) on a range of soil chemical and biological properties. Because this species is a prime candidate for the genetic enhancement of boreal forest landscapes and it has been introduced around the world, its impacts on soil microbiomes are likely of importance both locally and globally. In a 40-year common garden, we assessed how genetic increases in growth generated through controlled crossing of high-quality "plus" trees from across the central boreal zone of Sweden influenced a range of soil properties beneath the canopies. Properties included pH, carbon, nitrogen, nitrate, ammonium, phosphate, respiration rate, and the composition of microbial communities assessed via phospholipid fatty acids (PLFAs). We found that Norway spruce family significantly affected each of the seven chemical properties assessed, with differences of up to 140% among families, and that three of the seven were significantly correlated with mean family growth rate. We also found that fungal PLFAs varied significantly across Norway spruce families, but these differences were not strongly related to mean family growth rate. This study, representing just one cycle of selective breeding, suggests that genetic increases in tree growth rates may also be inadvertently altering soil communities and ecosystem services. Such alterations across forest landscapes may have unexpected implications for the function of forest ecosystems (i.e., nutrient cycling) as well as processes of global significance (i.e., carbon sequestration)

DISTRIBUTION OF WINTER BROWSING BY MOOSE: EVIDENCE OF LONG-TERM STABILITY IN NORTHERN SWEDEN

Predicting spatial distribution of large herbivore foraging is important for successful management, but accurate predictions remain elusive against a background of multiple causes modified by environmental stochasticity. Moose (Alces alces) might prefer to browse areas with high plant density, but if snow depth co-varies with plant density, this could restrict access to these sites and force use of sites with lower plant density and snow depth. Moose browsing was measured in 72 plots distributed within the subarctic birch (Betula spp.) forest landscape at Abisko in northern Sweden in 1996. In 2010, the same plots were revisited and the measurements repeated. A generalized linear model predicted moose browsing on birch in 2010 from the browsing pattern on birch measured in 1996. The model suggested that neither total density of willow and birch stems nor snow depth were influential of foraging distribution of birch at multiple spatial scales. The spatial scale at which clustering of browsing on birch occurred, coincided with the scale of clustering of birch and willow (Salix spp.) stems at distances of 1000-2500 m; at lesser distance browsing was distributed randomly. We concluded that moose demonstrate stability in spatial browsing patterns after 14 years which corresponds to 3-4 generations of moose, and that plant density represents a cue for moose only at certain scales. Predictability of feeding sites is valuable for long-term moose and forest management, and conservation planning

Spring phenology shows genetic variation among and within populations in seedlings of Scots pine (Pinus sylvestris L.) in the Scottish Highlands

Background: Genetic differentiation in phenotypic traits is often observed among forest tree populations, but less is known about patterns of adaptive variation within populations. Such variation is expected to enhance the survival likelihood of extant populations under climate change. Aims: Scots pine (Pinus sylvestris) occurs over a spatially and temporally heterogeneous landscape in Scotland. Our goal was to examine whether populations had differentiated genetically in timing of bud flush in response to spatial heterogeneity and whether variation was also maintained within populations. Methods: Two common-garden studies, involving maternal families of seedlings from 21 native pinewoods, were established and variation in the trait was measured at the beginning of the second growing season. Results: Populations showed genetic differences in the trait correlated with the length of growing season at their site of origin, but the majority of variation was observed within populations. Populations also differed in their levels of variation in the trait; a pattern that may be influenced by spatial variation in the extent of temporal climate variability. Conclusions: Our findings suggest that populations have adapted to their home environments and that they also have substantial ability to adapt in situ to changes in growing season length

Forest diversity effects on insect herbivores: do leaf traits matter?

Insect herbivore damage and abundance are often reduced in diverse plant stands. However, few studies have explored whether this phenomenon is a result of plant diversity effects on host plant traits. We explored indirect effects of tree species diversity on herbivory via changes in leaf traits in a long-term forest diversity experiment in Finland. We measured 16 leaf traits and leaf damage by four insect guilds (chewers, gall formers, leaf miners and rollers) on silver birch (Betula pendula) trees growing in one-, two-, three- and five-species mixtures. A decline in the frequency of birch in mixed stands resulted in reduced leaf area. This, in turn, mediated the reduction in chewing damage in mixed stands. In contrast, associational resistance of birch to leaf miners was not trait-mediated but driven directly by concurrent declines in birch frequency as tree species richness increased. Our results show that leaf trait variation across the diversity gradient might promote associational resistance, but these patterns are driven by an increase in the relative abundance of heterospecifics rather than by tree species richness per se. Therefore, accounting for concurrent changes in stand structure and key foliar traits is important for the interpretation of plant diversity effects and predictions of associational patterns

Weak isolation by distance and geographic diversity gradients persist in Scottish relict pine forest

Gene flow is one of the main factors shaping genetic diversity within and among tree populations, and occurs through pollen and seed dispersal. Recent findings of pollen-release asynchronies in distant populations of Scots pine (Pinus sylvestris L.) within Scotland suggest that gene dispersal among more distant populations might be less effective than previously thought. Limited gene dispersal is one of the major factors causing genetic structure for neutral markers, and pollen-release asynchrony could have driven isolation by distance (IBD) among Scottish populations. Previous studies of neutral markers found little differentiation among Scottish populations of Scots pine, however they did not consider IBD over the full Scottish range. We analysed data from 6 nuclear simple sequence repeats (SSR) and 5 chloroplast SSR loci in a total of 540 individuals of Scots pine from 18 populations across Scotland. Our aim was to assess contemporary levels and distribution of genetic variation and to test if the distribution of genetic diversity was consistent with IBD. We also analysed patterns of gene flow that could have contributed to the observed patterns of variation. Levels of genetic diversity were high, for both nuclear and chloroplast markers within populations, and there was no significant differentiation among populations. A weak signal of IBD was present. We found an increase in nuclear diversity towards the East along with greater gene flow in a West-East direction commensurate with the prevailing winds. Our findings suggest that this wind-driven gene flow is dominant over genetic drift and prevents differentiation among the Scottish populations. It may also counteract any pollen-release asynchronies among populations

Cumulative impact assessments of multiple host species loss from plant diseases show disproportionate reductions in associated biodiversity

Funding Information: This work was funded by BBSRC grant Protecting Oak Ecosystems (PuRpOsE): BB/N022831/1 with additional funding from the Scottish Government's Rural and Environment Research and Analysis Directorate 2016–2021 strategic research programme. The authors thank Nick Hodgetts for collating the lists of the bryophyte species associated with ash and oak, Ralph Harmer for conducting some of the site visits and the site owners for allowing us access to the sites. Katharine Preedy provided statistical advice. Steve Albon and Robin Pakeman kindly provided comments to improve earlier drafts.Peer reviewe