Variability in deer diet and plant vulnerability to browsing among forests with different establishment years of sika deer

Increased ungulate browsing alters the composition of plant communities and modifies forest ecosystems worldwide. Ungulates alter their diet following changes in availability of plant species; however, we know little about how browse selection and plant community composition change with different stages of deer establishment. Here, we provide insight into this area of study by combining multiple approaches: comparison of the understory plant community, analysis of records of browsing damage, and DNA barcoding of sika deer feces at 22 sites in forests in northern Japan varying in when deer were first established. The coverage of vegetation and number of plant species were only lower at sites where deer were present for more than 20 years, while the difference in plant coverage among deer establishment years varied among plant species. Deer diet differed across establishment years, but was more affected by the site, thereby indicating that food selection by deer could change over several years after deer establishment. Plant life form and plant architecture explained the difference in plant coverage across establishment years, but large variability was observed in deer diet within the two categories. Integrating these results, we categorized 98 plant taxa into six groups that differed in vulnerability to deer browsing (degree of damage and coverage). The different responses to browsing among plant species inferred from this study could be a first step in predicting the short- and long-term responses of forest plant communities to deer browsing.

Restoration of Natural Forests After Severe Wind Disturbance in a Cold, Snowy Region With a Deer Population: Implications From 15 Years of Field Experiments

Questions have been raised about the application of conventional post-windthrow forest practices such as salvage logging, site preparation, and afforestation in response to the increase in wind disturbance caused by climate change. In particular, it is necessary to identify effective forest management practices that consider the pressure from deer browsing in forests in cold, snowy regions because the population of ungulates is expected to increase. The impacts of legacy destruction, i.e., the destruction of advance regeneration, microsites, and soil structure, caused by conventional post-windthrow practices have rarely been assessed separately from the impacts of subsequent deer browsing on forest regeneration or evaluated based on sufficiently long monitoring periods to assess vegetation succession. This lack of studies is one reason that alternative forest management practices to salvaging and planting have not been proposed. We conducted a field experiment at a large-scale windthrow site with a deer population to (1) assess the impact of legacy destruction and deer browsing on vegetation biomass and species composition after 15 years and (2) identify the effects of legacy retention. The study design allowed us to distinguish between and measure the impact of legacy destruction and that of subsequent deer browsing during a 15-year period. The results revealed the following: (1) Salvage logging and site preparation suppressed the development of biomass of shrub and tree layers in forested areas where harvest residues were piled up and shifted the plant communities in these areas to herbaceous plant communities. (2) Subsequent deer browsing suppressed the development of the biomass of shrub and tree layers throughout the forested site and shifted herbaceous communities to ruderal communities dominated by alien species; and 3. Compared with salvaging and planting, legacy retention enabled the windthrow sites to more quickly develop into a stand with characteristics similar to that of a mature, natural forest. Forest management practices that consider the presence of deer are necessary. We propose a policy shift from planting trees after salvaging to leaving downed trees to regenerate natural forests, unless there is concern about insect damage to the remaining forestry land in the vicinity.

Intense Browsing by Sika Deer (Cervus Nippon) Drives the Genectic Differentiation of Hairy Nettle (Urtica Thunbergiana) Populations

Many studies have inferred the way in which natural selection, genetic drift and gene flow shape the population genetic structures, but very few have quantified the population differentiation under spatially and temporally varying levels of selection pressure, population fluctuation and gene flow. In Nara Park (6.6 km2; NP), central Japan, where several hundred sika deer (Cervus nippon) have been protected for more than 1,200 years, heavily- or moderately-haired nettle (Uritica thunbergiana) populations have evolved probably in response to intense deer browsing. Here, we analysed the genetic structure of two populations from NP and five from surrounding areas using amplified fragment length polymorphism markers. A total of 546 marker loci were genotyped from 210 individuals. A Bayesian method estimated 5.5% of these loci to be outliers, which are putatively under natural selection. Neighbour-joining, Bayesian clustering and principal coordinates analyses using all-loci, non-outlier loci and outlier loci datasets showed that the two populations from NP formed a cluster distinct from the surroundings. These results indicate the genome-wide differentiation of the populations from NP and the surroundings. Moreover, these imply that: (1) gene flow is limited between these populations and thus genetic drift is a major factor causing the differentiation; and (2) natural selection imposed by intense deer browsing has contributed to some extent to the differentiation. In conclusion, sika deer seems to have counteracted genetic drift to drive the genetic differentiation of hairy nettles in NP. This study suggests that a single herbivore species could lead genetic differentiation among plant populations.

Effect of biofuel waste, urea, deer browsing and vegetation control on Pinus banksiana seedling growth on a dry upland site in central Canada

Ash from biofuels and nitrogen fertilizer are increasingly being used as soil amendments. While this can increase tree growth, it can also increase mammalian grazing and competition with vegetation. We applied moderate amounts (1.5 t ha^-1 y^-1) of ash and 74 kg N ha^-1 y^-1 of urea in each of two years to a well-drained site in southeastern Manitoba, planted with <i>Pinus banksiana</i>. Subplots received deer browsing and/or vegetation control. The ash resulted in an increase in pH in the upper 15m of mineral soil from ca. 5.7 to 6.6, and the urea created short-term spikes in soil inorganic N (NH₄ and NO₃) levels. Urea combined with ash significantly increased seedling relative growth rates in the first two years, with seedlings being largest with urea, with or without ash. However, by the fourth year seedling growth and size did not differ between the amendments. Urea application increased browsing damage to 91 %, but only when vegetation was mowed. Browsing guards resulted in seedlings having 1.6 times greater shoot mass by the end of the fourth growing season. These results suggest that on sandy soils in the dry region of central Canada, <i>P. banksiana</i> may get little benefit from ash applications.