Across-species patterns of genetic variation in forest trees of Central Europe

The study focuses on geographical patterns of genetic variation at allozyme loci common for four main tree species of Central Europe (Norway spruce, silver fir, common beech and sessile oak). Moving-window averaging of four indicators of allelic richness and diversity (proportion of polymorphic loci, mean number of alleles per locus, effective number of alleles and expected heterozygosity) with window size of 50 × 50 km was used to identify the patterns. Moreover, local genetic divergence was assessed using the G _ST (Nei, Molecular population genetic and evolution, Amsterdam and Oxford, North-Holland, 1975) and D _j (Gregorius and Roberds, Theor Appl Genet 71:826–834, 1986) statistics for common beech and silver fir, where raw genotype data were available. Spatial patterns of diversity and allelic richness were quite similar. Romanian Carpathians were identified as the most important hotspot of genetic diversity and evolutionary divergence in Central Europe. Implications for genetic conservation are briefly discussed.     

Herbivory and plant competition reduce mountain beech seedling growth and establishment in New Zealand

The effect of herbivory and resource availability on the competitive ability of different plant species has been an area of intense debate amongst plant ecologists for at least two decades, but the interactive effects of herbivory and plant competition between woody and herbaceous plants are rarely studied and theory is poorly developed. This study used experimental manipulations on transplanted and naturally occurring mountain beech (Nothofagus solandri var. cliffortioides) seedlings to show the effects of deer browsing and competition from deer-induced, herbaceous turf communities on mountain beech regeneration in New Zealand. Differences in the species composition of these turfs had little effect on mountain beech seedling establishment, but turf removal increased seedling growth and survivorship, showing that competition with other plants had direct effects on mountain beech regeneration. Deer browsing reduced the establishment and growth of seedlings, but the size of this effect did not vary with light and nutrient availability. There was no immediate compositional response of turf communities following the removal of deer browsing. The addition of nutrients appeared to reduce the intensity of belowground competition (stem growth increased relative to root growth) and increase seedling mortality, but there was no effect of changing levels of light. These results showed simple and direct negative effects of deer browsing on mountain beech regeneration. Indirect negative effects on regeneration were caused by deer-induced turf communities. We found little evidence for interactive effects between herbivory, plant competition and the availability of light or nutrients on seedling regeneration, which suggests that these factors acted independently. Nomenclature: Beever et al. (1992); Parsons et al. (1995); Edgar and Connor (2000); and Brownsey and Smith-Dodsworth (2000). Raukaua simplex is described by Mitchell et al. (1997). Coprosma “taylorae” is referred to by Eagle (1986) and Halocarpus biformis, Phyllocladus alpinus, Podocarpus hallii and Podocarpus nivalis by Wilson and Galloway (1993).     

Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white‐tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5–6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures – none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m²) and greater woody species richness at the subplot scale (1 m²) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white‐tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing‐species richness curve, but have rarely been documented in forests.     

Genetic variability and the effect of habitat fragmentation in spotted suslik Spermophilus suslicus populations from two different regions

Endangered species worldwide exist in remnant populations, often within fragmented landscapes. Although assessment of genetic diversity in fragmented habitats is very important for conservation purposes, it is usually impossible to evaluate the amount of diversity that has actually been lost. Here, we compared population structure and levels of genetic diversity within populations of spotted suslik Spermophilus suslicus, inhabiting two different parts of the species range characterized by different levels of habitat connectivity. We used microsatellites to analyze 10 critically endangered populations located at the western part of the range, where suslik habitat have been severely devastated due to agriculture industrialization. Their genetic composition was compared with four populations from the eastern part of the range where the species still occupies habitat with reasonable levels of connectivity. In the western region, we detected extreme population structure (F _ST = 0.20) and levels of genetic diversity (Allelic richness ranged from 1.45 to 3.07) characteristic for highly endangered populations. Alternatively, in the eastern region we found significantly higher allelic richness (from 5.09 to 5.81) and insignificant population structure (F _ST = 0.03). As we identified a strong correlation between genetic and geographic distance and a lack of private alleles in the western region, we conclude that extreme population structure and lower genetic diversity is due to recent habitat loss. Results from this study provide guidelines for conservation and management of this highly endangered species.     

Species versus genotypic diversity of a nitrogen-fixing plant functional group in a metacommunity

Exploring species and genetic diversity interactions provides new opportunities for furthering our understanding of the ecology and evolution of population and community dynamics, and for predicting responses of ecosystems to environmental change. Theory predicts that species diversity within communities and genetic diversity within populations will covary positively, because either species and genetic diversity interact synergistically or they respond in parallel fashion to common habitat conditions. We tested the hypothesis of positive covariation between species and genotypic diversity in a metacommunity of the species-rich southwest Australian flora. We hypothesised that the connection between genotypic diversity and species diversity is strong within functional groups, but weak or non-existent if the species considered extend beyond the functional group. We show that allelic richness of Daviesia triflora, an ant-dispersed pea, covaries positively with the species richness of six co-occurring nitrogen-fixing legume species. No pattern can be detected between allelic richness of D. triflora and species richness of ant-dispersed species when four non-legumes are added. We also show that genetic diversity of D. triflora is not governed by the same environmental factors that determine the presence of a group of large-shrub/small-tree species in the same metacommunity. This study shows that species and genetic diversity covariation are more likely to be confined to within, rather than between, plant functional groups.     

The impact of habitat fragmentation on genetic structure of the Japanese sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) in southern Kantoh,revealed by mitochondrial D-loop sequences

In southern Kantoh, Japanese sika deer (Cervus nippon) are distributed discontinuously due to large urban areas and developed road networks. To assess the impact of habitat fragmentation on sika deer subpopulations, we examined mitochondrial D-loop sequences from 435 individuals throughout southern Kantoh. About 13 haplotypes were detected, and their distributions revealed spatial genetic structure. Significant genetic differentiation was observed among seven of eight subpopulations. We found no significant correlation between pairwise F _ST and geographical distance among subpopulations. Genetic diversity indices suggested that seven of eight subpopulations had probably experienced population bottlenecks in the recent past. Therefore, and in the light of the results of a nested clade analysis of these haplotypes, we conclude that recent fluctuations in population size and the interruption of gene flow due to past and present habitat fragmentation have played major roles influencing the spatial genetic structure of the sika deer population. This is the first evidence of spatial genetic population structure in the highly fragmented sika deer population in Honshu, Japan.     

High population differentiation in the rock-dwelling land snail (<Emphasis Type="Italic">Trochulus caelatus</Emphasis>) endemic to the Swiss Jura Mountains

Understanding patterns of genetic structure is fundamental for developing successful management programmes for isolated populations of threatened species. Trochulus caelatus is a small terrestrial snail endemic to calcareous rock cliffs in the Northwestern Swiss Jura Mountains. Eight microsatellite loci were used to assess the effect of habitat isolation on genetic population structure and gene flow among nine populations occurring on distinct cliffs. We found a high genetic differentiation among populations (mean F _ST = 0.254) indicating that the populations are strongly isolated. Both allelic richness and effective population size were positively correlated with the size of the cliffs. Our findings support the hypothesis that T. caelatus survived on ice-free cliffs during the Pleistocene glacier advancements from the Alps. Due to the establishment of beech and pine forest under recent, temperate climate conditions, dispersal between cliffs is no longer possible for rock-dwelling snails such as T. caelatus. Our results provide basic data for developing a conservation action plan for this endangered gastropod species.     

Consistent loss of genetic diversity in isolated cutthroat trout populations independent of habitat size and quality

Fragmentation and isolation of wildlife populations has reduced genetic diversity worldwide, leaving many populations vulnerable to inbreeding depression and local extinction. Nonetheless, isolation is protecting many native aquatic species from interactions with invasive species, often making reconnection an unrealistic conservation strategy. Isolation management is widely used to protect extant cutthroat trout (Oncorhynchus clarkii) populations from invasive species. Despite this, few studies have empirically examined how predictor variables including habitat length, population size, time since isolation and habitat quality, relate to levels of genetic diversity in isolated trout populations. We compared allelic richness of cutthroat trout across 14 microsatellite loci in two connected and 12 anthropogenically isolated populations of the Flathead River basin, Montana. Isolated populations in habitat fragments <8 km stream length had reduced genetic diversity, but diversity was not significantly related to any of our predictor variables. To broaden our scope, we analyzed seven geologically isolated populations from the same river basin occupying habitat fragments up to 18 km in length. These populations showed reduced diversity, regardless of fragment size. Furthermore, geologically isolated populations had significantly lower average allelic richness compared to streams recently isolated by anthropogenic activities. These results demonstrate a consistent loss of genetic diversity through time in isolated populations, emphasizing the need to explore strategies to minimize risks of inbreeding depression. Testing conservation theory and subsequent assumptions broadly across taxa is necessary to ensure efficacy of conservation efforts.     

Big game or big loss? High deer densities are threatening woody plant diversity and vegetation dynamics

Land-use change and current game management have favored an increasing population of wild ungulates (especially deer) in many regions of the Northern Hemisphere. Here, we assess the impact of high deer densities (>30 ind km^?2) on the highly diverse woody vegetation of Mediterranean ecosystems, where big game have been favored for the last decades. We examine whether prolonged deer browsing (over 30 years) affected plant composition, diversity and dynamics of the original (non-browsed) vegetation. Deer browsing led to an average decrease of 30.4 % in woody plant diversity (species richness), due to a lack of regeneration for the most preferred plant species. Species belonging to early stages of succession (mostly Labiatae and Cistaceae) were non-preferred by deer. Conversely, the most preferred species belonged mainly to late stages of plant succession. Deer impact on Mediterranean shrublands is causing biotic homogenization of plant communities and is threatening vegetation dynamics by forcing it to return to earlier succession stages. Strict deer population control favouring larger trophies but lower offspring numbers together with an adequate habitat management (increasing grass and acorn availability) would be the most efficient measures to reverse this diversity loss. Restoration work seems only appropriate for the most vulnerable species. We highlight the need of sampling deer-free areas with low or null historical browsing to assess the real impact of deer on woody plant diversity and vegetation dynamics.     

Different facets of tree sapling diversity influence browsing intensity by deer dependent on spatial scale

Browsing of tree saplings by deer hampers forest regeneration in mixed forests across Europe and North America. It is well known that tree species are differentially affected by deer browsing, but little is known about how different facets of diversity, such as species richness, identity, and composition, affect browsing intensity at different spatial scales. Using forest inventory data from the Hainich National Park, a mixed deciduous forest in central Germany, we applied a hierarchical approach to model the browsing probability of patches (regional scale) as well as the species‐specific proportion of saplings browsed within patches (patch scale). We found that, at the regional scale, the probability that a patch was browsed increased with certain species composition, namely with low abundance of European beech (Fagus sylvatica L.) and high abundance of European ash (Fraxinus excelsior L.), whereas at the patch scale, the proportion of saplings browsed per species was mainly determined by the species’ identity, providing a “preference ranking” of the 11 tree species under study. Interestingly, at the regional scale, species‐rich patches were more likely to be browsed; however, at the patch scale, species‐rich patches showed a lower proportion of saplings per species browsed. Presumably, diverse patches attract deer, but satisfy nutritional needs faster, such that fewer saplings need to be browsed. Some forest stand parameters, such as more open canopies, increased the browsing intensity at either scale. By showing the effects that various facets of diversity, as well as environmental parameters, exerted on browsing intensity at the regional as well as patch scale, our study advances the understanding of mammalian herbivore–plant interactions across scales. Our results also indicate which regeneration patches and species are (least) prone to browsing and show the importance of different facets of diversity for the prediction and management of browsing intensity and regeneration dynamics.     

Population size and habitat quality affect genetic diversity and fitness in the clonal herb <Emphasis Type="Italic">Cirsium dissectum</Emphasis>

Remaining populations of plant species in fragmented landscapes are threatened by declining habitat quality and reduced genetic diversity, but the interactions of these major factors are rarely studied together for species conservation. In this study, the interactions between population size, habitat quality, genetic diversity and fitness were investigated in 22 populations of the clonal herb Cirsium dissectum throughout the British Isles. Regression analysis was used to identify significant factors, and a structural equation model was developed to illustrate and integrate these interactions. It was found that smaller populations (measured as the total number of plants) had lower genetic diversity (proportion of polymorphic loci), and that reduced genetic diversity (allelic richness) had a negative impact on the survival of seedlings grown under standard conditions. Habitat quality also had a large effect on C. dissectum. Unmanaged sites with tall vegetation, no bare soil and higher nutrient levels had smaller populations of C. dissectum, but flowering was promoted. Flowering was suppressed in heavily grazed sites with short vegetation. Higher levels of bare soil and phosphorus both had a positive relationship with genetic diversity, but probably for distinctly different reasons: bare soil provides safe sites for establishment, whilst phosphorus may promote flowering and improve seed germination. In order to conserve C. dissectum, management needs to maintain site heterogeneity so that C. dissectum can flower and establishment gaps are still available for seedlings; when either component is reduced, negative feedbacks through reduced genetic diversity and individual fitness can be expected. This study therefore highlights the importance of considering both conservation genetics and habitat quality in the conservation of plant species.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

Habitat predictors of genetic diversity for two sympatric wetland‐breeding amphibian species

Population genetic diversity is widely accepted as important to the conservation and management of wildlife. However, habitat features may differentially affect evolutionary processes that facilitate population genetic diversity among sympatric species. We measured genetic diversity for two pond‐breeding amphibian species (Dwarf salamanders, Eurycea quadridigitata; and Southern Leopard frogs, Lithobates sphenocephalus) to understand how habitat characteristics and spatial scale affect genetic diversity across a landscape. Samples were collected from wetlands on a longleaf pine reserve in Georgia. We genotyped microsatellite loci for both species to assess population structures and determine which habitat features were most closely associated with observed heterozygosity and rarefied allelic richness. Both species exhibited significant population genetic structure; however, structure in Southern Leopard frogs was driven primarily by one outlier site. Dwarf salamander allelic richness was greater at sites with less surrounding road area within 0.5 km and more wetland area within 1.0 and 2.5 km, and heterozygosity was greater at sites with more wetland area within 0.5 km. In contrast, neither measure of Southern Leopard frog genetic diversity was associated with any habitat features at any scale we evaluated. Genetic diversity in the Dwarf salamander was strongly associated with land cover variables up to 2.5 km away from breeding wetlands, and/or results suggest that minimizing roads in wetland buffers may be beneficial to the maintenance of population genetic diversity. This study suggests that patterns of genetic differentiation and genetic diversity have associations with different habitat features across different spatial scales for two syntopic pond‐breeding amphibian species.     

Loss of genetic diversity in an outbreeding species: small population effects in the African wild dog <Emphasis Type="Italic">(Lycaon pictus</Emphasis>)

Genetic studies on the endangered African wild dog (Lycaon pictus) have primarily focused on the few remaining large and viable populations. However, investigations on the many isolated small African wild dog populations might also be informative for species management because the majority of extant populations are small and may contain genetic variability that is important for population persistence and for species conservation. Small populations are at higher risk of extinction from stochastic and deterministic demographic processes than larger populations and this is often of more immediate conservation concern than loss of genetic diversity, particularly for species that exhibit out-breeding behaviour such as long distance dispersal which may maintain gene flow. However, the genetic advantages of out-breeding behaviour may be reduced if dispersal is compromised beyond reserve borders (edge effects), further weakening the integrity of small populations. Mitochondrial DNA and 11 microsatellite genetic markers were used to investigate population genetic structure in a small population of out-breeding African wild dogs in Zambia, which occupies an historical dispersal corridor for the species. Results indicated the Zambian population suffered from low allelic richness, and there was significant evidence of a recent population bottleneck. Concurrent ecological data suggests these results were due to habitat fragmentation and restricted dispersal which compromised natural out-breeding mechanisms. This study recommends conservation priorities and management units for the African wild dog that focus on conserving remaining levels of genetic diversity, which may also be applicable for a range of out-breeding species.     

Reproductive strategies and isolation‐by‐demography in a marine clonal plant along an eutrophication gradient

Genetic diversity in clonal organisms includes two distinct components, (i) the diversity of genotypes or clones (i.e. genotypic richness) in a population and (ii) that of the alleles (i.e. allelic and gene diversity within populations, and differentiation between populations). We investigated how population differentiation and genotypic components are associated across a gradient of eutrophication in a clonal marine plant. To that end, we combined direct measurements of sexual allocation (i.e. flower and seed counts) and genotypic analyses, which are used as an estimator of effective sexual reproduction across multiple generations. Genetic differentiation across sites was also modelled according to a hypothesis here defined as isolation‐by‐demography, in which we use population‐specific factors, genotypic richness and eutrophication that are hypothesized to affect the source‐sink dynamics and thus influence the genetic differentiation between a pair of populations. Eutrophic populations exhibited lower genotypic richness, in agreement with lower direct measurements of sexual allocation and contemporaneous gene flow. Genetic differentiation, while not explained by distance, was best predicted by genotypic richness and habitat quality. A multiple regression model using these two predictors was considered the best model (R² = 0.43). In this study, the relationship between environment and effective sexual–asexual balance is not simply (linearly) predicted by direct measurements of sexual allocation. Our results indicate that population‐specific factors and the isolation‐by‐demography model should be used more often to understand genetic differentiation.     

Genetic diversity loss associated to high mortality and environmental stress during the recruitment stage of a coral reef fish

We investigated the short-term impact of environmental-induced stress on survival and neutral genetic diversity of recently settled juveniles of a damselfish, Dascyllus aruanus, using spatiotemporal caging experiments in various natural environmental conditions in Moorea (French Polynesia). Juveniles’ mortality was followed at five study sites and overall four experiments, mortality rates ranged from 0 to 45%. Mortality rate and average daily water temperature were positively correlated (P = 0.018). Juveniles’ mortality rate and allelic richness estimated from ten microsatellite loci were negatively correlated (P = 0.046). Together, an overdominance of heterozygotes was observed within hostile environments. These results suggest that an allelic richness loss may be expected as a direct consequence of unfavorable environmental conditions. Thus, a worrisome scenario on demographic and genetic consequences may be expected from habitat degradation in the context of global change and human pressure increases.     

Diverging trends between heterozygosity and allelic richness during postglacial colonization in the European beech

Variation at 12 polymorphic isozyme loci was studied in the European beech on the basis of an extensive sample of 389 populations distributed throughout the species range. Special emphasis was given to the analysis of the pattern of geographic variation on the basis of two contrasting measures of genetic diversity, gene diversity (H) and allelic richness, and to their relationship. Measures of allelic richness were corrected for variation in sample size by using the rarefaction method. As expected, maximum allelic richness was found in the southeastern part of the range (southern Italy and the Balkans), where beech was confined during the last ice age. Surprisingly, H was lower in refugia than in recently colonized regions, resulting in a negative correlation between the two diversity measures. The decrease of allelic richness and the simultaneous increase of H during postglacial recolonization was attributed to several processes that differentially affect the two diversity parameters, such as bottlenecks due to long-distance founding events, selection during population establishment, and increased gene flow at low population densities.     

Temporal shifts of DNA‐microsatellite allele profiles in roe deer (Capreolus capreolus L.) within three decades

DNA‐microsatellite polymorphism (four loci) was studied in 56 male roe deer (Capreolus capreolus) from a 900‐ha hunting territory in the Vosges du Nord Mountains (France), culled over 34 years (1956–1990). Changed allele frequencies at two loci within this period, and increased allelic diversity, were traced to a phase of reduced population density and subsequent immigration. Decadic population samples collected within 900‐ha were distinguished by higher genetic variability measures than were certain geographical samples across Central Europe (4–900 km). On average, the decadic cohorts were distinguished by a gene diversity index of G_ST = 0.0286, and a genetic distance of D = 0.0938, which reflect 54% (G_ST) and 69% (D) of the respective geographic (350 km) differentiation indices of roe deer in Central Europe. The importance of demography and population ecology effects for microevolution in a large mammal is demonstrated, as is the risk of artefact by composing population samples of deer over several years. Population genetic screening should cover various demes of roe deer from the same general region, and be based on many unlinked polymorphic loci, to minimize the distorting effects of genetic dynamics at the small spatial scale.     

Spatial and temporal genetic dynamics of the grasshopper Oedaleus decorus revealed by museum genomics

Analyzing genetic variation through time and space is important to identify key evolutionary and ecological processes in populations. However, using contemporary genetic data to infer the dynamics of genetic diversity may be at risk of a bias, as inferences are performed from a set of extant populations, setting aside unavailable, rare, or now extinct lineages. Here, we took advantage of new developments in next‐generation sequencing to analyze the spatial and temporal genetic dynamics of the grasshopper Oedaleus decorus, a steppic Southwestern‐Palearctic species. We applied a recently developed hybridization capture (hyRAD) protocol that allows retrieving orthologous sequences even from degraded DNA characteristic of museum specimens. We identified single nucleotide polymorphisms in 68 historical and 51 modern samples in order to (i) unravel the spatial genetic structure across part of the species distribution and (ii) assess the loss of genetic diversity over the past century in Swiss populations. Our results revealed (i) the presence of three potential glacial refugia spread across the European continent and converging spatially in the Alpine area. In addition, and despite a limited population sample size, our results indicate (ii) a loss of allelic richness in contemporary Swiss populations compared to historical populations, whereas levels of expected heterozygosities were not significantly different. This observation is compatible with an increase in the bottleneck magnitude experienced by central European populations of O. decorus following human‐mediated land‐use change impacting steppic habitats. Our results confirm that application of hyRAD to museum samples produces valuable information to study genetic processes across time and space.     

Suppression of Regeneration in New Zealand Mountain Beech Forests is Dependent on Species of Introduced Deer

We compared the impacts on forest regeneration of introduced sika (Cervus nippon) and red (Cervus elaphus) deer in New Zealand. Plot data were used to compare mountain beech (Nothofagus solandri var. cliffortioides) regeneration between a region with sika deer, and four regions without sika deer. All regions surveyed had red deer present. In the region where sika deer had been present for more than a decade, there was evidence of poor mountain beech seedling regeneration. In the four regions without sika deer, there was evidence of a strong regenerative response at stands with low occupancy by trees. When compared to larger deer species, sika deer have a digestive morphology allowing greater dietary versatility, which may result in them impeding forest regeneration where red deer do not. In contrast to mountain beech, some small-leaved shrub species may have been competitively advantaged by intensive browsing from sika deer. This is contra to a current view that small-leaved shrub species with interlacing branches were able to tolerate browsing from extinct ratite birds, but not introduced deer. Sika deer have been introduced into countries where other deer species are indigenous, such as Canada, Denmark, Great Britain, Czech Republic, Ireland and the United States of America. Because of their dietary advantage, sika deer may have a greater potential to impede forest regeneration and competitively exclude larger deer species, particularly at low basal area sites where impacts on tree regeneration are likely to be greatest.