Effects of red deer on tree regeneration and growth in Aorangi Forest, Wairarapa

New Zealand forests have been substantially modified by introduced red deer over the past century. New Zealand’s indigenous forest managers need to know if regeneration of palatable tree species can be restored following control or eradication of browsing ungulates. Aorangi Forest, Wairarapa, suffered dramatic changes in forest understorey composition by the 1950s after more than seven decades of colonisation by red deer (Cervus elaphus), feral goats (Capra hircus) and pigs (Sus scrofa). Since then feral goats have been eradicated, but red deer and pigs persist throughout Aorangi Forest despite ongoing recreational hunting. This study uses data from paired fenced and unfenced plots established at seven sites in Aorangi Forest between 1981 and 1987, and re-measured in 2004, to show the effects of ungulates on tree (≥ 2 cm diameter at breast height) regeneration and growth. Our results show that browsing by red deer has prevented regeneration of kanono (Coprosma grandifolia), a highly palatable, fast-growing sub-canopy hardwood tree. Deer reduced the growth of mahoe (Melicytus ramiflorus) trees, probably by directly browsing epicormic shoots. The regeneration of other less palatable sub-canopy trees (e.g. porokaiwhiri, Hedycarya arborea), and slower-growing canopy species (e.g. hinau, Elaeocarpus dentatus and rewarewa, Knightia excelsa) appears to have been unaffected by deer browsing. Accordingly, tree species composition does not appear to have been greatly affected by browsing from deer populations in Aorangi Forest over the past two decades.     

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.     

Introduced red deer reduce tree regeneration in Pureora Forest, central North Island, New Zealand

This study uses data from repeatedly measured forest monitoring plots (20 × 20 m) (n = 32) and nine ungulate exclosures (paired fenced and unfenced plots; 20 × 20 m) to show the effects of introduced ungulates on tree regeneration in Pureora Forest Park, central North Island, between 974 and 2002. Results show that introduced ungulates, particularly red deer (Cervus elaphus scoticus), have suppressed the regeneration of hardwood species such as Coprosma grandifolia, Elaeocarpus dentatus, Griselinia littoralis, Melicytus ramiflorus, Schefflera digitata and Weinmannia racemosa. These species were only common as saplings and small trees in the complete absence of ungulate browsing. The results of this study suggest that red deer will need to be culled to low densities to assure regeneration of palatable tree species in Pureora Forest.     

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).     

Windows of opportunity: white-tailed deer and the dynamics of northern hardwood forests of the northeastern US

Herbivory, lighting regimes, and site conditions are among the most important determinants of forest regeneration success, but these are affected by a host of other factors such as weather, predation, human exploitation, pathogens, wind and fire. We draw together >50 years of research on the Huntington Wildlife Forest in the central Adirondack Mountains of New York to explore regeneration of northern hardwoods. A series of studies each of which focused on a single factor failed to identify the cause of regeneration failure. However, integration of these studies led to broader understanding of the process of forest stand development and identified at least three interacting factors: lighting regime, competing vegetation and selective browsing by white-tailed deer (Odocoileus virginianus). The diverse 100–200 year-old hardwood stands present today probably reflect regeneration during periods of low deer density (<2.0 deer/km²) and significant forest disturbance. If this hypothesis is correct, forest managers can mimic these “natural windows of opportunity” through manipulation of a few sensitive variables in the system. Further, these manipulations can be conducted on a relatively small geographic scale. Control of deer densities on a scale of 500 ha and understory American beech (Fagus grandifolia) on a scale of <100 ha in conjunction with an even-aged regeneration system consistently resulted in successful establishment of desirable hardwood regeneration.     

Remnant Trees Affect Species Composition but Not Structure of Tropical Second-Growth Forest

Remnant trees, spared from cutting when tropical forests are cleared for agriculture or grazing, act as nuclei of forest regeneration following field abandonment. Previous studies on remnant trees were primarily conducted in active pasture or old fields abandoned in the previous 2–3 years, and focused on structure and species richness of regenerating forest, but not species composition. Our study is among the first to investigate the effects of remnant trees on neighborhood forest structure, biodiversity, and species composition 20 years post-abandonment. We compared the woody vegetation around individual remnant trees to nearby plots without remnant trees in the same second-growth forests (“control plots”). Forest structure beneath remnant trees did not differ significantly from control plots. Species richness and species diversity were significantly higher around remnant trees. The species composition around remnant trees differed significantly from control plots and more closely resembled the species composition of nearby old-growth forest. The proportion of old-growth specialists and generalists around remnant trees was significantly greater than in control plots. Although previous studies show that remnant trees may initially accelerate secondary forest growth, we found no evidence that they locally affect stem density, basal area, and seedling density at later stages of regrowth. Remnant trees do, however, have a clear effect on the species diversity, composition, and ecological groups of the surrounding woody vegetation, even after 20 years of forest regeneration. To accelerate the return of diversity and old-growth forest species into regrowing forest on abandoned land, landowners should be encouraged to retain remnant trees in agricultural or pastoral fields.     

Interactive effects of introduced herbivores and post‐flowering die‐off of bamboos in Patagonian Nothofagus forests

Question: In November 2000, Chusquea culeou, a bamboo species dominating Andean forest understories in southern Argentina and Chile, massively flowered and died over a north‐south distance of ca. 120 km. Because bamboo is the major forage for large herbivores in these forests, we examined the interactive influences of the bamboo die‐off and herbivory by introduced cattle on understory and tree regeneration. Location: Lanín National Park, Argentina. Methods: Permanent plots, in and outside livestock exclosures, were installed in a Nothofagus dombeyi forest in patches of flowered and non‐flowered C. culeou. Plots were monitored over four years for changes in understory composition and tree seedling densities and heights. Results: After the C. culeou die‐off, new establishment of N. dombeyi was low, both with and without herbivory. Livestock alone directly increased N. dombeyi seedling mortality through physical damage. However, tree seedling browse ratings and height growth were interactively affected by bamboo flowering and herbivory; unfenced plots in flowered bamboo patches had the shortest seedlings, highest browse ratings, and lowest tree seedling annual growth rates. Understory cover was higher where livestock were excluded, and this effect was intensified in the patches of flowered bamboo. Neither herbivory nor bamboo flowering resulted in major changes in species composition, with the exception of Alstroemeria aurea. Conclusion: Effects of livestock on N. dombeyi regeneration were contingent on flowering of C. culeou. Prior to introduction of livestock, N. dombeyi regeneration was probably successful beneath canopy gaps during windows of opportunity following bamboo die‐off, but now livestock impede tree regeneration. Herbivory during bamboo withering periods also produces more open understories, particularly affecting palatable heliophyllous herb species such as Alstroemeria aurea. The results underscore the importance of assessing herbivore impacts on tree regeneration during relatively short periods of potential tree regeneration immediately following rare bamboo flowering and die‐off.     

昆明西山半湿润常绿阔叶林及其〖HJ*3〗〖HJ*2〗次生演替群落的更新特征

对昆明西山国家森林公园处于不同演替阶段的云南油杉林、云南油杉-滇青冈混交林和滇青冈林内木本植物幼苗的种类组成、数量及更新方式等进行了调查分析。结果表明:在所调查的共144个25m2的样方中,共记录到木本植物幼苗32科45属共49种。随着演替的进行,幼苗总密度表现出先增加后降低的趋势,处于演替中期的云南油杉-滇青冈混交林内幼苗最丰富,平均密度达80株.25m-2。在不同演替阶段各植被类型中,实生和萌生两种更新方式同时存在,单一的更新方式可能使群落的更新面临较大的风险。幼苗发生了顶死或梢枯后,从其根颈处生出多个萌枝形成多干基株的更新方式结合了实生和萌生更新各自的优点,是幼苗在与环境长期作用过程中进化出的一种有效对策。     

Landscape structure affects food quality of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) evidenced by fecal nitrogen levels

Evaluating the quality of wildlife habitat is essential for understanding and predicting population dynamics in heterogeneous environments. We used fecal nitrogen levels as an indicator of habitat quality of sika deer (Cervus nippon) and explored important landscape elements influencing nitrogen levels, taking deer density into account. We established 92 plots differing in deer density and landscape structure on the Boso Peninsula, central Japan, and collected fecal samples along a 1-km transect at each plot. The regression models involving two independent variables, i.e., deer density and the length of forest edge within an area of 100 or 200 m from the transect, were selected based on the Akaike’s Information Criterion (AIC). Levels of fecal nitrogen were positively correlated with the length of the forest edge and negatively correlated with population density of deer. The area of 100 or 200 m from the transect most likely reflected the behavioral scale of the deer. Coverage of palatable understory vegetation increased with proximity to forest edge and decreased with deer density. Variability in the level of fecal nitrogen could thus be explained by food availability in the landscape. These results suggest that landscape alterations increase the carrying capacity of sika deer and thereby increase impacts upon the ecosystem.     

Long-term change in growth, mortality and regeneration of trees in Denny Wood, an old-growth wood-pasture in the New Forest (UK)

Long-term changes in stand composition and structure were recorded in Denny Wood (New Forest, UK) by means of a permanent transect covering 2 ha. Denny is an ancient, mixed deciduous wood-pasture dominated by beech (Fagus sylvatica), pedunculate oak (Quercus robur) and holly (Ilex aquifolium) whose canopy trees ranged in age from approximately 70 years to over 300 years when the study began in 1956. Individual trees, shrubs and saplings were mapped and measured at irregular intervals until 1996.

During the 40 years of observations, storms and drought disrupted the stand. Considerable volumes of dead wood accumulated, and canopy gaps extended to 30% of the transect area. Small trees and saplings were severely damaged by ponies and grey squirrels. Regeneration ceased after 1964, due principally to heavy grazing and browsing by deer and ponies. Despite the disturbances, most mortality was due to competitive exclusion within well-stocked parts of the stand.

Historical records from the 17th century onwards demonstrate a long-term change from oak dominance with groups of beech before 1800 to beech dominance in the late 20th century. The stand through which the transect now runs was enclosed in 1870, and this allowed beech to regenerate abundantly, but in the nearby unenclosed part of Denny Wood holly regenerated more abundantly than beech.

The patterns of growth, mortality and regeneration are compared with natural temperate deciduous woodland. The long-term relationship between beech and oak is likely to involve periodic oak regeneration after major disturbances, interspersed with steady increases in the proportion of beech. The implications for managing and monitoring the “Ancient and Ornamental Woods” of the New Forest are considered.     

Early-stage of invasion by beech bark disease does not necessarily trigger American beech root sucker establishment in hardwood stands

Two concomitant phenomena currently affect the dynamics of sugar maple-American beech (AB) stands in northeastern North America: beech bark disease (BBD), and increased AB understory density. Many studies suggest a causal link between the two phenomena, i.e., BBD favouring beech regeneration. But this link has yet to be experimentally demonstrated. To address the question, we compared regeneration composition between recently BBD-affected and -unaffected stands. A total of 109 stands were sampled; half were affected by BBD. Seedling and sapling density were assessed, together with the origin (seedling or sprout). While BBD affects stands in the eastern part of the study region, AB was observed in the understory across the entire region. No clear difference in AB sprout density between BBD-affected and -unaffected stands was observed while AB seedling density—as well as pooled AB seedling and sprout density were higher in unaffected stands. Findings suggests that BBD, in its early stage, is not a necessary trigger of AB understory establishment. Yet, AB sapling basal area generally was higher in stands affected by BBD, likely indicating a greater rate of AB understory development due to increased light availability beneath a more open crown canopy. That development can lead to AB understory dominance. This distinction—BBD not necessarily triggering AB root sucker establishment but favoring AB advance regeneration development—also questions the generalized perception that dense AB thickets necessarily originate from root suckers.     

Diet of ship rats following a mast event in beech (

Diet of 98 ship rats (Rattus rattus) was investigated by examination of stomach contents. Rats were trapped (as bycatch in stoat traps) from June to December 2006, in the Dart Valley near Lake Wakatipu, New Zealand, after a Nothofagus beech mast. Plant material was found in 43% of all stomachs, invertebrate fragments in 67%, feathers in 8%, hairs in 46% and skin with hairs attached in 12%, and animal (vertebrate and invertebrate) tissue in 76%. Hair samples examined by scanning electron microscopy were from mice, and rat and/or mouse DNA was found in all (10) samples subjected to DNA extraction. Mice may be an important food resource for rats following beech masts, when mouse population densities are high. Presence of feathers indicates predation of roosting or nesting birds, or scavenging. Consumption of beech seed beyond the time of germination may be due to caching of seed.     

Disturbance of forest by trampling: Effects on mycorrhizal roots of seedlings and mature trees of Fagus sylvatica

The effects of disturbance by recreational activities (trampling) on changes in soil organic matter (SOM) and on mycorrhizal roots of seedlings and mature trees were studied in four stands of a beech (Fagus sylvatica L.) forest near Basel, Switzerland. At each site, comparable disturbed and undisturbed plots were selected. Disturbance reduced ground cover vegetation and leaf litter. Beech seedlings had lower biomass after disturbance. Ergosterol concentration in seedling roots, an indicator of mycorrhizal fungi, was lower in two of the four disturbed plots compared to undisturbed plots; these two disturbed sites had especially low litter levels. Based on ergosterol measurements, mycorrhizas of mature trees did not appear to be negatively affected by trampling. Total fine roots and SOM were higher in the disturbed than in the undisturbed plots at three sites. At the fourth site, fine roots and SOM in the disturbed areas were lower than in the undisturbed areas most probably due to nutrient input following picnic activities. Principal component analysis revealed a close correlation between SOM and fine roots of mature trees as well as litter and seedling biomass. Trampling due to recreational activities caused considerable damage to the vegetation layer and in particular to the beech seedlings and their mycorrhizal fine roots, whereas, roots of mature trees were apparently resilient to trampling.     

Positive interactions between herbivores and plant diversity shape forest regeneration

The effects of herbivores and diversity on plant communities have been studied separately but rarely in combination. We conducted two concurrent experiments over 3 years to examine how tree seedling diversity, density and herbivory affected forest regeneration. One experiment factorially manipulated plant diversity (one versus 15 species) and the presence/absence of deer (Odocoileus virginianus). We found that mixtures outperformed monocultures only in the presence of deer. Selective browsing on competitive dominants and associational protection from less palatable species appear responsible for this herbivore-driven diversity effect. The other experiment manipulated monospecific plant density and found little evidence for negative density dependence. Combined, these experiments suggest that the higher performance in mixture was owing to the acquisition of positive interspecific interactions rather than the loss of negative intraspecific interactions. Overall, we emphasize that realistic predictions about the consequences of changing biodiversity will require a deeper understanding of the interaction between plant diversity and higher trophic levels. If we had manipulated only plant diversity, we would have missed an important positive interaction across trophic levels: diverse seedling communities better resist herbivores, and herbivores help to maintain seedling diversity.     

Gruppo VI: Bloenergetlca e Fotosintesi

Abstract

Understanding forest stand structural change is crucial for forest ecosystem research and management. The aim of our research was to identify structural and compositional changes and possible natural and anthropogenic causes of these changes in an old‐growth silver fir‐beech forest in the Dinaric mountains of Slovenia. Data on stand structure of the Rajhenavski Rog (51.14 ha) forest reserve originated from traditional methods, including five successive full callipering (1957–2007), two regeneration inventories and analysis of developmental phases (1984, 2007). A drastic decrease in the silver fir proportion in the last 50 years was probably caused by two indirect anthropogenic factors: polluted air and heavy browsing damage of silver fir regeneration. Nevertheless, other factors including reciprocal replacement of fir and beech and climate change also influenced the species composition, as similar processes were documented in the past with less air pollution. Furthermore, the alternation of fir and beech often developed asynchronously, regardless of ungulate density. Although the vertical profile of the old‐growth forest was opened due to the loss of fir trees during past decades, the establishment of more light‐demanding species was uncommon, primarily because the growing stock and forest climate were sustained. The results suggest a high stability of mixed Dinaric old‐growth forest ecosystems.     

Influence of site conditions on the survival of Fagus sylvatica seedlings in an old‐growth beech forest

Abstract. Following a full mast production in autumn 1995 in the old‐growth beech forest of la Tillaie (France), cupules, live and dead 1‐yr old seedlings were counted in September 1996 in 40 plots, representing a wide range of ecological conditions. The influence of geomorphology and forest stage on mast production and on seedling establishment and survival was studied. Maximum seedling abundance did not occur in areas with the largest cupule abundance, but this result was not statistically significant. The presence of a shallow sandstone table under Fontainebleau sand, where beech is far from its ecological optimum, positively influenced the production of mast but negatively influenced seedling survival. Contrary to expectation, the early biostatic rather than the late biostatic phase seemed to be the best stage for beech regeneration. Gaps also provide good environmental conditions for seedling establishment. Humus and light conditions, water availability and competition between beech individuals may explain the results. Light was the main factor influencing mast production in mature stages, and drought was an important factor causing seedling mortality. This study points out the importance of seedling survival for regeneration patterns in beech forests.     

Inter-specific competition from fallow deer Dama dama reduces habitat quality for the Italian roe deer Capreolus capreolus italicus

The roe deer of Mediterranean habitats in the central and southern parts of Italy has recently been recognised as a distinct subspecies, Capreolus capreolus italicus . A population of this endangered subspecies has been monitored in the Preserve of Castelporziano, near Rome, since 1988. We observed an abrupt population decline in 2000, which may severely threaten the sustainability of this population. We evaluated the hypothesis that competition by fallow deer may be a principal cause of this decline. By a new and innovative methodology, we modelled the spatial distribution of fallow deer density (FDD) in the study area to show that 1) habitat quality for roe deer was an inverse function of FDD, 2) habitat apportionment between fallow and roe deer increased as a function of FDD and – by applying structural equation modelling – 3) FDD was superior to habitat composition in explaining observed variations in home range size and probably in habitat quality for roe deer. This analysis is the first to document that inter-specific competition may influence the spatial behaviour of a deer species leading to poor phenotypic performance in the inferior competitor. We conclude that the conservation of this relict population would benefit by reducing fallow deer numbers at Castelporziano and from other measures aimed to decrease the level of inter-specific competition.     

Seedling regeneration,environment and management in a semi‐deciduous African tropical rain forest

Questions: How is seedling regeneration of woody species of semi‐deciduous rain forests affected by (a) historical management for combinations of logging, arboricide treatment or no treatment, (b) forest community type and (c) environmental gradients of topography, light and soil nutrients? Location: Budongo Forest Reserve, Uganda. Methods: Seedling regeneration patterns of trees and shrubs in relation to environmental factors and historical management types were studied using 32 0.5‐ha plots laid out in transects along a topographic gradient. We compared seedling species diversity, composition and distribution patterns along topographic gradients and within types of historical management regimes and forest communities to test whether environmental factors contributed to differences in species composition of seedlings. Results: A total of 85 624 woody seedlings representing 237 species and 46 families were recorded in this rain forest. Cynometra alexandri C.H. Wright and Lasiodiscus mildbraedii Engl. had high seedling densities and were widely distributed throughout the plots. The most species‐rich families were Euphorbiaceae, Fabaceae, Rubiaceae, Meliaceae, Moraceae and Rutaceae. Only total seedling density was significantly different between sites with different historical management, with densities highest in logged, intermediate in logged/arboricided and lowest in the nature reserve. Forest communities differed significantly in terms of seedling diversity and density. Seedling composition differed significantly between transects and forest communities, but not between topographic positions or historical management types. Both Chao‐Jaccard and Chao‐Sørensen abundance‐based similarity estimators were relatively high in the plot, forest community and in terms of historical management levels, corroborating the lack of significant differences in species richness within these groups. The measured environmental variables explained 59.4% of variance in seedling species distributions, with the three most important being soil organic matter, total soil titanium and leaf area index (LAI). Total seedling density was positively correlated with LAI. Differences in diversity of >2.0 cm dbh plants (juveniles and adults) also explained variations in seedling species diversity. Conclusions: The seedling bank is the major route for regeneration in this semi‐deciduous tropical rain forest, with the wide distribution of many species suggesting that these species regenerate continuously. Seedling diversity, density and distribution are largely a function of adult diversity, historical management type and environmental gradients in factors such as soil nutrient content and LAI. The species richness of seedlings was higher in soils both rich in titanium and with low exchangeable cations, as well as in logged areas that were more open and had a low LAI.     

An experimental study of the impacts of understorey forest vegetation and herbivory by red deer and rodents on seedling establishment and species composition in Waitutu Forest, New Zealand

Introduced mammalian herbivores are changing the structure and composition of New Zealand’s forest ecosystems and may modify forest succession after natural disturbances. We studied how introduced ungulates (red deer and feral pigs) and rodents (rats and house mice) affected the rate of recovery (i.e. the engineering resilience) of the forest understorey following artificial disturbance. We imposed disturbances by clearing understorey vegetation dominated by Blechnum ferns in forests on relatively fertile alluvium and elevated infertile marine terraces, and recorded recovery of vegetation (seedling establishment, species composition, cover and volume) in herbivore exclosures and controls. Seedlings quickly established on cleared plots: after 2 years, numbers of woody seedlings and ground cover of vascular plants relative to initial values were similar on cleared and uncleared treatments. Volume of plant biomass <2 m remained low on cleared subplots. Ungulates significantly reduced the re-establishment of woody seedlings ≥ 10 cm tall: only one seedling reached this height outside exclosures, compared with 29 seedlings inside. The number of seedlings <10 cm tall, expressed relative to numbers present pre-clearing, was not significantly affected by ungulates. The species composition of regenerating vegetation was more similar (Jaccard index) to pre-clearing understorey vegetation inside ungulate exclosures than outside. No consistent effect of rodents (primarily house mice) on seedling establishment or species composition was detected after 2 years, and rodent exclosures did not significantly affect survival of seedlings (Griselinia littoralis and Aristotelia serrata) planted as an index of rodent herbivory pressure. No significant differences in vegetation recovery were apparent between forest types. Rapid seedling recruitment in the absence of understorey vegetation and the presence of herbivores provided evidence that understorey vegetation competes with seedlings for light. Ungulate effects were consistent with other experiments that showed herbivores reduced the rate and altered the trajectory of vegetation regrowth after disturbance.