Landscape of fear in Europe: wolves affect spatial patterns of ungulate browsing in Białowieża Primeval Forest,Poland

Large carnivores can either directly influence ungulate populations or indirectly affect their behaviour. Knowledge from European systems, in contrast to North American systems, on how this might lead to cascading effects on lower trophic levels is virtually absent. We studied whether wolves Canis lupus via density‐mediated and behaviorally‐mediated effects on their ungulate prey species influence patterns of browsing and tree regeneration inside the Bia?owie?a National Park, Poland. Browsing intensity of tree saplings (height class < 150 cm), irrespective of tree species or forest type, was lower inside a wolf core area (50.5%) where predator presence is highest, than in the remainder of the wolf pack’s home range (58.3%). Additionally, browsing intensity was reduced when the amount of coarse woody debris (CWD), which can act as a ?ungulate escape impediment?, increased (within 5‐m radius) inside the wolf core area. No relationship existed outside the core area. As a result, the proportion of trees growing out of herbivore control increased more strongly with increasing amount of CWD inside compared to outside the wolf core area. This suggests that next to direct effects of wolves on ungulate density caused by a higher predation pressure inside the core area, risk effects are important and are enhanced by habitat characteristics. These results indicate that behaviorally‐mediated effects of predators on prey can become more important than density‐mediated effects in affecting lower trophic levels. This is the first study we are aware of, that shows CWD can create fine‐scale risk effects on ungulates with the potential for cascading effects of large predators on patterns of tree regeneration for a European forest system. This knowledge broadens the discussion on how the impact of large predators on ecosystem functioning depends on the physical landscape, by illustrating these effects for a system which largely contrasts in this respect to the North American systems.     

Managing for naturalness alone is not an effective way to preserve all the valuable natural features of the Białowieża Forest – a reply to Jaroszewicz et al.

Currently, Brzeziecki et al. 2016 (Journal of Vegetation Science 27: 460–467.) are using data from permanent study plots established in 1936 in Bia?owie?a National Park (NE Poland) to develop theoretical equilibrium tree size distributions and to then compare modelled and actual distributions with a view to assessing the population dynamics of the species involved. As part of their discussion, the authors address the question of possible consequences for the overall diversity of forest ecosystems under strict protection if long‐term trends relating to tree population densities and size structures are maintained. In the overall context of the above, the goal of the present paper is to respond to Jaroszewicz et al. (Journal of Vegetation Science 28: 218–222.) who suggest that the paper of Brzeziecki et al. (2016) is not representative for the whole Bia?owie?a National Park, and that – in this connection – strict protection should not be seen as a cause for concern. In this paper, we show that the data analysed by Brzeziecki et al. (2016) adequately characterize conditions in the wider Park. We also point out that the thorough scientific understanding of the long‐term dynamics of woodland communities under strict protection should indeed be taken into account as efforts are made to arrive at an effective conservation strategy capable of ensuring that the uniquely valuable features of the Bia?owie?a Forest are retained.     

Lack of demographic equilibrium indicates natural,large‐scale forest dynamics,not a problematic forest conservation policy – a reply to Brzeziecki et al.

Brzeziecki et al. 2016 (Journal of Vegetation Science 27: 460–467.) describe a decrease in population densities and proportion of younger individuals for several tree species in permanent research plots in the core zone of the Bia?owie?a National Park. They attribute insufficient tree recruitment inter alia to the strict protection. Although the authors performed a thorough analysis of tree population dynamics, the scales of the study mean that their far‐reaching conclusions on the causes and consequences of the lack of demographic equilibrium cannot be supported by the data. The inadequate spatial and temporal scales of the study did not allow for the observation of representative population dynamics. Furthermore, they did not compare the results obtained in the strictly protected area with known demographic dynamics of trees in surrounding managed forests or under other forms of nature conservation. Looking from the wider ecosystem perspective, it is clear that strict protection is not a cause for concern and, instead, that such a near‐natural forest manifests population dynamics at rather larger scales.     

Density and biomass of ungulates in Tsavo East National Park, Kenya

In view of the substantial vegetation changes in Tsavo National Park and their possible effects on the herbivore fauna, this study was undertaken to obtain quantitative information on densities of several ungulate species and their biomass. Road strip counts (belt transects) constituted the main method used, supplemented by aerial counts in parts of the study area. Densities of most ungulate species are rather low, the maximum of 2–25 per km² being attained by zebra in the southern poition of the study area. There are significant differences between two parts of the study area, related to differences in vegetation types. Mean biomass density is around 4000 kg/km², of which ?-¾ is made up of elephants. Total biomass density compares well with that in areas of similar climatic conditions but is much lower than that in moist savannas of western Uganda and Zaire with higher rainfall.     

Defecation rate in captive European bison, Bison bonasus

Data on the digestive characteristics of European bison, Bison bonasus (L.), are needed for studies of their role as the largest extant herbivore in Europe and a potential keystone species of the temperate forest ecosystem. Very little published data are available, particularly on the defecation rate which affects population estimates from dropping counts and also the individual seed deposition rate. We gathered data from a captive bison group kept at the Show Reserve of the Bia?owie?a National Park. Droppings accumulated in the enclosure over a 72-h period were counted in winter 2010. In addition, the group was observed over approximately 6-h periods three times in winter and 16 times in summer. The count of accumulated droppings over a 72-h period gave eight defecations per day. The summer direct observations recorded 7.5 defecations per day and winter observation 5.4 defecations per day. These estimates are within the range for other bovids of similar size. The difference between summer and winter observation-based estimates may be accounted for by a higher frequency of defecation in early morning and late afternoon, periods not covered in winter observations. Given the published density of seedlings emerging from droppings of the ～470 free-living bison in the nearby forest, eight defecations a day mean that seed deposition by European bison may contribute significantly to realize seed dispersal and plant establishment.     

The Seedling Ratio Method for determining ungulate impacts on forest understoreys: Utility in an Australian ecosystem

The Seedling Ratio Method was devised to assess the impacts of introduced ungulates on plant species richness in forest understoreys. The method has successfully assessed ungulate impacts on species richness in forests in New Zealand and Hawaii, which do not have native herbivores. We tested the three critical assumptions that underlie this method to investigate its potential for use in an Australian ecosystem. This study was conducted in the Yarra Ranges National Park (YRNP), Victoria, which had a high‐density population of an introduced deer species, Sambar (Cervus unicolor), in addition to several native herbivore species. One of the three key assumptions of the Seedling Ratio Method was not supported, whereas conventional differential exclosures showed clear and separate impacts by Sambar and native herbivores. We conclude that the Seedling Ratio Method could not provide a clear indication of browsing impact on forest understoreys in YRNP.     

The pattern of Dunlin Calidris alpina distribution and abundance in relation to habitat variation in the Flow Country of northern Scotland

Capsule Folivorous caterpillars constituted the majority of nestlings’ food in a primeval forest. Blue Tit broods only partially matched the caterpillar peak, and the mismatch did not affect food composition or nesting success.

Aims To describe factors influencing the timing of reproduction in Blue Tits under primeval conditions (Bia?owie?a National Park, Poland) and to check whether they schedule breeding so as to synchronize broods with a seasonal caterpillar peak.

Methods We gathered information on phenology of leaf development, seasonal availability of folivorous caterpillars (frass collection), timing of Blue Tit breeding, composition of its nestling food, and nest fate over a three-year period.

Results Caterpillars constituted c. 74% of nestling diet, but only 17–65% of broods matched the caterpillar peak in any season. Neither total nest loss, nor frequency of brood reduction depended on the level of mismatch. Caterpillar availability was probably adequate every year, regardless of the amount of mismatch, and no selective advantage of precise matching was detectable. Phenological events at all trophic levels occurred earlier in warmer springs. Egg-laying coincided with tree bud burst and appearance of caterpillars, but was not critically dependent on their timing.

Conclusion The observations are consistent with the view that Blue Tits under primeval conditions in Bia?owie?a National Park, Poland, breed as early as possible, rather than synchronizing their breeding with the caterpillar peak later in the season.     

Timing of breeding and nestling diet of Wood Warbler Phylloscopus sibilatrix in relation to changing food supply

Capsule Wood Warblers did not match their reproduction to the caterpillar peak

Aims To study the timing of Wood Warblers’ breeding and nestling diet in relation to caterpillar abundance under primeval conditions in the Bia?owie?a National Park, Poland.

Methods Observations of food brought by parents. Inter‐year and seasonal changes in availability of folivorous caterpillars were assessed by direct counts and caterpillar frass collection.

Results Maximum food requirements occurred two weeks after the peak of caterpillar abundance. The mismatch had no effect on nestlings’ development. Diet varied little across years and habitats, but varied strongly within a season. Following the decline of ‘green’ caterpillars, their proportion in the Wood Warbler diet strongly declined. Caterpillars were replaced by winged insects. Small nestlings received more spiders than older ones.

Conclusion Timing of Wood Warbler breeding in Bia?owie?a National Park was constrained by the females’ arrival time. Birds did not match their reproduction to the caterpillar peak. ‘Green’ caterpillars were the preferred food for nestlings; birds responded to the caterpillar decrease by feeding smaller proportions of them and switching to alternative prey, namely winged insects.     

Dark tree cavities – a challenge for hole nesting birds?

Holes provide the safest nest sites for birds, but they are an underutilized resource; in natural forests there are usually more holes than birds that could use them. Some bird species could be prevented from nesting in holes because of their inability to operate in the low light conditions which occur in cavities. As no visual system can operate in complete darkness some nest cavities could be too dark to be useable even by hole‐nesters. Thus, the light conditions within tree cavities could constrain both the evolution of the hole nesting habit, and the nest site choice of the hole‐nesting birds. These ideas cannot be tested because little is known about the light conditions in cavities. We took an opportunity provided by ongoing studies of marsh tits Poecile palustris and great tits Parus major breeding in a primeval forest (Bia?owie?a National Park, Poland) to measure illumination inside their nest cavities. We measured illuminance in cavities at daybreak, which is just after the parents commenced feeding nestlings. Only ca 1% of incoming light reached the level of the nest. Illuminance at nests of both species (median = 0.1–0.2 lx) fell within mesopic‐scotopic range, where colour vision is impaired. Measurements in model cavities showed strong declines in illumination with distance from the entrance, with light levels typically as low as 0.01 lx at 40 cm from the cavity entrance. Thus cavities can be very dark, often too dark for the use of colour vision, and we suggest that ‘lighting’ requirements can affect the adoption of specific nest sites by hole nesting birds. We discuss implications of the findings for understanding the adaptations for hole‐breeding in birds.     

Wood ants prefer conifers to broadleaved trees in mixed temperate forests

1. Wood ants, species of the subgenus Formica s. str., are one of the most important groups of insects in forest ecosystems due to their high abundance and activity.
2. We assessed the foraging pressure of Formica polyctena and Formica rufa, two dominant wood ant species in the Białowieża Forest. In addition, we compared coniferous and mixed stands in these respects.
3. In mixed stands F. polyctena visited more trees per plot than F. rufa. F. polyctena might forage a larger area due to higher population sizes, which further confirms higher foraging pressure of F. polyctena in comparison with F. rufa.
4. In our study area, host trees of the ant–aphid mutualism were larger in diameter than nonhost trees. The ants also preferred trees situated closer to their nests. In both mixed and coniferous stands, the most visited tree species was Norway spruce. Thus, we can conclude that Norway spruce seems to be preferred by the ant. The decline of living spruce in the Białowieża Forest may endanger their food source and impact the vitality of ant colonies.

     

Genes of the extinct Caucasian bison still roam the Białowieża Forest and are the source of genetic discrepances between Polish and Belarusian populations of the European bison,Bison bonasus

European bison (Bison bonasus) populations from both the Polish (PL) and the Belarusian (BY) sides of the Bia?owie?a Forest represent the Lowland genetic line (LB line) – progeny of the Lowland bison (Bison bonasus bonasus) that inhabited western, central, and south‐eastern Europe in historical times. During the species recovery, one of the founders was a descendant of the extinct Caucasian bison (Bison bonasus caucasicus) and its descendants formed the other genetic line – Lowland–Caucasian (LC). There have been justified suspicions that LB European bison in the former Soviet Union had undergone cross‐mating with the LC line. We performed a comparative genetic analyses on European bison from the BY and PL parts of the Bia?owie?a Forest, the LC line and extinct Caucasian bison, based on a set of 19 microsatellite markers and 1512 bovine single nucleotide polymorphism (SNP) markers, polymorphic in at least one of the studied populations. Although genetic variability (mean allele number and expected heterozygosity) for both populations were similar, the F_ST jack‐knifing and principal component analyses PCA revealed highly significant differences between PL and BY bison from the Bia?owie?a Forest. Examining DNA of the extinct Caucasian bison revealed that at least part of the genetic variants found in the BY, but not the PL, population were of Caucasian origin. The results indicate that the contemporary population of European bison from the BY part of the Bia?owie?a Forest should not be regarded as a LB line. The results also suggest that the actual global population size of the LB line European bison is only a half of its official status. Consideration of the presented results are crucial in determining management actions and policy decisions in order to conserve LB line bison within the Bia?owie?a Forest – its natural refuge. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 752–763.     

Effect of selective logging on tree and understory regeneration in miombo woodland in western Tanzania

Tree regeneration and understory response to selective cutting of Pterocarpus angolensis DC, and Sterculia quinqueloba Sim, was measured outside Katavi National Park, Tanzania. Contrary to expectations, a selective harvest had no effect on tree recruitment for either species in this miombo woodland. In unlogged plots along transects taken at increasing distance from a main road, the stand densities of young trees declined away from the road. Since anthropogenic activity along the road results in a high fire frequency but low native ungulate densities, the increased rate of tree regeneration near the road may result from either reduced grazing and browsing pressure by native ungulates, or reduced fire intensity and severity along roads. These effects appear to mask any possible effect of selective harvest on tree recruitment. Currently, there is no evidence of compensatory recruitment of trees of any species into the canopy to replace logged trees and this will lead to a gradual thinning in overstory stand density.     

Food supply and poaching limit giraffe abundance in the Serengeti

The iconic giraffe, an ecologically important browser, has shown a substantial decline in numbers across Africa since the 1990s. In Serengeti National Park, Tanzania, giraffes reached densities of 1.5–2.6 individuals km⁻² in the 1970s coincident with a pulse of Acacia tree recruitment. However, despite continued increases in woody cover between the 1980s and the 2000s, giraffe recruitment and survival rates have declined and density has dropped to only 0.3–0.4 giraffes km⁻². We used a decision table to investigate how four extrinsic factors may have contributed to these declines: food supply, predation, parasites, and poaching, which have all been previously shown to limit Serengeti ungulate populations. Lower recruitment likely resulted from a reduction in diet quality, owing to the replacement of preferred trees with unpalatable species, while decreased adult survival resulted from illegal harvesting, which appears to have had a greater impact on giraffe populations bordering the western and northern Serengeti. The Serengeti giraffe population will likely persist at low-to-moderate densities until palatable tree species regain their former abundance. Leslie matrix models suggest that park managers should meanwhile redouble their efforts to reduce poaching, thereby improving adult survival.     

Limits of pine forest distribution at the treeline: herbivory matters

Herbivores can affect future forest composition by feeding selectivity. At temperature-sensitive treelines, herbivory can exacerbate or constrain climate-driven distributional shifts in tree species. This study analyses the impact of herbivory in a Mediterranean treeline of widespread Pinus sylvestris and P. nigra pinewoods, testing whether herbivory damage reinforces or inhibits the climatic responses of these trees. We used naturally occurring sapling pairs of similar size and age of both species, thereby isolating plant characteristics from environmental effects in herbivore behaviour. Herbivory damage by ungulates proved higher than that caused by insects in saplings of both species. Low plant density and extreme abiotic conditions at the treeline could in part be responsible for the observed low incidence of insect herbivory. Ungulates preferred P. sylvestris over P. nigra, implying heavier browsing damage for a large number of P. sylvestris saplings, suffering reduced internode growth as a consequence. In addition, P. sylvestris could not compensate height-growth reductions due to browsing with higher growth rate than P. nigra. In fact, P. sylvestris showed similar or lower relative height growth with respect to P. nigra. Under a scenario of increasing aridity and maintenance of ungulate populations, the upward migration of P. sylvestris in its southern range could be restricted by higher drought vulnerability than P. nigra, a situation exacerbated by ungulate herbivory. Our results indicate that ungulate herbivory reinforces climatic response of coexisting P. sylvestris and P. nigra at treeline, favouring a potential change in community dominance towards Mediterranean P. nigra.     

Mesophication in temperate Europe: A dendrochronological reconstruction of tree succession and fires in a mixed deciduous stand in Białowieża Forest

1. The shift from shade‐intolerant species to shade‐tolerant mesophytic species in deciduous and mixed forests of the temperate zone is well described in studies from North America. This process has been termed mesophication and it has been linked to changes in fire regime. Fire suppression results in the cessation of establishment of heliophytic, fire‐dependent tree species such as oak (Quercus) and pine (Pinus). Due to the scarcity of old‐growth forests in Europe, data on long‐term compositional changes in mixed forests are very limited, as is the number of studies exploring whether fire played a role in shaping the dynamics.
2. The aim of this study was to reconstruct tree succession in a 43‐ha natural mixed deciduous forest stand in Bia?owie?a Forest (BF), Poland using dendrochronological methods. In addition, the presence of aboveground fire legacies (charred and fire‐scarred deadwood) enabled the fire history reconstruction.
3. Dendrochronological data revealed tree establishment (Quercus) back to the end of the 1500s and fires back to 1659. Under a regime of frequent fires until the end of the 18th century, only oak and pine regenerated, sporadically. A shift in the fire regime in the first half of the 19th century triggered oak and pine cohort regeneration, then gradually spruce (Picea) encroached. Under an increasingly dense canopy and less flammable conditions, regeneration of shade‐tolerant Carpinus, Tilia, and Acer began simultaneously with the cessation of oak and pine recruitment.
4. Synthesis. The study reports the first evidence of mesophication in temperate Europe and proves that fire was involved in shaping the long‐term dynamics of mixed deciduous forest ecosystems. Our data suggest that fire exclusion promoted a gradual recruitment of fire‐sensitive, shade‐tolerant species that inhibited the regeneration of oak and pine in BF.

     

Movements of European bison (Bison bonasus) beyond the Białowieża Forest (NE Poland): range expansion or partial migrations?

The European bison is the largest terrestrial mammal in Europe. After extinction in the wild at the beginning of the twentieth century, it was re-introduced to Bia?owie?a Forest and other woodlands in Eastern Europe. In this paper, we analysed the movements of European bison beyond the continuous woodlands of the Bia?owie?a Forest (NE Poland) between 1964 and 2010. In total, 1,117 direct observations of bison were collected. The number of males moving out of the forest fluctuated during the study period, whilst the number of females steadily increased. The number of male observations outside of the forest per annum was dependent on the population size and snow depth, whilst the number of cows in mixed groups moving outside of the forest was correlated with the population size only. Males were observed mainly alone (50 % of observations) or in small groups of two to three individuals (25 %); however, distribution of group size differed from those observed in the population. There was a significant difference between the direction of movement of males and females out of the forest—males moved mainly west and southwest, whilst females moved to the north. This was also significantly different from the expected movement direction. The mean distance of bison observations from the forest border was 1.8?±?0.13 km and did not differ significantly between sexes. After 1990, males were observed significantly farther away from the forest (2.2 km) than in the previous years (0.9 km). Most observations (94 % of bulls and 93 % of cows) were up to 5 km from the forest edge. The range of bison in the vicinity of the Bia?owie?a Forest was strictly seasonal. Most observations (78 % in males and 88 % in females) were recorded from November to April. Increasing utilisation of areas beyond the forest habitats may be driven by different factors but most probably it is related to range expansion and the bison's preference for open habitats. The strong seasonal pattern of bison movements indicate that the partial seasonal migrations were initiated in the Bia?owie?a population.     

Managing open habitats by wild ungulate browsing and grazing: A case‐study in North‐Eastern Germany

Question: Can wild ungulates efficiently maintain and restore open habitats? Location: Brandenburg, NE Germany. Methods: The effect of wild ungulate grazing and browsing was studied in three successional stages: (1) Corynephorus canescens‐dominated grassland; (2) ruderal tall forb vegetation dominated by Tanacetum vulgare; and (3) Pinus sylvestris‐pioneer forest. The study was conducted over 3 yr. In each successional stage, six paired 4 m²‐monitoring plots of permanently grazed versus ungrazed plots were arranged in three random blocks. Removal of grazing was introduced de novo for the study. In each plot, percentage cover of each plant and lichen species and total cover of woody plants was recorded. Results: Wild ungulates considerably affected successional pathways and species composition in open habitats but this influence became evident in alteration of abundances of only a few species. Grazing effects differed considerably between successional stages: species richness was higher in grazed versus ungrazed ruderal and pioneer forest plots, but not in the Corynephorus sites. Herbivory affected woody plant cover only in the Pioneer forest sites. Although the study period was too short to observe drastic changes in species richness and woody plant cover, notable changes in species composition were still detected in all successional stages. Conclusion: Wild ungulate browsing is a useful tool to inhibit encroachment of woody vegetation and to conserve a species‐rich, open landscape.     

Forest regeneration following ungulate removal in a montane Hawaiian wet forest

Most Hawaiian forests lack resiliency following disturbance due to the presence of non‐native and invasive plant and animal species. The montane wet forest within Hakalau Forest National Wildlife Refuge on Hawai'i island has a long history of ungulate disturbance but portions of the refuge were fenced and most ungulates excluded by the early 1990s. We examined patterns of regeneration within two 100 ha study sites in this forest following the removal of ungulates and in the absence of invasive woody tree species to determine, in part, if passive restoration techniques can be successful under these conditions. We characterized growth, mortality, and basal area (BA) changes for approximately 7,100 marked individuals of all native tree species present in two surveys over a 17–18‐year period within two hundred 30 m diameter forest plots. Considerable recruitment within plots of new trees of all species significantly changed size class distributions and erased deficits in small‐sized trees observed during the first survey, particularly for the codominant canopy tree, koa (Acacia koa). Overall, growth of established dominant 'ōhi'a trees (Metrosideros polymorpha) and recruitment of mid‐canopy trees contributed to increases in BA while high levels of mortality for large A. koa trees contributed to decreased BA. This resulted in a slight increase in BA between the two surveys (+1.9%). This study demonstrates that fencing and ungulate removal may have rescued the A. koa population by facilitating the first real pulse in recruitment in over a century, and that passive restoration can be a successful management strategy in this forest.     

Long‐term successional forest dynamics: species and community responses to climatic variability

Question: Are trees sensitive to climatic variability, and do tree species differ in their responses to climatic variability? Does sensitivity of forest communities to climatic variability depend on stand composition? Location: Mixed young forest at Walker Branch Watershed near Oak Ridge, East Tennessee, USA. Methods: Using a long‐term dataset (1967–2006), we analyzed temporal forest dynamics at the tree and species level, and community dynamics for forest stands that differed in initial species composition (i.e., chestnut oak, oak–hickory, pine, and yellow poplar stands). Using summer drought and growing season temperature as defined climate drivers, we evaluated relationships between forest dynamics and climate across levels of organization. Results: Over the four‐decade study period, forest communities underwent successional change and substantially increased in biomass. Variation in summer drought and growing season temperature contributed to temporal biomass dynamics for some tree species, but not for others. Stand‐level responses to climatic variability were related to the responses of component species, except in pine stands. Pinus echinata, the dominant species in pine stands, decreased over time due to periodic outbreaks of pine bark beetle (Dendroctonus frontalis). These outbreaks at Walker Branch could not be directly related to climatic conditions. Conclusions: The results indicate that sensitivity of developing forests to climatic variability is stand type‐dependent, and hence is a function of species composition. However, in the long term, direct effects of climatic variability on forest dynamics may be small relative to autogenic successional processes or climate‐related insect outbreaks. Empirical studies testing for interactions between forest succession and climatic variability are needed.     

Tree–Shrub Interactions During Early Secondary Forest Succession in Uganda

Abstract One of the greatest challenges for ecologists this century will be restoring forests on degraded tropical lands. This restoration will require understanding complex processes that shape successional pathways, including interactions between trees and other plants. Shrub species often quickly invade disturbed tropical lands, yet little is known about whether they facilitate or inhibit subsequent tree recruitment and growth. We examined how shrubs and other vegetation (e.g., vines, grasses, herbs) affect tree recruitment, survival, and growth during the first 6 years of forest succession in Kibale National Park, Uganda. The study was undertaken in two recently logged exotic softwood plantations. We studied the successional trajectories in two recently logged areas that varied in their initial densities of trees and shrubs. Analyses suggested tree seedling presence and density were not strongly related to shrub density or height during succession. Tree sapling presence and density were positively significantly related to shrub density and height. We found little response in the tree community to experimental shrub removal, and although removal of all nontree vegetation temporarily enhanced tree growth, the effect disappeared after 2 years. Some early‐successional trees benefited from reduced competition, whereas some mid‐successional trees benefited from the presence of other vegetation. Some specific tree species responded strongly to vegetation removal. We interpret our findings in light of designing manipulations promoting forest restoration for biodiversity conservation and conclude with four tentative guidelines: (1) manage at the species level, not the community level; (2) increase facilitation for seedlings, reduce competition for saplings; (3) be cautious of assumptions about plant interactions; and (4) be adaptable and creative with new strategies when manipulations fail.