Testing a facilitation model for ecosystem restoration: Does tree planting restore ground layer species in a grassy woodland?

Planning for the restoration of degraded ecosystems has a strong basis in facilitation successional theory, which, as applied in restoration practice, states that planting of structurally dominant tree species will assist the entry of other native species into a restored community. In Australia, tree planting has been widely applied in restoration of grassy woodland ecosystems. Trees have been postulated to reduce the cover and diversity of weed species, thus facilitating recolonization of native woodland species (indirect facilitation). The expected outcomes of this process include reduced species richness and abundance of exotic plant species and increased species richness and abundance/dominance of natives in areas beneath tree canopies, with these trends strengthening with time. To assess whether this was occurring, we carried out a comparative analysis of species assemblages found underneath and outside of planted tree canopies in sites replanted with juvenile canopy tree species 3–5 or 8–10 years previously. We sampled revegetated stands of Cumberland Plain Woodland, an endangered ecological community in Western Sydney, Australia. We found that neither the number nor abundance of native ground layer species beneath canopies increased as a result of trees being planted at sites of both ages. Where seed is limited, we predicted an increase in abundance of existing native species under planted tree canopies. On this point, the results were mixed and showed some natives with an increased abundance while others decreased. Exotic species richness showed the reverse of the expected pattern, being greater under tree canopies. These findings lend no support to the theory of indirect facilitation. We conclude that simple facilitation models may be inadequate to support planning of grassy woodland restoration and that those models incorporating successional time lags and restoration barriers are likely to be more informative about the development of communities initiated by tree planting.     

Influence of tree hollow characteristics on the diversity of saproxylic insect guilds in Iberian Mediterranean woodlands

Saproxylic diversity assessment is a major goal for conservation strategies in woodlands and it should consider woodland composition and configuration at site and tree level as key modelling factors. However, in Mediterranean woodlands little is known about the relation with the environmental factors that structure their assemblages, especially those linked to tree hollow microhabitats. We assessed the diversity of Syrphidae (Diptera) and Coleoptera saproxylic guilds that co-occurred in tree hollows located in three different Iberian Mediterranean woodlands in the Cabañeros National Park (Spain). Furthermore, we evaluated how differences in tree hollow microenvironmental variables (understood as the physical and biotic characteristics of a hollow and tree individual) influenced saproxylic guild diversity both within and among woodland sites. We found that woodland sites that provided greater heterogeneity of trees and hollow microhabitats determined higher saproxylic guild diversity. Nevertheless, certain species or even complete guilds can be favoured in woodlands where some hollow microhabitats predominate as a consequence of historical tree management. In general, hollow volume was the main determining factor for saproxylic guild richness and abundance in woodland sites, and large hollow volume was usually related to higher diversity, which highlighted the importance of multi-habitat hollow trees. Moreover, saproxylic guilds also responded to other different microenvironmental variables, which indicated different ecological preferences among guilds. The conservation of saproxylic insects in Iberian Mediterranean areas must be addressed to protect woodland sites that provide high diversity and large numbers of tree hollow microhabitats, and practices to enhance microhabitat heterogeneity should even be encouraged.     

Plant species as indicators of ancient woodland in northwestern Germany

Abstract. 315 isolated semi-natural and natural stands in NW Germany were investigated floristically, 285 stands of ancient woodlands and 30 recent ones. In the study area nearly all semi-natural and natural ancient woodlands are found on mesotrophic or eutrophic sites and can be assigned to the Fagetalia. 54 selected herbaceous and five woody Fagetalia-species have been tested in their association to ancient woodlands. 21 of the 59 selected plant species show a highly significant association to ancient woodlands and seven species show a weak significance. 31 plant species are not significantly correlated with ancient woodland sites, but 25 of them have a low frequency. The result shows that investigations of the historical ecology can help to understand floristic composition of present-day woodlands. Restriction of many woodland species to ancient woodlands, especially rare species, emphasizes the importance of woodlands with a long continuous history for the preservation of endangered species. Irrespective of ecological conditions, the restriction of plant species to ancient woodlands seems to be mainly caused by their low ability to colonize recent woodlands, especially isolated stands. Differences in the association to ancient woodlands between European countries are assumed to be a function of both time and degree of isolation of woodlands, of the dispersal mode of the plant species and of the availability of suitable habitats.     

Fox-hunting in England and Wales: its contribution to the management of woodland and other habitats

Hunting foxes with hounds has been a countryside pursuit in Britain since the 17th Century, but its effect nationally on habitat management is little understood by the general public. A survey questionnaire was distributed to 163 mounted fox hunts of England and Wales to quantify their management practices in woodland and other habitat. Ninety-two hunts (56%), covering 75,514 km², returned details on woodland management motivated by the improvement of their sport. The management details were verified via on-site visits for a sample of 200 woodlands. Following verification, the area of woodlands containing the management was conservatively estimated at 24,053 (±2241) ha, comprising 5.9% of woodland area within the whole of the area hunted by the 92 hunts. Management techniques included: tree planting, coppicing, felling, ride and perimeter management. A case study in five hunt countries in southern England examined, through the use of botanical survey and butterfly counts, the consequences of the hunt management on woodland ground flora and butterflies. Managed areas had, within the last 5 years, been coppiced and rides had been cleared. Vegetation cover in managed and unmanaged sites averaged 86% and 64%, respectively, and managed areas held on average 4 more plant species and a higher plant diversity than unmanaged areas (Shannon index of diversity: 2.25 vs. 1.95). Both the average number of butterfly species (2.2 vs. 0.3) and individuals counted (4.6 vs. 0.3) were higher in the managed than unmanaged sites.     

Where can managers effectively resist climate-driven ecological transformation in pinyon–juniper woodlands of the US Southwest?

Pinyon–juniper (PJ) woodlands are an important component of dryland ecosystems across the US West and are potentially susceptible to ecological transformation. However, predicting woodland futures is complicated by species-specific strategies for persisting and reproducing under drought conditions, uncertainty in future climate, and limitations to inferring demographic rates from forest inventory data. Here, we leverage new demographic models to quantify how climate change is expected to alter population demographics in five PJ tree species in the US West and place our results in the context of a climate adaptation framework to resist, accept, or direct ecological transformation. Two of five study species, Pinus edulis and Juniperus monosperma, are projected to experience population declines, driven by both rising mortality and decreasing recruitment rates. These declines are reasonably consistent across various climate futures, and the magnitude of uncertainty in population growth due to future climate is less than uncertainty due to how demographic rates will respond to changing climate. We assess the effectiveness of management to reduce tree density and mitigate competition, and use the results to classify southwest woodlands into areas where transformation is (a) unlikely and can be passively resisted, (b) likely but may be resisted by active management, and (c) likely unavoidable, requiring managers to accept or direct the trajectory. Population declines are projected to promote ecological transformation in the warmer and drier PJ communities of the southwest, encompassing 37.1%–81.1% of our sites, depending on future climate scenarios. Less than 20% of sites expected to transform away from PJ have potential to retain existing tree composition by density reduction. Our results inform where this adaptation strategy could successfully resist ecological transformation in coming decades and allow for a portfolio design approach across the geographic range of PJ woodlands.     

Bryophytes as indicators of ancient woodlands in Schleswig-Holstein (Northern Germany)

Ancient woodlands, with their long ecological continuity, frequently harbor a high number of typical, rare and threatened species, and are therefore of particular importance for nature conservation. To pinpoint these habitats, a common application is the use of plants as “ancient woodland indicators”. The occurrence of these particular species allows for evaluating the continuity of woodland cover in time. While lists of ancient woodland vascular plants have been derived for many regions, the identification and use of bryophytes as ancient woodland indicators has been widely neglected. This is a bit surprising because certain woodland bryophytes are very sensitive to varying environmental conditions or changes in land management. It therefore appeared promising to compile an ecologically grounded list of ancient woodland indicator bryophytes for practical use.In this study, we present a set of ancient woodland indicator bryophytes based on the analysis of datasets from the North German federal state of Schleswig-Holstein. To compile this list, we systematically evaluated the bryophyte distribution data from floristic surveys in relation to ancient woodland cover data from state-wide inventories. In this way, we were able to determine ancient woodland bryophytes using consistent and repeatable statistical methods.The presented list of 31 ancient woodland indicator bryophytes is ecologically sound and corresponds well with data from the sparse literature. We could distinguish two groups of ancient woodland indicator bryophytes. The first group is linked to base-rich, semi-natural deciduous woodlands with high soil and air humidity. The second group comprises acidophilic bryophytes that occur not only in acidic beech and oak woods, but also in acidic mixed or coniferous forests on ancient woodland sites. Apart from the ancient woodland indicator bryophytes, we could identify one group of recent woodland bryophytes and four groups of bryophytes that are more or less indifferent with respect to woodland continuity.Finally, we provide recommendations for the application of ancient woodland indicator bryophytes in nature conservation practice. Management suggestions for the conservation of the typical bryophyte diversity of ancient semi-natural woodlands are also given.     

Woodland structure,rather than tree identity,determines the breeding habitat of Willow Warblers Phylloscopus trochilus in the northwest of England

Capsule Woodland structure, rather than tree species, is the most important determinant of breeding habitat selection by Willow Warblers in North West England.

Aims To examine how habitat characteristics predict the occurrence of male Willow Warbler territories.

Methods Woodland structure (trunk density, trunk diameter, canopy cover and understory cover), tree species and food abundance were compared between woodland areas within and outside of male territories at a site in the UK.

Results Territories contained higher trunk numbers, had a narrow range of trunk diameters, and intermediate canopy cover. Food abundance did not differ with occupancy. Willow and alder were the most common trees within territories, in contrast to birch which has been found in previous studies. The habitat structure matches young woodlands, where birches often grow. However, at the study site the birches were large and mature, and therefore unsuitable. Moreover, woodland structure variables were better predictors of occupancy than any particular tree genera.

Conclusion The results indicate that vegetation structure, but not tree species or food availability, influence breeding habitat selection by Willow Warblers. The preferred structure is similar to coppice woodlands; therefore, the Willow Warbler decline may be linked to the loss of this traditional management across south England.     

Fragmentation in eucalypt woodlands promotes nest‐tree occupancy by a despotic species,the noisy miner (Manorina melanocephala)

The effects of habitat fragmentation as a threat to biodiversity are well known; decreased connectivity can potentially influence population processes and dynamics, resulting in smaller, isolated populations that may not function optimally. However, fragmentation may also increase the amount of edge or ecotone habitat available to open country species, benefiting their populations and enabling them to dominate remnant habitats. Noisy miners (Manorina melanocephala) are one such species, occupying eastern‐Australian eucalypt woodlands. They are considered a ‘despotic’ species, in that their presence negatively impacts woodland avifauna biodiversity due to their aggressive exclusion of other taxa from occupied areas. Despite this well‐known impact, little information exists on the patterns of nest‐tree occupancy by noisy miners within eucalypt woodlands. In the current study, we explored the patterns of nest‐tree occupancy by noisy miners across two successive years, aiming to identify preferences for breeding areas relative to vegetation structure. Our results show that both habitat fragmentation and the characteristics of individual eucalypt trees in an area influenced nest‐tree occupancy. Noisy miners constructed nests in trees near the edge of woodland patches more often than expected. Moreover, the nest tree chosen was a eucalypt that was significantly smaller than randomly selected trees from the surrounding area. The results highlight the importance of habitat management measures that may reduce the suitability of woodland patches as nesting sites for this species, in order to mitigate the severe effects of this despotic edge specialist.     

Contrasting responses of plants and pollinators to woodland disturbance

Preserving species diversity is critical to ensure ecosystem functioning; however, different components of diversity might respond to human disturbance in different ways. Similarly, trophic levels might have uncoupled responses to the same disturbance, thus ameliorating or aggravating the persistence of ecological communities. In this study, we analysed how the density, richness and evenness of flowers and pollinators respond to four levels of woodland thinning intensity (0, 30, 50 and 70% of woodland basal area removed) over 2 years in three contrasting sites. We found a mismatch in the response of flowers and pollinators to thinning. Flower density and richness had disparate responses, depending on the site and year, while evenness did not change with thinning. In contrast, pollinator density and richness, but not evenness, consistently increased with thinning among years and sites. These results suggest that thinning has a great influence on pollinators through changes in abiotic conditions and, perhaps, flower attractiveness rather than through small‐scale changes in flower density and richness. At the site where tree flowers were absent, bee pollinator community composition was impoverished, suggesting that trees provide important floral resources to pollinators. Our findings indicate that disturbance may diminish local plant abundance and richness, but pollinator abundance and richness are enhanced after intense thinning at small scales.     

Light acclimation of four native tree species in felling gaps within a tropical mountain rainforest

Ecuadorian mountain rainforests are declining dramatically due to deforestation. Exploitation of remaining forests has led to low abundances of native, valuable timber species. Enrichment planting of selected native tree species into forest gaps is a strategy that may increase their abundance and maintain biodiversity. However, the development of successful planting strategies requires knowledge of environmental demands on, and ecological requirements of, native species during their establishment. This knowledge is currently lacking for mid- and late-successional species in Central American forests. Two deciduous, mid-successional (Cedrela montana, Tabebuia chrysantha) and two evergreen, late-successional native tree species (Nectandra membranacea, Podocarpus sprucei) were planted into felling gaps. Photosynthetic performance and growth in height of these species were assessed along light gradients during seedling establishment to test whether species-specific light responses were related to plant successional traits. Both mid-successional species benefited from higher light levels in gaps up to 30% canopy openness_60°. In larger gaps, C. montana exhibited a significant decline in growth. As expected, growth of the late-successional species was only marginally increased at higher light levels. Nevertheless, the photosynthetic apparatus of N. membranacea displayed rapid acclimation to higher light conditions in gaps. Plant response to felling gaps may not always be predicted based on successional status. Our results suggest that the four investigated species may coexist in the same gap by occupying different niches along light gradients. This arrangement may offer an ecological basis to increase the abundance of valuable timber species through enrichment planting in Ecuador mountain rainforests.     

Lichens — the biodiversity value of western woodlands

Summary

The Atlantic broad-leaved woodlands of Britain are of international renown for their lichen floras. They are inhabited by 517 lichens, representing 28.3% of the total lichen flora and 73.2% of all British woodland lichens, and they are the main habitat for 165 species. Of these, 31 have a marked southern distribution and do not reach Scotland, whereas 26 species are found in Scotland, but not England or Wales. Their British Red-listed species are outnumbered by the 86 species for which Britain has International Responsibility.

Within the Atlantic broad-leaved woodlands, only 30 lichens show a preponderance for oak. With the exception of some ancient oakwoods in southern England, a high lichen biodiversity is rarely dependent on a dominance of oak in the woodland canopy, more usually it is the result of a long ecological continuity, often a varied tree and shrub composition, a varied canopy density, and good air quality. Consequently, the oak stands within former ‘industrial’ woodlands have a much lower lichen biodiversity compared with woodlands that have a history as ‘pasture woodland’ or, as with some ravine woodlands, have otherwise escaped intensive management.

The life-history of an oak tree is considered in relation to the niches it provides for lichen colonisation with time. Some management scenarios are provided with the enhancement of the lichen interest of former ‘industrial’ oakwoods as an objective.     

Establishment of tree seedlings in the understory of restoration plantations: natural regeneration and enrichment plantings

Little is known about the potential of restoration plantations to provide appropriate understory conditions to support the establishment of seeds arriving from neighboring native forests. In this article, we investigated how seedling establishment is affected in the understory of restoration sites of different ages and assessed some of the potential environmental factors controlling this ecological process. We first compared the density and richness of native tree seedlings among 10‐, 22‐, and 55‐year‐old restoration plantations within the Atlantic Forest region of southeastern Brazil. Then, we undertook a seed addition experiment in each study site, during the wet season, and compared seedling emergence, survival, and biomass on local versus old‐growth forest soil (transferred from a reference ecosystem), in order to test whether local substrate could hamper seedling establishment. As expected, the oldest restoration site had higher density and richness of spontaneously regenerating seedlings. However, seedling establishment was less successful both in the oldest restoration planting and using substrate transferred from a reference ecosystem, where emergence and survival were lower, but surviving seedlings grew better. We attribute these results to lower light availability for seedlings in the understory of the oldest site and speculate that higher incidence of pathogens on old‐growth forest soil may have increased seedling mortality. We conclude that the understory of young restoration plantations provides suitable microsite conditions at the early establishment phases for the spontaneous regeneration or enrichment planting of native trees.     

Perimeter forestry and landscape works at Rutland Water

The forestry management programme at Rutland Water is described. It incorporated management of existing old woodlands as well as establishment of new woodland and hedgerow areas together with amenity planting at car park and picnic sites. The focus of the new planting programme has been the water's edge zone within 2 m of top water level.     

Modelling fallen branch volumes in a temperate eucalypt woodland: implications for large senescent trees and benchmark loads of coarse woody debris

Fallen branches are a substantial component of coarse woody debris and a key ecological resource. The depletion of stocks of coarse woody debris since European settlement has contributed to the degradation of Australian grassy box woodlands, including the loss of biodiversity. Restoration options for remnant woodlands include the augmentation of coarse woody debris stocks. However, the extensive modification of grassy box woodlands has left few reference sites for establishing benchmarks to guide such restoration. In this paper we demonstrate a method for predicting fallen branch debris loads in the absence of reference sites, using data from a yellow box–red gum woodland. Our methodology is in two stages: first, the total volume of branch debris under individual trees was modelled; and second, these models were applied to groups of trees to predict stand‐level loads of fallen branch debris. Although the models were developed for yellow box–red gum woodlands, the methodology would be applicable to other communities that lack reference sites. Predicted benchmark loads of fallen branch debris for yellow box–red gum woodland were between 7.0 m³ ha^?1 and 11.9 m³ ha^?1. Large senescing trees contributed the bulk of fallen branch debris. Model predictions indicated a 100‐cm diameter at breast height (dbh) tree was 10 times more likely to produce debris than a 50‐cm dbh tree, and if debris was present a 100‐cm dbh tree produced approximately 10 times the volume of branch debris produced by a 50‐cm dbh tree. These results highlight the importance of large senescing trees for the production of fallen branch debris and support the keystone role of large trees within remnant woodlands, and the need to conserve these structures. Our results also support the active management of regrowth woodland stands to facilitate the progression of individual trees to maturity and senescence. In particular, thinning of regrowth stands may promote the growth of retained trees, ensuring they contribute to fallen branch debris stocks with a minimum time lag.     

松嫩草原榆树疏林对不同干扰的响应

榆树疏林是中国东北松嫩草原沙地植被演替的顶极群落,其对维持物种多样性,保持水土,涵养水源等都发挥着重要的功能。过去20a间,榆树疏林在人为干扰(农耕和放牧)下,其群落结构和物种组成发生了巨大变化。在1983年和2004年,对松嫩草原不同人为干扰下榆树疏林群落结构和物种组成等进行调查,结果显示:农耕和放牧一方面使松嫩草原榆树疏林群落多物种消失、生物量和多样性显著下降、植被结构简单化;另一方面,也是导致建群种——家榆低矮化,灌木化的主要因素。     

Dendroecology of Prosopis flexuosa woodlands in the Monte desert: Implications for their management

In the Monte desert of Argentina open woodlands of several species of Prosopis occur in areas with accessible underground water. The great latitudinal extent of the Monte (26–43°S) exhibits strong climatic gradients involving temperature, rainfall seasonality, and wind regime. Prosopis woodlands have been a source of subsistence for human communities for several centuries and continue to be exploited by the local inhabitants. The “mining” of this resource has led to severe desertification and consequent impoverishment of the local people. In order to suggest strategies for the better management and recuperation of these woodlands we studied the population structure and productivity of Prosopis flexuosa from multiple plots at Pipanaco (27°58′S), Telteca (32°20′S), and Ñacuñán (34°03′S). For each plot we measured the density of P. flexuosa trees, number of stems, basal diameter (DAB), height and canopy diameter of each tree. Tree ring data were used to determine the growth rates, annual wood production and biological rotation age for each area. The ecological structure of the woodlands differs between the three sites. Along this north–south transect, there is a decrease in adult tree density, mean basal diameter, mean tree height, canopy cover, productivity and total wood biomass. Consequently, the potential sustainable use of these woodlands varies. Only the northern, Pipanaco, woodlands have the potential for lumber production. In contrast, the short, multi-stem and low-productivity trees in the Telteca and Ñacuñán areas can only sustain a combination of local firewood production and activities such as extensive grazing by livestock. The present, uniform regulations for harvesting wood in these areas must be changed to acknowledge these differences in order to optimize wood production in, and conservation of, these woodlands.     

Effects of burning and rainfall on former agricultural land with remnant grassy woodland flora

Grassy woodlands have been extensively cleared for agricultural land uses; land managers need to know whether restoration of biodiversity on such sites requires further interventions beyond simply stopping agricultural land use. Cumberland Plain Woodland occurs on shale‐derived soils in western Sydney; former Cumberland Plain Woodland sites can range from grasslands cleared for agricultural use to regenerated woodlands. An experiment was established in Scheyville National Park to determine what effect repeated burning would have in this system. Four blocks were established (three in grassy areas, one in woodland) and plots in each block were either burnt in 2001 and 2005 or left unburnt. Native plant species richness was initially lower in the grassy blocks than in the woodland, and this ranking remained on unburnt plots over time. The first fire increased species richness of both natives and exotics on the grassy blocks, with the largest increases observed for native and exotic forbs, and lesser increases for grasses (native only), gramminoids and shrubs. Native species richness changed very little with burning in the woodland. Fire effects on species richness were still apparent 3 years later on the grassy blocks; the difference between the grassy blocks and the woodland was not significant on burnt plots at this stage. Changes in native species richness were far less after the second fire on the grassy blocks, with grasses and gramminoids showing increases; native species richness remained higher in the burnt treatment. The first fire reduced the initial differences in native species richness between the grassy blocks and the woodland, and the second fire maintained the benefit through time. Fire also increased exotic species richness; the proportion of total species as natives was not altered by the two fires. On unburnt grassy plots, native species richness and prior cumulative rainfall were positively related; a decline in native species richness on unburnt plots corresponded to increasingly drier conditions over the study.     

Explaining the saproxylic beetle diversity of a protected Mediterranean area

Saproxylic beetle diversity is high at the Cabañeros National Park (central Spain), where woodland habitats exhibit remarkable heterogeneity. Our aim was to explain the diversity of saproxylic beetles, focusing on species turnover among mature woodland types. We surveyed five woodland types that represented the heterogeneity of the park’s woodland habitats. Beetles were collected using window traps over a period of 20 months. The Jaccard Similarity Index was used as indirect value of beta diversity among woodlands and to test the relation between species turnover and geographical distance. We also identified the contribution of species turnover to landscape diversity by using a partitioning model. Moreover, the presence of mixed woodlands (more than one tree species) allowed us to attempt to valorise the effect of tree species (coupled with their historical management) on species turnover among woodlands. Finally, we looked for different saproxylic beetle preferences for habitat and tree species using an indicator value method. We found that saproxylic beetle species composition varied significantly among the studied woodlands. The variation in species turnover was independent from the distance among woodlands, which suggested that beetle dispersal abilities could not explain this high turnover. Tree species within woodlands were a key factor that increased diversity turnover in woodlands and, consequently, the diversity of the park. Moreover, we found saproxylic beetle species that had different habitat and tree species preferences. We conclude that woodland heterogeneity (highly affected by woodland composition) seems to be the driving force for saproxylic beetle diversity in this protected area.     

Understorey foraging and habitat selection by sheep in mixed Atlantic woodland

Abstract. The main objectives of this study were to investigate sheep foraging behaviour in mixed Atlantic woodland and to assess its impact on the forest understorey. We established 89 plots along four forest types: Fagus woodland, Quercus woodland, riparian gallery forest and conifer plantations. The presence of plant species in the forest understorey and their foraging damage was surveyed bimonthly from July 1996 to June 1997. In addition, we estimated the selection of woodland types by sheep through the pellet‐group count technique. The intensity of foraging by sheep was negligible for most of the plant species, however several species showed substantial damage in some woodland types. Among the species that were abundant and widespread in the entire study area, Rubus ulmifolius, graminoids and Ilex aquifolium were consumed most. Sheep selected only larch plantations, where grasses and Rubus were very abundant. This grazing behaviour reduced browsing damage of the understorey of woodland stands with higher conservation value, such as Quercus and Fagus woodlands.     

Elephant herbivory,frost damage and topkill in Kalahari sand woodland savanna trees

Questions: Which factors best predict the probability of elephant and frost damage in Kalahari sand woodland savanna? What is the association between tree mortality and the disturbance regime? Location: Western Zimbabwe. Methods: Elephant and frost damage, topkill, and whole‐plant mortality were quantified in ten common tree species in a Kalahari sand savanna in Zimbabwe. Individual trees were tagged in 20 plots and monitored over a two‐year period. A model selection approach was used to test the association between the probability of damage and size, prior damage, and neighbourhood effects, and to investigate the effect of damage on mortality. Results: Elephant damage differed strongly among species, and was not influenced by neighbourhood effects or prior disturbance. Frost damage also varied across species, and declined as a function of stem size and neighbourhood tree cover, and ‐ against expectations ‐ prior disturbance. Topkill increased as a function of elephant and frost damage, but was lower in previously damaged than in undamaged trees. Conclusions: Frost and elephant damage are influenced by community composition, and frost damage is also correlated with community structure and prior disturbance. Frost is an important and generally overlooked disturbance agent in southern African woodlands, where it may play a key role in association with other disturbance factors ‐ such as elephant herbivory ‐ that reduce woodland canopy cover.