Evaluation of a visual assessment method for tree condition of eucalypt floodplain forests

Summary For decades the floodplain forests of the River Murray have endured the effects of prolonged water stress. This has resulted in significant crown dieback and loss of condition. The Living Murray (TLM) initiative aims to restore the ecological health of six Icon Sites along the River. The two eucalypts River Red Gum (Eucalyptus camaldulensis) and Black Box (Eucalyptus largiflorens) that dominate the forests at five of the six Icon Sites are undergoing widespread decline. To enable effective management and restoration of these forests, we developed a standardised tree condition assessment method. Named the TLM tree condition assessment method, it utilises visual assessment of a range of tree crown variables (extent and density of the foliage in the crown, epicormic growth, new tip growth, reproductive activity, leaf die‐off, mistletoe infestation) and measurements of bark condition, diameter at breast height and dominance class. This article describes the TLM tree condition assessment method and assesses it for consistency between multiple observer teams after limited training. The level of observer agreement between six teams each comprised of two observers was assessed for seven of the ten variables. Intra‐class correlation was used to compare scores of 30 River Red Gum trees assessed on Gunbower Island on the River Murray. The level of agreement for all variables was statistically significant with six of seven variables having correlation coefficients over R = 0.5. The TLM tree condition assessment method was found to provide accurate estimates of a range of tree variables that can be used to determine tree condition. The TLM tree condition assessment method provides a valuable monitoring tool that can be used to assess management interventions, such as management flooding and silvicultural thinning.     

The decline of Greek fir (Abies cephalonica Loudon): Relationships with root condition

Abstract

The decline of Greek fir (Abies cephalonica Loudon) has been reported throughout Greece during the last three decades. The symptoms include crown dieback, needle discoloration and loss, death of twigs, branches or whole tree death. A number of causes for the fir decline have been proposed such as drought, climatic change, air pollution and pathogens or insects. Our studies have focused on the relationship between fir decline symptoms and root growth and condition. The crown and root condition of fir trees, at different stages of decline, were assessed and the results showed that root condition was related to needle loss and crown discoloration. Declining trees were characterized by a decreased portion of fine root biomass. No significant differences in total root length and number of roots for each root order were found. In conclusion, our studies showed that the crown condition of Greek firs is reflected on root condition.     

Does stand structure influence susceptibility of eucalypt floodplain forests to dieback?

Forest dieback is a worldwide problem that is likely to increase with climate change and increasing human demands for resources. Eucalyptus camaldulensis forests are an acute example of forest dieback, with 70% of the Victorian Murray River floodplain in some state of dieback. If we are to halt dieback in these floodplain forests, we need to understand what makes stands susceptible to dieback. Forest diebacks are often related to stand structure, with dieback more severe in senescent or high‐density stands. We determined whether certain stand structures make these forests more susceptible to dieback. We undertook an extensive survey of 176 stands across 100 000 ha of forest, covering the range of stand condition on this floodplain. Large and small trees (20‐, 40‐, 80‐ and 120‐cm diameter) showed a similar reduction in the probability of being alive with decreasing stand condition. A slight improvement in stand condition was found at higher densities and basal areas, which may reflect the higher productivity or younger age of these stands. Stand condition was moderately, positively correlated with longitude, with stand condition being higher in the east of the Murray River floodplain where flooding frequencies are currently higher. This suggests that dieback of these floodplain forests would be more effectively mitigated by increased water availability through flooding than by altering stand structure.     

Using forest growth trajectory modelling to complement BioCondition assessment of mine vegetation rehabilitation

The standard rehabilitation objective for open‐cut mines in Queensland is to establish a self‐sustaining native forest ecosystem. Consequently, mine regulators and managers need tools to project whether sites are likely or not to meet agreed completion criteria and to ensure timely remedial interventions. The Ecosystem Dynamics Simulator (EDS) is such a tool capable of modelling forest dynamics and projecting long‐term growth of woody species only. Here, the model was applied to rehabilitation sites aged between 5 and 22 years in Meandu open‐cut coal mine in southeast Queensland. EDS projected structural characteristics for trees (height, diameter, basal area, foliage projective cover and stem density) and tree species composition as a function of rehabilitation age. Projected stand growth attributes were assessed against BioCondition benchmarks developed from eucalypt (Eucalyptus/Corymbia) remnant forests adjacent to the mine. Growth trajectories indicated that sites with >30% eucalypt basal area composition were more likely to develop into eucalypt‐dominated self‐sustaining ecosystems compared with sites that were initially dominated by acacias (Acacia spp.). Projections suggested that some benchmark attributes such as number of large eucalypt trees would take more than 70 years to be met. The application of EDS provided a framework to support decisions on early remedial intervention and assess the risk associated with lease relinquishment.     

Crown dieback events as key processes creating cavity habitat for magellanic woodpeckers

Abstract Woodpeckers are considered keystone species for webs of cavity nesters and habitat and resource specialists that strongly depend on availability of trees suitable for cavity excavation. Most studies carried out in northern hemisphere temperate coniferous forests emphasize the importance of old growth stages of forests or large dead trees as habitat for cavity builders. We present a study of Nothofagus pumilio tree selection by the magellanic woodpecker (Campephilus magellanicus) that incorporates dendroecological data on long‐term growth trends of trees that provides new insights into the processes that create suitable habitat for cavity excavating species. We analysed 351 cavity and neighbouring control trees in terms of age and radial growth patterns, as well as external tree characteristics. In addition, from a subsample of these trees we developed tree‐ring chronologies for each group using standard methods in order to analyse potential differences in radial growth patterns between cavity and non‐cavity trees. Multivariate models that account for differences between paired cavities versus control trees indicated that growth decline and the degree of crown dieback were the primary variables explaining magellanic woodpecker tree selection for cavity building. In contrast to previous work, neither diameter (above a certain threshold) nor age, were important determinants of selection. Furthermore, trees that became present cavity are those that had synchronously declined in radial growth during the 1943–44 and 1956–57 droughts and the 1985–86 massive caterpillar defoliation. Insect outbreaks and extreme climatic events may episodically reduce vigour, induce partial crown mortality, trigger increased fungal attack and heart rot formation at different tree heights on the bole in a group of trees and thus increase availability of soft substrate and their likelihood of cavity excavation by the magellanic woodpecker. These results underscore the importance of drought/biotically‐induced canopy dieback events in creating habitat for woodpeckers and their dependent cavity users.     

Long‐term response of temperate canopy trees to removal of browsing from an invasive arboreal herbivore in New Zealand

Defoliation of forest tree canopies by herbivores and other agents, leading to tree mortality and reduced productivity, threatens the ecological stability of forests globally. This study shows that long‐term control of a mammalian arboreal folivore (brushtail possums; Trichosurus vulpecula Phalangeridae) reduces crown dieback and increases foliage cover in browsing‐damaged canopy trees. We monitored indices of possum density, possum browsing, tree foliage cover and crown dieback for 20 years following initiation of possum control in 1994 that repeatedly reduced possum densities to near zero every 5–6 years and kept the population below 35% of pre‐control levels over the entire period. Observable possum browsing was recorded on 20–49% of individuals of three palatable tree species at the time of first control. Those percentages fell to zero after control and never exceeded 2–10% for individual species over the next 19 years. We recorded significant increases in foliage cover attributable to recovery from defoliation by possums for all three species during the first 10 years. Large increases in foliage cover occurred on individuals that were heavily browsed in 1994 (mean increases: 36–89%), but mean population increases were modest (3–19%) because only 10–19% of trees were initially heavily browsed. Twenty‐year mortality rates were similar for plants with, or without, initial possum browsing, indicating no residual impact of pre‐control browsing on tree mortality. Times for full recovery of crown foliage cover varied from 10 years for the youngest trees and faster growing species to more than 20 years for mature individuals of the slowest growing species.     

Stable correlation structure among multiple plumage colour traits: can they work as a single signal?

The presence of multiple distinct ornamental traits in the same species is frequently explained by context‐specificity and different information content. However, the expression of multiple ornaments is often correlated, and such traits may therefore function as a single, integrated signal. Delayed use of an integrated signal relative to production requires temporal stability in integration, which has seldom been examined. We used autumn and spring reflectance data from the breast, breast stripe, and crown of great tits (Parus major) to assess the stability and mating implications of colour signal integration, as well as the repeatability of any integrated colour trait and its correlation with condition during moult. We found high levels of stability between seasons, years, sexes, and ages in the correlation patterns of colour measures across the three plumage areas. The first principal component colour axis described joint variation of ultraviolet (UV) reflectance on the crown and the breast stripe, thereby representing an among‐trait UV chroma axis. However, only breast yellow chroma showed condition‐dependence, whereas temporally consistent and significant assortative mating was restricted to crown UV chroma. Our results therefore do not support the idea that the overall UV chroma of the breast stripe and the crown is special in condition‐dependence and repeatability, or that it plays a specific role in mutual sexual selection as an integrated signal. The results show that stable association between display traits is an existing phenomenon. They also indicate that, even in the presence of correlated traits, functional trait integration among these requires further scrutiny. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 92–108.     

Estimating Canopy Structure in an Amazon Forest from Laser Range Finder and IKONOS Satellite Observations1

Canopy structural data can be used for biomass estimation and studies of carbon cycling, disturbance, energy balance, and hydrological processes in tropical forest ecosystems. Scarce information on canopy dimensions reflects the difficulties associated with measuring crown height, width, depth, and area in tall, humid tropical forests. New field and spaceborne observations provide an opportunity to acquire these measurements, but the accuracy and reliability of the methods are unknown. We used a handheld laser range finder to estimate tree crown height, diameter, and depth in a lowland tropical forest in the eastern Amazon, Brazil, for a sampling of 300 trees stratified by diameter at breast height (DBH). We found significant relationships between DBH and both tree height and crown diameter derived from the laser measurements. We also quantified changes in crown shape between tree height classes, finding a significant but weak positive trend between crown depth and width. We then compared the field‐based measurements of crown diameter and area to estimates derived manually from panchromatic 0.8 m spatial resolution IKONOS satellite imagery. Median crown diameter derived from satellite observations was 78 percent greater than that derived from field‐based laser measurements. The statistical distribution of crown diameters from IKONOS was biased toward larger trees, probably due to merging of smaller tree crowns, underestimation of understory trees, and overestimation of individual crown dimensions. The median crown area derived from IKONOS was 65 percent higher than the value modeled from field‐based measurements. We conclude that manual interpretation of IKONOS satellite data did not accurately estimate distributions of tree crown dimensions in a tall tropical forest of eastern Amazonia. Other methods will be needed to more accurately estimate crown dimensions from high spatial resolution satellite imagery.     

Quantifying the deciduousness of tropical forest canopies under varying climates

Abstract. Deciduousness is an important functional attribute of tropical trees, reflecting climatic conditions. Precisely quantifying and mapping deciduousness in tropical forests will be necessary for calibrating remote sensing images which attempt to assess canopy properties such as carbon cycling, productivity, or chlorophyll content. We thus set out to assess the degree of canopy deciduousness in three moist, semi‐deciduous tropical forests in central Panama. One site is a 6‐ha research plot near the Atlantic coast of Panama, where rainfall is 2830 mm/yr. The second site is a 50‐ha plot on Barro Colorado Island, near the center of the isthmus of Panama, where rainfall is 2570 mm/yr, and the final site is a 4‐ha plot near the Pacific coast of Panama, where rainfall is 2060 mm/yr. At each site, a random sample of trees from all canopy species (those with individuals ≥ 30 cm DBH) were visited and scored for deciduousness three times during the 1997 dry season. The estimated peak fraction of deciduous individuals in the canopy at the wetter site was 4.8%, at the intermediate site, 6.3%, and at the drier site, 24.3%. The estimated fraction of crown area deciduous peaked at 3.6%, 9.7%, and 19.1% at the wet, medium, and dry sites respectively. The percentage of canopy species that was deciduous –14%, 28%, and 41%–was much higher than the percentage of deciduous individuals, because not all individuals of deciduous species were deciduous. During the 1999 dry season, every individual of all the deciduous species was visited at the two drier sites, and the total number of deciduous trees observed closely matched the estimated numbers based on the smaller 1997 samples.     

Roost preferences and foraging ranges of the eastern forest bat Vespadelus pumilus under two disturbance histories in northern New South Wales,Australia

Little is known about the habitat requirements of Australian bats; however, this information is needed to make better‐informed decisions when systems are disturbed. This study contrasts the roosting and foraging ecology of the eastern forest bat Vespadelus pumilus (Vespertilionidae), one of Australia’s smallest bats, between two sites of differing disturbance history on the mid‐north coast of New South Wales. Lorne Flora Reserve (182 ha) is primarily old‐growth forest surrounded by regrowth forest and eucalypt plantations, while Swans Crossing is dominated by regrowth and eucalypt plantations established on part of an old dairy farm. A total of 38 bats were tracked during the maternity and mating seasons at the two sites. Roost preferences were determined by comparing trees used as roosts with those randomly available, while foraging bats were triangulated from fixed stations at night. Bats tracked at Lorne Flora Reserve typically roosted in hollows within large, mature trees and showed a strong preference for roosting and foraging (females only) within the Reserve. Lactating females at Swans Crossing roosted in hollows of remnant rainforest trees within a gully and dead eucalypts, while males often roosted in understorey trees (such as Acacia). Dead trees were frequently used as roosts at both sites. Under both disturbance histories, the mean distance of female maternity roosts from creeks was 20 m, indicating that riparian zones provide important roosting habitat for V. pumilus. However, roosts shifted to the mid‐slope prior to winter when bats mate. Retention of mature trees in a variety of topographic locations may allow behavioural adjustments with the seasons. Bats caught in the regrowth forest also foraged there, with foraging ranges averaging just 5.3 ha (n = 10), indicating that regrowth is used by this bat for both foraging and roosting.     

Inferring changes in foliar mass and area from foliage cover: A mechanistic model

Measuring foliar area or mass directly is destructive, and precludes long‐term, repeated observations of individual trees as they suffer or recover from foliar damage. Instead, foliage cover indices are often used as a proxy for foliar mass. Patterns of fluctuations in foliage cover indices can be used to infer qualitative changes in canopy health. However, foliage cover is not necessarily linearly related to foliar area or mass, and this may confound the detection of significant foliar damage, and comparisons of herbivore browse impacts between individual trees, tree species or sites. I derived a mechanistic model to quantify the relationship between foliar area or mass and foliage cover measured as the proportion of sky occluded by leaves. This one parameter model is close to linear for single‐tiered trees, but increasingly non‐linear for multi‐tiered trees. I compared the non‐linear model to a linear model using foliage cover data from an artificial defoliation experiment on two single‐tiered, sub‐canopy species and from simulated photographic images of single‐ and multi‐tiered canopies. The non‐linear model had lower errors than the linear model, and errors did not increase with foliage density (leaf area per unit area), variation (of leaf sizes within and between canopies) or leaf geometry. The non‐linear model can be easily parameterized from relatively low‐cost observations of foliage cover, independently of empirical measurements of foliar area or mass, and is applicable to a wide range of tree species. It should therefore help managers quantify how changes in foliage cover due to natural fluctuations or foliar damage affect foliar area and mass, and can be used to quantify parameters for models of browse impacts in mixed forest.     

Dieback and recovery in poplar and attack by the hornet clearwing moth, Sesia apiformis (Clerck) (Lepidoptera: Sesiidae)

A survey of 801 poplar trees in central east England in 1999 demonstrated a correlation between crown dieback and infestation by hornet clearwing moth (Sesia apiformis), induced by a period of drought in 1995-1996. To determine whether trees colonised by S. apiformis would subsequently deteriorate and die or whether they could recover despite infestation and damage to the stem, all trees in the original survey were re-assessed in 2001, 2003 and 2005. The repeat surveys showed that trees with 70% or less crown dieback in 1999 replaced their canopy and generally improved by 2005, irrespective of the numbers of S. apiformis in the stem, whereas trees that had 75% or more crown dieback in 1999 either died or declined further. The presence of S. apiformis did not prevent tree recovery, and there was little evidence that infestation slowed the rate of recovery. Populations of S. apiformis, measured in terms of the numbers of adult emergence holes visible in the base of the trees, decreased between 2001 and 2005 at the same time as the amount of dieback visible in the canopy of the poplars markedly decreased. However, the fall in numbers of emergence holes at this time reflected a decline in larval establishment 2-3 years earlier, and indicated that the moth population had responded to a more rapid restoration in the internal state of the trees, which was not reflected immediately by the gradual replacement of dead branches and reduction in dieback symptoms.     

Assessing canopy health of native eucalypt forests

Summary   The assessment of forest health is an essential part of the monitoring of ecological sustainability in managed native forests. In Australia, unfortunately, very limited quantitative information on forest health is actually obtained for management and reporting purposes. In this article, we summarize current approaches used in Australia to assess native forest health and some recent developments in the application of remotely acquired digital imagery for classifying canopy health. In a recent study examining Bell miner associated dieback (BMAD), high-resolution airborne imagery was successfully manipulated to present severity categories for BMAD affected canopy. The potential of remotely sensed imagery lies not in map production but in the statistical modelling capacity of this spatial information, particularly when added to climatic and terrain-based spatial data sets. There are several statistical approaches to modelling these spatial datasets and in this article, we discuss our approach to producing a preliminary BMAD model. The importance of ground-based assessments is also emphasized and we recommend tree crown condition as a key health attribute for the spatial modelling of forests. Although significant progress has been made in the application of remote sensing technologies, the structural complexity of native forests means that there are still technical issues that require resolving before this approach becomes operationally routine.     

Wood anatomy and carbon‐isotope discrimination support long‐term hydraulic deterioration as a major cause of drought‐induced dieback

Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought‐induced forest dieback and tree death. Here, we evaluate the relative role played by these two mechanisms in the long‐term by quantifying wood‐anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water‐use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long‐lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2–5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long‐term capacity of trees to withstand drought‐induced dieback.     

Impact of long-term,saline flooding on condition and reproduction of the clonal wetland tree, <Emphasis Type="Italic">Melaleuca ericifolia</Emphasis> (Myrtaceae)

Although water regime modification and salinity are recognised as significant threats to wetland ecosystems worldwide, the effects of long-term saline flooding (decades) on woody tree persistence are poorly quantified. We compared the condition, growth, structure and reproduction (sexual and asexual) of mature individuals of the clonal tree, Melaleuca ericifolia (Myrtaceae), that experienced continual (>30 years) flooding with trees that were only intermittently (approximately every 5 years) flooded. An index developed to assess the condition of multi-stemmed trees found that continually flooded trees were in significantly poorer condition than intermittently flooded trees, having lower crown cover, foliar cover and foliar density, and a higher incidence of dead stags and dieback. Annual stem growth correlated strongly with condition scores. Evidence for a trade-off between sexual and asexual reproductions was found; flooded trees were constrained in their vegetative lateral spread (<12 m dia.) and flowered more than intermittently flooded trees, which were not restricted in lateral spread (~30 m dia.). Flooded trees used intensively by the colonially roosting Australian Sacred Ibis (Threskiornis molucca) were in especially poor condition. These trees flowered infrequently and produced the lowest number of infructescences, but produced many new vegetative stems (ramets) within their centre. Although chronic flooding appeared to compromise the condition of M. ericifolia trees in Dowd Morass, their existence upon woody hummocks (~40 cm high) upon which they are able to produce new stems is likely to be a key mechanism in their persistence. It is unknown, however, how long this process can maintain the existing population. Production and maintenance of a large aerial seed bank by living mature trees under flooded conditions may allow M. ericifolia to regenerate sexually upon drawdown and may be important for population persistence in the longer term.     

The use of image analysis to monitor the germination of seeds of broccoli (Brassica oleracea) and radish (Raphanus sativus)

To study broccoli and radish seed germination under different temperature regimes the germination test has been used to assess final germination percentage, start and rate. This method has been integrated with a computer‐aided image analysis test which is more accurate in monitoring the extent of imbibition phases through the assessment of seed area increase and timing of radicle emergence detected on single seeds. In addition, seed area increase has been used also to establish a close relationship with radicle elongation rate in the time range when ‘visible germination’ is scored by a classical germination test. The results suggest that this image analysis parameter may be considered as a reliable seed imbibition marker to integrate the germination parameters obtained by a germination test.     

Mapping Tropical Forest Trees Using High‐Resolution Aerial Digital Photographs

The spatial arrangement of tree species is a key aspect of community ecology. Because tree species in tropical forests occur at low densities, it is logistically challenging to measure distributions across large areas. In this study, we evaluated the potential use of canopy tree crown maps, derived from high‐resolution aerial digital photographs, as a relatively simple method for measuring large‐scale tree distributions. At Barro Colorado Island, Panama, we used high‐resolution aerial digital photographs (~0.129 m/pixel) to identify tree species and map crown distributions of four target tree species. We determined crown mapping accuracy by comparing aerial and ground‐mapped distributions and tested whether the spatial characteristics of the crown maps reflect those of the ground‐mapped trees. Nearly a quarter (22%) of the common canopy species had sufficiently distinctive crowns to be good candidates for reliable mapping. The errors of commission (crowns misidentified as a target species) were relatively low, but the errors of omission (missed canopy trees of the target species) were high. Only 40 percent of canopy individuals were mapped on the air photographs. Despite failing to accurately predict exact abundances of canopy trees, crown distributions accurately reproduced the clumping patterns and spatial autocorrelation features of three of four tree species and predicted areas of high and low abundance. We discuss a range of ecological and forest management applications for which this method can be useful.     

Amazon Forest Structure from IKONOS Satellite Data and the Automated Characterization of Forest Canopy Properties

We developed an automated tree crown analysis algorithm using 1-m panchromatic IKONOS satellite images to examine forest canopy structure in the Brazilian Amazon. The algorithm was calibrated on the landscape level with tree geometry and forest stand data at the Fazenda Cauaxi (3.75° S, 48.37° W) in the eastern Amazon, and then compared with forest stand data at Tapajos National Forest (3.08° S, 54.94° W) in the central Amazon. The average remotely sensed crown width (mean ± SE) was 12.7 ± 0.1 m (range: 2.0–34.0 m) and frequency of trees was 76.6 trees/ha at Cauaxi. At Tapajos, remotely sensed crown width was 13.1 ± 0.1 m (range: 2.0–38.0 m) and frequency of trees was 76.4 trees/ha. At both Cauaxi and Tapajos, the remotely sensed average crown widths were within 3 percent of the crown widths derived from field measurements, although crown distributions showed significant differences between field-measured and automated methods. We used the remote sensing algorithm to estimate crown dimensions and forest structural properties in 51 forest stands (1 km²) throughout the Brazilian Amazon. The estimated crown widths, tree diameters (dbh), and stem frequencies differed widely among sites, while estimated biomass was similar among most sites. Sources of observed errors included an inability to detect understory crowns and to separate adjacent, intermingled crowns. Nonetheless, our technique can serve to provide information about structural characteristics of large areas of unsurveyed forest throughout Amazonia.     

Developmental Patterns of Tree Dimensions in a Neotropical Deciduous Forest1

For 26 tree species in very dry tropical forest in Mexico, the developmental trends of relationships among trunk diameter, tree height, and crown diameter were inferred from a one‐time measurement of dispersed individuals across the size range from saplings to large, mature trees. On hillside sites in this high diversity forest, maximum dimensions were usually <10‐m height, 4‐m crown diameter, and 0.3‐m trunk diameter. The relationship of height to trunk diameter was characterized by an asymptotic, three‐parameter model. Crown diameter was a linear function of trunk diameter. The parameter values for both models varied widely among the species. In general, the dispersion among species of the height–crown diameter relationship increased linearly with trunk diameter (up to 0.2 m). Arborescent cacti were distant from other species at all sizes, although they were well modeled using the same equations. Empirical and theoretical features and limitations of the present and previous models, including mechanical buckling and water‐stress theories, are considered.     

Patch dieback of Colophospermum mopane in a dysfunctional semi‐arid African savanna

Abstract Patch dieback occurred in an almost monospecific Colophospermum mopane (Kirk ex Benth.) Kirk ex J. Léonard woodland in the Northern Province, South Africa, following severe droughts in 1988–1989 and 1991–1992. Discrete patches of dieback and adjacent paired areas of ‘healthy’ vegetation lost an average of 87 and 13% of basal area to mortality, respectively. Whole trees mostly died on ‘dead’ plots, while single‐stem mortality prevailed on ‘live’ plots. Tree mortality decreased with increasing stem number per tree. Patch dieback did not occur on sandy soils. On fine‐textured soils, variation in soil type, topography or slope did not affect dieback. Dieback was influenced by vegetation structure, soil surface condition and soil chemistry. Intense intertree competition, shown by self‐thinning occurring prior to dieback, was a precondition for dieback. Intertree competition had heightened during the 30 years prior to dieback because of an increase in woody cover. Dieback patches had changed from functioning as sinks of water and sediment to sources of these as a result of loss of perennial herbaceous cover, decreased water retention on bared surfaces, and accelerated erosion during 50 years of livestock ranching. Vegetation had thus become increasingly drought‐prone, exacerbated in places by soils with a high sodium concentration. Dieback had occurred because the water requirements of C. mopane could no longer be met during drought years on the dysfunctional patches.