Impacts of cyclone Larry on the vegetation structure of timber plantations,restoration plantings and rainforest on the Atherton Tableland,Australia

Abstract We examined the impact of severe cyclone ‘Larry’ on the vegetation structure of monoculture and mixed species timber plantations, restoration plantings and reference sites in upland rainforests on the Atherton Tableland, north Queensland, Australia. Sites were initially assessed in 2000 and resurveyed in 2006, 6–8 months after the cyclone traversed the region. In both surveys, timber plantations had a relatively open canopy, grassy understorey and few shrubs or small‐sized trees; whereas restoration plantings had a relatively closed canopy, an understorey of bare ground, leaf litter and rainforest seedlings, a high density of small‐diameter trees and a moderate representation of special life forms characteristic of rainforest. Cyclone damage varied with tree size, site type, proximity to the cyclone and stem density. First, the proportion of trees that were severely damaged by the cyclone (major branches broken, stem snapped or pushed over) increased with the diameter of trees across all site types. Second, damage to larger‐sized trees (>10 cm d.b.h., >20 cm d.b.h.) was proportionally highest in monoculture plantations, intermediate in mixed species plantations and rainforest, and lowest in restoration plantings. Third, within site types, damage levels decreased with distance from the cyclone track and with stem density. There was no evidence that topographical position influenced damage levels, at least for timber plantations. We tentatively attribute the high levels of damage experienced by timber plantations to their relatively open structure and the large size of stems in plantations. Restoration plantings generally escaped severe damage by the cyclone, but their continued development towards rainforest conditions may require a coordinated monitoring and maintenance programme to address the potential threat of weed invasion.     

Wood density predicts plant damage and vegetative recovery rates caused by cyclone disturbance in tropical rainforest tree species of North Queensland,Australia

Abstract The ability to withstand disturbance (resistance) and the ability to recover biomass following disturbance (resilience) were investigated in Australian wet tropical rainforest tree species. These two attributes are expected to be negatively correlated, because investment of biomass in structural support (conferring resistance) results in trees exhibiting high wood densities and slow growth rates, and vice versa. We examined species’ responses to disturbance caused by a severe tropical cyclone to test this hypothesized trade‐off. We assessed cyclone damage in six species in three Mabi rainforest fragments on the Atherton Tablelands. Species differed in the proportion of individuals within four damage categories (minor damage, severe branch damage, snapped, uprooted). Resistance was positively related to wood density. We found a positive correlation between the proportion of trees experiencing minor damage only and wood density, supporting the hypothesized association between resistance and mechanical strength. Among the subset of trees in which stems snapped, rates of resprouting differed between species and were highest in low wood density species and lowest in species with highest wood density. Resilience, characterized as the ability to recover biomass following disturbance and estimated as growth rate standardized for stem diameter at breast height (g day^?1 · mm^?1), was negatively related to wood density. Thus, species with low wood densities were more likely to suffer stem and branch damage owing to cyclonic winds, but also demonstrated highest resprouting and fastest responses in terms of redeveloping biomass in the 8 months following disturbance. This suggests that a species’ position along the resistance–resilience spectrum can be predicted by mean wood density, which may allow managers to predict species’ responses to future cyclones. Our findings also provide mechanistic evidence for the ‘direct regeneration’ model of post‐cyclone succession, where response is characterized by resprouting and species composition is unchanged.     

Effects of Severe Tropical Cyclone Larry on rainforest vegetation and understorey microclimate near a road,powerline and stream

Abstract Severe Tropical Cyclone Larry damaged a large swathe of rainforest to the west of Innisfail in north‐eastern Queensland on 20 March 2006. Within the path of the most destructive core of the cyclone were sites previously established along human‐made (powerlines and highways) and natural (streams) linear canopy openings for a study of edge effects on adjacent rainforest plant communities and associated microclimates. Vegetation damage and understorey microclimate parameters were measured 6 months after the passage of Cyclone Larry and compared with results before the cyclone. We examined the spatial patterns of vegetation damage in relation to natural and artificial linear clearing edges and the vegetation structural factors influencing these patterns as well as resulting alterations to microclimate regimes experienced in the rainforest understorey. Vegetation damage was spatially patchy and not elevated near linear clearing edges relative to the forest interior and did not differ between edge types. Vegetation damage was influenced, albeit relatively weakly, by structural traits of individual trees and saplings, especially size (diameter at breast height, d.b.h.) and successional status: tree damage was greater in pioneer species and in larger trees, while sapling damage was greater in canopy tree species than in understorey tree or shrub species. Changes in the understorey microclimate mirrored the degree of damage to vegetation. Where vegetation damage appeared greater, the understorey microclimate was brighter, warmer, drier and windier than below less‐damaged areas of the forest canopy. Overall, understorey light availability, wind speed and the diurnal ranges of air temperature and vapour pressure deficit increased dramatically after Cyclone Larry, while pre‐cyclone edge gradients in light availability were lost and temperature and vapour pressure deficit gradients were reversed.     

Factors Affecting Tropical Tree Damage and Survival after Catastrophic Wind Disturbance

The structure and dynamics of cyclone‐prone tropical forests are driven in part by variation in tree species resistance to and survival after wind‐induced structural damage. We determined the factors associated with tree damage and 3‐yr survival following Category 5 Cyclone Olaf on the Polynesian island of Ta'u, American Samoa. Despite sustaining a high rate of severe damage (34.6% of all trees snapped, 23.0% uprooted), system resilience was high with 74.3 percent stem survival overall and an annual mortality rate of 7.9 percent compared with 2.1 percent in nearby undisturbed late successional forest. Three‐yr survival rate of trees sustaining severe damage was 63.1 percent, compared to about 89 percent for trees sustaining only branch loss or defoliation. Three‐yr survival differed according to damage type, 78.5 percent after snapping vs. 38.4 percent after uprooting. Species differed widely in resistance to and survival after snapping and uprooting. Several species and individual traits were associated with the probability of snapping or uprooting; however, wood density was the only species trait consistently, and negatively, associated with the probability of sustaining either damage type. Survival after snapping was negatively associated with the proportion of the tree snapped off, which was determined by individual tree architecture. Species growth rate was negatively associated with survival after uprooting, indicating the importance of shade tolerance for survival after uprooting. Thus, whereas species traits seemed to exclusively underpin resistance to and survival after uprooting, they only partly accounted for snapping resistance, and did not determine the intensity of snap damage or survival after snapping. Our results highlight the importance of considering each damage type separately when considering ecological trade‐offs.     

Do edge effects increase the susceptibility of rainforest fragments to structural damage resulting from a severe tropical cyclone?

Abstract If changes in the structural characteristics of rainforest at edges are caused by wind, then physical damage from a tropical cyclone might be greatest at edges or in small fragments that have a high proportion of edge. We tested whether this was true of a fragmented rainforest landscape impacted by a category 4 severe tropical cyclone in March 2006. Six structural variables (canopy cover, canopy height, cover of ground vegetation, leaf litter, stem density and counts of woody debris) were surveyed at 18 rainforest sites (six small linear remnants, and both edges and interiors of six large remnants) on the Atherton Tableland in north‐eastern Queensland, Australia. Data collected 7 and 12 months after the passage of Cyclone Larry were compared with an identical survey conducted 4 years prior to the cyclone. The cyclone had large effects across many components of forest structure. However, sites within 30 m of forest edges in small and large remnants were not impacted more than the interiors of large remnants. It is likely that the high wind intensity from severe tropical cyclones overrides the modest wind protection provided by surrounding forest. The cyclone's effects were highly patchy at local scales (0.5–1.0 km), leading to an increase in among‐site variation in forest structure and the disappearance of significant spatial autocorrelation among large remnant edge‐interior site pairs which had existed prior to the cyclone. The main effect of Cyclone Larry at these study sites was to increase the spatial heterogeneity of forest structure at local scales.     

Impacts of Cyclone Larry on arboreal folivorous marsupials endemic to upland rainforests of the Atherton Tableland,Australia

Abstract Intense cyclones might be expected to adversely affect populations of arboreal mammals, either directly or as a consequence of the destruction of food resources and other key habitat elements. However, such impacts have rarely been quantified. The present study examined the response of five species of arboreal folivorous marsupials to Severe Cyclone Larry at nine sites in upland rainforests of the Atherton Tableland, north‐east Australia. Sites were originally surveyed for folivores in 1995–1997, and then resurveyed in 2006, 6–8 months after Cyclone Larry had traversed the region. All sites showed evidence of structural damage to vegetation, but overall damage levels (assessed in terms of canopy cover, damage to trees, basal area of dead trees and volume of woody debris) decreased from east to west across the study region. The detectability of rainforest possums increased after the cyclone. For the most commonly observed species, the proportion of individuals observed >5 m from survey transects was correlated with the amount of structural damage to vegetation. To avoid confounding changes in detectability with changes in abundance, only observations close (<5 m) to transects were used to estimate folivore abundance before and after the cyclone. On this basis, there were no significant differences between pre‐ and post‐cyclone abundance estimates for any folivore species. Further, changes in folivore abundance after the cyclone were not correlated with damage to vegetation across sites. Cyclone Larry does not appear to have caused a catastrophic loss of key habitat resources for marsupial folivores at the sites surveyed. The high degree of folivory practiced by marsupial folivores may help make them resilient to cyclone impacts. These conclusions are more robust for three commonly observed folivore species (Hemibelideus lemuroides, Pseudochirulus herbertensis and Trichosurus vulpecula johnstonii) than for two less frequently encountered species (Pseudochirops archeri and Dendrolagus lumholtzi).     

Do cyclones and forest fragmentation have synergistic effects? A before–after study of rainforest vegetation structure at multiple sites

Abstract Ecological degradation within areas of remnant forest may be amplified if the effects of fragmentation interact with the effects of other environmental disturbances such as wind storms. We used before–after comparisons to assess the effects of Tropical Cyclone Larry on remnant and continuous rainforest in the Wet Tropics uplands of north‐eastern Australia. Vegetation structure was measured 3 years before the cyclone and 6 months afterwards, at eight continuous forest sites and eight remnants (6–37 ha), within 20 km of the cyclone's track. The cyclone caused extensive defoliation, felling and breakage of stems and branches (greatest among the trees >100 cm diameter which had around 50% stem loss), and increased litter and woody debris. Cyclone effects were strongly influenced by a site's spatial position (P = 0.005, 0.001 in multivariate analyses of overall damage). Maximum damage occurred 10–15 km south of the cyclone track, perhaps because of the additive effects of the west‐moving air at the southern eyewall combined with the cyclone's own rapid westward movement. Most fragments were south of the cyclone track, as a consequence of spatially selective deforestation practices, and therefore, showed greatest damage. However, once the effects of spatial position were considered, the independent differences in cyclone effects between fragments and continuous forest were lost (P = 0.23, 0.41 when north‐south distance was included as a covariate in analyses). The expected protection afforded by a continuous forest canopy seems to have disappeared in the face of extremely strong cyclonic winds and down‐draughts. Nevertheless, an interaction between fragmentation and disturbance may yet occur, during the period of post‐cyclone recovery, owing to the effects of landscape context on plant recruitment. For example, there was a higher diversity of exotic seedling germination in fragments, independent of the extent of cyclone damage.     

Correlating Stem Biomechanical Properties of Hawaiian Canopy Trees with Hurricane Wind Damage1

Hawaiian forests are subject to the effects of periodic hurricane conditions. Hurricane Iniki struck the island of Kauai, Hawaii on September 11, 1992 with winds exceeding 200 km/h and caused defoliation, felling of trees by snapping and uprooting, and standing tree mortality due excessive limb and leaf loss. The purpose of this study was to evaluate if measured wood mechanical characteristics could be correlated with stem failure of trees under windstorm conditions. A field survey indicated that post-hurricane stem condition (snapped, uprooted, or standing) differed among five common canopy species and was significantly correlated with stem apparent elastic modulus (relative flexibility). Species that tended to snap had significantly higher apparent elastic moduli than those that remained standing or were uprooted. Wood density and stem diameter were not significantly related to stem failure mode. Native trees had a higher percentage per species of standing individuals but also had increased uprooting. Nonnative tree species were more often snapped and fewer were standing after the hurricane. The higher incidence of stem failure for introduced canopy trees may increase the spread of alien understory species following wind disturbance events. These relationships provide a simple means to predict relative differences in stem failure due to high wind conditions and should be considered in planning reforestation efforts on the Hawaiian Islands.     

Using field survey and remote sensing to assess rainforest canopy damage following Cyclone Larry

Abstract We surveyed canopy damage in upland and lowland rainforests following Cyclone Larry, which severely impacted the Innisfail and Atherton Tableland regions in March 2006. An existing damage‐classification was used as a basis for field assessment of rainforest canopy damage. Our field measurements showed that the damage categories were not clearly separated. Upland and lowland sites significantly differed in tree fall variables and measures of forest structure. There was a difference in recruitment of disturbance indicator species owing to varying levels of coarse woody debris at sites. Aspect was not found to be a significant variable in predicting damage owing to complexity of topography and the cyclone wind field. Analysis of remotely sensed imagery indicated that only high damage levels could be reliably discerned. Areas of very rapid vegetation growth in severely damaged sites are most easily detected with vegetation indices based on both near infrared and short wave infrared data. Numbers of fallen trees and their trunk orientations can be reliably quantified using high resolution (sub‐metre) colour aerial photography. This permits some estimation of whether the wind field was unidirectional or locally vortical.     

Tree Species Traits but Not Diversity Mitigate Stem Breakage in a Subtropical Forest following a Rare and Extreme Ice Storm

Future climates are likely to include extreme events, which in turn have great impacts on ecological systems. In this study, we investigated possible effects that could mitigate stem breakage caused by a rare and extreme ice storm in a Chinese subtropical forest across a gradient of forest diversity. We used Bayesian modeling to correct stem breakage for tree size and variance components analysis to quantify the influence of taxon, leaf and wood functional traits, and stand level properties on the probability of stem breakage. We show that the taxon explained four times more variance in individual stem breakage than did stand level properties; trees with higher specific leaf area (SLA) were less susceptible to breakage. However, a large part of the variation at the taxon scale remained unexplained, implying that unmeasured or undefined traits could be used to predict damage caused by ice storms. When aggregated at the plot level, functional diversity and wood density increased after the ice storm. We suggest that for the adaption of forest management to climate change, much can still be learned from looking at functional traits at the taxon level.     

The impact of wind on trees in Australian tropical savannas: lessons from Cyclone Monica

Abstract The role of tropical cyclones in structuring tropical rainforests has long been recognized in the Americas and northern Australia. However, their role in the dynamics of Australian savannas has received little attention compared with the effects of fire and drought. In April 2006, Tropical Cyclone Monica caused trees to be snapped or uprooted across a swath about 50 km wide and 130 km inland in northern Australia. The proportion of trees damaged was assessed using on‐ground surveys and interpretation of aerial photographs along the cyclone path and along several perpendicular transects. From these data, a relationship between the damage to trees and estimated maximum gust speed was developed. From knowledge of gust recurrence, curves were then derived describing the changes with distance inland in the average recurrence interval for tree windthrow events. These relationships were consistent with the historical record of tree damage in the northwest of the Northern Territory. It was concluded that windthrow has profound, but hitherto unrecognized implications for ecological processes such as tree dynamics and fluxes of carbon and water in Australian tropical savannas.     

The impact of Tropical Cyclone Larry on bird communities in fragments of the endangered rainforest Type 5b

Abstract Bird surveys were conducted to assess the impact of a severe cyclone on bird communities in three fragments of the endangered rainforest Type 5b on the Atherton Tablelands of far north Queensland. Bird communities were surveyed using timed area searches in three sites in each of the three fragments and were undertaken prior to and following Tropical Cyclone Larry. Cyclone Larry caused short‐term changes in the abundance of some species of birds in Type 5b rainforest fragments. Two weeks after the storm, in two of three fragments surveyed, abundance of the frugivorous wompoo fruit‐dove (Ptilinopus magnificus) and figbird (Sphecotheres viridis) had decreased while the omnivorous Macleay's honeyeater (Xanthotis macleayana) and Lewin's honeyeater (Meliphaga lewinii) decreased in abundance in all three locations. Most insectivorous species increased in some sites and decreased in others following Cyclone Larry. Rapid recovery of bird communities to approximately their pre‐cyclone state after only 7 months appeared to reflect the capacity of species to either modify their foraging behaviour, switch foods, or to move within or between fragments or to other food sources in the landscape.     

Recruitment dynamics of invasive species in rainforest habitats following Cyclone Larry

Abstract In tropical forests, natural disturbance creates opportunities for species to claim previously utilized space and resources and is considered an important mechanism in the maintenance of species diversity. However, ecologists have long recognized that disturbance also promotes exotic plant invasions. Cyclones cause extensive defoliation, loss of major branches and multiple tree falls, resulting in a significantly more open canopy and increased light and heat levels in the understorey. The widespread and massive disturbance caused by cyclones provides ideal conditions for rapid recruitment and spread of invasive species. The ecological roles of invasive species in rainforest habitats following such a severe disturbance are poorly understood. Severe category 4 Cyclone Larry crossed the North Queensland coast in March 2006 causing massive disturbance to rainforest habitats from Tully to Cairns and west to the Atherton Tablelands. We established 10 plots in an area extensively damaged by this cyclone near El Arish in North Queensland. On each plot nine 2 × 2 m quadrats were established with three quadrats per plot in each of the following treatments: (i) complete debris removal down to the soil layer, (ii) removal of coarse woody debris only, and (iii) uncleared. We monitored recruitment, growth and mortality of all native and invasive species in the 90 quadrats every 3 months since the cyclone. Here we present the recruitment dynamics of invasive species across the study area in relation to the level of disturbance, the type of quadrat treatment, and the diversity and abundance of the native recruiting flora over the first 12 months post‐cyclone. Our results suggest that invasive species will mostly comprise a transient component of the flora in the early stages of the successional response. However, some species may have longer‐term effects on the successional trajectory of the rainforest and future forest composition and structure.     

热带雨林木质藤本植物叶片性状及其关联

热带雨林中木质藤本植物较为丰富。随着全球气候变化加剧,木质藤本植物的丰富度具有不断增加的趋势,有可能对热带森林的结构、功能和动态产生重要影响。然而,目前对木质藤本响应环境变化的机制所知甚少。本研究以13个科20种热带雨林常见木质藤本植物为材料,测定了冠层叶片的17个形态特征及结构性状,并分析了性状间的相互关系。结果表明,叶片相对含水量的种间变异最小（变异系数为5%）,而上表皮厚度的种间变异最大（变异系数为80%）,其它性状的种间变异系数为24%~61%。木质藤本植物的叶脉密度、叶片密度均与气孔密度呈显著正相关,叶片干物质含量与比叶面积呈显著负相关。与相同生境的树木相比,木质藤本的叶面积更小、气孔密度和叶片密度更低、比叶面积更高,但两种植物类群的叶片横切面组织结构厚度无显著差异。研究结果对理解木质藤本植物的生态适应性具有重要意义。     

Landscape‐scale impacts of Cyclone Larry on the forests of northeast Australia,including comparisons with previous cyclones impacting the region between 1858 and 2006

Abstract Cyclone Larry (category 4) was the most severe cyclone to impact on the Wet Tropics bioregion since the devastating 1918 Innisfail cyclone. Based on an analysis of earlier cyclones impacting on this region over the period 1856–2006, it was determined that Larry was a ‘1 in 50 year’ event. This paper provides an overview of the landscape‐scale impacts of Larry on the forest ecosystems of the Wet Tropics region, based on low‐level helicopter surveys 2 weeks after the event. Cyclone Larry has been described as a ‘midget’ cyclone. Severe forest damage only extended about 30 km from the central track of the cyclone while moderate to severe damage extended some 50 km. Moderate to slight canopy disturbance was rarely identified more than 75 km from the centre of the cyclone's track. Beyond 75 km, forest damage was restricted to exposed areas of elevated terrain and in places exposed to strong lee (gravity) waves from the west. The ecological role of cyclones as important disturbance agents affecting the structure and function of forest ecosystems in the region is discussed, followed by an evaluation of likely effects of climate change on cyclone frequency and intensity.     

Buttress trees in a 20-hectare tropical dipterocarp rainforest in Xishuangbanna,SW China

Aims Buttresses are prevalent and are important to many ecological processes in tropical rainforests but are overlooked in many rainforest studies. Based on a buttress survey in a 20-hectare plot, this study aims to answer the following questions: (I) Is buttress forming a fixed species characteristic? (ii) Is there any phylogenetic signal for buttress forming across a broad taxonomic scale? (iii) Is buttress forming an inherent feature or simply induced by environmental factors, and how is this relevant to the size of the tree?Methods We surveyed buttresses for all 95940 trees with diameter at breast height (DBH) ≥10mm in a 20-ha tropical dipterocarp rainforest in Xishuangbanna, SW China. The occurrence of buttresses was compared across different taxa and across different tree-size classes. A phylogenetic analysis was conducted among buttressed and non-buttressed species in order to understand the evolutionary background of buttress formation.Important findings This preliminary study showed that buttress trees are very abundant (making up 32% of trees with ≥100mm DBH) in this 20-ha tropical rainforest situated at the northern edge of the tropics. Fifty-one percent of the 468 tree species in the plot had stems that produced buttresses. Large trees were more likely to develop buttresses than smaller ones. We found that although buttress formation is not a fixed species characteristic, there is a strong phylogenetic signal for buttress formation in larger species.     

Effect of wood density and water permeability on wood decomposition rates of 32 Bornean rainforest trees

Aims A better understanding of wood litter decomposition is essential for predicting responses of forest ecosystems to global climate change. Recent studies suggest that chemical properties of wood litters, rather than physical ones such as wood density, are more important for interspecific differences in wood decomposition rates. However, empirical data are still limited, especially for tropical trees. In addition, decomposition rate of wood litter often varies with time, which makes interspecific comparison difficult. We studied the wood decomposition of 32 rainforest trees to elucidate (i) the degree of interspecific variation in wood decomposition rate of a given size and configuration and (ii) if initial wood density and water permeability are consistent predictors of the overall decomposition rate and its pattern over time.Methods A common garden decomposition experiment was conducted in a tropical rainforest in Malaysian Borneo for 32 native tree species. Small wood sticks were set on the forest floor and the weight loss was monitored monthly for 2.7 years.Important findings We found large variation in the wood decomposition rate (a 49-fold range), suggesting that we need to consider this variation when calculating community-level carbon dynamics of tropical rain forests. The physical traits of wood, i.e. wood density and water permeability, were related to wood decomposition rate and its pattern over time. Decomposition half-time related positively and negatively to initial wood density and water permeability, respectively. The time-dependent-rate model fitted better for 18 species (56% of the study species) that had higher water permeabilities than the others, suggesting that micelle porosity in wood relates to temporal changes in decomposition rate.     

Delayed induced responses of birch glandular trichomes and leaf surface lipophilic compounds to mechanical defoliation and simulated winter browsing

Changes in morphology and chemistry of leaf surface in response to herbivore damage may increase plant resistance to subsequent herbivore attack; however, there is lack of studies on induced responses of glandular trichomes and their exudates in woody plants and on effects of these changes on herbivores. We studied delayed induced responses in leaf surface traits of five clones of silver birch (Betula pendula Roth) subjected to various types of mechanical defoliation and simulated winter browsing. Glandular trichome density and concentrations of the majority of surface lipophilic compounds increased in trees defoliated during the previous summer. This induced response was systemic, since control branches in branch defoliated trees responded to the treatments similarly to defoliated branches, but differently from control trees. In contrast to defoliation treatments, simulated winter browsing reduced glandular trichome density on the following summer and had fewer effects on individual surface lipophilic compounds. Moreover, constitutive density of glandular trichomes was negatively correlated with induced total amount of lipophilic compounds per trichome, indicating a trade-off between constitutive and induced resistance in silver birch. Induced changes in leaf surface traits had no significant effect on leaf damage by chewers, miners and gall mites, but increased susceptibility of birch trees to aphids. However, leaf damage by chewers, miners and gall mites in defoliated (but not in control) trees was correlated with concentrations of some fatty acids and triterpenoids, although the direction of relationships varied among herbivore species. This indicates that induction of surface lipophilic compounds may influence birch resistance to herbivores. Our study thus demonstrated both specificity of elicitation of induced responses of birch leaf surface traits by different types of damage and specificity of the effects of these responses on different types of herbivores.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

The biomass of New England peppermint (Eucalyptus nova-anglica) in relation to insect damage associated with rural dieback

Two adjacent mature trees of New England peppermint (Eucalyptus nova-anglica) were harvested with the aid of a cherry picker to determine their biomass distribution and insect damage. One suffered from obvious symptoms of rural dieback and the other was healthy. Weights of foliage and wood were measured, and insect damage for all leaves and branches was quantified. For each tree 25% of the roots were extracted from the soil using a bulldozer and manual methods; they were then weighed and damage by insects estimated. The healthy tree lost more leaf surface area to insects (11% or 1.1 kg vs 9.2% or 0.3 kg); but the dieback tree had four times more wood affected by boring insects (19% cf. 5%); and only 20% root biomass remaining (92 kg cf. 488 kg). The accuracy of sampling techniques needed to measure defoliation and the consequences of insect damage to dieback of rural eucalypts are discussed.     

Ontogenetic changes in leaf traits of tropical rainforest trees differing in juvenile light requirement

Relationships between leaf traits and the gap dependence for regeneration, and ontogenetic changes therein, were investigated in juvenile and adult tropical rainforest tree species. The juveniles of the 17 species included in the study were grown in high light, similar to the exposed crowns of the adult trees. The traits were structural, biomechanical, chemical and photosynthetic. With increasing species gap dependence, leaf mass per area (LMA) decreased only slightly in juveniles and remained constant in adults, whereas punch strength together with tissue density decreased, and photosynthetic capacity and chlorophyll increased. Contrary to what has been mostly found in evergreen tropical rainforest, the trade-off between investment in longevity and in productivity was evident at an essentially constant LMA. Of the traits pertaining to the chloroplast level, photosynthetic capacity per unit chlorophyll increased with gap dependence, but the chlorophyll a/b ratio showed no relationship. Adults had a twofold higher LMA, but leaf strength was on average only about 50% larger. Leaf tissue density, and chlorophyll and leaf N per area were also higher, whereas chlorophyll and leaf N per unit dry mass were lower. Ranking of the species, relationships between traits and with the gap dependence of the species were similar for juveniles and adults. However, the magnitudes of most ontogenetic changes were not clearly related to a species’ gap dependence. The adaptive value of the leaf traits for juveniles and adults is discussed.