Effect of host-tree and environmental variables on honeydew production by scale insects (

Honeydew excreted by phloem-sap sucking scale insects (Ultracoelostoma sp.) living in the bark of beech (Nothofagus solandri:) trees growing at a high elevation (900 m) site in the Craigieburn range of Canterbury, New Zealand, was measured over four days during 1–10 May 1996. Average standing crop of honeydew sugar was 3.1 mg m-2, and ranged from 0.4 to 5.5 mg m-2. Daily production of honeydew sugar ranged from 0.2 to 1.5 mg insect-1 24 h-1, and 4.1 to 45.9 mg m-2 24 h-1. Honeydew production varied significantly between trees, and trees with the highest mean individual rates of production (mg sugar insect-1 24 h-1) tended to be those with the highest numbers of insects per unit bark area. Air temperature averaged over the 24 hours preceding each production sample explained 65% of the variability in 3-hourly honeydew production, showing that honeydew production is controlled by environmental and host-tree variables. Using this data, together with previously published estimates of carbon uptake and seasonal variability in honeydew standing crop, it was estimated that carbon contained in honeydew is equivalent to 1.8% of net primary production of beech trees at Craigieburn. Determining the effects of environmental variables on tree and insect physiology will allow formulation of an environmentally-driven process-based model of honeydew production.     

Soluble carbon production by honeydew scale insects in a New Zealand beech forest

We estimated the annual production of honeydew per unit land area of beech (Nothofagus spp.) forest by measuring the amount of honeydew produced in 24 h by scale insects (Ultracoelostoma spp.) (Hemiptera: Margarodidae) every month for 2 years. We used exclosures to prevent animals (notably Vespula wasps) removing honeydew, and we compared the standing crop of honeydew inside permanently closed exclosures with that outside exclosures. Honeydew production and the number of honeydew droplets was highly variable between individual trees, tree type, position on tree, and, exclosure type, and within and between years. The amount of honeydew available outside exclosures was significantly reduced in year 2, predominantly by Vespula wasps, even though wasp density was relatively low. Sugar composition also varied between tree type and between years. Up to 5% of the sugar was glucose, with varying proportions of fructose, sucrose and oligosaccharides. The surface area of trees infested with scale insects was estimated using allometric regression relationships between tree diameter and total surface area of tree trunk and branch material. These estimates were combined with measurements of tree diameter in 10-m radius circular plots to give a production estimate of between 3500 and 4500 kg dry weight honeydew ha^-1 year^-1. Using this data, combined with previously published estimates of carbon uptake, it was estimated that between 6 and 8% of net primary productivity was released as honeydew. Honeydew scale insects provide large amounts of biologically available carbon in the form of soluble sugar. It is a crucial resource for the above-ground system, and probably also for the below-ground system. We conclude that scale insects have the potential to function as keystone species in these forests.     

Competition between Honey-Bees (

Honeydew, the sugary exudate of the scale insect Ultracoelostoma brittini, is an important food source in black beech (Nothofagus solandri var. solandri) forests in the South Island of New Zealand. Two of the most prominent foragers of honeydew are honey bees (Apis mellifera) and wasps (Vespula germanica and V. vulgaris). Observations in the field and using a captive bee hive were used to investigate competition between bees and wasps feeding on honeydew. In laboratory trials, interference competition was often strong, and many cases of aggression were noted. In the forest, there was invariably enough room on the trees for bees and wasps to feed while rarely encountering one another. Over the whole year, environmental variables (especially low temperatures and rain), were found to constrain honey bee foraging to a greater degree than competition with wasps. Because the competition that did occur was primarily exploitation competition, reciprocal effects were likely to be felt. At Coopers Creek, bees may be reducing wasp densities, compared with the situation in Nelson—Marlborough where commercial hives are scarce. It may be possible to reduce wasp densities locally by increasing the number of bee hives in an area.     

Competition for honeydew between two social wasps in South Island beech forests,New Zealand

Summary Honeydew is a sugary secretion of beech scale insects (Ultracoelostoma spp.). Two introduced species of wasps forage on the sugar droplets in New Zealand beech forests. We hypothesize that competition between them may explain whyVespula germanica has become locally extinct in beech (Nothofagus) forest invaded byV. vulgaris. Changes in behaviour of the two wasp species in response to changes in the honeydew resource were monitored. Foraging and rainfall separately and together reduced the honeydew standing crop. In response to the standing crop decreasing, more wasps were found on honeydew trees, they became less active, spent more time lapping the tree surface, and ingested drops of honeydew at a slower rate.V. vulgaris was more active, and obtained drops and energy at a higher rate thanV. germanica. These behavioural differences may lead to competitive advantages affecting queen size and possibly survival.     

Invasive wasps,not birds,dominate in a temperate honeydew system

The Avian Convergence Hypothesis states that avian–honeydew associations are likely to develop when biogeographic and/or climatic factors limit the formation of ant–honeydew associations (the dominant association in tropical ecosystems). In this study we examine a honeydew‐influenced forest system in an island archipelago where ant diversity is low but invasive Vespula wasp species (Vespidae) are present. We found honeydew production was highly seasonal, with both standing crop and 24‐h production peaking in summer. When Vespula wasps were abundant (summer and autumn) they preferentially visited infested trees and fed regularly on honeydew droplets on infested branches. Two ant species occasionally fed on honeydew. No other insects or birds were observed feeding on honeydew during the study period. With the exception of Vespula, honeydew does not appear to be a preferred food source in this community, possibly because of the range of other food resources available in surrounding forest, farmland and gardens. The abundance of Vespula wasps at the site may also have disrupted bird–honeydew associations. We suggest the Avian Convergence Hypothesis could be restated to explicitly include both nectar availability and invasive social insects as both are likely to influence bird use of honeydew.     

Vertical stratification in the spatial distribution of the beech scale insect (Ultracoelostoma assimile) in Nothofagus tree canopies in New Zealand

Abstract. 1. The degree of infestation by New Zealand sooty beech scale insects (Ultracoelostoma assimile, Homoptera: Margarodidae) varies dramatically among adjacent southern beech trees (Nothofagus spp., Fagaceae), but has previously been assumed to be uniformly or randomly distributed within individual host trees. In this study, a full‐census survey was conducted from ground level to canopy level on 14 naturally occurring, canopy‐dominant red beech (Nothofagus fusca) trees (size range 38.7–107.6 cm diameter at breast height) to determine the degree of within‐tree heterogeneity in herbivore density. 2. The within‐tree distribution of the sooty beech scale was vertically stratified and highly heterogeneous, with the greatest densities occurring on bark surfaces in the canopy rather than on the trunk, and on the lower rather than upper sides of the branches. The spatial distribution was strongly negatively correlated with trunk and branch diameter, and increasing bark thickness (as a function of diameter) provides a plausible explanation for differences in the establishment and population density of sooty beech scale insects with trunk and branch size. Furthermore, there was a significant change in the spatial distribution of scale insect populations on trunks and branches of trees of increasing diameter at breast height. This indicates a strong temporal component to the spatial dynamics of the sooty beech scale insect driven by changing host phenology. Future studies on phytophagous insects infesting large host trees need to consider more explicitly changes in population dynamics through space and time. 3. Because of the high degree of within‐tree heterogeneity in population density, the total population size of scale insects on an individual tree could not be predicted from any measure of population density low on the trunk. However, the dry weight biomass of sooty mould fungi growing on the ground beneath infested trees was a remarkably accurate predictor of the total population size of scale insects. The use of sooty mould fungi as a relative measure of population size could be incorporated into studies of other honeydew‐producing hemipterans, since the growth of sooty mould is a distinctive feature synonymous with high concentrations of honeydew production worldwide.     

Distribution and abundance of endemic coelostomidiid scale insects (Hemiptera: Coelostomidiidae) in Auckland forests, New Zealand

Sap-feeding insects can excrete considerable quantities of sugar-rich honeydew. In New Zealand, South Island beech (Nothofagus spp.) forests are shaped by the extensive honeydew resource produced by two endemic coelostomidiid species (Ultracoelostoma assimile and U.?brittini) and geckos on northern islands are known to feed on the honeydew of a third endemic coelostomidiid, Coelostomidia zealandica. There are six other endemic coelostomidiid species in New Zealand that utilise a range of plant hosts but the ecological role of these species is poorly understood. A survey of mainland forests in the Auckland Ecological Region was conducted in February?April 2006 to investigate the distribution and abundance of coelostomidiids in this area. Three coelostomidiid species were detected in the survey (C.?zealandica, C.?pilosa and C.?wairoensis) and five new host?scale insect associations were identified. C.?zealandica was uncommon, C.?pilosa was widespread in broadleaved?podocarp forest but only formed light infestations, and C.?wairoensis was present in all teatree stands examined, often forming heavy infestations on k?nuka (Kunzea ericoides). Infested k?nuka trees had sooty moulds growing on them and exotic wasps were regularly seen feeding on C.?wairoensis honeydew. The extent and intensity of C. wairoensis infestation on k?nuka suggests it will have community-level impacts.     

Separating host-tree and environmental determinants of honeydew production by Ultracoelostoma scale insects in a Nothofagus forest

Abstract 1. Sugar‐rich honeydew excreted (‘produced’) by insects feeding on phloem sap is a key energy flow in a range of temperate and tropical ecosystems. The present study measured honeydew produced by Ultracoelostoma sp. (Homoptera: Coelostomidiidae) scale insects feeding on Nothofagus solandri var. solandri (Hook f.) Oerst. trees in a temperate evergreen forest in New Zealand. Simultaneous measurements of environmental variables and canopy photosynthesis were conducted to allow separation of host‐tree and environmental determinants of honeydew production. These relationships were further examined in experiments where canopy photosynthesis was manipulated by shading or plant nitrogen levels increased by foliar spray. 2. Rates of honeydew production varied nine‐fold from a maximum (± 1 SE) of 64.4 ± 15.2 mg dry mass m⁻² bark h⁻¹ in early summer (December) to a minimum of 7.4 ± 4.2 mg m⁻² h⁻¹ in winter (August). Rates of production measured 1.4 m from the base of the trees’ stems varied significantly with stem diameter, and were higher on medium‐sized (18 cm diameter) than small or large stems. 3. Rates of production were significantly related to environmental conditions over the hours preceding measurement (air temperature and air saturation deficit averaged over the preceding 24 and 12 h respectively). There was no evidence that rates of production were directly related to short‐term changes in the supply of carbohydrates from the canopy (either when compared with measurements of unmanipulated photosynthetic rate, or after sugar levels were manipulated by shading 80% of host‐trees’ leaf area), or to changes in phloem nitrogen content. 4. The results show that there is no clear effect of host‐tree carbon supply on honeydew production; if production is related to photosynthesis, the effect of this is much less important that the large and significant direct effect of environmental conditions on honeydew production.     

Preliminary evidence suggests that beech scale insect honeydew has a negative effect on terrestrial litter decomposition rates in

Honeydew production by New Zealand beech scale insects (Ultracoelostoma spp., Hemiptera: Margarodidae) is widely considered to have a positive influence on native animal abundance and ecosystem functioning. As a first assessment of whether there is a positive relationship between honeydew throughfall and litter decomposition rates, we placed experimental litter bags in each of 10 high and 10 low honeydew plots in mixed southern beech (Nothofagus) forest at each of two sites, Lakehead and Rotoiti, in the Nelson Lakes National Park. High and low honeydew plots were selected using sooty mould biomass on the forest floor as a surrogate for honeydew throughfall, as sooty mould biomass was shown to be strongly correlated (r = 0.906) with scale insect population size at the Rotoiti site. Contrary to our expectation, terrestrial litter decomposition was significantly lower in high honeydew plots than in low honeydew plots, at both Lakehead and Rotoiti. The presence of introduced wasps (Vespula spp., Hymenoptera: Vespidae) at the Lakehead site did not appear to have any significant effect on litter decomposition rates, despite the fact that wasps are thought to intercept much of the honeydew produced in this forest. Variance in litter decomposition rates between high and low honeydew treatments was predominantly determined by a direct relationship between sooty mould biomass and litter decomposition rate at the scale of individual litter bags. However, the mechanistic explanation for the observed relationship is unclear. Future studies should be directed towards quantifying the functional relationship between honeydew throughfall and growth rates of sooty mould, and their subsequent effects on abiotic conditions, microarthropod community dynamics and microbial activity rates in litter.     

Honeydew collection by the invasive garden ant Lasius neglectus versus the native ant L. grandis

Honeydew collection performed by the invasive ant Lasius neglectus and by the native ant L. grandis was compared. The invasive ant collected 2.09 kg of honeydew per tree while the native ant collected 0.82 kg. The aphid Lachnus roboris was visited by both ant species. In holm oaks colonized by L. neglectus, aphid abundance tended to increase and its honeydew production increased twofold. The percentage of untended aphids was lower in holm trees occupied by L. neglectus. As tending ants also prey on insects, we estimated the percentage of carried insects. The native ant workers carried more insects than the invasive ant. Both ant species preyed mainly on Psocoptera and the rarely tended aphid, Hoplocallis picta. We conclude that the higher honeydew collection achieved by L. neglectus was the consequence of (1) its greater abundance, which enabled this ant to tend more Lachnus roboris and (2) its greater level of attention towards promoting an increase of honeydew production. Handling editor: Heikki Hokkanen     

An episodic model of honeydew production in scale insects

Honeydew produced by sooty beech scale insects (Ultracoelostoma spp., Homoptera: Coelostomidiidae) is a keystone ecological process in New Zealand beech (Nothofagus spp., Nothofagaceae) forest. This work puts forward a model of honeydew production based on individual insects that presumes feeding and excretion are episodic processes driven by the insect rather than the passive processes that were previously assumed. The model is parameterized using existing data and then compared to an independent pre‐existing dataset. The model suggests that over a 12‐h period, on average the insects suck sap for 2 h, and excrete waste sap for 12 min. Resource uptake by the insects appears to be limited by the time required to process the sap, consistent with the observed relationship between honeydew production rates and ambient temperature. This implies that insect feeding rates may be ultimately limited by the low nitrogen content of phloem sap.     

Leaf dynamics and insect herbivory in a Eucalyptus camaldulensis forest under moisture stress

Abstract The influence of soil moisture content on leaf dynamics and insect herbivory was examined between September 1991 and March 1992 in a river red gum (Eucalyptus camaldulensis) forest in southern central New South Wales. Long-term observations of leaves were made in trees standing either within intermittently flooded waterways or at an average of 37. 5m from the edge of the waterways. The mean soil moisture content was significantly (P≤0.05) greater in the waterways than in the non-flooded areas. Trees in the higher soil moisture regime produced significantly larger basal area increments and increased canopy leaf area. This increase in canopy leaf area was achieved, in part, through a significant increase in leaf longevity and mean leaf size. Although a greater number of leaves was initiated and abscissed per shoot from the non-flooded trees, more leaves were collected from litter traps beneath the denser canopies of the flooded trees. Consumption of foliage by insects on the trees subjected to flooding compared to the non-flooded trees was not significantly different. However, the relative impact of insect herbivory was significantly greater on the non-flooded trees. Leaf chewing was the most common form of damage by insects, particularly Chryso-melidae and Curculionidae. No species was present in outbreak during this study. Leaf survival decreased as the per cent area eaten per leaf increased. In addition, irrespective of the level of herbivory, leaf abscission tended to be higher in E. camaldulensis under moisture deficit. The influence of soil moisture content on the balance between river red gum growth and insect herbivory is discussed.     

Antifeedant effects of plant lectins and an enzyme on the adult stage of the rice brown planthopper,Nilaparvata lugens

The feeding activity of adult rice brown planthopper,Nilaparvata lugens (Stal) on selected proteins incorporated into artificial diets was examined, based on quantitative analysis of honeydew excretion, in order to determine the mechanism of action of antimetabolic proteins towards homopteran pests. The lectinsGalanthus nivalis agglutinin (GNA), wheat germ agglutinin (WGA) and the enzyme soybean lipoxygenase (LPO), previously demonstrated to be toxic to this insect, reduced honeydew excretion levels of adultNilaparvata lugens over a 24 h period when incorporated into artificial diet at 0.1% {w:v}, indicating that these proteins acted as antifeedants. Of the proteins tested GNA was the most effective antifeedant, reducing honeydew droplet production by 96%, although after 24 h there was some recovery in the honeydew excretion levels and thus the insects appeared to tolerate the presence of the antifeedant with time. The lectinPisum sativum agglutinin (PSA), previously demonstrated to be non-toxic, showed no antifeedant properties.     

Density-dependent effects on the reproductive fitness of the New Zealand beech scale insect (Ultracoelostoma assimile) across multiple spatial scales

Abstract.  1. Intraspecific competition can be an important factor influencing the individual fitness of organisms. This study reports density-dependent effects on the fecundity of the beech scale insect ( Ultracoelostoma assimile , Hemiptera: Margarodidae) on naturally occurring, canopy-dominant red beech ( Nothofagus fusca , Fagaceae) trees in New Zealand. For the first time an increasing intensity of intraspecific competition at increasing spatial scales within individual host trees is demonstrated.
2. Beech scale insect 'tests' containing adult female scale insects and eggs were collected from the trunks of 10 red beech trees [17.7–48.5 cm in diameter at breast height (1.4 m), d.b.h.] with varying densities of scale insect infestation. The relationship between individual female fecundity and scale insect density at three spatial scales: local (within 5 cm), lower trunk (below 2 m above ground level), and whole tree, was tested.
3. Beech scale insect fecundity was density dependent, with total female egg load decreasing with increasing scale insect density. The strength of the density-dependent effect increased with increasing spatial scale, suggesting that scale insects are creating a tree-wide drain on the quality of phloem sap, rather than depleting nutrients from localised, high-density areas within trees. These results indicate that at high densities, the New Zealand beech scale insect can have a negative effect on the nutritional quality of the phloem of red beech, thus negatively affecting conspecifics elsewhere on individual host trees. The increasing intensity of the effect with increasing spatial scale within individual trees emphasises the importance of measuring density-dependent effects at the appropriate spatial scale.     

Effects of insects on primary production in temperate herbaceous communities: a meta-analysis

Abstract.  1. The effects of insects on primary production in temperate herbaceous communities were investigated in a meta-analysis. The following hypotheses were tested: (1) the effect of insects on primary production depends on community type, (2) the effect of insects on primary production varies as a function of productivity, (3) insects have a greater effect on primary production in communities with low species diversity, and (4) insects have a larger effect on primary production during outbreaks.
2. Data were collected from 24 studies in which insecticides were used to suppress insects in self-sown or pastoral communities. Effect sizes were calculated from sprayed and control plot standing crop or yield, expressed as the log response ratio, ln (sprayed plot phytomass/control plot phytomass).
3. There was a significant increase in primary production as a result of insect suppression. Forb-dominated communities showed a more variable response than graminoid communities. During outbreaks, insects had a greater negative impact on primary production. Effect size was unaffected by productivity or plant species richness.
4. Although insects lower primary production in a diversity of temperate herbaceous communities, the basic measures by which such communities are often described have little effect on the proportional impact that insects have on primary production. While outbreaks are significant predictors of higher negative impact on primary production, causes of outbreaks are not always related to traits of the plant community.     

Protection against herbivores of the myrmecophyte Leonardoxa africana (Baill.) Aubrèv. T3 by its principal ant inhabitant Aphomomyrmex afer Emery

Leonardoxa africana T3 is a myrmecophyte, a plant with specialized structures (domatia) that shelter ants. Adult trees are essentially all occupied by the ant Aphomomyrmex afer. One tree possesses one ant colony. Ants tend homopterans inside the domatia. The plant provides ants with nest sites and food via production of extrafloral nectar and via honeydew produced by homopterans. Workers patrol the young leaves, although their nectaries are not yet functional. This study was conducted to investigate the nature of the relationship between the plant and its ants. In order to determine whether ants protect the plant against herbivorous insects, we placed microlepidopteran larvae on young leaves of several trees, and measured the time until discovery of the larvae by the workers. We then studied the responses of workers as a function of insect size. We showed that workers patrolled the young leaves of the majority of trees. There was, however, inter-colony variability in intensity of patrolling. Workers attacked every larva they found, killing and eating the smaller ones, and chasing larger ones off the young leaf. Most of the phytophagous insects attacking young leaves of L. africana T3 were inventoried in this study. We showed that the larvae of microlepidopterans, one of the most important herbivores of this species, form part of the diet of A. afer. The function of the stereotyped behaviour of ant patrolling on young leaves may be in part to obtain insect protein to complement carbohydrate-rich nectar and honeydew, and in part to protect the host and thus increase its production of resources for ants. Our study shows that ants protect the tree against herbivores, and that even if this protection is less pronounced and more variable than that demonstrated for their sister species L. africana sensu stricto and Petalomyrmex phylax, the association between L. africana T3 and A. afer is a mutualism.     

紫胶虫蜜露对地表蚂蚁多样性的影响

2009年12月至2010年5月,采用陷阱法在云南省墨江县雅邑乡调查了紫胶林地表蚂蚁群落多样性,分析了紫胶虫蜜露对地表蚂蚁多样性的影响.结果表明:紫胶虫蜜露资源的有无及变动对地表蚂蚁群落物种组成、多度及多样性均产生影响.在紫胶林样地共采集蚂蚁标本4953头,隶属5亚科23属34种,在对照样地共采集蚂蚁标本2416头,隶属5亚科20属30种;紫胶林地表蚂蚁相对多度、物种丰富度(S)及ACE估计值均高于对照样地,地表蚂蚁常见种和指示种均与对照样地不同,表明放养紫胶虫改变了地表蚂蚁群落结构;紫胶虫成虫期蜜露分泌量高于幼虫期,其地表蚂蚁相对多度、S及ACE估计值也高于幼虫期,且两阶段的蚂蚁常见种和指示种显著不同.     

Comparative benthic plant ecology by SCUBA-monitored quadrats

The feasibility of permanent benthic quadrats checked by SCUBA divers was tested. Stations were set up on the bottom of six representative aquatic habitats, both marine and inland, including oligotrophic and eutrophic ponds, river, coastal lagoon, estuary, and open marine coast. Data include physical-chemical factors, species composition, number and height of shoots, and general observations. From volume/length constants, the standing crops per quadrat were calculated. Seasonal changes in environmental factors and plant standing crop are parallel-plotted for the six habitats, thus providing comparative information on stations. Benthic diel periodicity and light intensity periodicity at various depths were investigated also. In all habitats, maximum standing crop occurred in July and August, but species composition varied considerably. Length of time the water temperature exceeded 15C did not correlate with standing crop. Dissolved oxygen was never deficient, even under ice; and diurnal changes were slight. Light was diminished under ice, but 4–60 ft-c were recorded in midwinter. Lenght of day is about 2 hours shorter at 20 ft than at 6 ft. Onset of vernal benthic growth seems correlated more closely with increase in light intensity than with temperature. The permanent quadrats monitored by divers proved feasible in all habitats tested, and directly comparable data were obtained.Supported in part by a grant from the National Science Foundation.     

The value of insect pollination to yield of oilseed rape (Brassica rapa) in Bangladesh

Brassica rapa L., is a crop grown globally and studies have indicated that insect pollination can improve yields. However, the importance of insect pollination in this crop depends on cultivar reproductive biology, insect pollinator species and their abundances. In Bangladesh, the acreage of B. rapa for oilseed production has been expanding, but little is known about whether insects contribute to yield improvements. Using the commonly grown variety Tori-7, we found that plants with inflorescences left exposed to flower visiting insects (body width > 1.5 mm) had a 30.8% greater seed yield compared to those where insect flower visitors were excluded. Of 794 insect flower visiting individuals recorded from observation and trapping surveys conducted across four separate fields, Apis bees (four spp.) were the most abundant (66.1%), followed by flies Musca domestica (14.7%), Sepsis fulgens (6.3%) and hoverflies (3.0%; 3 spp; Syrphidae). Other flower visitors included ants, wasps, beetles, butterflies and moths. For the cultivar assessed, we calculated the economic value of oilseed rape was $US 87.5 million per annum in Bangladesh, of which the economic value of insect pollination was $US 26.92 million per annum. Thus, one in every four dollars earned by our Bangladeshi growers resulted from insect crop pollination. This demonstrates to growers the need to promote and protect insect pollination to optimize their economic returns.     

Water stress,leaf nutrients and defoliation: a model of dieback of rural eucalypts

Water stress, concentrations of foliar nutrients and damage of foliage by insects were studied over an eleven month period, for eight dieback eucalypts and eight closely matched healthy trees growing in close association on two grazing properties near Brisbane. Four healthy eucalypts in a neighbouring State Forest were also studied. The study region had suffered severe climatic stress between 1972 and 1976 when dieback was first observed by residents and when high populations of defoliating insects were observed. Dieback trees had a higher proportion of foliage damaged by insects and higher concentrations of foliar nutrients than did matched healthy trees. The study period was one of low climatic stress during which dieback and healthy trees followed similar seasonal patterns of predawn xylem pressure potential. Dieback trees developed lower daytime minimum xylem pressure potentials than did matched healthy trees, and differed in their stomatal responses. A model of initiation and development of rural eucalypt dieback is proposed. This may have general application to many non-specific rural diebacks in which heavy insect damage is implicated. The balance between rural eucalypts and their insect herbivores is precarious. Any factor capable of causing extensive defoliation, or an increase in foliar nitrogen, or an increase in populations of insect herbivores may upset this balance. A positive feedback loop may be activated, whereby the production of nitrogen rich epicormic foliage enhances a build up of insect populations. Repeated insect defoliation leads to tree dieback. Evidence in the literature supporting the model is reviewed and aspects requiring further research are outlined.