Phytophagous insect-woody sprout interactions in tropical eucalypt forest. II. Insect community structure

Abstract Sucking insects constituted 79% of all phytophagous insects collected from woody sprouts in the ground layer of a tropical eucalypt forest. Mobile insect groups such as non-psyllid Hemiptera and Orthoptera were relatively frequent in this environment compared to temperate, Eucalyptus-dominated vegetation. The high fire frequency of the tropical eucalypt forest may favour mobile insect groups. The capture of sucking insects and caterpillars peaked in dry season samples. Other patterns of abundance of phytophagous insect groups showed little consistency in their seasonal trends between host species or between vegetation types within host species. Disparities between chewing insect abundance in daytime samples and the damage chewing insects cause, may result from disproportionate consumption by large, mainly nocturnal insects, such as members of the Orthoptera. In this study, 21% of insect species were specialists on single plant species. This study suggested that insect abundance reflected the growth patterns of woody sprouts after regular burning, rather than that plant growth and development were tuned to the pressures of insect herbivory.     

Phytophagous insect communities in the canopies of three Eucalyptus forest types in south-eastern Australia

Intensive sampling of tree canopies for phytophagous insects was carried out in three contrasting eucalypt forest types comprised of species widely distributed in sub-alpine forests in Victoria and New South Wales (Eucalyptus delegatensis, E. dives, and E. pauciflora). The number of phytophagous insects present in the canopies of these forest types was low, with a seasonal average of 20 individuals per kg of foliage (dry weight). Numbers were much lower than expected from past literature reporting‘chronically high’levels of defoliation in eucalypt forests. Microlepidoptera, Geometridae, Chrysomelidae, and Curculionidae were the major leaf-chewing groups recorded. Most sap-feeders were either leafhoppers (Cercopidae and Cicadellidae) or in the superfamily Fulgoroidea. Psyllidae and gall-making species were rare. Leafhoppers made up a very large portion of the phytophagous insect communities in each forest type, particularly in the E. dives forest. Microlepidoptera was the most commonly encountered defoliator group in all three forest types making up 33-44% of the total count. Non-phytophagous arthropods accounted for 44-48% of all individuals encountered. The density of insect defoliators was greater in the lower crown than upper crown. The E. dives canopy supported many more phytophagous insects per unit weight of foliage, as well as more per hectare, than the other two forest types. The greatest number of phytophagous species was also encountered in E. dives canopy. The E. delegatensis canopy supported the lowest number of phytophagous insects per unit weight of foliage as Well as numbers per hectare. Abundance of insect defoliators in the eucalypt forest types in this study was similar to published figures of insect defoliators in northern temperate forests.     

Colonization of eucalypts in New Zealand by Australian insects

Since the 1860s, Australian insects have steadily colonized eucalypts in New Zealand. The rate of colonization has increased markedly over the last two decades. This increase may be related to increasing trade between the two countries. Currently there are 26 specialist eucalypt insect species and approximately 31 polyphagous insect species that can feed on Eucalyptus in New Zealand. The specialist eucalypt insects endemic to Australia have generally caused more damage than polyphagous or native insects. Eucalypt‐specific insects are dominated by sap sucking bugs, particularly psyllids, and defoliating Coleoptera and Lepidoptera. In some cases the major insect pest species have been those that are only occasional pests in Australia, for example Gonipterus scutellatus, Ctenarytaina eucalypti, Eriococcus coriaceus and Phylacteophaga froggatti. Some important insect pests have been rare, or even not described from Australia, prior to their appearance as a pest in New Zealand, for example Paropsis charybdis and Ophelimus eucalypti. Invading eucalypt insects are more likely to establish in the Auckland region than anywhere else in New Zealand.     

Australian biogeographical connections and the phylogeny of large genera in the plant family Myrtaceae

Aim To compare the phylogeny of the eucalypt and melaleuca groups with geological events and ages of fossils to discover the time frame of clade divergences. Location Australia, New Caledonia, New Guinea, Indonesian Archipelago. Methods We compare published molecular phylogenies of the eucalypt and melaleuca groups of the plant family Myrtaceae with geological history and known fossil records from the Cretaceous and Cenozoic. Results The Australasian eucalypt group includes seven genera, of which some are relictual rain forest taxa of restricted distribution and others are species‐rich and widespread in drier environments. Based on molecular and morphological data, phylogenetic analyses of the eucalypt group have identified two major clades. The monotypic Arillastrum endemic to New Caledonia is related in one clade to the more species‐rich Angophora, Corymbia and Eucalyptus that dominate the sclerophyll vegetation of Australia. Based on the time of rifting of New Caledonia from eastern Gondwana and the age of fossil eucalypt pollen, we argue that this clade extends back to the Late Cretaceous. The second clade includes three relictual rain forest taxa, with Allosyncarpia from Arnhem Land the sister taxon to Eucalyptopsis of New Guinea and the eastern Indonesian archipelago, and Stockwellia from the Atherton Tableland in north‐east Queensland. As monsoonal, drier conditions evolved in northern Australia, Arnhem Land was isolated from the wet tropics to the east and north during the Oligocene, segregating ancestral rain forest biota. It is argued also that the distribution of species in Eucalyptopsis and Eucalyptus subgenus Symphyomyrtus endemic in areas north of the stable edge of the Australian continent, as far as Sulawesi and the southern Philippines, is related to the geological history of south‐east Asia‐Australasia. Colonization (dispersal) may have been aided by rafting on micro‐continental fragments, by accretion of arc terranes onto New Guinea and by land brought into closer proximity during periods of low sea‐level, from the Late Miocene and Pliocene. The phylogenetic position of the few northern, non‐Australian species of Eucalyptus subgenus Symphyomyrtus suggests rapid radiation in the large Australian sister group(s) during this time frame. A similar pattern, connecting Australia and New Caledonia, is emerging from phylogenetic analysis of the Melaleuca group (Beaufortia suballiance) within Myrtaceae, with Melaleuca being polyphyletic. Main conclusion The eucalypt group is an old lineage extending back to the Late Cretaceous. Differentiation of clades is related to major geological and climatic events, including rifting of New Caledonia from eastern Gondwana, development of monsoonal and drier climates, collision of the northern edge of the Australian craton with island arcs and periods of low sea level. Vicariance events involve dispersal of biota.     

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.     

Dual colonization of Eucalyptus urophylla S.T. Blake by arbuscular and ectomycorrhizal fungi affects levels of insect herbivore attack

Abstract 1 Eucalypts are an important part of plantation forestry in Asia but, in south China, productivity is very low. This is due to infertile soils and lack of indigenous symbiotic mycorrhizal fungi. The genus Eucalyptus is unusual because it forms both arbuscular (AM) and ectomycorrhizal (ECM) associations. 2 Eucalyptus urophylla saplings were grown with and without AM (Glomus caledonium) and ECM (Laccaria laccata) fungi in a factorial design. Two experiments were performed: one to simulate nursery conditions and the other to simulate the early stages of plantation establishment. Plant growth was measured over 18 weeks and levels of insect attack were recorded. 3 The AM fungus reduced tree growth in the early stages, but the effect appeared to be transient. No effects of ECM were detected on tree growth, but the ectomycorrhiza reduced colonization by the arbuscular mycorrhiza. AM fungi appear to be rapid invaders of the root system, gradually being replaced by ECM. 4 Both fungal types affected levels of damage by insect herbivores. Most importantly, herbivory by the pest insects Anomala cupripes (Coleoptera) and Strepsicrates spp. (Lepidoptera) was decreased by ECM. 5 It is suggested that mycorrhizal effects on eucalypt insects may be determined by carbon allocation within the plant. Future studies of eucalypt mycorrhizas need to take into account the effects of the fungi on foliar‐feeding insects and also the effects of insect herbivory on mycorrhizal establishment.     

An omics evolutionary perspective on phytophagous insect–host plant interactions in Anastrepha obliqua: a review

Phytophagous insects have a close relationship with their host plants. For this reason, their interactions can lead to important changes in insect population dynamics and evolutionary trajectories. Next generation sequencing (NGS) has provided an opportunity to analyze omics data on a large scale, facilitating the change from a classical genetics approach to a more holistic understanding of the underlying molecular mechanisms of host plant use by insects. Most studies have been carried out on model species in Holarctic and temperate zones. In tropical zones, however, the effects of use of various host plants on evolutionary insect history is less understood. In the current review, we describe how omics methodologies help us to understand phytophagous insect–host plant interactions from an evolutionary perspective, using as example the Neotropical phytophagous insect West Indian fruit fly, Anastrepha obliqua (Macquart) (Diptera: Tephritidae), an economically important fruit crop pest in the Americas. Anastrepha obliqua could adopt a generalist or a specialist lifestyle. We first review the adaptive molecular mechanisms of phytophagous insects to host plants, and then describe the main tools to study phytophagous insect–host plant interactions in the era of omics sciences. The omics approaches will advance the understanding of insect molecular mechanisms and their influence on diversification and evolution. Finally, we discuss the importance of a multidisciplinary approach that integrates the use of omics tools and other, more classical methodologies in evolutionary studies.     

Reducing the impact of insect herbivory in eucalypt plantations through management of extrinsic influences on tree vigour

In this paper it is argued that concepts developed in ecologically derived insect–plant interaction models can contribute directly to the management of insect herbivory in eucalypt plantations. Common to most species of commercially planted eucalypt is their genetic potential for early rapid growth. Several plant defence theories predict that intrinsically fast growing plants are able to tolerate relatively high levels of herbivory. The risk of this strategy failing increases when plants are exposed to external stressful factors that reduce canopy growth and vigour. Results from a young Eucalyptus camaldulensis plantation stressed by moisture deficit and two young Eucalyptus dunnii plantations, stressed by flooding and weed competition, respectively, are summarized. In all three cases, the stress‐inducing agents reduced canopy growth rates and architecture so that the proportion of leaf tissue damaged by insects increased and the tree’s ability to tolerate this damage decreased. Therefore, alleviating tree stress through improved silvicultural practices or improved site selection techniques may indirectly reduce the impact of insect herbivory. In resource‐limiting environments, an alternative approach may be to plant eucalypt species that although slower growing, are predicted to have better defended foliage. Manipulation of these natural antiherbivore plant strategies are not exclusive of other management approaches, such as the need for routine surveillance of key pest insects or the genetic selection of natural insect resistance and selective chemical control techniques, but should be viewed as an overarching concept for plantation health.     

Experimental warming increases herbivory by leaf‐chewing insects in an alpine plant community

Climate warming is predicted to affect species and trophic interactions worldwide, and alpine ecosystems are expected to be especially sensitive to changes. In this study, we used two ongoing climate warming (open‐top chambers) experiments at Finse, southern Norway, to examine whether warming had an effect on herbivory by leaf‐chewing insects in an alpine Dryas heath community. We recorded feeding marks on the most common vascular plant species in warmed and control plots at two experimental sites at different elevations and carried out a brief inventory of insect herbivores. Experimental warming increased herbivory on Dryas octopetala and Bistorta vivipara. Dryas octopetala also experienced increased herbivory at the lower and warmer site, indicating an overall positive effect of warming, whereas B. vivipara experienced an increased herbivory at the colder and higher site indicating a mixed effect of warming. The Lepidoptera Zygaena exulans and Sympistis nigrita were the two most common leaf‐chewing insects in the Dryas heath. Based on the observed patterns of herbivory, the insects life cycles and feeding preferences, we argue that Z. exulans is the most important herbivore on B. vivipara, and S. nigrita the most important herbivore on D. octopetala. We conclude that if the degree of insect herbivory increases in a warmer world, as suggested by this study and others, complex interactions between plants, insects, and site‐specific conditions make it hard to predict overall effects on plant communities.     

Incongruence between chloroplast and species phylogenies in Eucalyptus subgenus Monocalyptus (Myrtaceae)

Seventy-eight polymorphic cpDNA (chloroplast DNA) characters were found in 13 closely related taxa from Eucalyptus series Amygdalinae (subgenus Monocalyptus) and seven potential outgroup taxa. The strict consensus of six cladograms generated from cpDNA data confirmed monophyly of Monocalyptus. However, cpDNA phylogeny within Monocalyptus was incongruent with taxonomic classification, being more related to geography, even when accessions were from divergent series. Monocalyptus cpDNA formed two major clades. On the island of Tasmania cpDNA was restricted to a single clade, exhibited very little variation, and was phylogenetically related to cpDNA found in central and western Victoria. In contrast, cpDNA of mainland monocalypt taxa was more variable, even within the Amygdalinae. Four out of six Tasmanian Amygdalinae species were polymorphic. The difference between cpDNA of replicates was often greater than differences between species from different series. The low level of cpDNA variation and extensive morphological intergradation between the Tasmanian endemics suggest recent speciation. However, the transfer of cpDNA through hybridization between lineages is the most likely explanation for the observed sharing of cpDNA across series. This study highlights that the geographical pattern to cpDNA variation in Eucalyptus may be an important source of information on past plant distributions in Australia.     

Insect pests of Tasmanian blue gum,Eucalyptus globulus globulus,in south‐western Australia: History,current perspectives and future prospects

Trends in, and potential causes of, insect pest problems of the Tasmanian blue gum, Eucalyptus globulus globulus, in south‐western Australia are reviewed. Historical evidence suggests that insect pest problems of E. g. globulus in south‐western Australia have greatly increased in the last 10 years, which corresponds to a time of rapid expansion of the blue gum industry in the region. Current major establishment pests include the African black beetle, Heteronychus arator, spring beetles, Liparetrus spp. and Heteronyx spp., and the wingless grasshopper, Phaulacridium vittatum. Current major pests of established trees are the Eucalyptus weevil, Gonipterus scutellatus, and chrysomelid beetles, Chrysophtharta spp. and Cadmus excrementarius. The occurrence of these insects on an introduced eucalypt is not unexpected because insect‐rich native eucalypt forests dominate the landscape where E. g. globulus plantations are grown. Insect damage may also be exacerbated because E. g. globulus is grown as a monoculture.     

Differences among five amaranth varieties (Amaranthus spp.) regarding secondary metabolites and foliar herbivory by chewing insects in the field

In this study, we determined the abundance of secondary metabolites present in leaves of five varieties of Amaranthus, described the community of chewing insects observed in the foliage and also quantified damage by folivore insects in the field. Three flavonoid glucosides (rutin, nicotiflorin and isoquercitin), nine phenolic compounds (coumaric, vanillic, caffeic, syringic, ferulic, sinapic, protocatechuic, salicylic and 4-hydroxybenzoic acid) and three betalains (amaranthine, iso-amaranthine and betanin) were found to be present in amaranth leaves. Flavonoids appeared in of all varieties analyzed, with rutin being the most important. Betalains occurred only in some varieties and at different proportions, and nine phenolic acids were observed in all the varieties, with the exception of sinapic acid. Significant differences in the chemical composition of the varieties were noted. A total of 17 species of chewing phytophagous insects were observed through visual counting in Amaranthus plants, with the order Coleoptera being the most important and having the highest diversity of species. The degree of herbivory differed significantly among the varieties. Multivariate regression analysis indicated that the eight analyzed compounds detected in the plants had significant linear relationships with herbivory in the field. However, to draw any conclusions relating the amount of any compound to the degree of herbivory damage is premature at this stage of the research.     

No detectable impacts of frequent burning on foliar C and N or insect herbivory in an Australian eucalypt forest

Question: What is the effect of frequent low intensity prescribed fire on foliar nutrients and insect herbivory in an Australian eucalypt forest? Location: Lorne State Forest (Bulls Ground Frequent Burning Study), mid‐north coast, New South Wales, Australia. Methods: Eighteen independent sites were studied representing three experimental fire regimes: fire exclusion (at least 45 years), frequently burnt (every 3 years for 35 years) and fire exclusion followed by the recent introduction of frequent burning (two fires in 6 years). Mature leaves were collected from the canopy of Eucalyptus pilularis trees at each site and analysed for nutrients and damage by invertebrate herbivores. Results: Almost 75% of all leaves showed some signs of leaf damage. The frequency of past fires had no effect on carbon and nitrogen content of canopy leaves. These results were consistent with assessments of herbivore damage where no significant differences were found in the amount of invertebrate herbivory damage to leaves across fire treatments. Conclusions: This eucalypt forest displayed a high degree of resilience to both frequent burning and fire exclusion as determined by foliar nutrients and damage by insect herbivores. Fire frequency had no detectable ecological impact on this aspect of forest health.     

Assessing density–damage relationships between water hyacinth and its grasshopper herbivore

Plants are variable in their responses to insect herbivory. Experimental increases in densities of phytophagous insects can reveal the type of plant response to herbivory in terms of impact and compensatory ability. The relationship between insect density and plant damage of a grasshopper, Cornops aquaticum Brüner (Orthoptera: Acrididae: Tetrataeniini), a candidate biological control agent, and an invasive aquatic plant, water hyacinth, Eichhornia crassipes Mart. Solms‐Laubach (Pontederiaceae), was investigated to assess potential damage to the weed. The impact of different densities of male and female grasshoppers on E. crassipes growth parameters was determined in a quarantine glasshouse experiment. Damage curves indicated that the relationship between plant biomass reduction and insect density was curvilinear whereas leaf production was linear. Female C. aquaticum were more damaging than males, causing high rates of plant mortality before the end of the trial at densities of three and four per plant. Feeding by C. aquaticum significantly reduced the total plant biomass and the number of leaves produced, and female grasshoppers caused a greater reduction in the number of leaves produced by water hyacinth plants than males. Grasshopper herbivory suppressed vegetative reproduction in E. crassipes, suggesting C. aquaticum could contribute to a reduction in the density and spread of E. crassipes infestations. The results showed that E. crassipes vigour and productivity decreases with an increase in feeding intensity by the grasshopper. Cornops aquaticum should therefore be considered for release in South Africa based on its host specificity and potential impact on E. crassipes.     

Top-down effects of insect herbivores during early succession: influence on biomass and plant dominance

We tested the hypothesis that phytophagous insects would have a strong top-down effect on early successional plant communities and would thus alter the course of succession. To test this hypothesis, we suppressed above-ground insects at regular intervals with a broad-spectrum insecticide through the first 3 years of old-field succession at three widely scattered locations in central New York State. Insect herbivory substantially reduced total plant biomass to a similar degree at all three sites by reducing the abundance of meadow goldenrod, Solidago altissima. As a result, Euthamia graminifolia dominated control plots whereas S. altissima dominated insecticide-treated plots by the third year of succession. S. altissima is the dominant old-field herbaceous species in this region but typically requires at least 5 years to become dominant. Past explanations for this delay have implicated colonization limitation whereas our data demonstrate that insect herbivory is a likely alternative explanation. A widespread, highly polyphagous insect, the xylem-tapping spittlebug, Philaenus spumarius, appeared to be the herbivore responsible for the reduction in standing crop biomass at all three sites. Insect herbivory typically caused little direct leaf tissue loss for the ten plant species we examined, including S. altissima. Consequently, the amount of leaf area removed was not a reliable indicator of the influence of insect herbivory on standing crop biomass or on early succession. Overall, we found a strong top-down effect of insect herbivores on biomass at several sites, so our results may be broadly applicable. These findings run counter to generalizations that top-down effects of herbivores, particularly insects, are weak in terrestrial systems. These generalizations may not apply to insects, such as spittlebugs, that can potentially mount an effective defense (i.e., spittle) against predators and subsequently reach relatively high abundance on common plant species. Our results suggest that insect herbivory may play an important but often overlooked role during early old-field succession. Received: 26 December 1998 / Accepted: 3 April 1999     

Benefits conferred by “timid” ants: active anti-herbivore protection of the rainforest tree Leonardoxa africana by the minute ant Petalomyrmex phylax

In this study, we demonstrate that an important benefit provided by the small host-specific ant Petalomyrmex phylax to its host plant Leonardoxa africana is efficient protection against herbivores. We estimate that in the absence of ants, insect herbivory would reduce the leaf area by about one-third. This contributes considerably to the fitness of the plant. Our estimates take into account not only direct damage, such as removal of leaf surface by chewing insects, but also the effects of sucking insects on leaf growth and expansion. Sucking insects are numerically predominant in this system, and the hitherto cryptic effects of ant protection against the growth-reducing effects of sucking insects accounted for half of the total estimated benefit of ant protection. We propose that the small size of workers confers a distinct advantage in this system. Assuming that resource limitation implies a trade off between size and number of ants, and given the small size of phytophagous insects that attack Leonardoxa, we conclude that fine-grained patrolling by a large number of small workers maximises protection of young leaves of this plant. Since herbivores are small and must complete their development on the young leaves of Leonardoxa, and since a high patrolling density is required for a fine-grained search for these enemies, numerous small ants should provide the most effective protection of young leaves of Leonardoxa. We also discuss other factors that may have influenced worker size in this ant. Received: 1 September 1996 / Accepted: 2 June 1997     

The constraints of selecting for insect resistance in plantation trees

• 1 High productivity in plantations of exotic tree species is achieved by management for fast growth in the absence of the full complex of co‐evolved insect herbivores. In the case of Eucalyptus, silvicultural selection for desirable wood traits is concomitant with a trade‐off against defence and a reduction of chemical and genetic diversity. These factors, combined with accidental introductions, rapid insect evolution and the emergence of new pests, increase the likelihood that future plantations will need insect pest management to maintain productivity.
• 2 Forestry researchers have suggested that selecting for resistant genotypes may be beneficial in insect control. There are, however, significant differences between long‐lived trees and annual crops that make this approach unlikely to be successful. This is illustrated using several examples of research into resistance to insect herbivores in trees.
• 3 Selection for resistance to insects in trees requires an assessment of trial plantations for heritable variation in insect damage and then a determination of the effect of variation in resistance on insect population parameters. Identifying rare resistant genotypes using markers is difficult because many factors interact to produce a resistant phenotype, and phytophagous insects have less intimate relationships with their host than pathogens, resulting in weak associations with genetic loci.
• 4 If resistant genotypes are identified, their widespread deployment in plantations might not provide satisfactory management of insect pests when the use of extensive monocultures is continued. In this paper, experiments are suggested that would explore the effectiveness of polycultures or chemotype mixtures with respect to ameliorating the damage of insects on plantation productivity. In addition, mitigating the effects of some insects on plantation productivity by maintaining vigour of fast‐growing eucalypts should be considered.

     

The effect of the taxon and geographic range size of host eucalypt species on the species richness of gall-forming insects

Abstract This field study was designed to test whether the taxonomic group and geographic range size of a host plant species, usually found to influence insect species richness in other parts of the world, affected the number of gall species on Australian eucalypts. We assessed the local and regional species richness of gall-forming insects on five pairs of closely related eucalypt species. One pair belonged to the subgenus Corymbia, one to Monocalyptus, and three to different sections of Symphyomyrtus. Each eucalypt pair comprised a large and a small geographic range species. Species pairs were from coastal or inland regions of eastern Australia. The total number of gall species on eucalypt species with large geographic ranges was greater than on eucalypt species with small ranges, but only after the strong effect of eucalypt taxonomic grouping was taken into account. There was no relationship between the geographic range size of eucalypt species and the size of local assemblages of gall species, but the variation in insect species composition between local sites was higher on eucalypt species with large ranges than on those with small ranges. Thus the effect of host plant range size on insect species richness was due to greater differentiation between more widespread locations, rather than to greater local species richness. This study confirms the role of the geographic range size of a host plant in the determination of insect species richness and provides evidence for the importance of the taxon of a host plant.     

Phytophagous insect-woody sprout interactions in tropical eucalypt forest. I. Insect herbivory

Abstract Foliar insect damage levels on woody sprouts in the ground layer of two tropical eucalypt forest communities on Melville Island were between 7.8 and 43.2%. Of eight common tree species, Eucalyptus confertiflora was damaged most by insects and Buchanania obovata and Terminalia ferdinandiana the least. Seasonal trends in insect damage were not consistent between plant species and were not always consistent between vegetation types for a particular plant species. The results of this study are not consistent with hypotheses suggesting that insect grazing is a critical determinant of tree species dominance or woody sprout dormancy.     

Bird predation enhances tree seedling resistance to insect herbivores in contrasting forest habitats

According to the associational resistance hypothesis, neighbouring plants are expected to influence both the insect herbivore communities and their natural enemies. However, this has rarely been tested for the effects of canopy trees on herbivory of seedlings. One possible mechanism responsible for associational resistance is the indirect impact of natural enemies on insect herbivory, such as insectivorous birds. But it remains unclear to what extent such trophic cascades are influenced by the composition of plant associations (i.e. identity of ‘associated’ plants). Here, we compared the effect of bird exclusion on insect leaf damage for seedlings of three broadleaved tree species in three different forest habitats. Exclusion of insectivorous birds affected insect herbivory in a species-specific manner: leaf damage increased on Betula pendula seedlings whereas bird exclusion had no effect for two oaks (Quercus robur and Q. ilex). Forest habitat influenced both the extent of insect herbivory and the effect of bird exclusion. Broadleaved seedlings had lower overall leaf damage within pine plantations than within broadleaved stands, consistent with the resource concentration hypothesis. The indirect effect of bird exclusion on leaf damage was only significant in pine plantations, but not in exotic and native broadleaved woodlands. Our results support the enemies hypothesis, which predicts that the effects of insectivorous birds on insect herbivory on seedlings are greater beneath non-congeneric canopy trees. Although bird species richness and abundance were greater in broadleaved woodlands, birds were unable to regulate insect herbivory on seedlings in forests of more closely related tree species.