The Seedling Community of Tropical Rain Forest Edges and Its Interaction with Herbivores and Pathogens1

Edge effects alter biotic interactions and forest regeneration. We investigated whether edge creation affected the seedling community and its interactions with herbivores and leaf‐fungal pathogens. In forest edges and interior sites in Chiapas, Mexico, we counted all woody seedlings and species (10–100 cm tall) present in 1‐m² plots, measured their size (height and leaf number), and examined them for the occurrence of herbivory and pathogen damage. We investigated relationships between levels of damage and size, species richness and density. Species composition and biotic damage varied greatly among sites and habitats (edge vs interior). Late‐successional species dominated the community, but richness was lower in interior sites and species similarity was greater among edges than among interiors. Nearly all species (95%) present at edges and interiors showed herbivory damage, whereas 76 percent of the species in edge plots and 68 percent in interior plots showed pathogen damage. Although leaf area damaged by herbivores was similar between habitats (average 9.2%), pathogen damage was three times greater in edge plots (1.85%) than in interior plots (0.57%). Size was positively related to biotic damage at both habitats. Relationships between herbivory and pathogen damage and between pathogen damage and leaf number were significant only for edges. Biotic damage was not related to density or species richness. Overall, plant community structure was similar between habitats, but biotic damage was enhanced at edges. Thus, disease spread at edges may arise as a threat to tropical rain forest vegetation.     

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.     

Induced biotic responses to herbivory and associated cues in the Amazonian ant-plant Maieta poeppigii

Ants inhabiting ant‐plants can respond to cues of herbivory, such as the presence of herbivores, leaf damage, and plant sap, but experimental attempts to quantify the dynamic nature of biotic defenses have been restricted to a few associations between plants and ants. We studied the relationship between certain features of the ant‐shrub Maieta poeppigii Cogn. (Melastomataceae) and the presence or absence of ant patrolling on the leaf surface in plants occupied by the ant Pheidole minutula Mayr (Hymenoptera: Formicidae). We also carried out field experiments to examine ant behavior following plant damage, and the potential cues that induce ant recruitment. These experiments included clipping of the leaf apex, as well as the presentation of a potential herbivore (live termite worker) and a foliar extract from Maieta on treatment leaves. The presence of ants patrolling the leaves of M. poeppigii is influenced by the number of domatia on the plant. Ant patrolling on the leaves of M. poeppigii was constant throughout a 24 h cycle, but the mean number of patrolling ants decreased from young to mature leaves, and from leaves with domatia to those without domatia. There was an overall increase in the number of ants on experimental leaves following all treatments, compared to control leaves. Visual and chemical cues associated with herbivory are involved in the induction of ant recruitment in the Maieta–Pheidole system. The continuous patrolling behavior of ants, associated with their ability to respond rapidly to foliar damage, may result in the detection and repellence/capture of most insect herbivores before they can inflict significant damage to the leaves.     

Performance and fate of tree seedlings on and around nests of the leaf‐cutting ant Atta cephalotes: Ecological filters in a fragmented forest

Habitat fragmentation is currently the most pervasive anthropogenic disturbance in tropical forests and some species of leaf‐cutting ants of the genus Atta (dominant herbivores in the neotropics) have become hyper‐abundant in forest edges where their nests directly impact up to 6% of the forest area. Yet, their impacts on the regeneration dynamics of fragmented forests remain poorly investigated. Here we examine the potential of Atta cephalotes nests to function as ecological filters impacting tree recruitment. Growth, survival and biomass partitioning of experimentally planted seedlings (six tree species) were examined at eight spatially independent A. cephalotes colonies in a large Atlantic Forest fragment. Seedling performance and fate (leaf numbers and damage) were monitored up to 27 months across three habitats (nest centre, nest edge and forest understorey). Plants at illuminated nest centres showed twice the gross leaf gain as understorey individuals. Simultaneously, seedlings of all species lost many more leaves at nests than in the forest understorey, causing a negative net leaf gain. Net leaf gain in the shaded understorey ranged from zero (Licania and Thyrsodium species) to substantial growth for Copaifera and Virola, and intermediate levels little above zero for Protium and Pouteria. Also seedling survival differed across habitats and species, being typically low in the centre and at the edge of nests where seedlings were often completely defoliated by the ants. Lastly, seedling survival increased strongly with seed size at nest edges while there was no such correlation in the forest. Our results suggest that Atta nests operate as ecological filters by creating a specific disturbance regime that differs from other disturbances in tropical forests. Apparently, Atta nests favour large‐seeded tree species with resprouting abilities and the potential to profit from a moderate, nest‐mediated increase in light availability.     

Do carbon-based defences reduce foliar damage? Habitat-related effects on tree seedling performance in a temperate rainforest of Chiloé Island, Chile

Carbon-based secondary compounds (CBSCs), such as phenols or tannins, have been considered as one of the most important and general chemical barriers of woody plants against a diverse array of herbivores. Herbivory has been described as a critical factor affecting the growth and survival of newly established tree seedlings or juveniles then, the presence of secondary metabolites as defences against herbivores should be a primary strategy to reduce foliar damage. We examined whether light-induced changes in leaf phenolic chemistry affected insect herbivory on seedlings of two rainforest tree species, Drimys winteri (Winteraceae) and Gevuina avellana (Proteaceae). Seedlings of both species were planted under closed canopy and in a canopy gap within a large remnant forest patch. Half of the seedlings in each habitat were disinfected with a wide-spectrum systemic insecticide and the other half were used as controls. Seedling growth, survival, and foliar damage (estimated by an herbivory index) due to insect herbivores were monitored over a period of 16 months (December 2001–April 2003). The total leaf content of phenols and condensed tannins were assessed in seedlings from both habitats. As expected, access to light induced a greater production of CBSCs in seedlings of both tree species, but these compounds did not seem to play a significant defensive role, as seedlings grown in gaps suffered greater leaf damage than those planted in forest interior. In addition, in both habitats, seedlings without insecticide treatment suffered a greater foliar damage than those with insecticide, especially 16 months after the beginning of the experiment. Canopy openness and herbivory had positive and negative effects, respectively, on seedling growth and survival in both tree species. In conclusion, despite the higher levels of defence in tree-fall gap, the higher densities of herbivore override this and lead to higher damage levels.     

Observations on the Ecology of Weaver Ants (Oecophylla smaragdina Fabricius) in a Thai Mangrove Ecosystem and Their Effect on Herbivory of Rhizophora mucronata Lam.

Ants of the genus Oecophylla are predators of other insects and are able to protect a variety of terrestrial plants against pest insects; however, observations on the ecology of these ants in mangrove forests are lacking. General observations on the ecology of Oecophylla smaragdina were carried out in a Thai mangrove forest to determine if these ants can protect their host plants in less favorable mangrove habitats. Leaf herbivory and the density of O. smaragdina ants were measured on Rhizophora mucronata trees at two sites. The results showed a negative correlation between ant density and herbivory. At both sites, the mean percent damaged leaf area was more than four times higher on trees without ants compared to “ant‐trees.” A significant negative correlation was found between tree mean percent leaf damage and the density of ants on the tree. Furthermore, on trees with ants, there was less herbivory on leaves close to ant nests compared to other leaves on the tree. Most damage was caused by chrysomelid beetles (62%) and sesarmid crabs (25%) and both types of herbivory were significantly reduced on ant‐trees.     

Global patterns of insect herbivory in gap and understorey environments,and their implications for woody plant carbon storage

Insect herbivory is thought to favour carbon allocation to storage in juveniles of shade‐tolerant trees. This argument assumes that insect herbivory in the understorey is sufficiently intense as to select for storage; however, understoreys might be less attractive to insect herbivores than canopy gaps, because of low resource availability and – at temperate latitudes – low temperatures. Although empirical studies show that shade‐tolerant species in tropical forests do allocate more photosynthate to storage than their light‐demanding associates, the same pattern has not been consistently observed in temperate forests. Does this reflect a latitudinal trend in the relative activity of insect herbivory in gap versus understorey environments? To date there has been no global review of the effect of light environment on insect herbivory in forests. We postulated that if temperature is the primary factor limiting insect herbivory, the effect of gaps on rates of insect herbivory should be more evident in temperate than in tropical forests; due to low growing season temperatures in the oceanic temperate forests of the Southern Hemisphere, the effect of gaps on insect herbivory rates should in turn be stronger there than in the more continental temperate climates of the Northern Hemisphere. We examined global patterns of insect herbivory in gaps versus understories through meta‐analysis of 87 conspecific comparisons of leaf damage in contrasting light environments. Overall, insect herbivory in gaps was significantly higher than in the understorey; insect herbivory was 50% higher in gaps than in understoreys of tropical forests but did not differ significantly between gaps and understories in temperate forests of either hemisphere. Results are consistent with the idea that low resource availability – and not temperature – limits insect herbivore activity in forest understoreys, especially in the tropics, and suggest the selective influence of insect herbivory on late‐successional tree species may have been over‐estimated.     

Insects and light interact to mediate vine colonization of fast growing Microberlinia bisulcata tree seedlings in gaps of an African rain forest

Vines thrive in lowland tropical forests, yet the biotic factors underlying their colonization of host tree seedlings and saplings remain surprisingly understudied. Insect herbivores presumably could influence this process, especially where disturbance has opened the canopy (i.e., gaps)—temporary areas of higher primary productivity favoring the recruitment of vines and trees and invertebrates in forests—but their impact on vine colonization has never been experimentally tested. Using data from an insect herbivore exclusion (mesh-netting cages) experiment conducted in an African rain forest (Korup, Cameroon), I logistically modeled the probability of vines colonizing seedlings of three co-dominant species (Microberlinia bisulcata vs. Tetraberlinia bifoliolata and T. korupensis) in paired shaded understory and sunny gap locations (41 blocks across 80 ha, starting n = 664 seedlings) in a 1–2-yr period (2008–2009). Vine colonization occurred almost exclusively in gaps, occurring on 16% of seedlings there. Excluding herbivores in gaps doubled colonization of the light-demanding and faster growing M. bisulcata but had negligible effects on the two shade-tolerant, slower growing and less palatable Tetraberlinia species, which together were twice as susceptible to vines under natural forest gap conditions (controls). When protected from herbivores in gaps, more light to individual seedlings strongly increased vine colonization of M. bisulcata whereas its well-lit control individuals supported significantly fewer vines. These results suggest vines preferably colonize taller seedlings, and because light-demanding tree species grow faster in height with more light, they are more prone to being colonized in gaps; however, insect herbivores can mediate this process by stunting fast growing individuals so that colonization rates becomes more similar between co-occurring slow and fast growing tree species. Further influencing this process might be associational resistance or susceptibility to herbivores linked to host species’ leaf traits conferring shade-tolerant ability as seedlings or saplings. A richer understanding of how vines differentially influence forest regeneration and species composition may come from investigating vine–tree–herbivore interactions across light gradients, ideally via long-term studies and intercontinental comparisons. Abstract in French is available with online material.     

Indirect effect of aquatic insect emergence on a terrestrial insect population through by birds predation

A manipulative field experiment was performed to determine the effect of birds, subsidized by aquatic insect emergence, on the insect herbivores in a riparian deciduous forest. Insectivorous birds were observed more frequently in the riparian forest than in upland forest away from the stream, utilizing both herbivorous insects feeding on the riparian vegetation and aquatic insects emerging from the stream as their prey. Field experiments revealed that the insect herbivore population in the riparian forest was more depressed by bird predation than that in the upland forest. This suggests that allochthonous prey input to the in situ prey population was responsible for a modification in the interaction between birds and herbivorous insects, resulting in a heterogeneous food web structure in the forest.     

Grasshoppers (Orthoptera: Acrididae) select vegetation patches in local‐scale responses to foliar nitrogen but not phosphorus in native grassland

Abstract Key elements such as nitrogen (N) and phosphorus (P) are often limiting relative to the nutritional needs of herbivores that feed on them. While N often limits insect herbivores in natural terrestrial ecosystems, the effect of P is poorly studied in the field, even though compelling hypotheses from the ecological stoichiometry literature predict its importance. We evaluated small‐scale spatial distributions of, and herbivory by, grasshoppers among neighboring plots that vary in foliar‐N and ‐P in tallgrass prairie. Grasshopper densities were 67% greater in N‐fertilized plots but detected no effect to grasshopper densities from P‐fertilizer. Leaf damage to the dominant grass Andropogon gerardii was 32% greater in N‐fertilized plots, but no response to foliar‐P was detected. Herbivore damage to a common forb, goldenrod (Solidago missouriensis), was not strongly linked by fertilizer treatments, although there was increased leaf damage in N‐fertilizer treatments when no P was applied (a significant N × P interaction). Under field conditions at local scales, we conclude that spatially heterogeneous distributions of grasshoppers are primarily affected by foliar‐N in host plants with little evidence that P‐levels contribute to the spatial patterns.     

Combined Effects of Host Plant Quality and Predation on a Tropical Lepidopteran: A Comparison between Treefall Gaps and the Understory in Panama

In tropical forests, light‐gaps created from treefalls are a frequent source of habitat heterogeneity. The increase in productivity, through gap formation, can alter food quality, predation and their impact on insect herbivores. We hypothesized that in gaps, herbivores would be less resource‐limited and more predator limited, whereas in the understory, we predicted the reverse. In this study, we investigate the combined effects of food quality and predation on the lepidopteran larva Zunacetha annulata feeding on its host plant Hybanthus prunifolius in two habitats; sunny treefall gaps and the shaded understory in Panama. In bioassays, Z. annulata feeding on sun leaves ate 22 percent less leaf area, grew 25 percent faster, and had higher pupal weights than larvae feeding on shade leaves. However, shade leaves had higher nitrogen content and specific leaf area. In gaps, predation was 26.4 percent compared to 13.8 percent in the understory. Larvae on understory plants traveled greater distances and spent more time searching and traveling than larvae on gap plants. These differences in behavior are consistent with lower predation risk and lower quality food in the understory. Using data from bioassays and field experiments we calculated 0.22 percent and 1.02 percent survival to adulthood for larvae in gaps and the understory, respectively. In conclusion, although these habitats were in close proximity, we found that larvae in the understory are more resource‐limited and larvae in gaps are more predator limited.     

Community composition and diversity of riparian forests regulate decomposition of leaf litter in stream ecosystems

Riparian forest plantings are a well‐established restoration technique commonly used to stabilize banks and intercept nutrient flow from adjacent agricultural fields. Tree species planted for these efforts may not reflect mature forest communities within the same region. Given contemporary research on links between biodiversity and ecosystem functioning, we conducted a leaf‐litter decomposition study to investigate how mixing of detrital resources that reflect forest community composition would regulate in‐stream leaf litter. Leaf litter bags containing material from a mature forest (Liriodendron tulipifera, Acer rubrum, Quercus rubra, full factorial treatments = 7) and a restored riparian forest (Cornus sericea, Fraxinus pennsylvanica, Platanus occidentalis, full factorial treatments = 7) were deployed in a stream reach that experienced riparian reforestation in 2004. Litter from the restored riparian community had less mass remaining (45.28 ± 2.27%) than that from the mature riparian community (54.95 ± 2.19%) after 5 weeks. In addition, mixed litter treatments in the restored riparian community had less mass remaining (40.54 ± 2.37%) than single‐species treatments (51.80 ± 4.05%), a pattern not observed in the mature forest community. Results highlight the importance of planting mixed‐species assemblages as this structure may regulate processes such as decomposition and food‐web structure, processes often not targeted in the restoration plans.     

Abundance and stratification of foliage arthropods in a lowland rain forest of Cameroon

Abstract.

• 1 Arthropod densities and apparent leaf damage were compared within the canopy ecotone and the shrub layer of a lowland rain forest in Cameroon, using a branch clipping method.
• 2 Most of the individuals collected consisted of ants (average 44%) and various herbivores (31%). Overall arthropod densities amounted to 17 individuals per sample, which, on average, consisted of 0.85 m² of foliage area. Arthropod densities were lower than on temperate foliage.
• 3 Arthropod densities were about 3 times higher in the canopy than within the shrub layer. In particular, ants and herbivores were significantly more abundant in the canopy than within the shrub layer. Usually, layer effects rather than site effects appeared to cause greater variance in arthropod abundance.
• 4 Arthropod species-richness, as estimated by the number of operational taxonomic units sorted, was higher in canopy samples than in samples obtained from the shrub layer. However, apparent leaf damage was higher within the shrub layer (10.9%) than on the canopy (5.2%).
• 5 Possible factors responsible for the high densities of ants and herbivores on the canopy and for the high leaf damage within the shrub layer are discussed.

     

Cutting More from Cut Forests: Edge Effects on Foraging and Herbivory of Leaf-Cutting Ants in Brazil

Edge-mediated changes in species composition are known to result in modified species interactions. Because of the crucial trophic position of herbivores and their far-reaching impact on plant communities, it is important to understand how edge influences herbivory. In the present paper, we investigated whether and how leaf-cutting ant foraging is altered in the forest edge, as this habitat is characterized by an increased proportion of pioneer species. We assessed basic foraging data as well as the herbivory rate ( i.e. , the proportion of the leaf material harvested by a colony in relation to the available leaf area in the foraging area) of Atta cephalotes colonies at the edge versus interior sites of a large remnant of the Atlantic forest in Northeast Brazil. Our results indicated clear edge effects on leaf-cutting ants: equally sized A. cephalotes colonies located at the forest edge removed about twice as much leaf area from their foraging grounds than interior colonies (14.3 vs. 7.8%/col/yr). This greater colony-level impact within the forest edge zone was a consequence of markedly reduced foraging areas (0.9 vs. 2.3 ha/col/yr) and moderately lower leaf area index in this habitat, whereas harvest rates were the same. Our results suggest that forest edges induce increased leaf-cutting ant herbivory, probably via the release of resource limitation. Together with the increase of leaf-cutting ant populations along forest edges, this may amplify environmental changes induced by habitat fragmentation.     

Predispersal predation of an understory rainforest herb Aphelandra aurantiaca (Acanthaceae) in gaps and mature forest

The opening of a canopy gap at Los Tuxtlas rainforest has an impact on populations of the understory herb Aphelandra aurantiaca: the ratio of recruited seedlings per reproductive individual is 1:17 in mature forest vs. gaps. Predation occurring before seed dispersal seems a plausible explanation for this observed difference. In a field experiment, in which insecticide was applied to plants growing in gaps and mature forest, we evaluated the extent to which herbivore damage to flowers, fruits, and seeds reduces the number of seeds available for seedling establishment. Under natural conditions, ∼30% of the flowers and >70% of the capsules of A. aurantiaca showed herbivore damage, but its impact changed depending on the type of forest habitat. Flower and fruit herbivores caused more damage in closed forest than in gaps, and this difference was even bigger under the insecticide treatment. Insecticide effectiveness varied depending on the type of forest patch. The highest herbivore impact on seeds was found in the mature forest without insecticide treatment, where most seeds were destroyed. The percentages of seed damage reported here show that predispersal predation is limiting seedling recruitment, especially in mature forest. Other possible explanations might be differences in insect composition, densities, and behavior between gaps and mature forest.     

Tree diversity drives associational resistance to herbivory at both forest edge and interior

Tree diversity is increasingly acknowledged as an important driver of insect herbivory. However, there is still a debate about the direction of associational effects that can range from associational resistance (i.e., less damage in mixed stands than in monocultures) to the opposite, associational susceptibility. Discrepancies among published studies may be due to the overlooked effect of spatially dependent processes such as tree location within forests. We addressed this issue by measuring crown defoliation and leaf damage made by different guilds of insect herbivores on oaks growing among conspecific versus heterospecific neighbors at forest edges versus interior, in two closed sites in SW France forests. Overall, oaks were significantly less defoliated among heterospecific neighbors (i.e., associational resistance), at both forest edge and interior. At the leaf level, guild diversity and leaf miner herbivory significantly increased with tree diversity regardless of oak location within stands. Other guilds showed no clear response to tree diversity or oak location. We showed that herbivore response to tree diversity varied among insect feeding guilds but not between forest edges and interior, with inconsistent patterns between sites. Importantly, we show that oaks were more defoliated in pure oak plots than in mixed plots at both edge and forest interior and that, on average, defoliation decreased with increasing tree diversity from one to seven species. We conclude that edge conditions could be interacting with tree diversity to regulate insect defoliation, but future investigations are needed to integrate them into the management of temperate forests, notably by better understanding the role of the landscape context.     

The comparative importance of species traits and introduction characteristics in tropical plant invasions

Aim We used alien plant species introduced to a botanic garden to investigate the relative importance of species traits (leaf traits, dispersal syndrome) and introduction characteristics (propagule pressure, residence time and distance to forest) in explaining establishment success in surrounding tropical forest. We also used invasion scores from a weed risk assessment protocol as an independent measure of invasion risk and assessed differences in variables between high‐ and low‐risk species. Location East Usambara mountains, Tanzania. Methods Forest transect surveys identified species establishing in disturbed and intact forest. Leaf traits (specific leaf area and foliar nutrient concentrations) were measured from leaves sampled in high‐light environments. Results A leaf traits spectrum was apparent, but species succeeding or failing to establish in either disturbed or intact forest were not located in different parts of the spectrum. Species with high invasion risk did not differ in their location on the leaf trait spectrum compared with low‐risk species but were more likely to be bird/primate‐dispersed. For 15 species establishing in forest quadrats, median canopy cover of quadrats where seedlings were present was correlated with a species value along the leaf trait spectrum. Species establishing in disturbed forest were planted in twice as many plantations and were marginally more likely to be bird‐ or primate‐dispersed than species failing to become established in disturbed forest. Establishment in intact forest was more likely for species planted closer to forest edges. Main conclusions Leaf and dispersal traits appear less important in the colonization of tropical forest than introduction characteristics. It appears, given sufficient propagule pressure or proximity to forest, alien species are much more likely to establish independently of leaf traits or dispersal syndrome in continental tropical forests.     

Effectiveness of protein-rich pea flour for the control of stored-product beetles

Abstract The association between visiting ants and the extrafloral nectaries (EFN)‐bearing shrub Hibiscus pernambucensis Arruda (Malvaceae) was investigated in two different coastal habitats – a permanently dry sandy forest and a regularly inundated mangrove forest. In both habitats the frequency of plants with ants and the mean number of ants per plant were much higher on H. pernambucensis than on non‐nectariferous neighbouring plants. In the sandy forest the proportion of live termite baits attacked by ants on H. pernambucensis was much higher than on plants lacking EFNs. In the mangrove, however, ants attacked equal numbers of termites on either plant class. Ant attendance to tuna/honey baits revealed that overall ant activity in the sandy forest is higher than in the mangrove area. The vertical distribution (ground vs. foliage) of ant activity also differed between habitats. While in the mangrove foraging ants were more frequent at baits placed on foliage, in the sandy forest ant attendance was higher at ground baits. Plants housing ant colonies were more common in the mangrove than in the sandy forest. Frequent flooding in the mangrove may have resulted in increased numbers of ant nests on vegetation and scattered ant activity across plant foliage, irrespective of possession of EFNs. Thus plants with EFNs in the mangrove may not experience increased ant aggression towards potential herbivores relative to plants lacking EFNs. The study suggests that the vertical distribution of ant activity, as related to different nest site distribution (ground vs. foliage) through a spatial scale, can mediate ant foraging patterns on plant foliage and probably affect the ants’ potential for herbivore deterrence on an EFN‐bearing plant species.     

Foliar nectar production and ant activity on a neotropical tree,Ochroma pyramidale

Summary In second growth forest in lowland Costa Rica, ants forage at the foliar nectaries of juvenile Ochroma pyramidale. The relationship between leaf development, foliar nectar production and ant visitation indicates that nectar secretion and ant maintenance are greatest following rapid leaf expansion. Nectar measurements in the glasshouse corroborate field measurements showing that nectar production on a sapling is continuous through time and correlated with distribution and abundance of ants within a sapling. The presence of two nectary types, leaf vein and petiolar, on the leaves of O. pyramidale results in the continual maintenance of ants on the leaf undersurface. Nectar production of a sapling increases with increasing leaf area resulting in greater number of ants per sapling. Energetic costs of nectar production and ant maintenance appear low, representing about one per cent of the total energy invested in leaves.Spatial and diurnal patterns of ant activity changed very little over the study period. Removal and exclusion of ants from saplings results in the utilization of foliar nectar by trigonid bees. A significant difference in leaf damage between ant-visited and unvisited saplings, coupled with ant behavioral characteristics, is consistent with the hypothesis that ants act as antiherbivore agents on Ochroma.     

Influence of leaf traits on the spatial distribution of insect herbivores associated with an overstorey rainforest tree

Summary The spatial distribution of insect herbivores associated with the Australian rainforest treeArgyrodendron actinophyllum (Sterculiaceae) was investigated by restricted canopy fogging. The foliage of this species was low in nitrogen and water content, and high in fibre content. Herbivore abundance was positively correlated with the amount of young foliage present within the samples and in adjacent samples, and with the nitrogen content of young leaves. In particular, the occurrence of phloem-feeders was correlated with the magnitude of translocation within the samples. The influence of leaf water content upon herbivore distribution was marginal, presumably because this factor is not limiting in rain-forest environments during the wet season, which usually coincides with the season of leaf-flush. Specific leaf weight, leaf size and foliage compactness had little or no apparent effect on herbivore distribution. Since the magnitude of leaf turnover affected both the quantity and the quality, as exemplified by translocation effects, of young foliage available, this factor may be critical to herbivores associated with evergreen rainforest trees which are particularly low in foliar nutrients, such asA. actinophyllum.