Differences in Survival and Growth Among Tropical Rain Forest Pioneer Tree Seedlings in Relation to Canopy Openness and Herbivory

Although differences in canopy openness, herbivory and their interaction may promote species coexistence, how these factors affect pioneer tree species and potentially limit growth, and survival has been poorly studied, particularly in tropical South Asia. We monitored the effect of canopy openness and herbivore damage on seedling survival and growth of 960 individuals of six pioneer tree species: Dillenia triquetra, Macaranga indica, Macaranga peltata, Schumacheria castaneifolia, Trema orientalis, and Wendlandia bicuspidata. Seedlings were placed in four gap‐understory positions—center, outer gap edge, inner forest edge, and understory—in four large, natural gaps within the Sinharaja World Heritage Reserve, Sri Lanka. Canopy openness positively affected survival probability beyond the 550‐d experiment, while herbivory decreased survival and was highest in understory conditions. The relative order of species survival stayed fairly consistent between gap‐understory positions and followed their known shade tolerance rankings. When averaged across all experimental conditions, T. orientalis had the lowest survival probability estimate beyond the 550‐d experiment (0.05), but the greatest capacity for growth where it successfully established, while the species with highest averaged survival probability (0.79), D. triquetra, showed the lowest growth. One species, W. bicuspidata, responded positively to herbivory by re‐sprouting. Coexistence of D. triquetra, T. orientalis, and W. bicuspidata can be explained by a trade‐off among species in survival, growth, and response to herbivory. In addition to variation in canopy light environment, herbivory may be important in determining pioneer species distribution through fine‐scale niche partitioning and should be carefully considered in reforestation efforts.     

Interactive effects of soil fertility and herbivory on Brassica nigra

Soil nutrient availability may affect both the amount of damage that plants receive from herbivores and the ability of plants to recover from herbivory, but these two factors are rarely considered together. In the experiment reported here, I examined how soil fertility influenced both the degree of defoliation and compensation for herbivory for Brassica nigra plants damaged by Pieris rapae caterpillars. Realistic levels of defoliation were obtained by placing caterpillars on potted host plants early in the life cycle and allowing them to feed until just before pupation on the designated plant. Percent defoliation was more than twice as great at low soil fertility compared to high (48.2% and 21.0%, respectively), even though plants grown at high soil fertility lost a greater absolute amount of leaf area (38.2 cm² and 22.1 cm², respectively). At both low and high soil fertility, total seed number and mean mass per seed of damaged plants were equivalent to those of undamaged plants. Thus soil fertility did not influence plant compensation in terms of maternal fitness. However, the pathways used to achieve compensation in seed production were different at low and high soil fertility. At low soil fertility, relative leaf growth rates (area added per inital area per day) of damaged plants were drastically reduced over the second week of caterpillar feeding. Damaged plants recovered the leaf area lost to herbivory in the two weeks following insect removal by increasing leaf relative growth rates above the levels seen for undamaged plants, but the replacement of leaf tissue lost to herbivory came at the expense of stem biomass. At high soil fertility, relative leaf growth rates of damaged plants were similar to those of undamaged plants both over the second week of caterpillar feeding and following caterpillar removal, and stem biomass was not affected by herbivory. These results suggest that higher levels of soil nutrients increased the ability of plants to stay ahead of their herbivores as they were being eaten. Because damaged plants at high soil fertility were able to maintain leaf growth rates to a greater extent than damaged plants at low soil fertility, they did not fall as far behind undamaged plants over the period of insect feeding and did not have as much catching up to do after feeding ended to compensate for herbivory.     

The effect of simulated herbivory on growth and nutrient status of focal and neighbouring early successional woody plant species

Defoliation through herbivory is well known to affect target plants and their associated belowground properties, but the response of plants and their soil environment to defoliation of their neighbours is less well understood. We performed a controlled shade‐house experiment involving three plant species that colonize New Zealand floodplains during primary succession, i.e. a palatable N₂‐fixing shrub (Carmichaelia odorata), a palatable deciduous small tree (Fuchsia excorticata) and a less palatable evergreen tree (Weinmannia racemosa). All species were grown in large pots for 40 months both singly and in two species pairs, and either one or both of the species grown in pairs were clipped to simulate herbivory. Responses of growth and foliar nutrient status to clipping varied strongly among species, with Carmichaelia having the largest response and Fuchsia having the smallest. Carmichaelia also enhanced soil microbial biomass and activity, and foliar N concentrations of Weinmannia. However, this did not translate to a net positive effect; instead Carmichaelia competitively reduced growth and foliar P concentrations of both other species. Most effects of Carmichaelia on the soil microflora, and growth and nutrient status of its neighbours, disappeared when Carmichaelia was clipped. Further, the effect of clipping Carmichaelia had a stronger impact on growth, soil activity and nutrient status of the other two species than did the clipping of those species. These results contradict expectations that N₂‐fixing plants should promote growth of other species in pioneer communities or that defoliation of N₂‐fixers exacerbate positive effects; in our study, defoliation of Carmichaelia merely mitigated the negative effects that it had on other species. They also suggest that interplay of competition and differential herbivory among coexisting plants has important implications for soil microflora and processes, relative nutrient acquisition and stoichiometry of coexisting plant species, and potentially plant community development.     

Growth,morphology and leaf characteristics after simulated herbivory in Chinese subtropical evergreen saplings

Growth, morphology and leaf characteristics were assessed in late spring following simulated autumnal defoliation in second-year saplings of three Chinese subtropical evergreen tree species.Castanopsis fargesii showed strong compensatory growth in terms of plant biomass after removal of both 50 and 75% of leaf biomass and slight compensatory growth after 90% defoliation. DefoliatedC. fargesii saplings had more leaves per unit shoot length than non-defoliated saplings. New leaves on defoliated plants were smaller and had higher per area nitrogen content than new leaves on non-defoliated plants.Pinus massoniana andElaeocarpus japonicus showed strong and no compensatory growth, respectively, after 50% defoliation. The strong compensatory growth inP. massoniana andC. fargesii may partly explain why these species predominate in the early and late successional phases of evergreen broad-leaved forests     

Environmental influences on growth and defence responses of the invasive shrub,Lonicera maackii,to simulated and real herbivory in the juvenile stage

Background and Aims

Tolerance and defence against herbivory are among the many mechanisms attributed to the success of invasive plants in their novel ranges. Because tolerance and defence against herbivory differ with the ontogeny of a plant, the effects of herbivore damage on plant fitness vary with ontogenetic stage and are compounded throughout a plant''s lifetime. Environmental stresses such as light and nutrient limitations can further influence the response of the plant. Much is known about the response of plants in the seedling and reproductive adult stages, but less attention has been given to the pre-reproductive juvenile stage.

Methods

Juvenile plants of the North American invasive Lonicera maackii were exposed to simulated herbivory under high and low light and nitrogen availability and growth, allocation patterns and foliar defensive chemistry were measured. In a second experiment, complete nutrient availability and damage type (generalist caterpillar or simulated) were manipulated.

Key Results

Juvenile plants receiving 50 % defoliation had lower total biomass and a higher root^:^shoot ratio than controls for all treatment combinations except low nitrogen/low light. Low light and defoliation increased root^:^shoot ratio. Light, fertilization and defoliation had little impact on foliar defensive chemistry. In the second experiment, there was a reduction in total biomass when caterpillar damage was applied. The root^:^shoot ratio increased under low soil fertility and was not affected by defoliation. Stem-diameter growth rates and specific leaf area did not vary by damage type or fertilization. Foliar protein increased through time, and more strongly in defoliated plants than in controls, while peroxidase activity and total flavonoids decreased with time. Overall, resource limitations were more influential than damage in the growth of juvenile L. maackii plants.

Conclusions

The findings illustrate that even when resources are limited, the tolerance and defence against herbivory of a woody invasive plant in the juvenile stage may contribute to the establishment and persistence of some species in a variety of habitats.     

Contrasting species responses to continued nitrogen and phosphorus addition in tropical montane forest tree seedlings

Global changes in nutrient deposition rates are likely to have profound effects on plant communities, particularly in the nutrient‐limited systems of the tropics. We studied the effects of increased nutrient availability on the seedlings of six tree species in montane forests of southern Ecuador in situ. After five years of continued N, P, or N+P addition, naturally grown seedlings of each of the two most common species at each elevation (1000, 2000, and 3000 m asl) were harvested for analyses of leaf morphology, nutrient content, herbivory, and tissue biomass allocation. Most species showed increased foliar N and P concentrations after addition of each respective element. Leaf tissue N:P ratios of >20 in the control plants of all species suggest that P is more growth‐limiting in these forests than N. Leaf morphological responses to nutrient addition were species and nutrient specific, with some species (Hedyosmum purparescens, Graffenrieda emarginata) exhibiting increased specific leaf area (SLA), and others (Graffenrieda harlingii) increased leaf area ratios (LAR). Pouteria torta (1000 m) had lower SLA and LAR after P addition. Increased herbivory was only evident in G. emarginata (after N and N+P addition). Only the species from 3000 m asl modified biomass allocation after nutrient addition. In general, N and N+P addition more strongly affected the species studied at the upper elevations, whereas P addition had a similar range of effects on the species at all elevations. We conclude that the responses of the studied tropical montane forest tree seedlings to chronic N and P addition are highly species‐specific and that successful adaptation to increased nutrient availability will depend on species‐specific morphological and physiological plasticity.     

Effect of defoliation on leaf growth,sexual expression and reproductive success of Cnidoscolus aconitifolius (Euphorbiaceae)

Several studies on herbivory have shown that plants may have different responses to foliar damage. Most studies have been done using hermaphroditic species, here we present, using a defoliation experiment, different types of response to foliar damage by the monoecious Cnidosculus acontifolius. We evaluated four treatments: natural herbivory (control), and 25%, 50% and 100% artificial defoliation. The results suggest that this species is capable of showing the following compensatory responses: (a) undercompensation in total production of flowers and in fruit set (damage >25%); (b) full compensation in seed weight (all levels of damage); and (c) overcompensation in leaf growth and in the proportion of female flower production (damage 25%). The results show that C. aconitifolius can exhibit a very fast and plastic response depending on the intensity of damage and the time since defoliation. The increase in foliar growth suggests a long-term tolerance response to hervibory, whereas the change in the proportion of female flowers produced does not appear to have a long-term effect due to the decrease of fruit production in all levels of damage. We suggest that the response in foliar growth and the change in sexual expression by C. aconitifolius, are associated with the high resource availability conditions prevalent in the population studied.     

Leaf damage effects on leaf expansion timing in Mallotus japonicus (Euphorbiaceae)

For many plant species, biotic factors determining the timing of leaf expansion have not been elucidated sufficiently. We investigated the effects of leaf damage on the timing of leaf defoliation and on the timing of leaf expansion in Mallotus japonicus (Euphorbiaceae). The degree of leaf herbivory of M. japonicus in the field was examined in summer, with subsequent investigation of the date of leaf defoliation in autumn and that of leaf expansion the following spring. Effects of artificial leaf damage on the timing of leaf defoliation and of leaf expansion were also examined in a greenhouse. In the field, the degree of leaf herbivory sustained by M. japonicus did not affect the timing of defoliation. However, the timing of leaf expansion the next spring advanced earlier, concomitantly with the increase of leaf damage during the previous year. Fifty per cent artificial leaf damage in M. japonicus saplings in summer had no marked effect on the timing of leaf defoliation in autumn. However, the effects of leaf damage on the timing of leaf expansion were apparent the following spring: the damaged plants expanded new leaves ca. 8 days earlier than control plants did. Plants that showed earlier leaf expansion tended to have a higher shoot/root ratio. Our results demonstrate that the plants advance the timing of their leaf expansion in response to leaf damage sustained during the previous year, suggesting that the shoot/root ratio is a determining factor.     

Lack of compensatory growth under phosphorus deficiency in grazing-adapted grasses from the Serengeti Plains

Summary Two shortgrass species (Sporobolus ioclados and Eustachys paspaloides) and two midgrass species (E. paspaloides and Pennisetum mezianum) from the Serengeti grasslands of Tanzania were grown under conditions of extreme phosphorus (P) deficiency. Production of each of these species is maintained or enhanced by defoliation under adequate nutrient supply (McNaughton et al. 1983). However, under the P-deficient conditions of our experiment, defoliation caused a reduction in biomass of all plant parts of each species. Green leaf biomass was reduced most strongly by defoliation, and crowns were least affected. Yield of biomass and nutrients to grazers (green leaves+clipped material) was enhanced by weekly defoliation in the shortgrass grazing-adapted species, whereas yield to producers (live biomass and nutrients retained by the plant) and yield to decomposers (litter) were strongly reduced by defoliation in all species. Phosphate absorption capacity (V _max) measured on excised roots was enhanced by defoliation in the grazing-adapted Sporobolus, but, due to low affinity (high K _m) of roots of defoliated plants for phosphate, absorption rate was not greatly altered at low solution concentrations. Phosphate absorption capacity was reduced or unaffected by defoliation in other species. We conclude that under conditions of P deficiency, plants are unable to acquire the nutrients necessary to replenish large nutrient losses to grazers. In low-nutrient environments, compensatory growth (stimulation of production by grazing) is not a viable strategy. Therefore, in these environments plants respond evolutionarily to herbivores by developing chemical or morphological defenses.     

Interactive effects of clipping and nutrient availability on the compensatory growth of a grass species

Resource availability is an important factor affecting the capacity of compensatory growth after grazing. We performed a greenhouse experiment with Poa bulbosa, a small perennial grass of the Mediterranean and Central Asian grasslands, to test the importance of nutrient availability for compensatory growth after clipping. We also compared the results with predictions of the limited resource model (LRM). Plants were grown at low and high fertilization levels and subjected to a clipping treatment. Contrary to the LMR, we found that in Poa plants compensatory growth occurred under the high fertilization level, while it did not occur under the low level. The LMR predicts a higher tolerance for grazing in the stressful environment. Our plants showed a significant decrease in their relative growth rates (RGR) after clipping. Although the plants allocated a 32–188% greater fraction of the mass to lamina growth after clipping, this greater allocation to the leaves did not fully compensate for the initial reduction in leaf area ratio (LAR). A sensitivity analysis showed for the clipped plants under the high fertilization treatment, that changes in leaf allocation (f _lam) enabled the plants to compensate for a part of the potential loss caused by defoliation. Probably, the increased biomass allocation comes largely from the bulbs. We conclude that the inconsistency of the LRM with our results originates in the lack of compensatory mechanisms in the model. To better understand how environmental conditions affect tolerance to herbivory, the effects of compensatory growth should be taken into account.     

Interspecific variation and ontogenetic change in antiherbivore defense in myrmecophytic Macaranga species

The present study examined whether or not coexisting congeneric plant species have different defense strategies against herbivores, and the intensity of defense changes ontogenetically. We focused on nine myrmecophytic Macaranga species and estimated the intensity of non-biotic and biotic defense by the degree of leaf damage in ant-free and ant-occupied plants, respectively. Ant colonization of myrmecophytic Macaranga species occurred in the early stage of plant development (5–50 cm-tall seedlings). Following the colonization, damage by leaf eaters was minimized and stable during the ontogenetic development of the host plants due to protection by ants. In ant-free trees, however, herbivore damage was immense in seedlings and decreased as trees grew. Interspecific comparison of leaf damage and herbivore fauna supported that coexisting congeneric plants differ in their types of non-biotic (chemical/structural) defense: without ant protection, Macaranga beccariana, for example, was somewhat resistant to leaf eaters but susceptible to gall-makers, Macaranga trachyphylla was heavily infested by generalist leaf eaters, and Macaranga winkleri was exploited by ant-predatory birds. Despite these variations in chemical/structural defense, ant-colonized plants were generally well defended by ants against all kinds of herbivores. This suggests that the individual host-specific ant mutualists are well adapted to deter the chemically or structurally adapted herbivores. These results imply that in the history of diversification in the Macaranga–ant–herbivore system, a sequence of mutual counter adaptation took place not only between plants and herbivores but also between ants and herbivores.     

Life history variation in Arabidopsis lyrata across its range: effects of climate,population size and herbivory

For plants with wide distributional areas, covering a wide range of ecologically distinct habitats, evolutionary divergence can lead to substantial phenotypic variation across a species’ range. These intraspecific trait differences can be very informative about the nature of the selective environment as they potentially reflect different environmental selection pressures while controlling for other species characteristics. In this study, multiple regression and structural equation models were used to examine the relative importance of environmental, ecological, population size and population density effects for variation in growth, reproduction and leaf morphology among 36 populations of the perennial plant Arabidopsis lyrata ssp. petraea across its northwest European range. Substantial variation in temperature, soil nutrient levels and herbivory was observed across the species’ range. In addition, large differences in flowering percentage and individual seed production were found. Leaf morphology varied considerably, with a substantial amount of variation in specific leaf area and trichome density among populations. Structural equation modeling suggested that this species is sensitive to small population sizes, eutrophication and herbivory. Reproductive output was negatively related to herbivory. In addition population size was negatively associated with soil nutrient concentrations. Leaf morphology was shown to be mainly associated with temperature and herbivory. Lower specific leaf areas and lower trichome densities were related to colder areas and high trichomes densities were related to high levels of herbivory. These model results are consistent with the interpretation that, in addition to changing environmental effects across its range, ecological effects such as herbivory contribute to the large variation in life history and morphology of this species. The results reveal a strong negative effect of herbivory on the reproductive output of this species, not only via direct effects of herbivory on flowers and seeds, but also indirectly via a shift in life history strategy.     

Linking above-ground and below-ground effects in autotrophic microcosms: effects of shading and defoliation on plant and soil properties

Although factors affecting plant growth and plant carbon/nutrient balance – e.g., light availability and defoliation by herbivores – may also propagate changes in below‐ground food webs, few studies have aimed at linking the above‐ground and below‐ground effects. We established a 29‐week laboratory experiment (～one growing season) using autotrophic microcosms to study the effects of light and defoliation on plant growth, plant carbon/nutrient balance, soil inorganic N content, and microbial activity and biomass in soil. Each microcosm contained three substrate layers – mineral soil, humus and plant litter – and one Nothofagus solandri var. cliffortioides seedling. The experiment constituted of the presence or absence of two treatments in a full factorial design: shading (50% decrease in light) and artificial defoliation (approximately 50% decrease in leaf area in the beginning of the growing season). At the end of the experiment a range of above‐ground and below‐ground properties were measured. The shading treatment reduced root and shoot mass, root/shoot ratio and leaf production of the seedlings, while the defoliation treatment significantly decreased leaf mass only. Leaf C and N content were not affected by either treatment. Shading increased NO ₃–N concentration and decreased microbial biomass in humus, while defoliation did not significantly affect inorganic N or microbes in humus. The results show that plant responses to above‐ground treatments have effects which propagate below ground, and that rather straightforward mechanisms may link above‐ground and below‐ground effects. The shading treatment, which reduced overall seedling growth and thus below‐ground N use and C allocation, also led to changes in humus N content and microbial biomass, whereas defoliation, which did not affect overall growth, did not influence these below‐ground properties. The study also shows the carbon/nutrient balance of N. solandri var. cliffortioides seedlings to be highly invariant to both shading and defoliation.     

Resource allocation in an annual herb: Effects of light,mycorrhizal fungi,and defoliation

Concurrent interactions and the availability of resources (e.g., light) affect the cost/benefit balance during mutualistic and antagonistic interactions, as well as plant resource allocation patterns. Mycorrhizal interactions and herbivory concur in most plants, where mycorrhizae can enhance the uptake of soil nutrients by plants as well as consuming a large fraction of the plant's carbon, and defoliation usually reduces light interception and photosynthesis, thereby causing direct losses to the hosts of mycorrhizal fungi. Both types of interactions affect the carbon budget of their host plants and thus we predict that the relative costs of herbivory and mycorrhizal colonization will increase when photosynthesis is reduced, for instance in light limited environments. We conducted a greenhouse experiment using Datura stramonium to investigate the effects of defoliation and mycorrhizal inoculation on the resource allocation patterns in two different light environments. Defoliated plants overcompensated in terms of leaf mass in both light environments, but total seed mass per fruit was negatively affected by defoliation in both light environments. Mycorrhizal inoculation had a positive effect on vegetative growth and the leaf nitrogen content, but defoliation negates the benefit of mycorrhizal interactions in terms of the leaf nitrogen content. In general, D. stramonium compensated for the relative costs of concurrent mycorrhizal interactions and defoliation; plants that lacked both interactions exhibited the same performance as plants with both types of interactions.     

Mycorrhiza and seedling establishment in a subarctic meadow: Effects of fertilization and defoliation

Abstract. Question: How does changing resource availability induced by fertilization and defoliation affect seedling establishment and mycorrhizal symbiosis in a subarctic meadow? Location: 610 m a.s.l., Kilpisjärvi (69°03’N, 20°50’E), Finland. Methods: A short‐term full‐factorial experiment was established, with fertilization and defoliation of natural established vegetation as treatments. Seeds of two perennial herbs Solidago virgaurea and Gnaphalium norvegicum were sown in natural vegetation and their germination and growth followed. At the final harvest we measured the response in terms of arbuscular mycorrhizal (AM) colonization, biomass and nitrogen concentration of the seedlings and the established vegetation. Results: Germination rate was negatively affected by defoliation in the unfertilized plots. The shoot biomass of S. virgaurea seedlings was reduced by the defoliation and fertilization treatments, but not affected by their interaction. In G. norvegicum, the germination rate and the seedling shoot biomass were negatively correlated with moss biomass in the plots. In the established plants the arbuscular colonization rate was low and defoliation and fertilization treatments either increased or did not affect the colonization by AM fungi. In the seedlings, the colonization rate by AM fungi was high, but it was not affected by treatments. Both seedlings and established plants were colonized by dark septate fungi. Conclusions: Reduction of plant biomass by herbivores can have different effects on seedling growth in areas of high and low soil nutrient availability. The weak response of AM colonization to defoliation and fertilization suggests that AM symbiosis is not affected by altering plant resource availability under the conditions employed in this study.     

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.     

A comparison of artificial defoliation techniques using canola (<Emphasis Type="Italic">Brassica napus</Emphasis>)

We conducted two experiments that investigated how the method and location of artificial defoliation influenced growth, reproduction, and allocation in canola, Brassica napus. In one experiment, 0%, 25%, or 50% of leaf area was removed by cutting circular holes at three possible locations: concentrated at either the base of leaves or at their tips, or dispersed throughout leaf blades. Plants fully compensated for such damage; reproduction and allocation were unaffected by either defoliation intensity or wound location. In a second experiment, we again initiated three intensities of defoliation: non-damaged plants served as controls, while others had 25% or 50% of their leaf areas removed. The method of removal in the second experiment consisted of cutting either multiple, similar-sized, circular holes or single, contiguous patches of a leaf blade. At the highest defoliation intensity reproductive output and allocation were significantly less in plants treated with the former method than the latter, even though an equivalent initial amount of leaf area was removed in both treatments. We conclude that simulated herbivory studies must account for not only how much of the plant is damaged, but also the pattern of leaf damage itself, since both factors contribute to a plant’s physiological and ecological responses to grazing.     

Additive effects of genotype,nutrient availability and type of tissue damage on the compensatory response of Salix planifolia ssp. planifolia to simulated herbivory

Plant responses to herbivory include tolerance (i.e. compensatory growth) and defense. Several factors influence the tolerance of a plant following herbivory, including plant genetic identity, site nutrient availability, and previous and/or concurrent herbivory. We studied the effects of these factors on the compensatory response of Salix planifolia ssp. planifolia, a shrub species common in the boreal and subarctic regions of North America. We cloned several genets of S. planifolia and submitted them to simulated root and/or leaf herbivory while varying the nutrient availability. Simulated leaf herbivory was more detrimental to the plant than simulated root herbivory, reducing both above- and below-ground tissue production. Leaf demography was unaffected by either simulated herbivory treatment. There was some compensatory growth following simulated leaf and root herbivory, but only the root compartment responded to increased nutrient availability. Simulated leaf herbivory increased leaf transpiration and reduced stomatal resistance, suggesting increased carbon fixation. The unexpected finding of the experiment was the absence of interactions among factors (genotype, nutrient availability and type of tissue damage) on the compensatory response of S. planifolia. These factors thus have additive effects on the species' compensatory ability.     

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.     

Effects of herbivory on growth and survival of seedlings of a rainforest tree,Alphitonia whitei (Rhamnaceae)

This study examined the effects of insect herbivory on growth and mortality of seedlings of a mid- successional rainforest tree, Alphitonia whitei Braid. Two experiments were conducted in which seedlings were exposed to 0% and 50% natural defoliation by insect herbivores and placed in light gaps in simple notophyll vine forest at Paluma, near Townsville, North Queensland. In the first experiment, insect herbivory significantly increased mortality of 2-month-old seedlings. Smaller seedlings had significantly greater mortality rates than larger seedlings, irrespective of herbivory. A significantly greater proportion of smaller seedlings died from being smothered by fallen leaves and soil as a result of digging by vertebrates than for larger seedlings. In the second experiment, the effects of seedling age were examined by comparing 2-month and 4-month-old seedlings. Mortality rates were significantly influenced by seedling age, with eight times greater survival of older seedlings than of younger seedlings. Although insect herbivory was correlated with a significant decrease in shoot mass and a significant increase in root:shoot ratio, there was no effect of insect herbivory on seedling survival in the second experiment. Thus, mortality rates were greater for seedlings if they were young or small (which indirectly results from insect herbivory), because small, young seedlings were more susceptibile to other mortality factors, such as burial by fallen debris and digging by vertebrates.