Seedling Herbivory Research Articles

Vertebrate herbivores are the main cause of seedling mortality in a logged African rainforest—implications for forest restoration

AbstractHigh seedling mortality can limit tropical forest regeneration after disturbance, such as selective logging. However, causes of seedling mortality are often poorly understood although this information is critical for designing appropriate restoration methods. We studied seedling mortality in a rainforest in Uganda where natural recovery has failed after selective logging carried out 40 years earlier and the sites are dominated by herbaceous vegetation. We established an experiment where seeds of 12 indigenous tree species were sown into different‐sized logging gaps in order to capture factors that cause early seedling mortality. We examined whether seedling mortality was influenced by vegetation cover (VC), vertebrate herbivores (VHs), gap size, or forest compartment and recorded causes of mortality. We found that seedling mortality was in general high but showed high tree species‐specific variation (ranging from 19 to 92% six months after emergence). VHs, particularly rodents, were the most important cause of mortality. Rotting caused mortality particularly in smaller gaps, but elephant trampling, drying, being washed away by rain and falling debris were only minor mortality causes. Dense VC was not a considerable barrier to seedling survival, but by providing shelter for rodents, it indirectly increased seedling herbivory. Our results show that high seedling mortality on logged sites can limit forest regeneration and rodents in particular can be a significant factor causing high seedling mortality.

Low tolerance to simulated herbivory in Hawaiian seedlings despite induced changes in photosynthesis and biomass allocation.

Seedling herbivory is an important factor underlying plant community diversity and structure. While considerable research has characterized seedling defence in terms of resistance, very little is known about seedling tolerance of herbivory. Moreover, few studies have attempted to identify mechanisms of tolerance across a range of plant species. Seedling tolerance of simulated herbivory was tested in a diverse pool of ten Hawaiian plant species, including several lobeliad species (family Campanulaceae), a grass, a herb and common woody trees and shrubs. Tolerance was measured as the relative survival and growth of damaged plants receiving 50% defoliation with simultaneous jasmonic acid application compared with undamaged control plants, assessed 1·5 and 5weeks after damage. Putative mechanisms of tolerance were measured, including photosynthetic parameters, light use efficiency, and biomass allocation reflecting growth priorities, and analysed using species-level regression analyses on tolerance indices. No species fully tolerated 50% defoliation at either harvest date, and simulated herbivory significantly reduced shoot as well as root biomass. Lobeliad species had particularly low tolerance. Species varied considerably in size, biomass allocation parameters and their constitutive (pre-damage) and induced (post-damage) photosynthetic parameters. However, only constitutive levels of non-photochemical quenching were significantly related to tolerance, indicating that species with more efficient light use (and less heat dissipation) are better at tolerating damage than species with high levels of heat dissipation. Native Hawaiian plants expressed low tolerance to a conservative level of simulated herbivory. Root growth decreased in response to damage, but this was not associated with greater tolerance, suggesting this response may be due to allocation constraints following defoliation and not due to adaptive plasticity. Conservation of native island plants threatened by invasive herbivores should prioritize protection for seedlings for improved regeneration and the persistence of native plants in disturbed habitats.

Changes in spatial patterns of sika deer distribution and herbivory of planted seedlings: a comparison before and after deer population control by culling

Sika deer (Cervus nippon) sometimes cause extensive damage to planted tree seedlings. To evaluate the effects of culling on the spatial distribution patterns of sika deer and browse damage to planted seedlings, we compared the data collected before and after experimental culling in a cool, temperate, mixed forest on Kyushu Island, Japan. Experimental culling, conducted in an area of 1 km2, removed five, four and two sika deer in April, June and October 2011, respectively. During the year before culling, the spatial pattern of the number of sika deer caught on camera corresponded to the predicted sika deer density. Sika deer immediately browsed planted seedlings after the initial planting. The cumulative number of browsed seedlings increased over time, especially in winter. The spatial pattern of the cumulative number of sika deer caught on camera corresponded to that of browsed seedlings at the year's end. During the year when culling was conducted, the number of sika deer caught on camera decreased around the center of the study site where the culling was conducted and the number of browsed seedlings decreased. During the year following culling, the cumulative number of browsed seedlings was very similar to that in the year before the culling, while the same low number of sika deer was caught on camera. These results indicate that the effects of deer culling resulted in decreased levels of sika deer appearance and browse damage for more than 1 year and for several months, respectively.

A possible ontogenetic trade-off between defense and tolerance in response to simulated herbivory in seedlings and saplings of Araucaria angustifolia

When facing herbivory, plants can defend themselves and/or tolerate the inflicted damage. Unlike defense, tolerance does not prevent herbivory, but enables plants to compensate for damage. Resource allocation theories assume that plants have a limited pool of resources and that those allocated to one function or structure cannot be used by another. Also, due to their limited photosynthetic area and root biomass, seedlings are expected to invest less in defenses than later plant stages. Seedlings of the conifer species Araucaria angustifolia (Brazilian pine) were shown to be quite tolerant to shoot damage, but their ability to defend themselves from herbivores is not yet known. In the present study we tested whether the species is equally tolerant to shoot damage at the seedling and early sapling stages. We also looked for a possible ontogenetic trade-off between tolerance and defense along early plant ontogeny, and compared the allocation of starch reserves between those two developmental stages in response to the inflicted damage. To simulate herbivory, we severed the shoots from seedlings and saplings (~80 % of shoot mass removed). Shoot replacement by sprouting was observed in both groups, although the number of sprouts produced in saplings was greater than in seedlings. Damage resulted in increased mobilization of seed starch in seedlings. In saplings, underground reserves were apparently deployed in response to simulated herbivory. While seedlings had a greater ability to compensate for tissue loss, saplings were more able to chemically defend themselves, suggesting a possible ontogenetic trade-off between tolerance and defense.

Weed seedling herbivory by field cricket Teleogryllus emma (Orthoptera: Gryllidae) in relation to the depth of seedling emergence

This study aimed to determine whether seedling herbivory by the field cricket, Teleogryllus emma (Ohmachi et Matsuura), promoted the mortality of Italian ryegrass (Lolium multiflorum Lam.), a non‐native grass weed in Japan, whether the impact of seedling herbivory differed depending on the depth of seedling emergence and the influence of cricket density on seedling mortality. First, the seedlings at emergence depths of 0, 1 and 2 cm were exposed to the cricket for 2 days in plastic cups and the amount of seedling mortality by herbivory was calculated. The level of mortality of 60 seedlings that emerged from seeds on the soil surface was 92.4%, significantly higher than that of the seedlings that emerged from seeds at depths of 1 cm (10.8%) and 2 cm (9.6%). Second, the seedlings at the emergence depths of 0 and 1 cm were exposed to the crickets at four different densities (zero, one, two and three individuals per 5700 cm2) for 5 days in plastic containers and the amount of seedling mortality by herbivory was calculated. The level of mortality by herbivory of the seedlings that emerged from 100 seeds on the soil surface was higher (33.3–61.3%) than that of the seedlings that emerged from seeds at a depth of 1 cm (4.7–13.1%) in the containers with one‐to‐three crickets. The level of seedling mortality tended to increase with an increasing cricket density. These results suggest that seedling herbivory by crickets is an important factor in weed mortality, particularly in no‐till fields and in field margins, where the seeds are more prevalent near the soil surface.

Trait-based plant mortality and preference for native versus non-native seedlings by invasive slug and snail herbivores in Hawaii

Non-native herbivores may alter plant communities through their preferential consumption of seedlings of different species. We assessed seedling herbivory by two invasive gastropod species in Hawaii, the giant African snail (Achatina fulica) and the Cuban brown slug (Veronicella cubensis). We hypothesized that six native species would suffer greater gastropod herbivory than four non-native species, and that species with short stature, thin leaves, and lacking physical defenses would suffer the greatest mortality from gastropods. Herbivory was measured during 13-day preference trials using enclosures that each contained four different woody species (two native, two non-native) and were assigned to one of three treatments: giant African snail, Cuban brown slug, or control (no gastropod). Discriminant function analysis was used to predict gastropod-induced seed- ling mortality from a suite of seedling characteristics. Native species did not always experience greater herbivory than non-natives species, and seedling mortality was 0-100 %. Native Pipturus albidus and Clermontia parviflora suffered 100 % mortality from V. cubensis herbivory, and P. albidus, Psychotria hawaiiensis, and Myrsine lessertiana suffered C80 % mortality from A. fulica. Two non-natives (Fraxinus uhdei, Clidemia hirta), and two natives (Metrosideros polymorpha, Diospyros sandwicensis), suffered little damage and no mortality. Non-native Ardisia elliptica suffered 10-30 % gastropod mortality, and non-native Psidium cattleianum mortality was 0-50 %. Leaf thickness best predicted species mortality caused by slugs and snails; some thicker-leaved species suffered most. Invasive snails and slugs threaten some native and non-native seedlings by directly consuming them. Current and future plant community structure in Hawaii may in part reflect the feeding preferences of invasive gastropods.

Similar tree seedling responses to shrubs and to simulated environmental changes at Pyrenean and subarctic treelines

Background: Climate, land-use and disturbance regimes are key drivers of treeline dynamics worldwide, but local and regional spatio-temporal patterns indicate that additional factors play an important role. Some studies suggest that shrub-tree interactions control tree seedling recruitment patterns across the treeline ecotone, but little is known about the generality of this interaction. Aims: We established an experiment in a treeline ecotone in the central Pyrenees to investigate the role of such interactions and other environmental factors on tree seedling growth and survival. It was based on a similar experiment we completed recently in the subarctic Scandes. By comparing local phenomena between both experiments we assessed the generality of the findings across regions with different biogeographic histories and species characteristics. Methods: We followed the survival and growth of transplanted Pinus uncinata seedlings during three growing seasons in a multi-factorial design (forest vs. treeline, +/– shrub removal, +/– temperature increase and +/– nutrient addition). Results: There was better seedling growth at the treeline compared to the forest, and the presence of shrubs prevented seedling winter damage and herbivory. Both temperature and nutrient increase enhanced seedling performance. Conclusions: Although some particular details were exclusive to the Pyrenees or to the Scandes, the similarities indicated general patterns and help to interpret the underlying mechanisms of treeline dynamics in both regions. This convergence of responses points to the potential of the development of a robust mechanistically founded predictive framework for scaling up shrub impacts on treeline dynamics to other regions.

Recruitment limitations in Primula vulgaris in a fragmented landscape

Plant recruitment is limited by dispersal, if seeds cannot arrive at potential recruitment sites, and by establishment, due to a low availability of safe sites for recruitment. Seed-sowing experiments, scarcely applied along gradients of landscape alteration, are very useful to assess these limitations. Habitat loss and fragmentation may foster recruitment limitations by affecting all the processes from seed dispersal to seedling establishment. In this study, we perform a seed-sowing experiment to disentangle the importance of dispersal and establishment limitations in different stages of recruitment of the perennial herb Primula vulgaris in fragmented forests of the Cantabrian Range (Northwestern Spain). We evaluated the influence of ecological gradients resulting from habitat loss and fragmentation (modifications of habitat amount at the landscape and microhabitat scales, changes in the species’ population size, changes in seed predation and seedling herbivory) on seedling emergence, survival and early growth. We found strong evidence of dispersal limitation, as seedling emergence was very low in experimental replicates where no seeds were added. This limitation was independent of landscape alterations, as we found no relation with any of the ecological gradients studied. Establishment limitations at the germination phase were also unrelated to ecological gradients, probably because these limitations are more related to fine-scale environmental gradients. However, further monitoring revealed that seedling survival after summer and winter periods and seedling growth were conditioned by landscape alteration, as we found effects of habitat amount at the landscape and microhabitat scales, of presence of populations of P. vulgaris and of seedling herbivory. These effects were complex and sometimes opposite to what can be expected for adult plants, revealing the presence of different requirements between life stages.

Long-term performance and herbivory of tree seedlings planted into primary and secondary forests of Central Amazonia

Abstract:Plant survival and growth in tropical rain forest are affected by different biotic and abiotic forces. As time elapses and plants grow the relative importance of such forces as regeneration inhibitors and/or facilitators may change according to habitat and species. To detect within- and among-species divergences in performance over time in different habitats we followed, for nearly a decade, the survival, growth and herbivory of seedlings of the native tree species: Chrysophyllum pomiferum, Micropholis venulosa and Pouteria caimito. In Central Amazonia, young seedlings were planted into old-growth and secondary forests dominated by Vismia spp. One year after planting, C. pomiferum ranked first (i.e. fast growth, fewer dead and less herbivory) for both habitats, followed by M. venulosa and P. caimito. Initial trends changed over time. In the long term, M. venulosa ranked first for both habitats, followed by C. pomiferum and P. caimito ranked consistently lowest. Within-species divergences in growth and herbivory were greater in secondary forest. Initial seedling responses cannot always be used to predict species persistence in the long term. Contrary to previous estimations, old-growth-forest species can persist under Vismia spp. stands, at least when planted.

Herbivory and habitat association of tree seedlings in lowland evergreen rainforest on white-sand and terra-firme in the upper Rio Negro

Background: It has been proposed that the interaction between herbivory and soil nutrient availability drives habitat association of tree species in Peruvian Amazonia. Nevertheless, there is no empirical evidence that this interaction holds across other Amazonian regions. Aims: We address this knowledge gap by testing whether the interaction between herbivory and soil nutrient contributes to habitat association of tree species in white-sand and terra-firme forests in the upper Rio Negro, Brazil. Methods: We conducted a reciprocal transplanting field experiment in which we controlled for the presence of herbivores. We tested for differences in tree-seedling growth and herbivory among seven white-sand and seven terra-firme habitat-specialist species. Additionally, we assessed whether tree seedlings differed in their functional traits. Results: We found no empirical evidence that an interaction between herbivory and soil nutrients shapes habitat association in white-sand and terra-firme forests of the upper Rio Negro. Tree seedlings showed higher mortality when growing in their non-typical habitat. Growth and herbivory were similar regardless of the presence or absence of herbivore protection and type of soil. Conclusions: We suggest that the overall differences in soil nutrient status between white-sand and terra-firme forests in the upper Rio Negro are insufficient to trigger an interaction between herbivory and soil nutrient availability.

Phylogenetic ecology applied to enrichment planting of tropical native tree species

Enrichment planting within established plantations or secondary forests is a common strategy to enhance forest recovery, given that later successional forest species tend to have low dispersal and limited recruitment into these sites. It is difficult, however, to predict how species of seedlings will perform when planted under different overstory species. The field of phylogenetic ecology offers tools to help guide the selection of seedlings, drawing on the evolutionary conservatism of important functional traits. We evaluated the survival, growth, foliar disease, and herbivory of various native tropical tree seedlings at different evolutionary distances from monospecific stands of trees beneath which they were planted. We expected that seedlings planted under conspecific overstory trees would have low survival and growth and high percent foliar damage (as predicted by the Janzen–Connell Hypothesis), and that seedling performance would improve steadily with phylogenetic distance between seedling and overstory species. We found that seedlings planted under conspecific canopies had lower survivorship, reduced growth, and greater foliar damage than seedlings planted under canopies of different tree species. An overall increase in seedling performance with greater phylogenetic distance between seedling and overstory species was dominated by the contrast in performance between conspecific pairs and seedlings planted beneath extra-familial overstory species; but lack of available congeneric pairing limited inference about interactions among close relatives. Most pathogenic fungi isolated from enrichment-planted seedlings also caused disease when inoculated on the overstory tree species where the seedlings had been planted; this is consistent with overstory trees being an important reservoir of pathogens that affect seedlings. We conclude that enrichment planting with species more distantly related to those that dominate the canopy should enhance seedling’s performance. Closer analysis at the congeneric level is warranted because of expected strong biotic interactions at close phylogenetic distances.

Within and between plant variation of 4β-hydroxiwithanolide E in cape gooseberry (Physalis peruviana; Solanaceae)

4β-Hydroxywithanolide E is a compound with antifeedant activity against certain species of insect herbivores. This compound has been found in roots, leaves, calyxes, and fruits of Physalis peruviana plants. In this study, we quantified by HPLC the concentration of 4β-hydroxywithanolide E in different organs of P. peruviana plants in different developmental stages. Also, we determined the effect of simulated herbivory on concentration of this compound in different parts of the plant. In seedlings, 4β-Hydroxywithanolide E had a higher concentration in leaves than in roots. Conversely, in mature plants, the concentration was lower in leaves than in roots. In reproductive parts, calyxes had higher concentration than fruits. On the other hand, there was no change in the concentration of 4β-Hydroxywithanolide E in P. peruviana organs after simulated herbivory in either seedlings or mature plants.

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.

Feathermoss seedbeds facilitate black spruce seedling recruitment in the forest–tundra ecotone (Labrador, Canada)

As climate warms, conifers are expected to expand their ranges into alpine tundra where ecological factors such as seedbed availability, and post‐dispersal seed and seedling predation may control local recruitment. Seedbed composition may influence microhabitat, nutrients, physical structure, and predation level and, therefore, affect the success of conifer recruitment, thereby providing the template for future expansion. In the boreal forest, seedbed–seedling competition dominates such that seedbed removal increases black spruce recruitment. In the harsher climate of the Mealy Mountains boreal forest–tundra ecotone (Labrador, Canada) the Stress gradient hypothesis (SGH) predicts that facilitation may dominate seedbed–seedling interactions. This study investigated potential mechanisms of seedbed facilitation (temperature, water, nutrients, physical protection) in three conifer seedbeds (Pleurozium schreberi, Cladonia spp., bare soil) and examined whether seed predation and/or seedling herbivory varied among seedbeds over three years. Seed emergence was low overall (< 10% on all treatments), but highest on Pleurozium (6.3%), followed by bare ground (4.6%) and Cladonia (0.3%). Facilitation was observed between Pleurozium and black spruce as seedling height increase (31%) and survival (55%) were highest; herbivory, seed predation and overwinter mortality were lowest compared to both Cladonia and bare ground seedbeds. Unlike in the closed canopy boreal forest, seedlings recruited poorly on bare soil as seedling height increase and survival were 20.5% and 26%. Temperature and water availability were similar across seedbeds, while nutrient availability was higher on Pleurozium. The physical structure of Pleurozium likely protects first to third‐year seedlings from temperature extremes and predators. As climate warms and seed availability increases, Pleurozium may facilitate black spruce recruitment and treeline expansion.

Spatial distribution and performance of native and invasive Ardisia (Myrsinaceae) species in Puerto Rico: the anatomy of an invasion

Comparisons between native and invasive congeners are potentially useful approaches for identifying characteristics that promote invasiveness. Those traits for which an invasive exhibits superior ecological performance are likely to contribute to its invasiveness. We tested the hypothesis that invasive tree species have better ecological performance in early life cycle stages than native species in forests where they coexist. We studied locally sympatric populations of the invasive Ardisia elliptica and the native A. obovata (Myrsinaceae) in Puerto Rico. We compared spatial distribution, herbivory and growth in seedlings, seed germination in the field and under controlled conditions, and fruit production. We found the distribution of each species was aggregated in the three categories of size (seedlings, juveniles and adults) and the populations partially overlapped. The invasive species was the most abundant species in every category of size. The two species did not differ in percentage of leaf area consumed and seedlings of both species had the same relative growth rate (RGR) in the forest. However, the invasive species had higher germination success in the field, faster mean germination time in the lab and higher fruit production. It appears that the success of A. elliptica is not through escape from pathogens or herbivores, but by a better performance in fruiting and seed germination in the forest.

Herbivory of tropical rain forest tree seedlings correlates with future mortality

Tree seedlings in tropical rain forests are subject to both damage from natural enemies and intense interspecific competition. This leads to a trade-off in investment between defense and growth, and it is likely that tree species specialized to particular habitats tailor this balance to correspond with local resource availability. It has also been suggested that differential herbivore impacts among tree species may drive habitat segregation, favoring species adapted to particular resource conditions. In order to test these predictions, a reciprocal transplant experiment in Sabah, Malaysia, was established with seedlings of five species of Dipterocarpaceae. These were specialized to either alluvial (Hopea nervosa, Parashorea tomentella) or sandstone soils (Shorea multiflora, H. beccariana), or were locally absent (S. fallax). A total of 3000 seedlings were planted in paired gap and understory plots in five sites on alluvial and sandstone soils. Half of all seedlings were fertilized. Seedling growth and mortality were recorded in regular samples over 3.5 years, and rates of insect herbivore damage were estimated from censuses of foliar tissue loss on marked mature leaves and available young leaves. Greater herbivory rates on mature leaves had no measurable effects on seedling growth but were associated with a significantly increased likelihood of mortality during the following year. In contrast, new-leaf herbivory rates correlated with neither growth nor mortality. There were no indications of differential impacts of herbivory among the five species, nor between experimental treatments. Herbivory was not shown to influence segregation of species between soil types, although it may contribute toward differential survival among light habitats. Natural rates of damage were substantially lower than have been shown to influence tree seedling growth and mortality in previous manipulative studies.

Cryptic seedling herbivory by nocturnal introduced generalists impacts survival, performance of native and exotic plants

Although much of the theory on the success of invasive species has been geared at escape from specialist enemies, the impact of introduced generalist invertebrate herbivores on both native and introduced plant species has been underappreciated. The role of nocturnal invertebrate herbivores in structuring plant communities has been examined extensively in Europe, but less so in North America. Many nocturnal generalists (slugs, snails, and earwigs) have been introduced to North America, and 96% of herbivores found during a night census at our California Central Valley site were introduced generalists. We explored the role of these herbivores in the distribution, survivorship, and growth of 12 native and introduced plant species from six families. We predicted that introduced species sharing an evolutionary history with these generalists might be less vulnerable than native plant species. We quantified plant and herbivore abundances within our heterogeneous site and also established herbivore removal experiments in 160 plots spanning the gamut of microhabitats. As 18 collaborators, we checked 2000 seedling sites every day for three weeks to assess nocturnal seedling predation. Laboratory feeding trials allowed us to quantify the palatability of plant species to the two dominant nocturnal herbivores at the site (slugs and earwigs) and allowed us to account for herbivore microhabitat preferences when analyzing attack rates on seedlings. The relationship between local slug abundance and percent cover of five common plant taxa at the field site was significantly negatively associated with the mean palatability of these taxa to slugs in laboratory trials. Moreover, seedling mortality of 12 species in open-field plots was positively correlated with mean palatability of these taxa to both slugs and earwigs in laboratory trials. Counter to expectations, seedlings of native species were neither more vulnerable nor more palatable to nocturnal generalists than those of introduced species. Growth comparison of plants within and outside herbivore exclosures also revealed no differences between native and introduced plant species, despite large impacts of herbivores on growth. Cryptic nocturnal predation on seedlings was common and had large effects on plant establishment at our site. Without intensive monitoring, such predation could easily be misconstrued as poor seedling emergence.

Crecimiento, supervivencia y herbivoría de plántulas de Brosimum alicastrum (Moraceae), una especie del sotobosque neotropical

Growth responses, survival, and herbivory, on seedlings of Brosimum alicastrum were studied in a neotropical Mexican forest. We selected 122 seedlings and divided them into three groups assigned to defoliation treatments: control or 0 (n=21), 50 (n=51) and 90% (n=50). Every 4 months during two years we measured seedling growth (in terms of relative growth rate in biomass, leaf area growth, produced leaves and height growth) and survival. In addition, we evaluated every 12 months pathogen damage and insect herbivory using a 2 mm(-2) grid. Separately, we estimated mammal herbivory in 3-month old seedlings that were selected within a plot of 500 m x 10 m (N=1095). Pathogen damage and insect herbivory were evaluated within the same plot in 113 seedlings. We found that 50% defoliated seedlings showed compensatory responses in all growth parameters. Relative growth rate and height growth also had a compensatory response in seedlings at 90% defoliation. Relative growth rate and leaf area growth gradually decreased with time although height growth seedling showed an opposite pattern. Leaves produced were not affected by time. Estimated seedling survival probability increased with defoliation to a maximum of 97%, decreasing at 24 month to 37%. Mammal herbivory was more frequent and severe than herbivory caused by pathogens and insects. In some cases, mammal herbivory produced total defoliation. Compensatory growth in leaf area growth, produced leaves and height growth seedling suggest a synergic compensatory mechanism expressed in a whole-plant growth biomass (relative growth rate). Compensation and survival results suggest trade-offs at the leaf level, such as leaf area growth and produced leaves versus chemical defenses, respectively.

Beneficial effect of spider presence on seedling recruitment of the tropical rainforest tree Dipteryx oleifera (Fabaceae)

This research presents evidence showing that: (1) lower arthropod herbivory correlates with seedling survival, and (2) spider presence correlates with lower arthropod herbivory, seedling growth, and seedling survival of the tropical rainforest tree species Dipteryx oleifera in eastern Nicaragua. The study was conducted from January 2005 to January 2006 in a 6.37 ha permanent plot established in 2002. Seedling height, spider behavior and presence on seedlings, and percentage of leaf area lost due to arthropod herbivory were measured. Arthropod herbivory was assessed from digital photographs of each seedling within the permanent plot. Seedling fate was followed in order to determine its correlation with spider presence, initial seedling size, and arthropod herbivory. A GLM showed that seedling survival correlated negatively with lower levels of arthropod herbivory (<20%), while seedlings with higher levels of herbivore damage experienced mortalities close to 100%. Results from another GLM suggests that seedling mean height (aprox. 8 cm) would be increased by approximately 1.5 cm for each year that spiders were present on seedlings and would be decreased 0.75 cm in height for each percent unit of arthropod herbivory. We also report a trend toward lower arthropod herbivory in seedlings colonized by spiders with aggressive traits, presumably because more aggressive spiders better defended seedlings against herbivorous arthropods than less aggressive spiders.

Canopy cover mediates interactions between a specialist caterpillar and seedlings of a neotropical tree

1 Light availability may be crucial for understanding dynamics of plant–herbivore interactions in temperate and tropical forest communities. This is because local light availability can influence both tree seedling tolerance and susceptibility to herbivory – yet how they mediate levels of insect herbivory that vary with the density of host population is virtually unknown. Here we tested predictions of three key, non-mutually exclusive hypotheses of plant–herbivore interactions: the Limiting Resource Model (LRM), the Plant Vigour Hypothesis (PVH), and the Janzen-Connell Mechanism (JCM). 2 In an Amazonian forest, we planted Swietenia macrophylla seedlings (c. 5 months old) into natural canopy gaps and the shaded understorey and simulated the damage patterns of the specialist herbivore moth, Steniscadia poliophaea, by clipping seedling leaves. Over the next 8 months, we monitored seedling performance in terms of growth and survivorship and also quantified herbivory to new young leaves on a seasonal basis. 3 In support of the LRM, severe leaf damage (≥ 50%) was lethal for Swietenia macrophylla seedlings in the understorey, but in gaps only reduced seedling growth. In support of the PVH, gap seedlings suffered greater post-simulated herbivory (up to 100% defoliation) by S. poliophaea caterpillars than their understorey counterparts. 4 Adding a novel dimension to the Janzen–Connell hypothesis, we found that early wet season herbivory of seedlings in gaps increased with conspecific adult density within a 125-m radius; whereas in the understorey only those seedlings within 50 m of a Swietenia tree were attacked by caterpillars. 5 Synthesis. These results suggest lepidopterans that need young leaves for food may forage more widely in forests to find seedlings in light-rich canopy gaps. Moths may achieve this successfully by being first attracted to gaps, and then searching within them for suitable hosts. A conceptual model, integrating conspecific adult tree density with light-driven changes in seedling tolerance/vigour and their susceptibility to herbivory and mortality, is presented. Spatial variation in the light available to tree seedlings often affects their tolerance and vigour, which may have important consequences for leaf-chewing insects and the scale of density-dependent herbivory in forests.