Testing for enemy‐mediated density‐dependence in the mortality of seedlings: field experiments with five Neotropical tree species

The coexistence of plant species in species‐rich tropical forests can be promoted by specialised enemies acting in a negatively density‐dependent manner. While survival of tropical tree seedlings is often negatively density‐dependent, the causes have rarely been identified. We tested whether insects and plant pathogens cause density‐dependent seedling recruitment and survival in five forest tree species in Belize, Central America. We manipulated densities of seeds or newly germinated seedlings in small (1 m² or 0.25 m²) plots close to fruiting conspecific trees. Using a factorial design, we excluded enemies from subsets of the plots with fungicides and insecticides. Seed germination (for two species) and early seedling survival (for all species) were monitored at approximately weekly intervals for up to eight weeks, during the period when plants are likely to be most susceptible to natural enemies. In Terminalia amazonia, seed germination was negatively density‐dependent and the proportion of seeds germinating increased when insects were excluded. However, the magnitude of the insecticide effect was independent of density. The only significant density effect for survival of young seedlings was in Acacia polyphylla; counter to expectation, seedling survival was higher at high densities. In a few cases pesticide application had a significant effect on seedling survival, but in only one case (Terminalia amazonia) was a significant pesticide × density interaction detected. Our results caution against generalising from studies conducted on a single species at a single time and place and illustrate the challenges of experimentally testing for enemy‐mediated negative density‐dependence. Experimental outcomes are likely to depend on the spatial scale at which the principal enemies disperse and respond to plant density, and the timescales over which they act. Gathering information on these variables will improve our understanding of the natural histories of tropical forest species and help inform the design of future experiments.     

Impacts of logging on density-dependent predation of dipterocarp seeds in a South East Asian rainforest

Much of the forest remaining in South East Asia has been selectively logged. The processes promoting species coexistence may be the key to the recovery and maintenance of diversity in these forests. One such process is the Janzen-Connell mechanism, where specialized natural enemies such as seed predators maintain diversity by inhibiting regeneration near conspecifics. In Neotropical forests, anthropogenic disturbance can disrupt the Janzen-Connell mechanism, but similar data are unavailable for South East Asia. We investigated the effects of conspecific density (two spatial scales) and distance from fruiting trees on seed and seedling survival of the canopy tree Parashorea malaanonan in unlogged and logged forests in Sabah, Malaysia. The production of mature seeds was higher in unlogged forest, perhaps because high adult densities facilitate pollination or satiate pre-dispersal predators. In both forest types, post-dispersal survival was reduced by small-scale (1 m(2)) conspecific density, but not by proximity to the nearest fruiting tree. Large-scale conspecific density (seeds per fruiting tree) reduced predation, probably by satiating predators. Higher seed production in unlogged forest, in combination with slightly higher survival, meant that recruitment was almost entirely limited to unlogged forest. Thus, while logging might not affect the Janzen-Connell mechanism at this site, it may influence the recruitment of particular species.     

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 test of the escape and colonization hypotheses for zoochorous tree species in a Western Amazonian forest

In order to assess the importance of seed dispersal (escape and colonization hypotheses), I used transplant experiments for seeds and seedlings of 5–11 plant species with fleshy fruits in a lowland tropical forest (Tinigua National Park, Colombia). I controlled seed density, distance to parental tree, and habitat type. I monitored seed removal, seedling survival, and seedling growth during the first year of development for an average of 554 seeds and 169 seedlings for each species. I supplemented the experimental results with measurements of natural recruitment. I found little support for the escape hypothesis during the seed and seedling stages. For six species that showed differences in seed removal associated with distance, five showed highest removal away from, than close to parent trees, suggesting predator satiation. Seedling survival during the first year was not consistently associated with low densities and long distances from parent trees. For the majority of species, seedlings did not survive flooding in low basins, and there was growth advantage for most plant species in canopy gaps. These differences imply advantages for seed dispersal to adequate habitats, as predicted by the colonization hypothesis. In contrast to experiments, strong negative distance-dependent effects were evident when analyzing natural recruitment patterns. The ratio between saplings and seedlings was higher away from parent trees for the species with enough recruitment to be analyzed and this suggests that a negative distance-dependent effect may also occur after seedling establishment. This pattern is suspected for several other species, but an analysis with some of the other most common trees showed a variety of negative, neutral, and positive distance dependent effects. This study emphasizes the importance of long-term studies to asses the role of seed dispersal.     

Experimental evidence for a phylogenetic Janzen-Connell effect in a subtropical forest

Observational evidence increasingly suggests that the Janzen-Connell effect extends beyond the species boundary. However, this has not been confirmed experimentally. Herein, we present both observational and experimental evidence for a phylogenetic Janzen-Connell effect. In a subtropical forest in Guangdong province, China, we observed that co-occurring tree species are less phylogenetically related than expected. The inhibition effects of neighbouring trees on seedling survival decreased with increasing phylogenetic distance between them. In a shade-house experiment, we studied seedling survival of eight species on soil collected close to Castanopsis fissa relative to their survival on soil close to their own adult trees, and found that this relative survival rate increased with phylogenetic distance from C. fissa. This phylogenetic signal disappeared when seedlings were planted in fungicide-treated soil. Our results clearly support negative effects of phylogenetically similar neighbouring trees on seedling survival and suggest that these effects are caused by associated host-specific fungal pathogens.     

Patterns of temporal and enemy niche use by a community of leaf cone moths (Caloptilia) coexisting on maples (Acer) as revealed by metabarcoding

The diversity of herbivorous insects is often considered a function of host plant diversity. However, recent research has uncovered many examples of closely related herbivores using the same host plant(s), suggesting that partitioning of host plants is not the only mechanism generating diversity. Herbivores sharing hosts may utilize different parts of the same plant, but such resource partitioning is often not apparent; hence, the factors that allow closely related herbivores to coexist are still largely undetermined. We examined whether partitioning of phenology or natural enemies may explain the coexistence of leaf cone moths (Caloptilia; Gracillariidae) associated with maples (Acer; Sapindaceae). Larval activity of 10 sympatric Caloptilia species found on nine maple species was monitored every 2–3 weeks for a total of 13 sampling events, and an exhaustive search for internal parasitoid wasps was conducted using high‐throughput sequencing. Blocking primers were used to facilitate the detection of wasp larvae inside moth tissue. We found considerable phenological overlap among Caloptilia species, with two clear peaks in July and September–October. Coexisting Caloptilia species also had largely overlapping parasitoid communities; a total of 13 chalcid and ichneumon wasp species attacked Caloptilia in a nonspecific fashion at an overall parasitism rate of 46.4%. Although coexistence may be facilitated by factors not accounted for in this study, it appears that niche partitioning is not necessary for closely related herbivores to stably coexist on shared hosts. Co‐occurrence without resource partitioning may provide an additional axis along which herbivorous insects attain increased species richness.     

The fine-scale population dynamics of spruce budworm: survival of early instars related to forest condition

Abstract.  1. A lagged, density-dependent relationship between survival of early instars and host-tree condition is revealed during outbreaks of spruce budworm, Choristoneura fumiferana Clem. Persistent damage to hosts leads to deterioration of the stand.
2. Resource limitation affects survival during early-instar dispersal of spruce budworm. Impediments to distinguishing these events with estimates of survival were overcome with a simple model that describes the dispersal and survival processes. The model was used to analyse a recent 15-year population series from Black Sturgeon Lake and two historical datasets from Green River, in Canada.
3. Defoliation-induced damage to the trees resulted in increased losses of spring-emerging larvae that are dispersing in search of feeding sites. Losses were further exacerbated by biotic factors such as maternal fecundity, rates of infection by the pathogen, Nosema fumiferanae , and by weather-related effects on the foraging period.
4. Survival of early-stage budworm larvae in persistent outbreaks declined and the likelihood of other density-related factors such as rate of mortality from natural enemies increased. These results may reconcile outstanding differences in interpretation of the role of the forest resource in spruce budworm population dynamics and point to a common process linking the dynamics of other well-known budworm species.     

Associational resistance mediated by natural enemies

Abstract.  1. Associational resistance theory suggests that the association of herbivore-susceptible plant species with herbivore-resistant plant species can reduce herbivore density on the susceptible plant species. Several casual mechanisms are possible but none has so far invoked natural enemies. Associational resistance mediated by natural enemies was tested for by examining densities of a gall fly, Asphondylia borrichiae (Diptera: Cecidomyiidae), and levels of parasitism on two closely related seaside plants, Borrichia frutescens and Iva frutescens , when alone and when co-occurring.
2. Both Borrichia and Iva grow alone or together on small offshore islands in Florida. Each host plant species has its own associated race of fly, but both races of fly are attacked by the same four species of parasitoids. Borrichia normally has a higher density of galls than Iva , and galls are larger on Borrichia than on Iva .
3. Gall size, gall abundance, parasitism levels, and parasitoid community composition were quantified on both Borrichia and Iva on islands where each species grew alone or together. Some islands were then manipulated by adding Borrichia to islands supporting only Iva , and by adding Iva to islands supporting only Borrichia . Subsequent gall densities and gall parasitism levels on the original native species were then examined.
4. On both natural and experimentally manipulated islands, gall densities on Iva were significantly lowered by the presence of Borrichia . This is because bigger parasitoid species that were common on Borrichia galls, which are bigger, spilled over and attacked the smaller Iva galls. Thus, parasitism rates on Iva were higher on islands where Borrichia co-occurred than on islands where Borrichia were absent. Most parasitoids from Iva were too small to successfully attack the large Borrichia galls and so gall density on Borrichia was unaffected by the presence of Iva .     

Competition and dispersal in the pea aphid: clonal variation and correlations across traits

Abstract.  1. The presence of an across-species trade-off between dispersal ability and competitive ability has been proposed as a mechanism that facilitates coexistence. It is not clear if a similar trade-off exists within species. Such a trade-off would constrain the evolution of either trait and, given appropriate selection pressures, promote local adaptation in these traits.
2. This study found substantial levels of heritable variation in competitive ability of the pea aphid, Acyrthosiphon pisum Harris (Homoptera: Aphididae), measured in terms of relative survival when reared with a single clone of the vetch aphid, Megoura viciae Buckton (Homoptera: Aphididae).
3. Pea aphids can move to new patches by either flying (longer distance dispersal) or walking (local dispersal) from plant to plant. There was considerable clonal variation in dispersal ability, measured in terms of the proportion of winged offspring produced, and ability to survive away from their host plant.
4. Winged individuals showed longer off-plant survival times than wingless forms of the same pea aphid clone.
5. There was no evidence of a relationship between clonal competitive ability and either measure of dispersal ability, although the power of the test is limited by the number of pea aphid clones used in the trial.
6. However, there was a positive correlation between clonal fecundity and the proportion of winged offspring produced. Although speculative, it is suggested that clones that are more likely to either overwhelm their host plant or attract higher numbers of natural enemies as a result of having higher fecundity are more likely to produce winged morphs.     

Does low nutritional quality act as a plant defence? An experimental test of the slow‐growth,high‐mortality hypothesis

Abstract.  1. The slow-growth, high-mortality hypothesis was experimentally tested in this study by investigating the effects of plant quality and natural enemies on leaf-miner growth, performance, and survivorship. Two leaf miners ( Acrocercops albinatella and Brachys tesselatus ) occurring on the turkey oak Quercus laevis were studied using a factorial design that manipulated predation/parasitism pressure and plant nutritional quality.
2. Forty trees were randomly divided into four treatments: (1) control plants (nutrients and natural enemies unaltered); (2) nutrients added, natural enemies unaltered; (3) nutrients unaltered, natural enemies reduced; and (4) nutrients added and natural enemies reduced. Water content, leaf toughness, tannin concentration, and foliar nitrogen were quantified monthly for each plant, and mine growth and survivorship were assessed by tracing mines on a 2–3-day interval and by following the fates of 50 mines per species per treatment combination.
3. Fertilised plants exhibited significantly higher amounts of nitrogen, but no significant differences among treatments were observed for water content, leaf toughness, and tannin concentration. These results only partially support the slow-growth, high-mortality hypothesis, as mines were significantly smaller and developed faster on fertilised plants, but neither fertilisation nor natural enemy exclusion significantly affected mine survivorship or mortality caused by natural enemies.     

Does plant diversity increase top–down control of herbivorous insects in tropical forest?

Higher trophic level interactions are key mediators of ecosystem functioning in tropical forests. A rich body of theory has been developed to predict the effects of plant diversity on communities at higher trophic levels and the mechanisms underlying such effects. The 'enemies hypothesis’ states that predators exert more effective top–down control of herbivorous insects with increasing plant diversity. Support for this hypothesis has been found in temperate forests and agroecosystems, but remains understudied in tropical forests. We compared incidence of attacks of different natural enemies using artificial caterpillars in a tropical forest landscape and investigated the role of plant community structure (i.e. species richness, composition and density), and the role of forest fragmentation (i.e. patch size, edge distance and canopy openness) on predation intensity. Plant community effects were tested with respect to three vegetation strata: trees, saplings and herbs. Observed predation was substantially due to ants. Predation rates increased with plant species richness for trees and herbs. Density of saplings, herb cover and herb species composition were important factors for predation. No significant patterns were found for fragmentation parameters, suggesting that forest fragmentation has not altered predation intensity. We conclude that in tropical forests, top–down control of herbivorous insects in the understory vegetation is affected by a combination of plant diversity, plant species composition and structural features of the plant community.     

Habitat fragmentation changes the adaptive value of seed mass for the establishment of a tropical canopy tree

To cope with the limiting light conditions in the rain forest understory, many tropical tree species have evolved large seeds that provide the emerging seedlings with nutritional reserves. Habitat fragmentation might change the adaptive value of seed size by modifying the biotic and physical conditions of the forest understory. We experimentally assessed the potential of fragmentation to alter how seed mass affects seedling survival, vigor, and attack by natural enemies of the tropical tree Nectandra ambigens. Seeds from different mother trees (families) were individually weighed and sown in experimental sites established in continuous forest and in forest fragments. Seedling survival, vigor, and damage by herbivores and pathogens were recorded periodically. While seedlings derived from larger seeds had higher survival rates in both habitats, seedling survival and vigor were significantly greater in forest fragments, and the seedlings also suffered fewer attacks by natural enemies. We found genetic variance for seed mass among families with a heritability value (h²) of 0.66, and we found evidence for selection on seed size. Average seed size differed between dead and living seedlings in three sites. In one fragment, seed size was selectively neutral in relation to survival. Overall, selection for seed size promoted survival and thus appears to affect the distribution of this trait. The maintenance of genetic variance could be related to the stochastic nature of the formation of light gaps. Our results highlight the importance of evaluating the adaptive value of traits susceptible to environmental changes for conservation purposes.     

Herbivore attack in Casearia nitida influenced by plant ontogenetic variation in foliage quality and plant architecture

Traits influencing plant quality as food and/or shelter for herbivores may change during plant ontogeny, and as a consequence, influence the amount of herbivory that plants receive as they develop. In this study, differences in herbivore density and herbivory were evaluated for two ontogenetic stages of the tropical tree Casearia nitida. To assess plant ontogenetic differences in foliage quality as food for herbivores, nutritional and defensive traits were evaluated in saplings and reproductive trees. Predatory arthropods were quantified and the foraging preferences of a parasitoid wasp of the genus Zacremnops were assessed. In addition, survival rates of lepidopteran herbivores (Geometridae) were evaluated experimentally. Herbivore density was three times higher and herbivory was 66% greater in saplings than in reproductive trees. Accordingly, concentrations of total foliar phenolics were higher in reproductive trees than in saplings, whereas leaf toughness, water and nitrogen concentration did not vary between ontogenetic stages. Survival rates of lepidopteran larvae exposed to natural enemies were equivalent in reproductive trees and saplings. Given the greater herbivore density on saplings, equal survival rates implied a greater foraging effort of predators on reproductive trees. Furthermore, observed foraging of parasitoid wasps was restricted to reproductive trees. I propose that herbivore density, and as a consequence, leaf damage were lower in reproductive trees than in saplings due to both traits influencing food quality, and architectural or unmeasured indirect defensive traits influencing foraging preference of natural enemies of herbivores.     

Is selection of host plants by Plagiodera versicolora based on plant‐related performance?

Plant-related performance may be one of the most important factors in the selection of host plants by insect herbivores. We investigated the importance of plant-related performance in host selection by the willow leaf beetle, Plagiodera versicolora (Laicharting) (Coleoptera: Chrysomelidae), on four willow species: Salix chaenomeloides Kimura, Salix eriocarpa Fr. et Sav., Salix   integra Thunb., and Salix serissaefolia Kimura (Salicaceae). Bagging experiments in the field revealed that the performance of P. versicolora adults and larvae differed significantly among willow species under enemy-free conditions and at constant densities. Egg clutch and larval abundance were positively related to adult abundance. Plagiodera versicolora adults did not discriminate strongly among willow species for feeding and oviposition. Larval performance did not differ among willow species in the presence of natural enemies, suggesting that interspecific differences in host quality were overridden by mortality from natural enemies. Adult and egg clutch abundance of P. versicolora changed seasonally despite the temporal stability of adult and larval performance under enemy-free field conditions. Thus, plant-related performance of P. versicolora adults and larvae may contribute little to population growth and temporal dynamics of host use in P. versicolora . Potential factors that reduce discrimination of P. versicolora among host willow species are discussed.     

Seed dispersal and establishment of endangered plants on Oceanic Islands: the Janzen-Connell model, and the use of ecological analogues

Background

The Janzen-Connell model states that plant-specific natural enemies may have a disproportionately large negative effect on progeny close to maternal trees. The majority of experimental and theoretical studies addressing the Janzen-Connell model have explored how it can explain existing patterns of species diversity in tropical mainland areas. Very few studies have investigated how the model''s predictions apply to isolated oceanic islands, or to the conservation management of endangered plants. Here, we provide the first experimental investigation of the predictions of the Janzen-Connell model on an oceanic island, in a conservation context. In addition, we experimentally evaluate the use of ecological analogue animals to resurrect the functional component of extinct frugivores that could have dispersed seeds away from maternal trees.

Methodology/Principal Findings

In Mauritius, we investigated seed germination and seedling survival patterns of the critically endangered endemic plant Syzygium mamillatum (Myrtaceae) in relation to proximity to maternal trees. We found strong negative effects of proximity to maternal trees on growth and survival of seedlings. We successfully used giant Aldabran tortoises as ecological analogues for extinct Mauritian frugivores. Effects of gut-passage were negative at the seed germination stage, but seedlings from gut-passed seeds grew taller, had more leaves, and suffered less damage from natural enemies than any of the other seedlings.

Conclusions/Significance

We provide the first experimental evidence of a distance-dependent Janzen-Connell effect on an oceanic island. Our results potentially have serious implications for the conservation management of rare plant species on oceanic islands, which harbour a disproportionately large fraction of the world''s endemic and endangered plants. Furthermore, in contrast to recent controversy about the use of non-indigenous extant megafauna for re-wilding projects in North America and elsewhere, we argue that Mauritius and other oceanic islands are ideal study systems in which to empirically explore the use of ecological analogue species in restoration ecology.     

The full path of Janzen-Connell effects: genetic tracking of seeds to adult plant recruitment

The Janzen-Connell (J-C) model (Janzen 1970; Connell 1971) has been a dominant yet controversial paradigm for forest community dynamics for four decades, especially in the tropics. With increasing distance from the parent plant, the density of dispersed seeds decreases and, because of a reduced impact of distance- and density-responsive seed and seedling enemies, propagule survival increases, resulting in peak recruitment at some distance from the parent and little recruitment near adult conspecifics. This spacing generates gaps near adult trees for the recruitment of heterospecifics, enhancing species coexistence and species richness. Field studies, primarily focused on seeds and young seedlings, have repeatedly demonstrated increasing survival with increasing distance from parents or decreasing density of propagules (e.g. Clark & Clark 1984; Gilbert et al. 1994; Swamy & Terborgh 2010). Yet a meta-analysis of distance-dependent propagule survival failed to support a general pattern of survival increasing with distance from adult conspecifics, suggesting that there is no need for further experimental tests of the J-C hypothesis in terms of diversity enhancement-results are species-specific, not general (Hyatt et al. 2003). However, a lack of consistent experimental results is not surprising. The outcome of tests of the hypothesis can vary as a function of many factors that can affect successive recruitment stages differently (Schupp 1992; Hyatt et al. 2003; Swamy & Terborgh 2010). This highlights a critical gap-a full test of the J-C model requires data demonstrating that effects carry over to recruitment of new reproductive adults, yet few studies have gone beyond early stages. There is strong inferential evidence that adult trees can show the imprint of J-C effects (e.g. Nathan et al. 2000; Howe & Miriti 2004), and focal individual modelling has clearly demonstrated that J-C effects can operate from sapling through adult stages in a significant number of species (Peters 2003). It is likely that such results are not unusual, but there have been few attempts to demonstrate J-C spacing at the adult stage. In this issue of Molecular Ecology, Steinitz et al. (2011) studied the Mediterranean pine Pinus halepensis (Aleppo pine) and combined a unique situation with an innovative approach to provide the most elegant demonstration yet that adult recruits are spaced further from parents than expected from the initial seed distribution, clear evidence of a J-C effect carrying over to reproductive adults. A major advancement of this study is that it incorporates estimates of the initial patterns of seed dispersal and parentage analysis of adult-offspring relationships, illustrating the value of combined field and genetic approaches.     

Isolation of Enterobacter cowanii from Eucalyptus showing symptoms of bacterial blight and dieback in Uruguay

Aims:  This study was performed to identify bacterial strains isolated simultaneously with Pantoea species from Eucalyptus trees showing symptoms of bacterial blight and dieback in Uruguay.
Methods and Results:  Several molecular techniques including 16S rRNA and rpoB gene sequencing and DNA–DNA hybridization were used to characterize the Gram-negative, facultatively anaerobic, slime-producing bacterial strains isolated along with Pantoea species from Eucalyptus . Hypersensitivity reactions (HR) and pathogenicity tests were performed on tobacco and Eucalyptus seedlings, respectively. The isolates clustered closely with the type strain of Enterobacter cowanii in both phylogenetic trees constructed. The DNA–DNA similarity between the isolates and the type strain of Ent. cowanii ranged from 88% to 92%. A positive HR was observed on the tobacco seedlings, but no disease symptoms were visible on the inoculated Eucalyptus seedlings.
Conclusions:  Enterobacter cowanii was isolated from trees with symptoms of bacterial blight although strains of this bacterial species do not appear to be the causal agent of the disease.
Significance and Impact of the Study:  This study provides the first report of Ent. cowanii isolated from Eucalyptus . Its presence in Eucalyptus tissue suggests that it is an endophyte in trees showing symptoms of blight.     

Linking intra‐specific trait variation and plant function: seed size mediates performance tradeoffs within species

Substantial intra‐specific trait variation exists within plant communities, and in theory this variation could influence community dynamics. Although recent research has focused on intra‐specific variation in traits themselves, it is the influence of this variation on plant performance that makes intra‐specific trait variation relevant to ecological dynamics within or among species. Understanding the links between trait and performance variation, and the role of traits in mediating relationships among multiple components of performance, is critical for assessing the importance of intra‐specific trait variation for community dynamics. Seed size is thought to affect aspects of plant performance including fecundity, seedling growth, dispersal and tolerance of natural enemies. For two tropical tree species, we assessed how seed size was related to performance variation within each species and determined whether intra‐specific trait variation mediates intra‐specific performance tradeoffs. We used field seed rain collection to characterize size‐dependent outcomes of dispersal, sowed seeds of known size in soil collected near or far from conspecifics to characterize susceptibility to soil pathogens, and monitored growth of seedlings from seeds of known size. We found that intra‐specific seed size variation caused intra‐specific performance variation. The degree of trait‐based performance variation was consistently smaller than the degree of trait variation, and seed size influenced different components of performance for each species. One species exhibited a tradeoff in which small seeds had a fecundity advantage (more seedlings per unit reproductive mass) but produced smaller seedlings, whereas the other species exhibited a tradeoff in which small seeds dispersed to areas of low conspecific density but were less tolerant of density‐responsive natural enemies. Our results indicate that a single trait can influence multiple components of performance and can mediate different tradeoffs in co‐occurring species. Complex and heterogeneous effects of a single trait in multidimensional niche space may favour inter‐specific niche differentiation and coexistence.     

Oviposition and feeding preference of Acrolepiopsis assectella Zell. (Lep., Acrolepiidae)

Abstract:  The leek moth, Acrolepiopsis assectella (Zell.), is a recently discovered exotic species in eastern Ontario and western Quebec. This Allium spp. (Asparagales, Alliaceae) specialist can cause up to 40% crop damage. A no-choice experiment was used to determine the relationship between oviposition behaviour and larval survival of the leek moth as the phylogenetic distance from the preferred host Allium ampeloprasum var. porrum L. increased. Results indicate that oviposition preference and larval survival of the leek moth declined as the phylogenetic distance from the preferred host increased. These results support the conclusion that the leek moth is a specialist feeder on closely related Allium spp. although the strength of this preference may decline as the motivation to oviposit increases. This may indicate that the leek moth is able to use closely related novel hosts as temporary refuges if the preferred host plant is unavailable.     

Phytophagy on phylogenetically isolated trees: why hosts should escape their relatives

Hosts belonging to the same species suffer dramatically different impacts from their natural enemies. This has been explained by host neighbourhood, that is, by surrounding host-species diversity or spatial separation between hosts. However, even spatially neighbouring hosts may be separated by many million years of evolutionary history, potentially reducing the establishment of natural enemies and their impact. We tested whether phylogenetic isolation of oak hosts from neighbouring trees within a forest canopy reduces phytophagy. We found that an increase in phylogenetic isolation by 100 million years corresponded to a 10-fold decline in phytophagy. This was not due to poorer living conditions for phytophages on phylogenetically isolated oaks. Neither species diversity of neighbouring trees nor spatial distance to the closest oak affected phytophagy. We suggest that reduced pressure by natural enemies is a major advantage for individuals within a host species that leave their ancestral niche and grow among distantly related species.