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Adaptive trade-offs underlie the specialisation that permits habitat partitioning in species rich plant communities. We investigated the influence of the trade-offs that determine differences in growth and survival among six species of neotropical pioneer trees in gaps in semideciduous forest in Panama. Seedlings of Miconia argentea, Cecropia insignis, Luehea seemannii, Trema micrantha, Ochroma pyramidale and Croton bilbergianus were planted into artificial small (25 m2), medium (64 m2) and large (225 m2) gaps in secondary forest in the Barro Colorado Nature Monument. Trema and Ochroma suffered ≥50% mortality across all gap sizes, while Cecropia had high mortality only during the dry season and in the small gaps, and Miconia and Croton suffered low to zero mortality across all environments. The highest growth rates in large gaps were attained by Cecropia seedlings and in the smaller gaps by Miconia seedlings, although there were indications that Trema and Ochroma required gaps that were larger than any used in this study. Variation in growth and mortality could not be attributed to differences in foliar herbivore damage. Instead, there was strong evidence of a trade-off between maximum growth in the wet season and the ability to survive seasonal drought, particularly in small gaps. We conclude that variation in allocation in response to multiple limiting resources may be as important as allocation to growth and defence in determining the habitat preferences of neotropical pioneers. [ABSTRACT FROM AUTHOR]

We examined the hypothesis that high concentrations of secondary compounds in leaf litter of some tropical riparian tree species decrease leaf breakdown by inhibiting microbial and insect colonization. We measured leaf breakdown rates, chemical changes, bacterial, fungal, and insect biomass on litterbags of eight species of common riparian trees incubated in a lowland stream in Costa Rica. The eight species spanned a wide range of litter quality due to varying concentrations of nutrients, structural and secondary compounds. Leaf breakdown rates were fast, ranging from 0.198 d−1 ( Trema integerrima) to 0.011 d−1 ( Zygia longifolia). Processing of individual chemical constituents was also rapid: cellulose was processed threefold faster and hemicellulose was processed fourfold faster compared to similar studies in temperate streams. Leaf toughness ( r = −0.86, P = 0.01) and cellulose ( r = −0.78, P = 0.02) were the physicochemical parameters most strongly correlated with breakdown rate. Contrary to our initial hypothesis, secondary compounds were rapidly leached (threefold faster than in temperate studies), with all species losing all secondary compounds within the first week of incubation. Cellulose was more important than secondary compounds in inhibiting breakdown. Levels of fungal and bacterial biomass were strongly correlated with breakdown rate (fungi r = 0.64, P = 0.05; bacteria r = 0.93, P < 0.001) and changes in structural compounds (lignin r = −0.55, P = 0.01). Collector−gatherers were the dominant functional group of insects colonizing litterbags, in contrast to temperate studies where insect shredders dominate. Insect biomass was negatively correlated with breakdown rate ( r = −0.70, P = 0.02), suggesting that insects did not play an important role in breakdown. Despite a wide range of initial concentrations of secondary compounds among the eight species used, we found that secondary compounds were rapidly leached and were less important than structural compounds in determining breakdown rates. [ABSTRACT FROM AUTHOR]

Bischofia javanica Blume, an exotic tree, dominates many forest areas of the Bonin Islands in the western Pacific of Japan. The aim of this study was to test the hypothesis that the success of B. javanica (a mid-successional plant species) is related to its high acclimation capacity to sudden light increase due to canopy gap formation. We compared its ecophysiological response to simulated canopy opening with those of native species of different successional status: Trema orientalis Blume, Schima mertensiana (Sieb, et Zucc.) Koidz, Elaeocarpus photiniaefolius Hook.Et Arn. and Ardisia sieboldii Miquel. In all species, transfer of leaves developed in shade (5.3% of full sun) to full sun resulted in a substantial initial reduction in the dark-adapted quantum yield of photosystem II (Fv/Fm). T. orientalis, a pioneer plant species, showed the least reduction (38%), whereas E. photiniaefolius and A. sieboldii, both late-successional plant species, demonstrated large reductions (about 80%). In all four native species, Fv/Fm in shade leaves gradually recovered following transfer, but B. javanica recovered more fully and rapidly than the other species. Unlike Fv/Fm, the chlorophyll content in all species did not recover following the initial decline. This indicates that the recovery of quantum yield (Fv/Fm) was independent of the reduction in chlorophyll. Among all the species, B. javanica showed the highest (1) increase in maximum photosynthetic rate of shade leaves after transfer, (2) production of newly formed sun leaves, and (3) increase in relative growth rate. Ecophysiological characters of B. javanica in simulated canopy openings indicated rapid photosynthetic acclimation in existing shade leaves by minimizing photoinhibition and a rapid deployment of new sun leaves with high photosynthetic capacity. Because its habitats on these Pacific Islands are prone to typhoon disturbance, the successful invasion of B. javanica may lie in the congruence of its acclimation potential and the frequent gap events. [ABSTRACT FROM AUTHOR]

Measuring biodiversity is difficult. This has led to efforts to seek taxa whose species richness correlates with the species richness of other taxa. Such indicator taxa could then reduce the time and cost of assessing the biodiversity of the more extensive community. The search for species richness correlations has yielded mixed results, however. This may be primarily because of the lack of functional relationships between the taxa studied. Trematode parasites are highly promising bioindicators. Diverse assemblages of larval trematode parasites are easily sampled in intermediate host snails. Through their life cycles these parasites are functionally coupled with the surrounding free-living diversity of vertebrate and invertebrate animals. It has been shown that larval trematodes in snails correlate positively with bird diversity and abundance. Here, we explore whether trematodes also correlate with standard measures of fishes, and large and small benthos, for 32 sites in three wetlands. We found associations between trematodes and benthic communities that were not consistent across wetlands. The associations were, however, consistently positive for large benthic species richness and density. Some of the contrasting associations between trematode and benthos may be explained by negative associations between large and small benthos. We found no associations with fish communities (probably because of the inadequacy of standard “snapshot” sampling methods for highly mobile fishes). The results support further exploration of trematodes as bioindicators of diversity and abundance of animal communities. [ABSTRACT FROM AUTHOR]

Understanding the multiple biotic and abiotic controls of aboveground biomass (AGB) is important for projecting the consequences of global change and to effectively manage carbon storage. Although large-scale studies have identified the major environmental and biological controls of AGB, drivers of local-scale variation are less well known. Additionally, involvement of multiple causal paths and scale dependence in effect sizes potentially confounds comparisons among studies differing in methodology and sampling grain. We tested for scale dependence in evidence supporting selection, complementarity and environmental factors as the main determinants of AGB variation over a 50 ha study extent in subtropical China, modelling this at four sampling grains (0.01, 0.04, 0.25 and 1 ha). At each grain, we used piecewise structural equation models to quantify the direct and indirect effects of environmental (topographic and edaphic properties) and forest attributes (structure, diversity and functional traits) on AGB, while controlling for spatial autocorrelation. Direct scale-invariant effects on AGB were evident for structure and community-mean traits, supporting dominance of selection effects. However, diversity had strong indirect effects on AGB via forest structure, particularly at larger sampling grains (≥ 0.25 ha), while direct effects only emerged at the smallest grain size (0.01 ha). The direct and indirect effects of edaphic and topographic factors were also important for explaining both forest attributes and AGB across all scales. Although selection effects appeared to be more influential on ecosystem function, ignoring indirect causal pathways for diversity via structural attributes risks overlooking the importance of complementarity on ecosystem functioning, particularly as sampling grain increases. [ABSTRACT FROM AUTHOR]

Grasses are recognized as a critical regeneration barrier in tropical pastures, yet the effects of rodents and rodent–grass interactions are not well understood. As selective foragers, rodents could shape tree communities, moderating biodiversity in regenerating tropical landscapes. We utilized a fully crossed two-way factorial design to examine the effect that grasses, rodents, and their interaction had on tree seedling establishment in pasture habitat. We followed two separate tree cohorts for 1 year each within the experimental framework. Multiple cohorts were used to better represent successional tree species variation and responses. Trees species were characterized by a gradient of seed masses and as pioneer or persistent successional type. Both cohort seedlings were altered when rodents were present compared to control treatments. In Cohort 1, rodents adversely affected seedlings of persistent tree species only in the absence of grass. In Cohort 2, seedlings of persistent tree species were decimated by rodents in the absence or presence of grass. In both cohorts, seedlings of persistent species established better in grass treatments, while seedlings of pioneer tree species were strongly suppressed. Tree species seed mass positively correlated with seedling establishment across all treatments except no grass–rodent treatments. Strong suppression of tree seedlings by rodents (Sigmodon toltecus) is a novel result in tropical land recently released from agriculture. One implication is that selective foraging by rodents on large-seeded persistent tree species may be facilitated by the removal of grass. Another implication is that temporary rodent control in pastures may permit higher establishment of deep-forest persistent species. [ABSTRACT FROM AUTHOR]

Species that span large latitudinal gradients face strong differences in voltinism and in winter conditions within their range. Latitudinal gradients in winter survival and especially their underlying mechanisms and association with voltinism patterns are poorly studied. We tested in the damselfly Enallagma cyathigerum whether high-latitude populations were better in dealing with the longer winters compared to central- and low-latitude populations and whether this was associated with changes in voltinism. We thereby evaluated whether higher initial levels and/or lower reductions during winter of energy storage (measured as fat content) and investment in immune function [measured as the activity of phenoloxidase (PO)], and/or stronger compensatory responses in food intake contributed to the higher winter survival in high-latitude populations. To this end, we simulated a long high-latitude winter at 4 °C under manipulated food conditions. Across food levels, winter survival was highest in Swedish larvae, intermediate in Belgian larvae, and lowest in Spanish larvae, indicating latitude-specific thermal adaptation that could be partly linked to differences in voltinism. The semi-voltine Swedish larvae were growing slower before winter and as a result accumulated the highest fat content and PO activity when the winter started compared to the univoltine, faster growing Belgian, and Spanish larvae. Fat content and PO activity declined during the winter, yet equally across latitudes, and were not buffered by compensatory food intake. Our data identified possible underlying physiological mechanisms of winter survival and support the hypothesis that widespread latitude-associated voltinism shifts may be a selective factor contributing to latitudinal shifts in winter survival. [ABSTRACT FROM AUTHOR]

The relative importance of host-plant resources, natural enemies or their interactions in controlling the population of galling insects and their parasitism is poorly known for tropical gallers. In this study, we assessed the impacts of plant quality and density of host trees in regulating the densities of a galler species, the cecidomyiid leaf galler (Cecidomyiini sp. 1EJV) and its parasitoids and inquilines on Neoboutonia macrocalyx trees in Uganda. We manipulated the nutritional quality (or vigour) and the resource concentration with four levels each of fertilization and the group size of host tree. We then recorded the effects of these treatments on the growth rate and total leaf area of host plants, the density of gallers and their mortality by parasitoids and inquilines. Higher levels of fertilization and host density resulted in significantly higher total leaf area than did ambient nutrient levels, and lowest tree densities, respectively. Fertilization also caused significant change in the growth rate of leaf area. Both higher fertilization and host density caused higher density of gallers. Total leaf area was positively associated with galler density, but within galled replicates, the galled leaves were larger than the ungalled leaves. Although highest levels of fertilization and density of host trees caused significant change in the densities of parasitoids, the rate of parasitism did not change. However, tree-density manipulations increased the rate of inquilinism, but on a very low level. Our results demonstrate a trophic cascade in the tropical galler and its parasitoids as a response to bottom-up effects. [ABSTRACT FROM AUTHOR]

Infection-induced changes in a host's thermal physiology can represent (1) a generalized host response to infection, (2) a pathological side-effect of infection, or (3), provided the parasite's development is temperature-dependent, a subtle case of host manipulation. This study investigates parasite-induced changes in the thermal biology of a first intermediate host infected by two castrating trematodes (genera Maritrema and Philophthalmus) using laboratory experiments and field surveys. The heat tolerance and temperatures selected by the snail, Zeacumantus subcarinatus, displayed alterations upon infection that differed between the two trematodes. Upon heating, snails infected by Maritrema sustained activity for longer durations than uninfected snails, followed by a more rapid recovery, and selected higher temperatures in a thermal gradient. These snails were also relatively abundant in high shore localities in the summer only, corresponding with seasonal elevated microhabitat temperatures. By contrast, Philophthalmus-infected snails fell rapidly into a coma upon heating and did not display altered thermal preferences. The respective heat tolerance of each trematode corresponded with the thermal responses induced in the snail: Maritrema survived exposure to 40°C, while Philophthalmus was less heat tolerant. Although both trematodes infect the same tissues, Philophthalmus leads to a reduction in the host's thermal tolerance, a response consistent with a pathological side effect. By contrast, Maritrema induces heat tolerance in the snail and withstood exposure to high heat. As the developmental rate and infectivity of Maritrema increase with temperature up to 25°C, one adaptive explanation for our findings is that Maritrema manipulates the snail's thermal responses to exploit warm microhabitats. [ABSTRACT FROM AUTHOR]

Predators can have important impacts on host-parasite dynamics. For many directly transmitted parasites, predators can reduce transmission by removing the most heavily infected individuals from the population. Less is known about how predators might influence parasite dynamics in systems where the parasite relies on vectors or multiple host species to complete their life cycles. Digenetic trematodes are parasitic flatworms with complex life cycles typically involving three host species. They are common parasites in freshwater systems containing aquatic snails, which serve as obligate first intermediate hosts, and multiple trematode species use amphibians as second intermediate hosts. We experimentally examined the impact of predatory salamanders ( Ambystoma jeffersonianum) and trematode parasites ( Echinostoma trivolvis and Ribeiroia ondatrae) on short-term survival of wood frog tadpoles ( Rana sylvatica) in 150-L outdoor pools. Two trematode species were used in experiments because field surveys indicated the presence of both species at our primary study site. Parasites and predators both significantly reduced tadpole survival in outdoor pools; after 6 days, tadpole survival was reduced from 100% in control pools to a mean of 46% in pools containing just parasites and a mean of 49% in pools containing just predators. In pools containing both infected snails and predators, tadpole survival was further reduced to a mean of 5%, a clear risk-enhancement or synergism. These dramatic results suggest that predators may alter transmission dynamics of trematodes in natural systems, and that a complete understanding of host-parasite interactions requires studying these interactions within the ecological framework of community interactions. [ABSTRACT FROM AUTHOR]

The manner in which parasite intensity and aggregation varies with host age can provide insights into parasite dynamics and help identify potential means of controlling infections in humans and wildlife. A significant challenge is to distinguish among competing mechanistic hypotheses for the relationship between age and parasite intensity or aggregation. Because different mechanisms can generate similar relationships, testing among competing hypotheses can be difficult, particularly in wildlife hosts, and often requires a combination of experimental and model fitting approaches. We used field data, experiments, and model fitting to distinguish among ten plausible drivers of a curvilinear age-intensity relationship and increasing aggregation with host age for echinostome trematode infections of green frogs. We found little support for most of these proposed drivers but did find that the parsimonious explanation for the observed age-intensity relationship was seasonal exposure to echinostomes. The parsimonious explanation for the aggregated distribution of parasites in this host population was heterogeneity in exposure. A predictive model incorporating seasonal exposure indicated that tadpoles hatching early or late in the breeding season should have lower trematode burdens at metamorphosis, particularly with simulated warmer climates. Application of this multi-pronged approach (field surveys, lab experiments, and modeling) to additional parasite-host systems could lead to discovery of general patterns in the drivers of parasite age-intensity and age-distribution relationships. [ABSTRACT FROM AUTHOR]

host's first line of defense in response to the threat of parasitic infection is behavior, yet the efficacy of anti-parasite behaviors in reducing infection are rarely quantified relative to immunological defense mechanisms. Larval amphibians developing in aquatic habitats are at risk of infection from a diverse assemblage of pathogens, some of which cause substantial morbidity and mortality, suggesting that behavioral avoidance and resistance could be significant defensive strategies. To quantify the importance of anti-parasite behaviors in reducing infection, we exposed larval Pacific chorus frogs ( Pseudacris regilla) to pathogenic trematodes ( Ribeiroia and Echinostoma) in one of two experimental conditions: behaviorally active (unmanipulated) or behaviorally impaired (anesthetized). By quantifying both the number of successful and unsuccessful parasites, we show that host behavior reduces infection prevalence and intensity for both parasites. Anesthetized hosts were 20-39% more likely to become infected and, when infected, supported 2.8-fold more parasitic cysts. Echinostoma had a 60% lower infection success relative to the more deadly Ribeiroia and was also more vulnerable to behaviorally mediated reductions in transmission. For Ribeiroia, increases in host mass enhanced infection success, consistent with epidemiological theory, but this relationship was eroded among active hosts. Our results underscore the importance of host behavior in mitigating disease risk and suggest that, in some systems, anti-parasite behaviors can be as or more effective than immune-mediated defenses in reducing infection. Considering the severe pathologies induced by these and other pathogens of amphibians, we emphasize the value of a broader understanding of anti-parasite behaviors and how co-occurring stressors affect them. [ABSTRACT FROM AUTHOR]

We explore processes of seed immigration and seedling recruitment before an experimental rainforest restoration matures enough to affect either. Twenty-four 30 × 30-m plots were fenced in 12 ha of pasture in 2006. Seeds were collected in ninety-six 1-m seed traps; recruits were censused in ~12,000 m in which establishment was allowed. We tested effects of distance from forest, living trees, and stumps of trees cut during site preparation on seed rain in 2007 and effects of these and soil depth on recruits through June 2008. Seed fall and recruitment were not correlated with distance to forest 90-400 m away, nor to living shade trees outside the 160 × 485-m experimental grid. Recruitment differed for animal- and wind-dispersed species in a topographically complex landscape. Recruitment of wind-dispersed species was random with respect to soil depth or distance to recent stumps. Recruitment of animal-dispersed species was multimodal; partial correlations with number of stumps within 30 m of plots were significant with soil depth held constant ( P < 0.025), as were correlations of recruitment with soil depth with number of stumps held constant ( P < 0.01). Animal-dispersed recruits were often not conspecifics of adults that had been cut, indicating a legacy of attraction by fruiting trees of animals bearing seeds from distant sources. Ecological implications are that recruitment in pastures released from grazing reflects a mix of widely scattered wind-dispersed pioneers and, where animal-dispersed trees exist, multi-modal and decidedly non-random recruitment of pioneer and later successional animal-dispersed trees from seed banks. [ABSTRACT FROM AUTHOR]

Characterizing the causes of spatial and temporal variation in parasite-induced mortality under natural conditions is crucial to better understanding the factors driving host population dynamics. Our goal was to quantify this variation in the amphipod Paracalliope novizealandiae, a second intermediate host of the trematode, Maritrema novaezealandensis. If infection and development of trematode metacercariae are benign, we expected mature metacercariae to accumulate within amphipods inhabiting high infestation areas. In field samples, intensity levels of mature metacercariae decreased linearly when amphipods harbored >5 immature metacercariae. This finding is consistent with the hypothesis that the parasite can be detrimental at high intensities of infection. Short-term field experiments showed that host survival also declines with the intensity of new infections and drops below 80% when early stage metacercariae reach 10 amphipod−1. However, parasite effects varied over space and time. High-shore amphipods suffered an increased risk of infection in the summer and a lower likelihood of survival: there was a 10–30% decrease in survivorship for any given infection intensity at high- versus low-shore locations. We also tested for differences in the susceptibility of naive and exposed populations using transplant experiments, and found that naive amphipods acquired greater parasite loads (on average, 4.7 vs. 2.8 metacercariae amphipod−1). Because survival decreases rapidly with infection intensity of both early- and late-stage metacercariae, naive populations would suffer considerably if the parasite were to increase its range. Our results indicate that trematode infections cause high mortality in amphipods during summer months under natural conditions, and emphasize that the effects of parasitism vary at local spatial scales and with exposure history. [ABSTRACT FROM AUTHOR]

Conservation strategies depend on our understanding of the ecosystem and community dynamics. To date, such understanding has focused mostly on predator–prey and competitor interactions. It is increasingly clear, however, that parasite–host interactions may represent a large, and important, component of natural communities. The need to consider multiple factors and their synergistic interactions if we are to elucidate the contribution of anthropogenic factors to loss in biodiversity is exemplified by research into present-day amphibian declines. Only recently has the role of factors such as trematode parasite infections been incorporated into studies of the population and community dynamics of aquatic systems. We argue that this is due, at least in part, to difficulties faced by aquatic ecologists in sifting through the complex systematics that pervade the parasite literature. We note that two trematode species are of dominant importance with regard to North American larval anuran communities, and provide in this review a clear explanation of how to distinguish between the infective stages of these two parasites. We describe the general biology and life history of these parasites, as well as what is known about their effect on larval anurans, and the interactive effects of environmental stressors (typically anthropogenic in nature) and parasites on larval anurans. We hope that this review will convince the reader of the potential importance of these parasites to aquatic communities in general, and to amphibian communities specifically, and will also provide the information necessary for aquatic ecologists to more frequently consider the role of these parasites in their studies of aquatic ecology. [ABSTRACT FROM AUTHOR]

The similarity in species composition between two communities generally decays as a function of increasing distance between them. Parasite communities in vertebrate definitive hosts follow this pattern but the respective relationship in intermediate invertebrate hosts of parasites with complex life cycles is unknown. In intermediate hosts, parasite communities are affected not only by the varying vagility of their definitive hosts (dispersing infective propagules) but also by the necessary coincidence of all their hosts in environmentally suitable localities. As intermediate hosts often hardly move they do not contribute to parasite dispersal. Hence, their parasite assemblages may decrease faster in similarity with increasing distance than those in highly mobile vertebrate definitive hosts. We use published field survey data to investigate distance decay of similarity in trematode communities from three prominent coastal molluscs of the Eastern North-Atlantic: the gastropods Littorina littorea and Hydrobia ulvae, and the bivalve Cerastoderma edule. We found that the similarity of trematode communities in all three hosts decayed with distance, independently of local sampling effort, and whether or not the parasites used the mollusc as first or second intermediate host in their life cycle. In H. ulvae, the halving distance (i.e. the distance that halves the similarity from its initial similarity at 1 km distance) for the trematode species using birds as definitive hosts was approximately two to three times larger than for species using fish. The initial similarities (estimated at 1 km distance) among trematode communities were relatively higher, whereas mean halving distances were lower, compared to published values for parasite communities in vertebrate hosts. We conclude that the vagility of definitive hosts accounts for a high similarity at the local scale, while the strong decay of similarity across regions is a consequence of the low probability that all necessary hosts and suitable environmental conditions coincide on a large scale. [ABSTRACT FROM AUTHOR]

With ecosystems increasingly supporting multiple invasive species, interactions among invaders could magnify or ameliorate the undesired consequences for native communities and ecosystems. We evaluated the individual and combined effects of rusty crayfish ( Orconectes rusticus) and Chinese mystery snails [ Bellamya (= Cipangopaludina) chinensis] on native snail communities ( Physa, Helisoma and Lymnaea sp.) and ecosystem attributes (algal chlorophyll a and nutrient concentrations). Both invaders are widespread in the USA and commonly co-occur within northern temperate lakes, underscoring the importance of understanding their singular and joint effects. An outdoor mesocosm experiment revealed that while the two invaders had only weakly negative effects upon one another, both negatively affected the abundance and biomass of native snails, and their combined presence drove one native species to extinction and reduced a second by >95%. Owing to its larger size and thicker shell, adult Bellamya were protected from crayfish attack relative to native species (especially Physa and Lymnaea), suggesting the co-occurrence of these invaders in nature could have elevated consequences for native communities. The per capita impacts of Orconectes (a snail predator) on native snails were substantially greater than those of Bellamya (a snail competitor). Crayfish predation also had a cascading effect by reducing native snail biomass, leading to increased periphyton growth. Bellamya, in contrast, reduced periphyton biomass, likely causing a reduction in growth by native lymnaeid snails. Bellamya also increased water column N:P ratio, possibly because of a low P excretion rate relative to native snail species. Together, these findings highlight the importance of understanding interactions among invasive species, which can have significant community- and ecosystem-level effects. [ABSTRACT FROM AUTHOR]

Trees forage for light through optimal leaf display. Effective leaf display is determined by metamer traits (i.e., the internode, petiole, and corresponding leaf), and thus these traits strongly co-determine carbon gain and as a result competitive advantage in a light-limited environment. We examined 11 metamer traits of sun and shade trees of 38 coexisting moist forest tree species and determined the relative strengths of intra- and interspecific variation. Species-specific metamer traits were related to two variables that represent important life history variation; the regeneration light requirements and average leaf size of the species. Metamer traits varied strongly across species and, in contrast to our expectation, showed only modest changes in response to light. Intra- and interspecific responses to light were only congruent for a third of the traits evaluated. Four traits, amongst which leaf size, specific leaf area (SLA), and leaf area ratio at the metamer level (LAR) showed even opposite intra- and interspecific responses to light. Strikingly, these are classic traits that are thought to be of paramount importance for plant performance but that have completely different consequences within and across species. Sun trees of a given species had small leaves to reduce the heat load, but light-demanding species had large leaves compared to shade-tolerants, probably to outcompete their neighbors. Shade trees of a given species had a high SLA and LAR to capture more light in a light-limited environment, whereas shade-tolerant species have well-protected leaves with a low SLA compared to light-demanding species, probably to deter herbivores and enhance leaf lifespan. There was a leaf-size-mediated trade-off between biomechanical and hydraulic safety, and the efficiency with which species can space their leaves and forage for light. Unexpectedly, metamer traits were more closely linked to leaf size than to regeneration light requirements, probably because leaf-size-related biomechanical and vascular constraints limit the trait combinations that are physically possible. This suggests that the leaf size spectrum overrules more subtle variation caused by the leaf economics spectrum, and that leaf size represents a more important strategy axis than previously thought. [ABSTRACT FROM AUTHOR]

The up-regulation of secondary metabolic pathways following herbivore attack and the subsequent reduction in herbivore performance have been identified in numerous woody plant species. Eucalypts constitutively express many secondary metabolites in the leaves, including terpenes and formylated phloroglucinol compounds (FPCs). We used clonal ramets from six clones of Eucalyptus grandis and two clones of E. grandis × camaldulensis to determine if methyl jasmonate (MeJA) treatment could induce changes in the foliar concentrations of either of these groups of compounds. We also used bioassays to determine if any changes in the performance of larvae of Paropsis atomaria, a chrysomelid leaf beetle, could be detected in treated ramets versus the untreated controls, thus indicating whether MeJA induced the up-regulation of defences other than terpenes or FPCs. We found no significant effects of MeJA treatment on either the foliar concentrations of terpenes and FPCs or on herbivore performance. We did, however, detect dramatic differences in larval performance between Eucalyptus clones, thereby demonstrating large variations in the levels of constitutive defence. Larval feeding on clones resistant to P. atomaria resulted in high first instar mortality and disruption of normal gregarious feeding behaviour in surviving larvae. Histological examination of larvae feeding on a resistant clone revealed damage to the midgut consistent with the action of a toxin. These findings concur with mounting evidence that most evergreen perennial plants lack foliar-induced defences and suggest that constitutively expressed secondary metabolites other than those commonly examined in studies of interactions between insect herbivores and Eucalyptus may be important in plant defence. [ABSTRACT FROM AUTHOR]

We examined 15 traits in leaves and stems related to leaf C economy and water use for 32 co-existing angiosperms at ridge sites with shallow soil in the Bonin Islands. Across species, stem density was positively correlated to leaf mass per area (LMA), leaf lifespan (LLS), and total phenolics and condensed tannins per unit leaf N (N-based), and negatively correlated to leaf osmotic potential and saturated water content in leaves. LMA and LLS were negatively correlated to photosynthetic parameters, such as area-, mass-, and N-based assimilation rates. Although stem density and leaf osmotic potential were not associated with photosynthetic parameters, they were associated with some parameters of the leaf C economy, such as LMA and LLS. In the principal component (PCA) analysis, the first three axes accounted for 74.4% of total variation. Axis 1, which explained 41.8% of the total variation, was well associated with parameters for leaf C and N economy. Similarly, axis 2, which explained 22.3% of the total variation, was associated with parameters for water use. Axis 3, which explained 10.3% of the total variation, was associated with chemical defense within leaves. Axes 1 and 2 separated functional types relatively well, i.e., creeping trees, ruderal trees, other woody plants, C3 shrubs and forbs, palms, and CAM plants, indicating that plant functional types were characterized by similar attributes of traits related to leaf C and N economy and water use. In addition, when the plot was extended by two unrelated traits, leaf mass-based assimilation rates and stem density, it also separated these functional types. These data indicate that differences in the functional types with contrasting plant strategies can be attributed to functional integration among leaf C economy, hydraulics, and leaf longevity, and that both leaf mass-based assimilation rates and stem density are key factors reflecting the different functions of plant species. [ABSTRACT FROM AUTHOR]

Population density and infection with parasites often are important factors affecting the growth and development of individuals. How these factors co-occur and interact in nature should have important consequences for individual fitness and higher-order phenomena, such as population dynamics of hosts and their interactions with other species. However, few studies have examined the joint effects of density and parasitism on host growth and development. We examined the co-influences of rearing density and parasitism, by the trematode Echinostoma trivolvis, on the growth and development of larval frogs, Rana (= Lithobates) pipiens. We also examined the potential role of parasite-mediated intraspecific competition by observing how unparasitized individuals performed when housed with other unparasitized tadpoles, versus housing with a combination of unparasitized and parasitized hosts. Mean mass and mean developmental stage were reduced under high rearing densities. The presence of parasitized conspecifics had no significant effect, but there was a significant interaction of density and parasitism presence on host mass, due to the fact that parasitized conspecifics grew poorly at high densities. Unparasitized individuals reared with parasitized and unparasitized conspecifics fared no better than unparasitized individuals reared only with one another. This result indicates that infected hosts compete as much as uninfected hosts for resources, even though infected individuals have reduced mass under high-density conditions. Resource acquisition and resource allocation are different processes, and parasitism, if it only affects the latter, might not have a discernible impact on competitive interactions. [ABSTRACT FROM AUTHOR]

A seed size–seed number trade-off exists because smaller seeds are produced in greater number but have a lower probability of establishment. This reduced establishment success of smaller-seeded species may be determined by biophysical constraints imposed by scaling rules. Root and shoot diameter, root growth extension rate ( R GER) and shoot length at death for dark-grown seedlings are predicted to scale with the cube root of seed embryo and endosperm mass ( m). We confirmed this expectation for ten neotropical gap-dependent tree species with an embryo and endosperm dry mass >1 mg. However, for nine smaller seeded species ( m < 1 mg) with photoblastic germination, root and shoot diameters were larger than expected, and consequently, R GER was slower than expected. The maximum shoot thrust of seedlings from seeds with masses ≥1 mg was comparable to the estimated force required to displace overlying litter, supporting the hypothesis that photoblastic behaviour only occurs in seeds with insufficient shoot thrust to displace overlying leaves. Using the model soil water, energy and transpiration to predict soil drying in small and large gaps, we showed that: (1) gaps that receive a significant amount of direct sunlight will dry more quickly than small gaps that do not, (2) compared to the wet-season, soil that is already dry at depth (i.e. the dry-season) will dry faster after rainfall (this drying would most likely kill seedlings from small seeds) and (3) even during the wet-season, dry periods of a few days in large gaps can kill shallow-rooted seedlings. We conclude that the smaller the seed, the more vulnerable its seedling would be to both covering by litter and soil drying because it can only emerge from shallow depths and has a slow R GER. Consequently, we suggest that these allometrically related factors contribute to the reduced establishment success of smaller-seeded species that underpins the seed size–seed number trade-off. [ABSTRACT FROM AUTHOR]

A range of traits, including metabolic costs, biomass allocation and seed reserves, may contribute to interspecific variation in the shade tolerance of tree seedlings. In addition, shade tolerance may be affected by differential responses of species to soil resource availability at low light. We used a custom-built whole-plant gas-exchange chamber to quantify instantaneous whole-plant light compensation point (WPLCP) and to parameterize whole-plant daily C gain models for seedlings of eight temperate deciduous tree species. We examined the relationship of WPLCP to growth, biomass allocation and gas-exchange under high and low light and nutrient availabilities and compared it to WPCLP of naturally recruited saplings. For species showing a response, both increased light and nutrient availability resulted in increased WPLCP. However, species’ responses to resource availability did not correspond closely with shade tolerance as has generally been predicted. Variation in WPLCP within species was best predicted by whole-plant dark respiration rates, leaf-level light compensation point and leaf mass per area. Among species, seed size was a strong negative correlate of WPLCP, explaining 66% of the variation. Species with the lowest WPLCP maintained lower growth rates across treatments but greater biomass in the low-light treatment compared with more light-demanding species. These data suggest that a number of traits, in particular metabolic costs and seed size, contribute to WPLCP. However, gas-exchange-based WPLCP was 1.5–3.5 times lower than corresponding growth-based field estimates of WPLCP, suggesting that other factors such as biotic interactions or ontogenetic shifts in whole-plant light requirements may substantially increase species’ WPLCP under natural conditions. [ABSTRACT FROM AUTHOR]

Spatial heterogeneities in the abundance of free-living organisms as well as in infection levels of their parasites are a common phenomenon, but knowledge on parasitism in invertebrate intermediate hosts in this respect is scarce. We investigated the spatial pattern of four dominant trematode species which utilize a common intertidal bivalve, the cockle Cerastoderma edule, as second intermediate host in their life cycles. Sampling of cockles from the same cohort at 15 sites in the northern Wadden Sea (North Sea) over a distance of 50 km revealed a conspicuous spatial heterogeneity in infection levels in all four species over the total sample as well as among and within sampling sites. Whereas multiple regression analyses indicated the density of first intermediate upstream hosts to be the strongest determinant of infection levels in cockles, the situation within sites was more complex with no single strong predictor variable. However, host size was positively and host density negatively correlated with infection levels and there was an indication of differential susceptibility of cockle hosts. Small-scale differences in physical properties of the habitat in the form of residual water at low tide resulted in increased infection levels of cockles which we experimentally transferred into pools. A complex interplay of these factors may be responsible for within-site heterogeneities. At larger spatial scales, these factors may be overridden by the strong effect of upstream hosts. In contrast to first intermediate trematode hosts, there was no indication for inter-specific interactions. In other terms, the recruitment of trematodes in second intermediate hosts seems to be largely controlled by pre-settlement processes both among and within host populations. [ABSTRACT FROM AUTHOR]

Chemical communication among freshwater organisms is an adaptation to improve their coexistence. Here,we focus on the chemical cues secreted by the freshwater gastropod Lymnaea stagnalis, which are known to stimulate behavioural responses of Trichobilharzia ocellata (Plathelminthes, Digenea, Trematoda) miracidia. Such responses are commonly claimed to influence transmission positively, but in response to chemical cues miracidia randomly change their swimming direction. This kind of response does not necessarily increase transmission, because miracidia may be trapped at the periphery of very large snail odour-clouds, which may prevent them from approaching the snail. On the other hand, the odour clouds may be too small to improve host-localisation. To shed light on these scenarios, the spreading of molecules released around L. stagnalis (active space) was visualised by recording host-finding responses of T. ocellata miracidia when they approached snails. Behavioural responses of miracidia indicated the spreading of compounds forming an attractive active space only around the host-snail L. stagnalis, but not around sympatric non-host-snail species. The active space increased approximately linearly with the time the snail rested at the same spot and within 5 min it reached a volume of more than 30 times that of the snail. We also demonstrated in a large-scale experiment, that the active space of L. stagnalis significantly increases the transmission success of T. ocellata miracidia. Additionally, the microhabitat selection of T. ocellata miracidia was studied, demonstrating that peripheral locations near the water surface were preferred, which are also preferred sites of L. stagnalis. Improved chemoperception and microhabitat selection may have been a consequence of coevolution with snails and benefited miracidia, which became efficient transmissive stages. [ABSTRACT FROM AUTHOR]

We hypothesized that pioneer and late successional species show different morphological and physiological responses in water use after gap formation. The magnitude of the responses was compared between two pioneer species (Macaranga giganteaandTrema orientalis) and four late successional species (Shoreasp.), in an experiment in which saplings were transferred from shade to sun. Although transpiration demand increased following the transfer, root hydraulic conductivity (Lpr) decreased.Lpr was sensitive to brief treatments with HgCl2 (a specific inhibitor of aquaporins). This allowsLpr to be divided into two components: cell-to-cell and apoplastic pathways. TheLpr of cell-to-cell pathway decreased in all species following the transfer, relating to aquaporin depression in roots. Following the transfer, leaf osmotic potentials at full hydration decreased and both leaf mass per area [leaf mass/leaf area (LMA)] and fine-root surface area/leaf surface area (root SA/leaf SA) increased in almost all species, allowing saplings to compensate for the decrease inLpr. Physiologically, pioneer species showed larger decreases inLpr and more effective osmotic adjustment than late successional species, and morphologically, pioneer species showed larger increases in root SA/leaf SA and LMA. Water balance at the whole-plant level should be regulated by coupled responses between the aboveground and the belowground parts. Interspecific differences in responses after gap formation suggest niche differentiation in water use between pioneer and late successional species in accordance with canopy-gap size. [ABSTRACT FROM AUTHOR]

Zoochory is the most common mode of seed dispersal for the majority of plant species in the tropics. Based on the assumption of tight plant-animal interactions several hypotheses have been developed to investigate the origin of life history traits of plant diaspores and their dispersers, such as species-specific co-evolution, the low/high investment model (low investment in single fruits but massive fruiting to attract many different frugivores versus high investment in single fruits and fruit production for extended periods to provide food for few frugivores), and the evolution of syndromes which represent plant adaptations to disperser groups (e.show $132#g. birds, mammals, mixed). To test these hypotheses the dispersal strategies of 34 tree species were determined in the littoral forest of Sainte Luce (SE-Madagascar) with the help of fruit traps and tree watches. The impact of fruit consumers on the seeds was determined based on detailed behavioral observations. Phenological, morphological and biochemical fruit traits from tree species were measured to look for co-variation with different types of dispersal. No indication for species-specific co-evolution could be found nor any support for the low/high investment model. However dispersal syndromes could be distinguished as diaspores dispersed by birds, mammals or both groups (mixed) differ in the size of their fruits and seeds, fruit shape, and seed number, but not in biochemical traits. Five large-seeded tree species seem to depend critically on the largest lemur, Eulemur fulvus collaris, for seed dispersal. However, this does not represent a case of tight species-specific co-evolution. Rather it seems to be the consequence of the extinction of the larger frugivorous birds and lemurs which might also have fed on these large fruits. Nevertheless these interactions are of crucial importance to conserve the integrity of the forest. [ABSTRACT FROM AUTHOR]

The Ituri Forest, Democratic Republic of Congo (formerly Zaire) is an example of a closed canopy forest showing extreme depletion in 13C. δ13C values for plants from the canopy top, from gaps in the canopy, and from the subcanopy average -29.0±1.7‰, -30.4±0.9‰, and -34.0±1.5‰, respectively. The δ13C of forest mammals show these differences, with the subcanopy browsers (okapi, dwarf antelope) having δ13C values for tooth enamel much more negative than subcanopy frugivores who derive their food from the canopy top, and from folivores and omnivores living in gap or clearing areas. Nitrogen isotopes in plants from this ecosystem have an average δ15N value of 5.4±1.8‰ and do not show significant differences at the 95% confidence interval between plants from the canopy top, from gaps in the canopy, and from the subcanopy. The δ18OSMOW values of surface waters in the study area are between -2.0 and -2.7. The δ18OPDB for tooth enamel ranged from -3 to +7‰. [ABSTRACT FROM AUTHOR]

A large number of herbaceous and woody plants from tropical woodland, savanna, and monsoon forest were analysed to determine the impact of environmental factors (nutrient and water availability, fire) and biological factors (microbial associations, systematics) on plant δ15N values. Foliar δ15N values of herbaceous and woody species were not related to growth form or phenology, but a strong relationship existed between mycorrhizal status and plant δ15N. In woodland and savanna, woody species with ectomycorrhizal (ECM) associations and putative N2-fixing species with ECM/arbuscular (AM) associations had lowest foliar δ15N values (1.0–0.6‰), AM species had mostly intermediate δ15N values (average +0.6‰), while non-mycorrhizal Proteaceae had highest δ15N values (+2.9 to +4.1‰). Similar differences in foliar δ15N were observed between AM (average 0.1 and 0.2‰) and non-mycorrhizal (average +0.8 and +0.3‰) herbaceous species in woodland and savanna. Leguminous savanna species had significantly higher leaf N contents (1.8–2.5% N) than non-fixing species (0.9–1.2% N) indicating substantial N acquisition via N2 fixation. Monsoon forest species had similar leaf N contents (average 2.4% N) and positive δ15N values (+0.9 to +2.4‰). Soil nitrification and plant NO3- use was substantially higher in monsoon forest than in woodland or savanna. In the studied communities, higher soil N content and nitrification rates were associated with more positive soil δ15N and plant δ15N. In support of this notion, Ficus, a high NO3- using taxa associated with NO3- rich sites in the savanna, had the highest δ15N values of all AM species in the savanna. δ15N of xylem sap was examined as a tool for studying plant δ15N relations. δ15N of xylem sap varied seasonally and between differently aged Acacia and other savanna species. Plants from annually burnt savanna had significantly higher δ15N values compared to plants from less frequently burnt savanna, suggesting that foliar 15N natural abundance could be used as marker for assessing historic fire regimes. Australian woodland and savanna species had low leaf δ15N and N content compared to species from equivalent African communities indicating that Australian biota are the more N depauperate. The largest differences in leaf δ15N occurred between the dominant ECM Australian and African savanna (miombo) species, which were depleted and enriched in 15N, respectively. While the depleted δ15N of Australian ECM species are similar to those of previous reports on ECM species in natural plant communities, the 15N-enriched δ15N of African ECM species represent an anomaly. [ABSTRACT FROM AUTHOR]

Large frugivores are considered to be important seed dispersers for many tropical plant species. Their roles as seed dispersers are not well known in Southeast Asia, where degraded landscapes typically lack these animals. Interactions between 259 (65 families) vertebrate-dispersed fruits and frugivorous animals (including 7 species of bulbul, 1 species of pigeon, 4 species of hornbill, 2 species of squirrel, 3 species of civet, 2 species of gibbon, 1 species of macaque, 2 species of bear, 2 species of deer, and 1 species of elephant) were studied for 3 years in a tropical seasonal forest in Khao Yai National Park, Thailand. The purpose was to examine the dietary overlaps among the large frugivores and the characteristics of fruits they consumed. Most fruit species are eaten by various kinds of frugivores; no close relationship between a particular fruit and a frugivore was found. The number of frugivore groups that served a given plant species was negatively correlated with seed size. Additionally, the fruit/seed diameters consumed by bulbuls were significantly smaller than consumed by the other nine groups. These trends of fruit characteristics were consistent with those observed elsewhere in Southeast Asia: small fruits and large, soft fruits with many small seeds are consumed by a wide spectrum of frugivores while larger fruits with a single large seed are consumed by relatively few potential dispersers. Importantly, these large, single-seed fruits are not consumed by the small frugivores that thrive in small forest fragments and degraded areas in Southeast Asia. To insure the natural seed dispersal process in the forest, an evaluation of all frugivore groups in the forest is urgently needed in Southeast Asia. [ABSTRACT FROM AUTHOR]

This study presents novel evidence that 15N natural abundance can be used as a robust indicator to detect pollutant nitrogen in natural plant communities. Vegetation from the heavily polluted industrial area of Cubatão in São Paulo State, SE Brazil, was strongly 15N depleted compared to plants at remote sites. Historic herbarium samples from Cubatão were significantly less 15N depleted than extant plants, indicating that 15N depletion of vegetation is associated with present-day nitrogen pollution in Cubatão. The heavy load of nitrogenous atmospheric pollutants in Cubatão provides a nitrogen source for plants, and strongly 15N depleted air NH3 is likely to contribute to plant and soil 15N depletion. Epiphytic plants from Cubatão were extremely 15N depleted (average –10.9‰) contrasting with epiphytes at remote sites (averages –1.0‰ and –3.0‰). Nitrogen isotope composition of vegetation provides a tool to determine input of pollutant nitrogen into plant communities. The strong isotopic change of epiphytes suggests that epiphytes are particularly sensitive biomonitors for atmospheric pollutant nitrogen. [ABSTRACT FROM AUTHOR]

Evidence concerning mechanisms hypothesized to explain species coexistence in hyper-diverse communities is reviewed for tropical forest plants. Three hypotheses receive strong support. Niche differences are evident from non-random spatial distributions along micro-topographic gradients and from a survivorship-growth tradeoff during regeneration. Host-specific pests reduce recruitment near reproductive adults (the Janzen-Connell effect), and, negative density dependence occurs over larger spatial scales among the more abundant species and may regulate their populations. A fourth hypothesis, that suppressed understory plants rarely come into competition with one another, has not been considered before and has profound implications for species coexistence. These hypotheses are mutually compatible. Infrequent competition among suppressed understory plants, niche differences, and Janzen-Connell effects may facilitate the coexistence of the many rare plant species found in tropical forests while negative density dependence regulates the few most successful and abundant species. [ABSTRACT FROM AUTHOR]

Spatial variation in parasitism is commonly observed in intermediate host populations. However, the factors that determine the causes of this variation remain unclear. Increasing evidence has suggested that spatial heterogeneity in parasitism among intermediate hosts may result from variation in recruitment processes initiated by definitive hosts. I studied the perching and habitat use patterns of wading birds, the definitive hosts in this system, and its consequences for the recruitment of parasites in snail intermediate hosts. Populations of the mangrove snail, Cerithidea scalariformis, collected from mangrove swamps on the east coast of central Florida are parasitized by a diverse community of trematode parasites. These parasites are transmitted from wading birds, which frequently perch on dead mangrove trees. I tested the hypothesis that mangrove perches act as transmission foci for trematode infections of C. scalariformis and that the spatial variation of parasitism frequently observed in this system is likely to emanate from the distribution of wading birds. On this fine spatial scale, definitive host behaviors, responding to a habitat variable, influenced the distribution, abundance and species composition of parasite recruitment to snails. This causal chain of events is supported by regressions between perch density, bird abundance, bird dropping density and ultimately parasite prevalence in snails. Variation between prevalence of parasites in free-ranging snails versus caged snails shows that while avian definitive hosts initiate spatial patterns of parasitism in snails through their perching behaviors, these patterns may be modified by the movement of snail hosts. Snail movement could disperse their associated parasite populations within the marsh, which may potentially homogenize or further increase parasite patchiness initiated by definitive hosts. [ABSTRACT FROM AUTHOR]

During deposit feeding on benthic micro-algae, the siphon of buried tellinid bivalves like Macoma balthica is vulnerable to nipping by plaice Pleuronectes platessa and other flatfish. We experimentally tested the hypothesis that siphon nippers facilitate predation on the entire bivalve (by shorebirds, for example) by inducing a decrease in burying depth. Three experiments in May and in September, during which likely siphon nippers (juvenile plaice P. platessa) were allowed to feed on M. balthica siphons, demonstrated that the bivalves indeed lost siphon mass and came closer to the surface. However, the strength of the burying response and the speed of recovery of siphons after nipping varied greatly between experiments, partly as a consequence of body condition-related differences in initial burying depth. Our experimental study confirms that siphon nippers can enhance the availability of bivalve prey to probing predators, and will thus facilitate avian predation on intertidal flats. [ABSTRACT FROM AUTHOR]

The interspecific variability of sunlit leaf carbon isotope composition (δ13C), an indicator of leaf intrinsic water-use efficiency (WUE, CO2 assimilation rate/leaf conductance for water vapour), was investigated in canopy trees of three lowland rainforest stands in French Guiana, differing in floristic composition and in soil drainage characteristics, but subjected to similar climatic conditions. We sampled leaves with a rifle from 406 trees in total, representing 102 species. Eighteen species were common to the three stands. Mean species δ13C varied over a 6.0‰ range within each stand, corresponding to WUE varying over about a threefold range. Species occurring in at least two stands displayed remarkably stable δ13C values, suggesting a close genetic control of species δ13C. Marked differences in species δ13C values were found with respect to: (1) the leaf phenology pattern (average δ13C=–29.7‰ and –31.0‰ in deciduous-leaved and evergreen-leaved species, respectively), and (2) different types of shade tolerance defined by features reflecting the plasticity of growth dynamics with respect to contrasting light conditions. Heliophilic species exhibited more negative δ13C values (average δ13C=–30.5‰) (i.e. lower WUE) than hemitolerant species (–29.3‰). However, tolerant species (–31.4‰) displayed even more negative δ13C values than heliophilic ones. We could not provide a straightforward ecophysiological interpretation of this result. The negative relationship found between species δ13C and midday leaf water potential (Ψwm) suggests that low δ13C is associated with high whole tree leaf specific hydraulic conductance. Canopy carbon isotope discrimination (ΔA) calculated from the basal area-weighed integral of the species δ13C values was similar in the three stands (average ΔA=23.1‰), despite differences in stand species composition and soil drainage type, reflecting the similar proportions of the three different shade-tolerance types among stands. [ABSTRACT FROM AUTHOR]

The effects of biological invasions are most evident in isolated oceanic islands such as the Hawaiian Archipelago, where invasive plant species are rapidly changing the composition and function of plant communities. In this study, we compared the specific leaf area (SLA), leaf tissue construction cost (CC), leaf nutrient concentration, and net CO2 assimilation ( A) of 83 populations of 34 native and 30 invasive species spanning elevation and substrate age gradients on Mauna Loa volcano in the island of Hawaii. In this complex environmental matrix, where annual precipitation is higher than 1500 mm, we predicted that invasive species, as a group, will have leaf traits, such as higher SLA and A and lower leaf CC, which may result in more efficient capture of limiting resources (use more resources at a lower carbon cost) than native species. Overall, invasive species had higher SLA and A, and lower CC than native species, consistent with our prediction. SLA and foliar N and P were 22.5%, 30.5%, and 37.5% higher, respectively, in invasive species compared to native ones. Light-saturated photosynthesis was higher for invasive species (9.59 μmol m−2 s−1) than for native species (7.31 μmol m−2 s−1), and the difference was larger when A was expressed on a mass basis. Leaf construction costs, on the other hand, were lower for the invasive species (1.33 equivalents of glucose g−1) than for native species (1.37). This difference was larger when CC was expressed on an area basis. The trends in the above traits were maintained when groups of ecologically equivalent native and invasive species (i.e., sharing similar life history traits and growing in the same habitat) were compared. Foliar N and P were significantly higher in invasive species across all growth forms. Higher N may partially explain the higher A of invasive species. Despite relatively high N, the photosynthetic nitrogen use efficiency of invasive species was 15% higher than that of native species. These results suggest that invasive species may not only use resources more efficiently than native species, but may potentially demonstrate higher growth rates, consistent with their rapid spread in isolated oceanic islands. [ABSTRACT FROM AUTHOR]

Parasitism may be an important factor determining the geographic distribution of closely related species. A habitat-specific risk of parasitism may lead to exclusion of susceptible host types from parasite-rich environments, and promote speciation if it leads to reproductive isolation between susceptible and resistant types. We surveyed populations of the freshwater snail Lymnaea peregra for differences in habitat distribution and trematode parasitism between its two distinct shell morphs, L. ovata and L. peregra. We surveyed 58 populations (43 L. ovata, 15 L. peregra). At each location we recorded an array of habitat characteristics that were summarized using a nonlinear principal components analysis. This yielded two orthogonal habitat score variables. Discriminant analysis with these habitat dimensions indicated that the snail morphs differed in their habitat distribution. L. ovata preferred larger, more permanent natural habitats surrounded by forests, while L. peregra was found more often at a higher altitude, in nonpermanent habitats, often surrounded by meadows. The snails were parasitized by four cercarial types of castrating trematodes. The morphs had a similar prevalence of infection by each of the parasite types, with one exception: monostomid cercariae were found at a higher prevalence in L. ovata than in L. peregra. However, monostomes were rare parasites, and the difference in prevalence of infection was not significant when only populations with monostomes were compared. Our results indicate that variation in the overall prevalence of infection seems to be independent of snail morph, and do not support the idea that a difference in the rate of parasitism might explain differences in the habitat distribution of these snail morphs. [ABSTRACT FROM AUTHOR]

The suitability of the natural 15N abundance and of total N concentration of leaves as indicators of the type of plant N nutrition in a rain forest of French Guiana were tested. Leaf samples from primary legume species, non-legumes (pioneer species) and from the non-N2-fixing species Dicorynia guianensis were analyzed. Both δ15N and total leaf N varied widely (−1 ?δ15N (‰) ? 7 and 1 ? leaf N(%) ? 3.2) suggesting possible distinctions between diazotrophic and non-fixing plants. The δ15N also revealed two statistically distinct groups of non-N2-fixing species (δ15N = 5.14 ± 0.3 vs δ15N = 1.65 ± 0.17) related to the different ecological behaviors of these species in the successional processes. We conclude that the δ15N signature of plant leaves combined with their total N concentration may be relevant indicators for identifying functional groups within the community of non-N2-fixing species, as well as for detecting diazotrophy. Despite the variability in the δ15N of the non-N2-fixing species, N2-fixing groups can still be identified, provided that plants are simultaneously classified taxonomically, by their leaf δ15N and total N concentration and by the presence or absence of nodules. The variability in the δ15N of the non-fixing species is discussed. [ABSTRACT FROM AUTHOR]

The level of host exploitation is expected, under theory, to be selected to maximise (subject to constraints) the lifetime reproductive success of the parasite. Here we studied the effect of two castrating trematode species on their intermediate snail host, Potamopyrgus antipodarum. One of the trematode species, Microphallus sp., encysts in the snail host and the encysted larvae “hatch” following ingestion of infected snails by birds. The other species, Notocotylus gippyensis, by contrast, releases swimming larvae; ingestion of the snail host is not required for, and does not aid, transmission to the final host. We isolated field-collected snails for 3 months in the laboratory, and followed the survival of infected and uninfected snails under two conditions: not fed and fed ad libitum. Mortality of the infected hosts was higher than mortality of the uninfected ones, but the response to starvation treatment was parasite species specific. N. gippyensis induced significantly higher mortality in starved snails than did Microphallus. Based on these results, we suggest that host exploitation by different species of trematodes may depend on the type of transmission. Encysting in the snail host may select for a reduced rate of host exploitation so as to increase the probability of transmission to the final host. [ABSTRACT FROM AUTHOR]

Nitrogen relations of natural and disturbed tropical plant communities in northern Australia (Kakadu National Park) were studied. Plant and soil N characteristics suggested that differences in N source utilisation occur at community and species level. Leaf and xylem sap N concentrations of plants in different communities were correlated with the availability of inorganic soil N (NH+ 4 and NO− 3). In general, rates of leaf NO− 3 assimilation were low. Even in communities with a higher N status, including deciduous monsoon forest, disturbed wetland, and a revegetated mine waste rock dump, levels of leaf nitrate reductase, xylem and leaf NO− 3 levels were considerably lower than those that have been reported for eutrophic communities. Although NO− 3 assimilation in escarpment and eucalypt woodlands, and wetland, was generally low, within these communities there was a suite of species that exhibited a greater capacity for NO− 3 assimilation. These “high- NO− 3 species” were mainly annuals, resprouting herbs or deciduous trees that had leaves with high N contents. Ficus, a high-NO− 3 species, was associated with soil exhibiting higher rates of net mineralisation and net nitrification. “Low-NO− 3 species” were evergreen perennials with low leaf N concentrations. A third group of plants, which assimilated NO− 3 (albeit at lower rates than the high-NO− 3 species), and had high-N leaves, were leguminous species. Acacia species, common in woodlands, had the highest leaf N contents of all woody species. Acacia species appeared to have the greatest potential to utilise the entire spectrum of available N sources. This versatility in N source utilisation may be important in relation to their high tissue N status and comparatively short life cycle. Differences in N utilisation are discussed in the context of species life strategies and mycorrhizal associations. [ABSTRACT FROM AUTHOR]

Host-parasite interactions that result in host castration are evolutionarily similar to predator-prey interactions because both interactions terminate reproduction for the host or prey. Yet, host-parasite interactions differ from predator-prey interactions in that infected hosts remain alive and potentially can make adjustments to their life-history strategy before castration is complete. Here we exposed juvenile snails ( Potamopyrgus antipodarum) to infection by a digenetic trematode ( Microphallus sp.) in order to determine whether: (1) pre-reproductive individuals could be infected, (2) individuals that were exposed to infection shifted resources to early reproduction (fecundity compensation), and (3) infected individuals exhibit altered growth rates relative to uninfected individuals. We found that juveniles are susceptible to infection; hence P. antipodarum could be selected for earlier maturation in populations where the risk of infection is high. We also found that fecundity compensation does not occur in this snail. Finally, we found that Microphallus-infected snails exhibit altered growth rates; individuals infected as juveniles have lower growth rates and are smaller than uninfected snails. These results suggest that growth is altered by infection of a trematode parasite but reproduction in uninfected snails is not induced by exposure to trematode eggs. [ABSTRACT FROM AUTHOR]

Functional aspects of biodiversity were investigated in a lowland tropical rainforest in French Guyana (5°2′N, annual precipitation 2200 mm). We assessed leaf δ15N as a presumptive indicator of symbiotic N2 fixation, and leaf and wood cellulose δ13C as an indicator of leaf intrinsic water-use efficiency (CO2 assimilation rate/leaf conductance for water vapour) in dominant trees of 21 species selected for their representativeness in the forest cover, their ecological strategy (pioneers or late successional stage species, shade tolerance) or their potential ability for N2 fixation. Similar measurements were made in trees of native species growing in a nearby plantation after severe perturbation (clear cutting, mechanical soil disturbance). Bulk soil δ15N was spatially quite uniform in the forest (range 3–5‰), whereas average leaf δ15N ranged from −0.3‰ to 3.5‰ in the different species. Three species only, Diplotropis purpurea, Recordoxylon speciosum (Fabaceae), and Sclerolobium melinonii (Caesalpiniaceae), had root bacterial nodules, which was also associated with leaf N concentrations higher than 20 mg g−1. Although nodulated trees displayed significantly lower leaf δ15N values than non-nodulated trees, leaf δ15N did not prove a straightforward indicator of symbiotic fixation, since there was a clear overlap of δ15N values for nodulated and non-nodulated species at the lower end of the δ15N range. Perturbation did not markedly affect the difference δ15Nsoil − δ15Nleaf, and thus the isotopic data provide no evidence of an alteration in the different N acquisition patterns. Extremely large interspecific differences in sunlit leaf δ13C were observed in the forest (average values from −31.4 to −26.7‰), corresponding to intrinsic water-use efficiencies (ratio CO2 assimilation rate/leaf conductance for water vapour) varying over a threefold range. Wood cellulose δ13C was positively related to total leaf δ13C, the former values being 2–3‰ higher than the latter ones. Leaf δ13C was not related to leaf δ15N at either intraspecific or interspecific levels. δ13C of sunlit leaves was highest in shade hemitolerant emergent species and was lower in heliophilic, but also in shade-tolerant species. For a given species, leaf δ13C did not differ between the pristine forest and the disturbed plantation conditions. Our results are not in accord with the concept of existence of functional types of species characterized by common suites of traits underlying niche differentiation; rather, they support the hypothesis that each trait leads to a separate grouping of species. [ABSTRACT FROM AUTHOR]

Using field surveys and experimental infections, we investigated the influence of a trematode parasite on life history traits of adult Lymnaea elodes snails. We found that parasitism significantly affected the growth, fecundity, and survival of host snails. Within five of the six natural L. elodes populations we sampled, shell length of echinostome-infected hosts was significantly greater than for uninfected conspecifics. Furthermore, we show that gigantism occurs among experimentally infected snails due to an accelerated growth rate and size-selective mortality following an Echinostoma revolutum infection. The fecundity of infected snails sharply decreased beginning at 3 weeks post exposure (PE) and all egg production eventually ceased for most hosts by 5–6 weeks PE. Energy constraints, imposed by parasite development, alter the host energy budget. Early in the infection, parasite depletion of host energy reserves reduces host reproduction, but sufficient resources remain to allow accelerated host growth. Mortality was increased among host snails at two distinct stages: shortly after exposure and several weeks after cercariae were first released. We did not observe tissue degradation in snails during the first 4 weeks after exposure to the parasite, but destruction of host tissues was noted among snails dying later in the infection. [ABSTRACT FROM AUTHOR]

Snail hosts of different ages constitute different resource environments for larval trematodes because each individual has a particular energy budget and energy allocation pattern at the time of infection. Effects of monomiracidial trematode infections on shell growth, body dry weight and reproductive effort of the snail host were compared between controls and infected juvenile and adult snails during the prepatent and patent periods. The results demonstrate phenotypic plasticity in the host response. There is an age/size-specific effect characterized by limitation of growth rate when snails are infected as juveniles and reduction of reproductive effort when snails are infected as adults, and a time-specific effect with early enhancement of growth rate and reproductive effort for infected juvenile and adult snails respectively during prepatency, before reduction and cessation during patency. The parasite is considered as a generalist in its energy exploitation strategy of the snail host, but taking into account the differential host responses observed relative to the host age, the possibility of a host preference for juvenile or adult snails is discussed. [ABSTRACT FROM AUTHOR]

This study investigates some factors affecting the foraging activity of the leaf-cutting ant Atta laevigata. The study was conducted on an abandoned farm near Manaus, in Brazilian Amazonia, where forest was beginning to regenerate. I determined how temporal changes in the structure of the woody plant community (linked to the regeneration process) affected the leaf harvesting activity of this Neotropical ant. A 0.39-ha plot was established, and woody plants existing and emerging in this plot were identified, tagged, measured and mapped. At intervals of approximately 14 days all marked plants were checked to determine which had been attacked by A. laevigata during that period. Woody plant density doubled over the 18-months of the study. However, the number of plants attacked by A. laevigata, controlling for seasonal differences in ant activity, was independent of the number available for attack. The number of plants attacked was also independent of the number of new plants emerged in the previous 2 months. Apart from changes in total plant density, the relative abundance of most species also changed with time. Data on the composition of the plant community and on the composition of the ants' diet at different time intervals were subject to ordination analysis. The analyses revealed that successional changes in the composition of the woody plant community did significantly affect the composition of the leaf diet of A. laevigata. Variations in the ants' diet correlated most strongly with variations in the abundance of Bellucia imperialis. As this increased in abundance its relative contribution to the ants' diet increased. In addition, I observed a small decline in the diversity of plant resources exploited by the ants. B. imperialis was one of the species most preferred by A. laevigata, suggesting that variations in the diversity of its diet were related to the abundance of preferred species. The chances of an individual plant being attacked was independent of whether or not the plant had been attacked during the previous 5 months. This suggests that plant resistance against ant herbivory is not affected by previous damage. Two other lines of evidence also pointed to this. First, there was no significant difference in the leaf area cut (as measured during assays of leaf preference with A. laevigata) between leaves of plants that had been subject to artificial defoliation 3 mo earlier and those of control plants. Second, the number of attacks by A. laevigata on individuals of a given species usually followed a random distribution, even though a uniform or clumped distribution would have been expected under the hypothesis that damage by leaf-cutting ants affects the plant's resistance to further damage. [ABSTRACT FROM AUTHOR]

The effect of a change in the environment on reproductive and somatic energy allocation in an iteroparous freshwater clam Anodonta piscinalis was studied at different time of the seasonal reproductive cycle. Environmental change was produced by reciprocal transplant experiments among sites of varying productivity. In addition, clams were caged at high density to reduce the availability of resources. Transplanting females before fertilization (from May to June), or during the early development of the brood (from July to August) had no detectable effect on reproductive output. Early-season environment, however, affected body mass and percent fat content of females from two populations. This suggests that maintaining the level of reproductive allocation when resources are reduced early in the season leads to lower allocation to somatic growth and biochemical storage. Transplanting females late in the season (from September to November) had a substantial effect on reproductive output and body mass, but not on fat content, suggesting that late in the season allocation to biochemical energy storage is important. Hence, late in the season reproductive allocation may be adjusted to prevailing conditions in preparation for winter. Indeed, over-wintering site had a significant effect on percent fat content, body mass and shell growth when females were kept in a new environment from September to March. Variation among transplant sites in female body mass matched the estimated productivity of the sites, suggesting that it occurred in response to differences in the productivity of the habitats. The results emphasize the importance of taking seasonal changes in the priorities of energy allocation and the seasonality of reproductive processes into account when developing or testing models of optimal energy allocation. [ABSTRACT FROM AUTHOR]

Larval digenetic trematodes commonly castrate their first intermediate hosts, and should therefore impose strong selection on the timing and mode of host reproduction. Here we examine spatial variation in infection by trematodes in the freshwater snail Potamopyrgus antipodarum. Snails were collected at 11 different sites at Lake Alexandrina on the South Island of New Zealand from transects that ran perpendicular to the shore and across several different habitat types (from 0 to 8 m deep). Logistic regression was used to analyze the relationships between the frequency of trematode infection and snail size, habitat type, and transect location. On average, the probability of infection increased 3.3 times with each 1 mm increase in shell length. Prevalence of infection by the most common species of trematode, Microphallus sp., was highest in the shallow-water habitats where its final hosts (waterflow) spend most of their time. Prevalence of infection by another parasite, Telogaster ophistorchis (final host: eels) increased with depth, but because Microphallus was much more common, total infection by all trematodes decreased with depth. The effects of transect location were minor for Telogaster, but there was significant variation in Microphallus prevalence among transects, especially in the shore-bank habitat. Taken together, these results suggest that the risk of infection is spatially variable, but generally higher in shallow-water habitats, which may explain the greater frequency of sexual individuals as well as earlier reproduction among individuals near shore. [ABSTRACT FROM AUTHOR]