Habitat patch shape, not corridors, determines herbivory and fruit production of an annual plant

Habitat corridors confer many conservation benefits by increasing movement of organisms between habitat patches, but the benefits for some species may exact costs for others. For example, corridors may increase the abundance of consumers in a habitat to the detriment of the species they consume. In this study we assessed the impact of corridors on insect herbivory of a native plant, Solanum americanum, in large-scale, experimentally fragmented landscapes. We quantified leaf herbivory and assessed fruit production as a proxy for plant fitness. We also conducted field surveys of grasshoppers (Orthoptera), a group of abundant, generalist herbivores that feed on S. americanum, and we used exclosure cages to explicitly link grasshopper herbivory to fruit production of individual S. americanum. The presence of corridors did not increase herbivory or decrease plant fruit production. Likewise, corridors did not increase grasshopper abundance. Instead, patches in our landscapes with the least amount of edge habitat and the greatest amount of warmer "core" area had the highest levels of herbivory, the largest cost to plant fruit production as a result of herbivory, and the most grasshoppers. Thus habitat quality, governed by patch shape, can be more important than connectivity for determining levels of herbivory and the impact of herbivory on plant fitness in fragmented landscapes.     

Role of herbivory in controlling phytoplankton abundance: annual pigment budget for a temperate marine fjord

An annual pigment budget was constructed for Dabob Bay, Washington (USA) by comparing the downward vertical loss of phytoplankton pigments (chlorophyll and pheopigments) to the production of chlorophyll within the euphotic zone. The vertical flux of pigments was measured with sediment traps deployed at intervals of 1 to 6 wk over a 2.5 yr period yielding 763 d of trap exposure (28 November 1978–16 June 1981). The production rate of chlorophyll was calculated from measurements of algal specific growth rates, chlorophyll (chl) crops, primary production (as carbon) and appropriate C: chl ratios. Sixty one to 77% of the annual chlorophyll production was accounted for by the vertical flux of pheopigments resulting from herbivorous zooplankton grazing (macrozooplankton). Algal sinking, represented by downward chlorophyll flux, accounted for only 5 to 6% of the annual chlorophyll production. The remaining fraction of chlorophyll production was estimated to be consumed by small herbivores (microzooplankton), whose fecal material contributes to the suspended pool of pheopigments found in the euphotic zone. The suspended pheopigments are continuously removed by photodegradation. In Dabob Bay, the major process controlling phytoplankton abundance is zooplankton grazing and it appears that the ultimate fate of most phytoplankton is to be consumed by herbivores.     

Successional changes in plant resistance and tolerance to herbivory

Despite considerable research on plant defenses, we know very little about how temporal changes in the environment may influence resistance and tolerance levels, or the costs and benefits of these defense strategies for long-lived plant species. We hypothesized that, in successional habitats, predictable environmental changes should favor strong plasticity in defense phenotypes and that the costs, benefits, and levels of tolerance and resistance will change with environmental context. Using a widely distributed, old-field perennial, late goldenrod (Solidago altissima), we conducted a field experiment to test these predictions. We planted goldenrod genets exhibiting varying levels of resistance and tolerance into three early-successional and three late-successional fields (approximately three and 15 years in age, respectively) and experimentally measured resistance and tolerance levels and their associated costs and selection coefficients. We found a significant effect of successional stage but no effect of genotype or stage-genotype interaction on defense levels. Genets planted in early-successional fields appeared to be more resistant and less tolerant to herbivory than those same genets planted in late-successional fields. There were significant trade-offs between resistance and tolerance in early-successional fields but not in late-successional fields. Each late-successional field exhibited a significant cost or selection gradient for resistance, but there was no general pattern of resistance costs or selection gradients specific to a successional stage class. In contrast, there was evidence of stage-specific costs of tolerance; late-successional fields exhibited significant costs of tolerance whereas early-successional fields did not. There was no evidence of direct selection for or against tolerance in either stage. Our results suggest that defense phenotypes might change in qualitative ways during succession. High resistance in early stages may be attributed to associational effects of the early-successional community, reducing the probability of damage, and despite a cost of tolerance in late stages, tolerance may be beneficial in mitigating the effects of both herbivory and environmental stresses (i.e., low light availability) that limit fitness in these fields. This study provides experimental evidence that succession can strongly influence defense phenotypes and promote temporal variability in relative resistance and tolerance levels.     

Turbinaria ornata as an herbivory refuge for associate algae

Habitat associations are an integral part of coral reef community structure. Commonly, one organism lives in such close association within or near another that a spatial refuge occurs, whereby one of the organisms provides protection to the other. This is often the result of defenses of the host deterring an associate organism’s consumers. In Moorea, French Polynesia, the range and abundance of the brown macroalga, Turbinaria ornata, have increased drastically since 1980 such that dense aggregations of this macroalga are a dominant component of the backreef habitat. Turbinaria ornata is both mechanically and chemically defended from herbivores. Other species of macroalgae grow within aggregations of Turbinaria and may benefit from these defenses. This study investigates whether aggregations of Turbinaria create a refuge from herbivory for associate macroalgae. When Turbinaria aggregations were removed experimentally, there was a significant increase in the number of associate algal species. Moreover, an herbivory assay using the palatable local alga Acanthophora spicifera identified herbivory as the mechanism for lower diversity on bommies lacking Turbinaria aggregations. The local increase in algal richness due to the refuge from herbivory afforded by Turbinaria may be an important contribution to macroalgal and community dynamics on reefs in Moorea, French Polynesia.     

Role of climate and competitors in limiting fitness across range edges of an annual plant

It is often assumed that the geographic distributions of species match their climatic tolerances, but this assumption is not frequently tested. Moreover, few studies examine the relative importance of abiotic and biotic factors for limiting species ranges. We combined multiple approaches to assess the extent to which fitness of a widespread native annual legume, Chamaecrista fasciculata, decreases at and beyond its northern and western range edges, and how this is influenced by the presence of neighbors. First, we examined plant fitness and the effect of neighbors in natural populations at different geographic range locations for three years. Fitness decreased toward the northern range edge, but not the western edge. Neighbor removal had a consistently positive effect on seedpod production across all years and sites. Second, we established experimental populations at sites within the range, and at and beyond the northern and western range edges. We tracked individual fitness and recorded seedling recruitment in the following year (a complete generation) to estimate population growth rate. Individual fitness and population growth declined to near zero beyond both range edges, indicating that C. fasciculata with its present genetic composition will not establish in these regions, given conditions currently. We also carried out a neighbor removal treatment. Consistent with the natural populations, neighbors reduced seedpod production of reproductive adults. However, neighbors also increased early-season survival, and this positive effect early in life history resulted in a net positive effect of neighbors on lifetime fitness at most range locations. Our data show that the population growth rate of C. fasciculata includes values above replacement, and populations are well adapted to conditions up to the edge of the range, whereas the severely compromised fitness at sites beyond the edge precludes immediate establishment of populations and thereby impedes adaptation to these conditions.     

Crab herbivory regulates plant facilitative and competitive processes in Argentinean marshes

Interactions among plants have been hypothesized to be context dependent, shifting between facilitative and competitive in response to variation in physical and biological stresses. This hypothesis has been supported by studies of the importance of positive and negative interactions along abiotic stress gradients (e.g., salinity, desiccation), but few studies have tested how variation in biotic stresses can mediate the nature and strength of plant interactions. We examined the hypothesis that herbivory regulates the strength of competitive and facilitative interactions during succession in Argentinean marshes dominated by Spartina densiflora and Sarcocornia perennis. Spartina densiflora is preferred by the dominant herbivore in the system, the crab Chasmagnathus granulatus. We experimentally manipulated crab herbivory, plant structure, and shade, and we found that, when herbivory was low in the spring and summer, competitive interactions between plants were dominant, but in the fall, when herbivory was highest, facilitative interactions dominated, and Spartina densiflora survival was completely dependent upon association with Sarcocornia perennis. Moreover, experimental removal of Sarcocornia perennis across recently disturbed tidal flats revealed that, while Sarcocornia perennis positively affected small Spartina densiflora patches by decreasing herbivory, as patch size increases and they can withstand the impact of herbivory, competitive interactions predominated and Spartina densiflora ultimately outcompeted Sarcocornia perennis. These results show that herbivory can mediate the balance between facilitative and competitive processes in vascular plant communities and that the strength of consumer regulation of interactions can vary seasonally and with patch size.     

Constitutive and induced defenses to herbivory in above- and belowground plant tissues

A recent surge in attention devoted to the ecology of soil biota has prompted interest in quantifying similarities and differences between interactions occurring in above- and belowground communities. Furthermore, linkages that interconnect the dynamics of these two spatially distinct ecosystems are increasingly documented. We use a similar approach in the context of understanding plant defenses to herbivory, including how they are allocated between leaves and roots (constitutive defenses), and potential cross-system linkages (induced defenses). To explore these issues we utilized three different empirical approaches. First, we manipulated foliar and root herbivory on tobacco (Nicotiana tabacum) and measured changes in the secondary chemistry of above- and belowground tissues. Second, we reviewed published studies that compared levels of secondary chemistry between leaves and roots to determine how plants distribute putative defense chemicals across the above- and belowground systems. Last, we used meta-analysis to quantify the impact of induced responses across plant tissue types. In the tobacco system, leaf-chewing insects strongly induced higher levels of secondary metabolites in leaves but had no impact on root chemistry. Nematode root herbivores, however, elicited changes in both leaves and roots. Virtually all secondary chemicals measured were elevated in nematode-induced galls, whereas the impact of root herbivory on foliar chemistry was highly variable and depended on where chemicals were produced within the plant. Importantly, nematodes interfered with aboveground metabolites that have biosynthetic sites located in roots (e.g., nicotine) but had the opposite effect (i.e., nematodes elevated foliar expression) on chemicals produced in shoots (e.g., phenolics and terpenoids). Results from our literature review suggest that, overall, constitutive defense levels are extremely similar when comparing leaves with roots, although certain chemical classes (e.g., alkaloids, glucosinolates) are differentially allocated between above- and belowground parts. Based on a meta-analysis of induced defense studies we conclude that: (1) foliar induction generates strong responses in leaves, but much weaker responses in roots, and (2) root induction elicits responses of equal magnitude in both leaves and roots. We discuss the importance of this asymmetry and the paradox of cross-system induction in relation to optimal defense theory and interactions between above- and belowground herbivory.     

亚热带阔叶林植物叶片虫食特征研究

植物与昆虫是森林生态系统的重要组成成分,两者通过长期的协同进化形成了密切的相互关系,在森林生态系统中发挥着承上启下的作用,连接了初级生产和高级消费,是森林生态系统中重要的动态中心。植物叶片的虫食特征是植物与昆虫相互作用关系的重要表征。选取福建梅花山和浙江天童两地76种常绿阔叶林植物为研究对象,采用野外调查与室内统计分析相结合的方式对植物叶片的虫食率和虫食频度进行了研究,以期了解亚热带常绿阔叶林植物叶片所面临的食叶昆虫取食压力。结果表明:76种植物平均虫食率为7.21%,虫食频度为32.95%。多数植物的叶片虫食率低于10%,叶片虫食频度主要分布在10%-60%之间。乔木种与灌木种、优势植物与伴生植物之间的叶片虫食率和虫食频度均不存在显著差异(P>0.05)。超过60%的叶片虫食率和虫食频度发生在展叶期。福建梅花山常绿阔叶林植物的叶片虫食率(P=0.012)和虫食频度(P=0.74)均高于浙江天童。植物幼叶的虫食率随着海拔的升高而下降。以上结果表明,常绿阔叶林植物的叶片虫食强度介于热带雨林(11.1%)和温带森林(7.1%)之间,展叶期是叶片虫食发生的主要阶段,表现出过渡性特征;叶片的虫食在不同生活型和优势程度的植物间存在差异;叶片虫食率随纬度的升高和海拔的上升而降低。     

Reduced pollinator service and elevated pollen limitation at the geographic range limit of an annual plant

Mutualisms are well known to influence individual fitness and the population dynamics of partner species, but little is known about whether they influence species distributions and the location of geographic range limits. Here, we examine the contribution of plant-pollinator interactions to the geographic range limit of the California endemic plant Clarkia xantiana ssp. xantiana. We show that pollinator availability declined from the center to the margin of the geographic range consistently across four years of study. This decline in pollinator availability was caused to a greater extent by variation in the abundance of generalist rather than specialist bee pollinators. Climate data suggest that patterns of precipitation in the current and previous year drove variation in bee abundance because of its effects on cues for bee emergence in the current year and the abundance of floral resources in the previous year. Experimental floral manipulations showed that marginal populations had greater outcross pollen limitation of reproduction, in parallel with the decline in pollinator abundance. Although plants are self-compatible, we found no evidence that autonomous selfing contributes to reproduction, and thus no evidence that it alleviates outcross pollen limitation in marginal populations. Furthermore, we found no association between the distance to the range edge and selfing rate, as estimated from sequence and microsatellite variation, indicating that the mating system has not evolved in response to the pollination environment at the range periphery. Overall, our results suggest that dependence on pollinators for reproduction may be an important constraint limiting range expansion in this system.     

Gastropod herbivory in response to elevated CO2 and N addition impacts plant community composition

In this study, the influence of elevated carbon dioxide (CO2) and nitrogen (N) deposition on gastropod herbivory was investigated for six annual species in a California annual grassland community. These experimentally simulated global changes increased availability of important resources for plant growth, leading to the hypothesis that species with the most positive growth and foliar nutrient responses would experience the greatest increase in herbivory. Counter to the expectations, shifts in tissue N and growth rates caused by N deposition did not predict shifts in herbivore consumption rates. N deposition increased seedling N concentrations and growth rates but did not increase herbivore consumption overall, or for any individual species. Elevated CO2 did not influence growth rates nor have a statistically significant influence on seedling N concentrations. Elevated CO2 at ambient N levels caused a decline in the number of seedlings consumed, but the interaction between CO2 and N addition differed among species. The results of this study indicate that shifting patterns of herbivory will likely influence species composition as environmental conditions change in the future; however, a simple trade-off between shifting growth rates and palatability is not evident.     

Hydrologic regime and herbivory stabilize an alternative state in Yellowstone National Park.

A decline in the stature and abundance of willows during the 20th century occurred throughout the northern range of Yellowstone National Park, where riparian woody-plant communities are key components in multiple-trophic-level interactions. The potential causes of willow decline include climate change, increased elk browsing coincident with the loss of an apex predator, the gray wolf, and an absence of habitat engineering by beavers. The goal of this study was to determine the spatial and temporal patterns of willow establishment through the 20th century and to identify causal processes. Sampled willows established from 1917 to 1999 and contained far fewer young individuals than was predicted from a modeled stable willow population, indicating reduced establishment during recent decades. Two hydrologically distinct willow establishment environments were identified: fine-grained beaver pond sediments and coarse-grained alluvium. Willows established on beaver pond sediment earlier in time, higher on floodplain surfaces, and farther from the current stream channel than did willows on alluvial sediment. Significant linear declines from the 1940s to the 1990s in alluvial willow establishment elevation and lateral distance from the stream channel resulted in a much reduced area of alluvial willow establishment. Willow establishment was not well correlated with climate-driven hydrologic variables, but the trends were consistent with the effects of stream channel incision initiated in ca. 1950, 20-30 years after beaver dam abandonment. Radiocarbon dates and floodplain stratigraphy indicate that stream incision of the present magnitude may be unprecedented in the past two millennia. We propose that hydrologic changes, stemming from competitive exclusion of beaver by elk overbrowsing, caused the landscape to transition from a historical beaver-pond and willow-mosaic state to its current alternative stable state where active beaver dams and many willow stands are absent. Because of hydrologic changes in streams, a rapid return to the historical state may not occur by reduction of elk browsing alone. Management intervention to restore the historical hydrologic regime may be necessary to recover willows and beavers across the landscape.     

Consequences of niche overlap for ecosystem functioning: an experimental test with pond grazers

While the number of studies investigating the effects of species diversity on ecosystem properties continues to expand, few have explicitly examined how ecosystem functioning depends quantitatively on the degree of niche complementarity among species. We report the results of a microcosm experiment where similarity in habitat use among aquatic snail species was evaluated as a predictor of changes in community and ecosystem properties due to increasing species richness. Replicate microcosms with all possible one- and two-species combinations of a guild of six snail species were stocked with identical initial snail biomass. Microcosms with two species of snails had greater final snail biomass, lower attached algae biomass, and less total organic matter than monocultures. Snail species differed in their use of five distinct habitat types in the microcosms. Similarity in habitat use between a species pair was negatively related to the magnitude of change (e.g., deltaEF [change in ecosystem function]) in dissolved oxygen. periphyton biomass, and accrual of organic matter with a change in diversity. However, using the most stringent criterion for complementarity effects (e.g., Dmax [proportional deviation of the total polyculture yield from the highest yielding monoculture]), a relationship between species' niche similarity and changes in function with increasing species richness was only observed for dissolved oxygen. The identity of snail species present in the microcosms had strong effects on total organic matter, snail biomass, dissolved oxygen, periphyton biomass, and sedimentation rate. In this study, herbivore identity, sampling effects, and niche complementarity all appear to contribute to species richness effects on pond ecosystem properties and community structure. The analytical approach employed here may profitably be used in other systems to quantify the role of niche complementarity in species richness-ecosystem function relationships.     

Toxicokinetic-toxicodynamic modelling in an individual based context—Consequences of parameter variability

Toxicokinetic-toxicodynamic (TKTD) models simulate the time-course of toxicant concentration in the organism and toxicity at the level of the organism. A link between TKTD models that simulate survival and individual based models for populations (IBMs) is proposed which allows TKTD parameters to vary between individuals. The TKTD-IBM predicts different survival in response to toxicants when TKTD parameters vary amongst individuals compared to the survival predicted with fixed TKTD parameters. The model with fixed parameters represents the concept of stochastic death whereas the model with variable parameters behaves, at least partly, according to the individual tolerance distribution concept. The whole set of TKTD parameters of an individual can be interpreted as constituting “individual tolerance”.     

Genetic sub-structure and intermediate optimal outcrossing distance in the marine angiosperm <Emphasis Type="Italic">Zostera marina</Emphasis>

The spatial distribution of genetic variability depends on the spatial patterns of clonal and sexual reproduction, gene flow, genetic drift and natural selection. Species with restricted dispersal may exhibit genetic structuring within populations with immediate neighbours being close relatives, and may show differentiation among populations. Genetic structuring of a species may have important genetic, evolutionary and ecological consequences including distance-dependent mating success. In this study we used microsatellite markers to show that clones of Zostera marina in a population in the Ria Formosa, Portugal, were aggregated and covered distances of up to 3–4 m. Clones within 4 m of each other exhibited significant and positive coancestry values, reflecting the limited seed dispersal of this species. Hand-pollinations between near (0–10.9 m), intermediate (11–32 m) and far (15 km) individuals resulted in similar levels of seed set, although the near pollinations had higher, although not statistically significant, levels of seed abortion during maturation. Seeds from intermediate-distance pollinations had a significantly higher proportion of seeds germinate and shorter germination time than both the near and far seeds. Similarly, the average number of seedlings produced per pollination, used as an overall estimate of fitness, was significantly greater for the intermediate distance when compared to both near and far pollinations. These results suggest that the genetic structuring observed may result in both inbreeding and outbreeding depression, which gives rise to an intermediate optimal outcrossing distance.     

Seasonal and annual variability in an epifaunal community in the German Bight

While the benthic infauna of the North Sea has been studied intensively over the past decades, few studies have focused on the larger mobile epifauna. Studies carried out to date have described the distribution of epifaunal communities over the whole of the North Sea, but variability within the identified communities, which occurs on a much smaller scale, has so far remained unstudied. This is the first study to describe seasonal and annual variability of an epifaunal assemblage in the German Bight area, where environmental conditions are highly variable. The benthic community was sampled with a 2 m beam trawl from 1998 to 2001. The echinoderms Ophiura albida and Asterias rubens and the crustacean Pagurus bernhardus were the dominant species caught throughout the study period. Overall the species composition of the catches was relatively consistent, while abundances of dominant species fluctuated considerably between sampling periods. Differences between sampling periods were not only influenced by the abundances of dominant species, but also by less dominant species such as Ophiura ophiura, Astropecten irregularis, Corystes cassivelaunus, Crangon crangon and Aphorrahis pespelicani. The abundances of these species varied annually and seasonally in the assemblage. Clear differences between summer and winter in the species composition, abundance and biomass were identified. Annual and seasonal changes were most likely linked to migratory movements of epifauna into and out of the area under investigation. Temporal changes in species composition and abundance correlated best with water temperatures, while the spatial distribution of the total biomass over the whole sampling period was correlated with sediment characteristics. Anthropogenic influences such as fishing activity and chronic large-scale eutrophication are thought to have influenced the community on a long-term basis, but have been considered unlikely causes for the short-term variability described by this study.Communicated by J.P. Thorpe, Port Erin     

Tissue type matters: selective herbivory on different life history stages of an isomorphic alga

Selective grazing by herbivores can have large effects on the population dynamics and community structure of primary producers. However, the ecological impacts of within-species herbivore preference for tissues of different phases (e.g., ploidy levels) or reproductive status remain relatively poorly known, especially among algae and other species with free-living haploid (gametophyte) and diploid (sporophyte) phases. We tested for herbivore selectivity among tissue types of the isomorphic (identical haploid and diploid free-living stages) red alga Mazzaella flaccida. Laboratory feeding assays demonstrated that the snail Tegula funebralis exhibited more than a threefold preference for gametophyte reproductive tissue over other tissue types, due to morphological differences. In contrast, the urchin Strongylocentrotus purpuratus did not distinguish as clearly between gametophytes and sporophytes; but it did prefer sporophyte reproductive to nonreproductive tissue, due to differences in water-soluble chemicals. Field surveys of grazer damage on M. flaccida blades were consistent with these laboratory preferences, with more damage found on gametophytes than sporophytes and reproductive than nonreproductive tissues. Differential fecundity can contribute to a skew in relative frequencies of phases in the field, and our results suggest that differential grazing by snails may contribute to this pattern and thus play a role in algal population biology.     

Variation in herbivore-mediated indirect effects of an invasive plant on a native plant

Theory predicts that damage by a shared herbivore to a secondary host plant species may either be higher or lower in the vicinity of a preferred host plant species. To evaluate the importance of ecological factors, such as host plant proximity and density, in determining the direction and strength of such herbivore-mediated indirect effects, we quantified oviposition by the exotic weevil Rhinocyllus conicus on the native wavyleaf thistle Cirsium undulatum in midgrass prairie on loam soils in the upper Great Plains, USA. Over three years (2001-2003), the number of eggs laid by R. conicus on C. undulatum always decreased significantly with distance (0-220 m) from a musk thistle (Carduus nutans L.) patch. Neither the level of R. conicus oviposition on C. undulatum nor the strength of the distance effect was predicted by local musk thistle patch density or by local C. undulatum density (<5 m). The results suggest that high R. conicus egg loads on C. undulatum near musk thistle resulted from the native thistle's co-occurrence with the coevolved preferred exotic host plant and not from the weevil's response to local host plant density. Mean egg loads on C. undulatum also were greater at sites with higher R. conicus densities. We conclude that both preferred-plant proximity and shared herbivore density strongly affected the herbivore-mediated indirect interaction, suggesting that such interactions are important pathways by which invasive exotic weeds can indirectly impact native plants.