Directional change in upland tundra plant communities 20‐30 years after seismic exploration in the Canadian low‐arctic

Question: What is the disturbance response of low‐arctic plant communities two to three decades after seismic exploration. Location: Mackenzie River Delta, low‐arctic, northwestern Canada. Methods: Plant communities in two upland tundra vegetation types were compared between winter seismic lines, created between 1970 and 1986, and adjacent “reference” tundra. Also, we used aerial surveys to quantify the total area impacted by visible linear features. Results: Vascular plant cover was significantly higher, and lichen cover significantly lower, on seismic lines than in reference tundra. The increase in vascular plant cover was attributable to deciduous shrubs and graminoids. There were significant differences in plant community composition between seismic lines and reference tundra but no differences in species diversity or richness. Betula glandulosa and Arctagrostis latifolia were significant indicator species for seismic lines, while Saussurea angustifolia was a significant indicator for reference tundra. Based on the aerial surveys, these effects apply to at least 90% of seismic lines from two‐dimensional programs in these habitat types during the 1970s. Conclusions: Vegetation composition and structure on 20‐30‐year‐old seismic lines differs from reference upland tundra despite no persistent differences in organic layer depth or depth to permafrost. We propose that this reflects: (1) successional redevelopment following changes in soil conditions and nutrient availability arising from the disturbance, and/or (2) disturbance‐initiated succession towards a community reflecting current climatic conditions.     

Vegetation responses in Alaskan arctic tundra after 8 years of a summer warming and winter snow manipulation experiment

We used snow fences and small (1 m²) open‐topped fiberglass chambers (OTCs) to study the effects of changes in winter snow cover and summer air temperatures on arctic tundra. In 1994, two 60 m long, 2.8 m high snow fences, one in moist and the other in dry tundra, were erected at Toolik Lake, Alaska. OTCs paired with unwarmed plots, were placed along each experimental snow gradient and in control areas adjacent to the snowdrifts. After 8 years, the vegetation of the two sites, including that in control plots, had changed significantly. At both sites, the cover of shrubs, live vegetation, and litter, together with canopy height, had all increased, while lichen cover and diversity had decreased. At the moist site, bryophytes decreased in cover, while an increase in graminoids was almost entirely because of the response of the sedge Eriophorum vaginatum. These community changes were consistent with results found in studies of responses to warming and increased nutrient availability in the Arctic. However, during the time period of the experiment, summer temperature did not increase, but summer precipitation increased by 28%. The snow addition treatment affected species abundance, canopy height, and diversity, whereas the summer warming treatment had few measurable effects on vegetation. The interannual temperature fluctuation was considerably larger than the temperature increases within OTCs (<2°C), however. Snow addition also had a greater effect on microclimate by insulating vegetation from winter wind and temperature extremes, modifying winter soil temperatures, and increasing spring run‐off. Most increases in shrub cover and canopy height occurred in the medium snow‐depth zone (0.5–2 m) of the moist site, and the medium to deep snow‐depth zone (2–3 m) of the dry site. At the moist tundra site, deciduous shrubs, particularly Betula nana, increased in cover, while evergreen shrubs decreased. These differential responses were likely because of the larger production to biomass ratio in deciduous shrubs, combined with their more flexible growth response under changing environmental conditions. At the dry site, where deciduous shrubs were a minor part of the vegetation, evergreen shrubs increased in both cover and canopy height. These changes in abundance of functional groups are expected to affect most ecological processes, particularly the rate of litter decomposition, nutrient cycling, and both soil carbon and nitrogen pools. Also, changes in canopy structure, associated with increases in shrub abundance, are expected to alter the summer energy balance by increasing net radiation and evapotranspiration, thus altering soil moisture regimes.     

The response of Alaskan arctic tundra to experimental warming: differences between short- and long-term responses

Global climate models predict continued rapid warming for most of the Arctic throughout the next century. To further understand the response of arctic tundra to climate warming, four sites in northern Alaska were warmed for five to seven consecutive growing seasons using open‐top chambers. Sites were located in dry heath and wet meadow communities near Barrow (71°18′N, 156°40′W) and Atqasuk (70°29′N, 157°25′W). Change in plant community composition was measured using a point frame method. During the period of observation, species richness declined in control plots by up to 2.7 species plot^?1. Responses to warming varied by site but similar trends included increased canopy height (?0.1 to 2.3 cm) and relative cover of standing dead plant matter (1.5–6.0%) and graminoids (1.8–5.8%) and decreased species diversity (0.1–1.7 species plot^?1) and relative cover of lichens (0.2–9.1%) and bryophytes (1.4–4.6%) (parentheses enclose the range of average values for the sites). The response to warming was separated into an initial short‐term response assessed after two growing seasons of warming and a secondary longer‐term response assessed after an additional three to five growing seasons of warming. The initial responses to warming were similar in the four sites, while the secondary responses varied by site. The response to warming was greater at Barrow than Atqasuk because of a greater initial response at Barrow. However, the long‐term response to warming was projected to be greater at Atqasuk because of a greater secondary response at Atqasuk. These findings show that predictions of vegetation change due to climate warming based on manipulative experiments will differ depending on both the duration and plant community on which the study focuses.     

Biomanipulation as a useful water quality management tool in deep stratifying reservoirs

In six deep, soft-water reservoirs, ranging from oligotrophic to eutrophic, fishery management has been guided by the use of biomanipulation to improve water quality and opportunities for recreational fishing. As evidenced by the establishment of larger-bodied daphnids, a low level of zooplanktivory could be maintained in the newly filled Grosse Dhünn and refilled Brucher and Lingese Reservoirs only by regular stocking of piscivores (Sander lucioperca, Esox lucius, Salmo trutta lacustris, Onchorhynchus mykiss) in combination with size and bag limitations for the recreational fisheries. However, in the mesotrophic Pre-Reservoir Grosse Dhünn, Bever Reservoir and the newly filled slightly eutrophic Wupper Reservoir it took between 8 and 10 years before the predator-resistant zooplankton community responded to management with a switch to larger daphnids. Except for oligotrophic Grosse Dhünn Reservoir where angling is prohibited, the expected enhancement of piscivore biomass through stocking not least was prevented by anglers. However, growth of perch (Perca fluviatilis) benefited from the changed fishery management relying upon stocking piscivores only allowing them to reach the size of piscivory. The appearance of larger daphnids in Pre-Reservoir Grosse Dhünn and Wupper Reservoir resulted in the biomass of the Daphnia spring peak to increase and occur earlier causing prolonged clear water conditions. Although the larger daphnids increased transparency, total summer mean chlorophyll concentrations in the euphotic zone only decreased in the slightly eutrophic reservoirs due to reduced phosphorus availability following unchanged external phosphorus loadings. Reduced phosphorus availability in these reservoirs caused a real oligotrophication. Although the edible seston fraction was controlled “top down” the results did not support the trophic cascade hypotheses because total phytoplankton remained controlled “bottom up”, admittedly triggered by “top down” forces. In general, the results support the importance of indirect (non-lethal) effects as the driving forces for the successful biomanipulations, particularly in slightly eutrophic reservoirs. Not least, the delayed zooplankton response provides an indication that the underlying change of internal feedbacks was not driven by external forces to stabilize the reservoirs trophic state in these deep stratifying reservoirs. Handling editor: D. Ryder     

Taller and larger: shifts in Arctic tundra leaf traits after 16 years of experimental warming

Understanding plant trait responses to elevated temperatures in the Arctic is critical in light of recent and continuing climate change, especially because these traits act as key mechanisms in climate‐vegetation feedbacks. Since 1992, we have artificially warmed three plant communities at Alexandra Fiord, Nunavut, Canada (79°N). In each of the communities, we used open‐top chambers (OTCs) to passively warm vegetation by 1–2 °C. In the summer of 2008, we investigated the intraspecific trait responses of five key species to 16 years of continuous warming. We examined eight traits that quantify different aspects of plant performance: leaf size, specific leaf area (SLA), leaf dry matter content (LDMC), plant height, leaf carbon concentration, leaf nitrogen concentration, leaf carbon isotope discrimination (LCID), and leaf δ¹⁵N. Long‐term artificial warming affected five traits, including at least one trait in every species studied. The evergreen shrub Cassiope tetragona responded most frequently (increased leaf size and plant height/decreased SLA, leaf carbon concentration, and LCID), followed by the deciduous shrub Salix arctica (increased leaf size and plant height/decreased SLA) and the evergreen shrub Dryas integrifolia (increased leaf size and plant height/decreased LCID), the forb Oxyria digyna (increased leaf size and plant height), and the sedge Eriophorum angustifolium spp. triste (decreased leaf carbon concentration). Warming did not affect δ¹⁵N, leaf nitrogen concentration, or LDMC. Overall, growth traits were more sensitive to warming than leaf chemistry traits. Notably, we found that responses to warming were sustained, even after many years of treatment. Our work suggests that tundra plants in the High Arctic will show a multifaceted response to warming, often including taller shoots with larger leaves.     

Food web changes in arctic ecosystems related to climate warming

Sedimentary records from three Canadian High Arctic ponds on Ellesmere Island, spanning the last several thousand years, show major shifts in pond communities within the last ～200 years. These paleolimnological data indicate that aquatic insect (Diptera: Chironomidae) populations rapidly expanded and greatly increased in community diversity beginning in the 19th century. These invertebrate changes coincided with striking shifts in algal (diatom) populations, indicating strong food‐web effects because of climate warming and reduced ice‐cover in ponds. Predicted future warming in the Arctic may produce ecological changes that exceed the large shifts that have already occurred since the 19th century.     

Changes in high arctic tundra plant reproduction in response to long‐term experimental warming

We provide new information on changes in tundra plant sexual reproduction in response to long‐term (12 years) experimental warming in the High Arctic. Open‐top chambers (OTCs) were used to increase growing season temperatures by 1–2 °C across a range of vascular plant communities. The warming enhanced reproductive effort and success in most species; shrubs and graminoids appeared to be more responsive than forbs. We found that the measured effects of warming on sexual reproduction were more consistently positive and to a greater degree in polar oasis compared with polar semidesert vascular plant communities. Our findings support predictions that long‐term warming in the High Arctic will likely enhance sexual reproduction in tundra plants, which could lead to an increase in plant cover. Greater abundance of vegetation has implications for primary consumers – via increased forage availability, and the global carbon budget – as a function of changes in permafrost and vegetation acting as a carbon sink. Enhanced sexual reproduction in Arctic vascular plants may lead to increased genetic variability of offspring, and consequently improved chances of survival in a changing environment. Our findings also indicate that with future warming, polar oases may play an important role as a seed source to the surrounding polar desert landscape.     

Demographic effects of warming,elevated soil nitrogen and thinning on the colonization of a perennial plant

Global change is causing significant modifications to native plant communities. These effects can be direct through changes in productivity, or indirect through the spread of invading species. Identifying vital traits important for individual species’ response to environmental variation could be useful for making predictions about how entire communities may respond to global change. I studied the effects of factors associated with global change on the demography of an experimentally introduced species, Pityopsis aspera. In a Florida old-field, I investigated how warming, increased soil nitrogen and thinning of the extant plant community affected survival, growth and reproduction of P. aspera using a life table response experiment. The estimated population growth rate (λ) of P. aspera was reduced by nitrogen addition, as a result of decreased fecundity. However, λ increased in response to the warming treatment, as a result of increased fecundity. In the presence of thinning, both warming and nitrogen served to increase λ as a result of an increase in the growth of young individuals. This experiment illustrates how different vital rates contribute to the population level responses of an experimentally introduced plant to warming, and nitrogen deposition. Results also show how these demographic responses may occur via indirect effects through established species. This work highlights the importance of studying interactions among temperature, soil nitrogen and demography across the entire life cycle in order to capture the complex and, often, non-additive relationships mediating global change effects.     

Size‐balanced community reorganization in response to nutrients and warming

It is widely accepted that global warming will adversely affect ecological communities. As ecosystems are simultaneously exposed to other anthropogenic influences, it is important to address the effects of climate change in the context of many stressors. Nutrient enrichment might offset some of the energy demands that warming can exert on organisms by stimulating growth at the base of the food web. It is important to know whether indirect effects of warming will be as ecologically significant as direct physiological effects. Declining body size is increasingly viewed as a universal response to warming, with the potential to alter trophic interactions. To address these issues, we used an outdoor array of marine mesocosms to examine the impacts of warming, nutrient enrichment and altered top‐predator body size on a community comprised of the predator (shore crab Carcinus maenas), various grazing detritivores (amphipods) and algal resources. Warming increased mortality rates of crabs, but had no effect on their moulting rates. Nutrient enrichment and warming had near diametrically opposed effects on the assemblage, confirming that the ecological effects of these two stressors can cancel each other out. This suggests that nutrient‐enriched systems might act as an energy refuge to populations of species under metabolic constraints due to warming. While there was a strong difference in assemblages between mesocosms containing crabs compared to mesocosms without crabs, decreasing crab size had no detectable effect on the amphipod or algal assemblages. This suggests that in allometrically balanced communities, the expected long‐term effect of warming (declining body size) is not of similar ecological consequence to the direct physiological effects of warming, at least not over the six week duration of the experiment described here. More research is needed to determine the long‐term effects of declining body size on the bioenergetic balance of natural communities.     

Quality or quantity: the direct and indirect effects of host plants on herbivores and their natural enemies

Resource quality (plant nitrogen) and resource quantity (plant density) have often been argued to be among the most important factors influencing herbivore densities. A difficulty inherent in the studies that manipulate resource quality, by changing nutrient levels, is that resource quantity can be influenced simultaneously, i.e. fertilized plants grow more. In this study we disentangled the potentially confounding effects of plant quality and quantity on herbivore trophic dynamics by separately manipulating nutrients and plant density, while simultaneously reducing pressure from natural enemies (parasitoids) in a fully factorial design. Plant quality of the sea oxeye daisy, Borrichia frutescens, a common coastal species in Florida, was manipulated by adding nitrogen fertilizer to increase and sugar to decrease available nitrogen. Plant density was manipulated by pulling by hand 25 or 50% of Borrichia stems on each plot. Because our main focal herbivore was a gall making fly, Asphondylia borrichiae, which attacks only the apical meristems of plants, manipulating plant nitrogen levels was a convenient and reliable way to change plant quality without impacting quantity because fertilizer and sugar altered plant nitrogen content but not plant density. Our other focal herbivore was a sap-sucker, Pissonotus quadripustulatus, which taps the main veins of leaves. Parasitism of both herbivores was reduced via yellow sticky traps that caught hymenopteran parasitoids. Plant quality significantly affected the per stem density of both herbivores, with fertilization increasing, and sugar decreasing the densities of the two species but stem density manipulations had no significant effects. Parasitoid removal significantly increased the densities of both herbivores. Top-down manipulations resulted in a trophic cascade, as the density of Borrichia stems decreased significantly on parasitoid removal plots. This is because reduced parasitism increases gall density and galls can kill plant stems. In this system, plant quality and natural enemies impact per stem herbivore population densities but plant density does not.     

Long‐term experimental warming alters community composition of ascomycetes in Alaskan moist and dry arctic tundra

Arctic tundra regions have been responding to global warming with visible changes in plant community composition, including expansion of shrubs and declines in lichens and bryophytes. Even though it is well known that the majority of arctic plants are associated with their symbiotic fungi, how fungal community composition will be different with climate warming remains largely unknown. In this study, we addressed the effects of long‐term (18 years) experimental warming on the community composition and taxonomic richness of soil ascomycetes in dry and moist tundra types. Using deep Ion Torrent sequencing, we quantified how OTU assemblage and richness of different orders of Ascomycota changed in response to summer warming. Experimental warming significantly altered ascomycete communities with stronger responses observed in the moist tundra compared with dry tundra. The proportion of several lichenized and moss‐associated fungi decreased with warming, while the proportion of several plant and insect pathogens and saprotrophic species was higher in the warming treatment. The observed alterations in both taxonomic and ecological groups of ascomycetes are discussed in relation to previously reported warming‐induced shifts in arctic plant communities, including decline in lichens and bryophytes and increase in coverage and biomass of shrubs.     

A test for top-down cascade in a detritus-based food web by litter-dwelling web spiders

A top-down cascade in detritus-based food webs has the potential to alter ecosystem functioning but its prevalence in terrestrial systems remains unclear. We manipulated the density of the dominant web spider Neriene brongersmai for 1.5 years to see whether the cascading effect would propagate down to the litter decomposition rate in the forest-floor of a Japanese cedar forest. Decreasing spider abundance increased collembolan density, but it did not change the biomass of microorganisms or the litter decomposition rate. We propose possible explanations for the absence of a top-down cascade and discuss the difficulty in generalizing the presence and absence of a cascading effect in soil ecosystems.     

Critical indirect effects of climate change on sub‐Antarctic ecosystem functioning

Sub‐Antarctic islands represent critical breeding habitats for land‐based top predators that dominate Southern Ocean food webs. Reproduction and molting incur high energetic demands that are sustained at the sub‐Antarctic Prince Edward Islands (PEIs) by both inshore (phytoplankton blooms; “island mass effect”; autochthonous) and offshore (allochthonous) productivity. As the relative contributions of these sustenance pathways are, in turn, affected by oceanographic conditions around the PEIs, we address the consequences of climatically driven changes in the physical environment on this island ecosystem. We show that there has been a measurable long‐term shift in the carbon isotope signatures of the benthos inhabiting the shallow shelf region of the PEIs, most likely reflecting a long‐term decline in enhanced phytoplankton productivity at the islands in response to a climate‐driven shift in the position of the sub‐Antarctic Front. Our results indicate that regional climate change has affected the balance between allochthonous and autochthonous productivity at the PEIs. Over the last three decades, inshore‐feeding top predators at the islands have shown a marked decrease in their population sizes. Conversely, population sizes of offshore‐feeding predators that forage over great distances from the islands have remained stable or increased, with one exception. Population decline of predators that rely heavily on organisms inhabiting the inshore region strongly suggest changes in prey availability, which are likely driven by factors such as fisheries impacts on some prey populations and shifts in competitive interactions among predators. In addition to these local factors, our analysis indicates that changes in prey availability may also result indirectly through regional climate change effects on the islands' marine ecosystem. Most importantly, our results indicate that a fundamental shift in the balance between allochthonous and autochthonous trophic pathways within this island ecosystem may be detected throughout the food web, demonstrating that the most powerful effects of climate change on marine systems may be indirect.     

Initial effects of experimental warming on above- and belowground components of net ecosystem CO2 exchange in arctic tundra

The Arctic contains extensive soil carbon reserves that could provide a substantial positive feedback to atmospheric CO₂ concentrations and global warming. Evaluation of this hypothesis requires a mechanistic understanding of the in situ responses of individual components of tundra net ecosystem CO₂ exchange (NEE) to warming. In this study, we measured NEE, total ecosystem respiration and respiration from below ground in experimentally warmed plots within Alaskan acidic tussock tundra. Soil warming of 2-4°C during a single growing season caused strong increases in total ecosystem respiration and belowground respiration from moss-dominated inter-tussock areas, and similar trends from sedge-dominated tussocks. Consequently, the overall effect of the manipulation was to substantially enhance net ecosystem carbon loss during mid-summer. Components of vascular plant biomass were closely correlated with total ecosystem respiration and belowground respiration in control plots of both microsites, but not in warmed plots. By contrast, in the warmed inter-tussock areas, belowground respiration was most closely correlated with organic-layer depth. Warming in tussock areas was associated with increased leaf nutrient pools, indicating enhanced rates of soil nutrient mineralisation. Together, these results suggest that warming enhanced net ecosystem CO₂ efflux primarily by stimulating decomposition of soil organic matter, rather than by increasing plant-associated respiration. Our short-term experiment provides field evidence to support previous growth chamber and modelling studies indicating that arctic soil C reserves are relatively sensitive to warming and could supply an initial positive feedback to rising atmospheric CO₂ concentrations/changing climate.     

Detecting changes in arctic tundra plant communities in response to warming over decadal time scales

Detecting the response of vegetation to climate forcing as distinct from spatial and temporal variability may be difficult, if not impossible, over the typical duration of most field studies. We analyzed the spatial and interannual variability of plant functional type biomass from field studies in low arctic tussock tundra and compared these to climate change simulations of plant community composition using a dynamic tundra vegetation model (ArcVeg). Spatial heterogeneity of peak season live aboveground biomass was estimated using field samples taken from low arctic tundra at Ivotuk, Alaska (68.5°N, 155.7°W) in 1999. Coefficients of variation for live aboveground biomass at the 1 m² scale ranged from 14.6% for deciduous shrubs, 18.5% for graminoids and 25.3% for mosses to over 57% for forbs and lichens. Spatial heterogeneity in the ArcVeg dynamic vegetation model was simulated to be greater than the field data, ranging from 37.1% for deciduous shrubs to 107.9% for forbs. Disturbances in the model, such as caribou grazing and freezing–thawing of soil, as well as demographic stochasticity, led to the greater variability in the simulated results. Temporal variances of aboveground live biomass over a 19-year period using data from Toolik Lake, AK fell within the range of field and simulation spatial variances. However, simulations using ArcVeg suggest that temporal variability can be substantially less than site-scale spatial variability. Field data coupled with ArcVeg simulations of climate change scenarios indicate that some changes in plant community composition may be detectable within two decades following the onset of warming, and shrubs and mosses might be the key indicators of community change. Model simulations also project increasing landscape scale spatial heterogeneity (particularly of shrubs) with increasing temperatures.     

Competition,predation and flow rate as mediators of direct and indirect effects in a stream food chain

Using semi-natural stream channels, we estimated the effects of competition and predation exerted by juvenile and adult exotic rainbow trout (Oncorhynchus mykiss) on the diel activity pattern of juvenile native Atlantic salmon (Salmo salar), a secondary consumer. We also evaluated the direct and indirect effects of competition, predation and abiotic factors (water depth and velocity) on the growth rate of salmon, the biomass of invertebrate grazers (primary consumers) and the biomass of periphytic algae (primary producers; chlorophyll a). The presence of chemical cues emanating from adult predatory trout reduced the daily activity of juvenile Atlantic salmon. In contrast, competition imposed by juvenile rainbow trout forced Atlantic salmon to be more active during the day, even if adult rainbow trout were also present. We found no effect of either competition or of predatory cues on the growth rate of Atlantic salmon, and no evidence of indirect effects on either the biomass of invertebrates or the biomass of chlorophyll a. In contrast, we demonstrated that this food chain (fish--invertebrate grazers--periphytic algae) was under the control of a critical abiotic factor, the water velocity, and of bottom-up processes. We concluded that the exotic species directly increases the risk of predation of the native Atlantic salmon, but behavioral compensation probably limits the effects on growth rate. The competition and predation imposed by the invaders had no indirect effects on lower trophic levels. Top-down effects may have been mitigated by the dominant influence of water velocity controlling all components of the food chain and by elevated levels of primary production.     

Ecological consequences of ontogenetic shifts in predator diet: Seasonal constraint of a behaviorally mediated indirect interaction

Predators play an important role in structuring assemblages through direct and cascading indirect effects. While there has been recent interest in how the strength and direction of trophic cascades vary spatially, seasonal variability in trophic links is seldom considered. In North Carolina, recruitment-failure of bay scallops typically occurs following the spring but not the fall spawning despite the presence in each of these seasons of predatory blue crabs. One explanation for this pattern is that in the fall, seasonally abundant predators of blue crabs reduce the foraging efficiency of crabs on scallops and thus the overall magnitude of top-down effects. Quantification of bay scallop consumption by blue crabs in closed mesocosms with or without pinfish supported the hypothesis that seasonally abundant adult pinfish indirectly increase survivorship of bay scallop recruits in fall by reducing predation by blue crabs. Despite voracious consumption of bay scallops during both the day and night in mesocosms to which only small blue crabs were added, blue crabs in mesocosms with visually-foraging adult pinfish consumed bay scallops only by night. Juvenile pinfish that dominate estuarine populations in spring did not impede consumption of bay scallops by blue crabs. In mesocosms from which animals could not emigrate, the addition of neither adult nor juvenile pinfish increased the mortality of blue crabs, indicating a behaviorally mediated interaction. Blue crabs restricted by adult pinfish to nocturnal feeding did not compensate for lost feeding time by increasing their night-time consumption of bay scallops. These results strongly suggest that greater survivorship of bay scallops in fall than spring is due to adult pinfish, potential predators of small blue crabs, restricting blue crab foraging to hours of dark. In spring, when pinfish are small and incapable of consuming blue crabs, blue crabs consume bay scallops by day and by night. Such seasonal variation in the number of trophic links in a system may have important evolutionary implications. By timing reproduction to occur in fall when the pinfish-crab-scallop cascade is in operation, bay scallops maximize recruitment.     

Intraguild indirect effects through trophic cascades between stream-dwelling fishes

1. To examine the strength and role of indirect effects through trophic cascades by omnivorous fish on algal biomass in streams, we introduced one of four fish species (ayu Plecoglossus altivelis altivelis, pike gudgeon Pseudogobio esocinus esocinus, Japanese dace Tribolodon hakonensis and pale chub Zacco platypus) in experimental pools. We also investigated the indirect effects of gudgeon, dace and chub on the growth of ayu. 2. We chose the four fish species across a continuum of omnivory. Ayu fed mainly on benthic algae, and gudgeon predominantly on invertebrates. Dace and chub fed on both algae and invertebrates, but dace preyed on invertebrates more than chub. 3. The presence of gudgeon, dace and chub reduced the number of algal-feeding invertebrates and increased the algal biomass through trophic cascades. Consequently, ayu growth rate over the experimental period in pools with one of the three fish species was 25.9-44.1 times greater than the growth rate in pools with only ayu. 4. The positive indirect effect on ayu growth was large for gudgeon and dace and small for chub, whereas the addition of ayu reduced ayu growth considerably due to intraspecific competition. 5. The relative intensity of indirect effects on ayu growth through trophic cascades was predictable from the food overlap between ayu and the other fishes. However, the similar strength of indirect effects by gudgeon and dace that fed differently on algae and invertebrates suggests that feeding behaviour, prey preference and trait-mediated indirect interactions were also important in the prediction.     

Bottom-up and top-down regulation of decomposition in a tropical forest

The soil nutrients, microbes, and arthropods of tropical forests are patchy at multiple scales. We asked how these three factors interact to generate patterns of decomposition in 450 100 cm² litterbags arrayed along a 50 m ridge top in a Panama rainforest. We tested top-down (via grazing by microbivores like collembola and diplopods) and bottom-up (via added N and P) effects on the decomposition of cellulose. By using a 1,000-fold gradient in mesh size we generated a two-fold gradient in arthropod grazing. Microbivore grazing first retarded then ultimately enhanced decomposition rates. Micropulses of N and P (simulating concentrated urine) enhanced neither decomposition rates nor microbivores but increased the abundance of predacious ants. Decomposition rates also varied across the ridge, and were lowest in a plot with the deepest litter and highest soil moisture. These data generate the working hypothesis that N and P cascade upward at grains of 100 cm² to enhance a major predator in the litter; predators then absorb any increases in microbivores attracted to the extra fungal growth. These population interactions are in turn embedded in mesoscale variability generated by individual tree canopies that drive changes in litter quality and soil moisture. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.