Monoterpene chemodiversity of ponderosa pine in relation to herbivory and bark beetle colonization

Summary. Ponderosa pine, Pinus ponderosa Laws. (Pinaceae), forests in Arizona have suffered from a nine-year period of drought and bark beetle, Ips lecontei Swaine (Coleoptera: Scolytidae), outbreaks. Abiotic and biotic stress in ponderosa pine results in the induced synthesis of certain monoterpenes that may in turn affect bark beetle behavior and survival. In this study, we investigate whether induced monoterpene production could result in a different monoterpene composition that remains stored in the needles or the trunk resin of the tree. Needle and resin samples in addition to trunk cores were collected from ponderosa pines at three locations in Arizona. Ungulate browsing induced a significant increase in limonene (P=0.010) and in chemodiversity (P=0.009), a measure of the evenness of distribution among the monoterpenes present in needles. We compared the level of ‘stress’ of the trees by measuring the thickness of annual rings in living trees and those that were killed by bark beetles. Where drought occurred, the spacing of annual rings from the last 10 years of trees killed by bark beetles was significantly smaller (P=0.020) compared to living trees. There was no difference in the monoterpene composition between the core sections of closest spacing of annual rings (stressed years) compared to the sections of widest spacing, which indicates that monoterpenes are distributed evenly throughout the extended resin system. In the area where the degree of drought was less overall, none of the individual monoterpenes present in the resin was related to bark beetle killed trees. However, about half the living pines had resin in which one of the major monoterpenes (α-pinene, Δ³-carene, and limonene) was absent, and these trees had a lower monoterpene chemodiversity compared to trees killed by bark beetles. Trees with these three major monoterpenes, corresponding to the average relative proportion in living pines at that location, may sustain higher selection and colonization by bark beetles.     

Soil responses to management, increased precipitation, and added nitrogen in ponderosa pine forests.

Forest management, climatic change, and atmospheric N deposition can affect soil biogeochemistry, but their combined effects are not well understood. We examined the effects of water and N amendments and forest thinning and burning on soil N pools and fluxes in ponderosa pine forests near Flagstaff, Arizona (USA). Using a 15N-depleted fertilizer, we also documented the distribution of added N into soil N pools. Because thinning and burning can increase soil water content and N availability, we hypothesized that these changes would alleviate water and N limitation of soil processes, causing smaller responses to added N and water in the restored stand. We found little support for this hypothesis. Responses of fine root biomass, potential net N mineralization, and the soil microbial N to water and N amendments were mostly unaffected by stand management. Most of the soil processes we examined were limited by N and water, and the increased N and soil water availability caused by forest restoration was insufficient to alleviate these limitations. For example, N addition caused a larger increase in potential net nitrification in the restored stand, and at a given level of soil N availability, N addition had a larger effect on soil microbial N in the restored stand. Possibly, forest restoration increased the availability of some other limiting resource, amplifying responses to added N and water. Tracer N recoveries in roots and in the forest floor were lower in the restored stand. Natural abundance delta15N of labile soil N pools were higher in the restored stand, consistent with a more open N cycle. We conclude that thinning and burning open up the N cycle, at least in the short-term, and that these changes are amplified by enhanced precipitation and N additions. Our results suggest that thinning and burning in ponderosa pine forests will not increase their resistance to changes in soil N dynamics resulting from increased atmospheric N deposition or increased precipitation due to climatic change. Restoration plans should consider the potential impact on long-term forest productivity of greater N losses from a more open N cycle, especially during the period immediately after thinning and burning.     

Nitrogen spatial heterogeneity influences diversity following restoration in a ponderosa pine forest, Montana.

The resource heterogeneity hypothesis (RHH) is frequently cited in the ecological literature as an important mechanism for maintaining species diversity. The RHH has rarely been evaluated in the context of restoration ecology in which a commonly cited goal is to restore diversity. In this study we focused on the spatial heterogeneity of total inorganic nitrogen (TIN) following restoration treatments in a ponderosa pine (Pinus ponderosa)/Douglas-fir (Pseudotsuga menziesii) forest in western Montana, USA. Our objective was to evaluate relationships between understory species richness and TIN heterogeneity following mechanical thinning (thin-only), prescribed burning (burn-only), and mechanical thinning with prescribed burning (thin/burn) to discern the ecological and management implications of these restoration approaches. We employed a randomized block design, with three 9-ha replicates of each treatment and an untreated control. Within each treatment, we randomly established a 20 x 50 m (1000 m2) plot in which we measured species richness across the entire plot and in 12 1-m(2) quadrats randomly placed within each larger plot. Additionally, we measured TIN from a grid consisting of 112 soil samples (0-5 cm) in each plot and computed standard deviations as a measure of heterogeneity. We found a correlation between the net increase in species richness and the TIN standard deviations one and two years following restoration treatments, supporting RHH. Using nonmetric multidimensional scaling ordination and chi-squared analysis, we found that high and low TIN quadrats contained different understory communities in 2003 and 2004, further supporting RHH. A comparison of restoration treatments demonstrated that thin/burn and burn-only treatments created higher N heterogeneity relative to the control. We also found that within prescribed burn treatments, TIN heterogeneity was positively correlated with fine-fuel consumption, a variable reflecting burn severity. These findings may lead to more informed restoration decisions that consider treatment effects on understory diversity in ponderosa pine/Douglas-fir ecosystems.     

Litter and dead wood dynamics in ponderosa pine forests along a 160-year chronosequence.

Disturbances such as fire play a key role in controlling ecosystem structure. In fire-prone forests, organic detritus comprises a large pool of carbon and can control the frequency and intensity of fire. The ponderosa pine forests of the Colorado Front Range, USA, where fire has been suppressed for a century, provide an ideal system for studying the long-term dynamics of detrital pools. Our objectives were (1) to quantify the long-term temporal dynamics of detrital pools; and (2) to determine to what extent present stand structure, topography, and soils constrain these dynamics. We collected data on downed dead wood, litter, duff (partially decomposed litter on the forest floor), stand structure, topographic position, and soils for 31 sites along a 160-year chronosequence. We developed a compartment model and parameterized it to describe the temporal trends in the detrital pools. We then developed four sets of statistical models, quantifying the hypothesized relationship between pool size and (1) stand structure, (2) topography, (3) soils variables, and (4) time since fire. We contrasted how much support each hypothesis had in the data using Akaike's Information Criterion (AIC). Time since fire explained 39-80% of the variability in dead wood of different size classes. Pool size increased to a peak as material killed by the fire fell, then decomposed rapidly to a minimum (61-85 years after fire for the different pools). It then increased, presumably as new detritus was produced by the regenerating stand. Litter was most strongly related to canopy cover (r2 = 77%), suggesting that litter fall, rather than decomposition, controls its dynamics. The temporal dynamics of duff were the hardest to predict. Detrital pool sizes were more strongly related to time since fire than to environmental variables. Woody debris peak-to-minimum time was 46-67 years, overlapping the range of historical fire return intervals (1 to > 100 years). Fires may therefore have burned under a wide range of fuel conditions, supporting the hypothesis that this region's fire regime was mixed severity.     

Frequent fire alters nitrogen transformations in ponderosa pine stands of the inland northwest

Recurrent, low-severity fire in ponderosa pine (Pinus ponderosa)/interior Douglas-fir (Pseudotsuga menziesii var. glauca) forests is thought to have directly influenced nitrogen (N) cycling and availability. However, no studies to date have investigated the influence of natural fire intervals on soil processes in undisturbed forests, thereby limiting our ability to understand ecological processes and successional dynamics in this important ecosystem of the Rocky Mountain West. Here, we tested the standing hypothesis that recurrent fire in ponderosa pine/Douglas-fir forests of the Inland Northwest decreases total soil N, but increases N turnover and nutrient availability. We compared soils in stands unburned over the past 69-130 years vs. stands exposed to two or more fires over the last 130 years at seven distinct locations in two wilderness areas. Mineral soil samples were collected from each of the seven sites in June and July of 2003 and analyzed for pH, total C and N, potentially mineralizable N (PMN), and extractable NH4+, NO3-, PO4(-3), Ca+2, Mg+2, and K+. Nitrogen transformations were assessed at five sites by installing ionic resin capsules in the mineral soil in August of 2003 and by conducting laboratory assays of nitrification potential and net nitrification in aerobic incubations. Total N and PMN decreased in stands subjected to multiple fires. This loss of total N and labile N was not reflected in concentrations of extractable NH4+ and NO3-. Rather, multiple fires caused an increase in NO3 sorbed on ionic resins, nitrification potential, and net nitrification in spite of the burned stands not having been exposed to fire for at least 12-17 years. Charcoal collected from a recent fire site and added to unburned soils increased nitrification potential, suggesting that the decrease of charcoal in the absence of fire may play an important role in N transformations in fire-dependent ecosystems in the long term. Interestingly, we found no consistent effect of fire frequency on extractable P or alkaline metal concentrations. Our results corroborate the largely untested hypothesis that frequent fire in ponderosa pine forests increases inorganic N availability in the long term and emphasize the need to study natural, unmanaged sites in far greater detail.     

Making a stand: five centuries of population growth in colonizing populations of Pinus ponderosa

The processes underlying the development of new populations are important for understanding how species colonize new territory and form viable long-term populations. Life-history-mediated processes such as Allee effects and dispersal capability may interact with climate variability and site-specific factors to govern population success and failure over extended time frames. We studied four disjunct populations of ponderosa pine in the Bighorn Basin of north-central Wyoming to examine population growth spanning more than five centuries. The study populations are separated from continuous ponderosa pine forest by distances ranging from 15 to >100 km. Strong evidence indicates that the initial colonizing individuals are still present, yielding a nearly complete record of population history. All trees in each population were aged using dendroecological techniques. The populations were all founded between 1530 and 1655 cal yr CE. All show logistic growth patterns, with initial exponential growth followed by a slowing during the mid to late 20th century. Initial population growth was slower than expectations from a logistic regression model at all four populations, but increased during the mid-18th century. Initial lags in population growth may have been due to strong Allee effects. A combination of overcoming Allee effects and a transition to favorable climate conditions may have facilitated a mid-18th century pulse in population growth rate.     

Community occupancy responses of small mammals to restoration treatments in ponderosa pine forests, northern Arizona, USA

In western North American conifer forests, wildfires are increasing in frequency and severity due to heavy fuel loads that have accumulated after a century of fire suppression. Forest restoration treatments (e.g., thinning and/or burning) are being designed and implemented at large spatial and temporal scales in an effort to reduce fire risk and restore forest structure and function. In ponderosa pine (Pinus ponderosa) forests, predominantly open forest structure and a frequent, low-severity fire regime constituted the evolutionary environment for wildlife that persisted for thousands of years. Small mammals are important in forest ecosystems as prey and in affecting primary production and decomposition. During 2006-2009, we trapped eight species of small mammals at 294 sites in northern Arizona and used occupancy modeling to determine community responses to thinning and habitat features. The most important covariates in predicting small mammal occupancy were understory vegetation cover, large snags, and treatment. Our analysis identified two generalist species found at relatively high occupancy rates across all sites, four open-forest species that responded positively to treatment, and two dense-forest species that responded negatively to treatment unless specific habitat features were retained. Our results indicate that all eight small mammal species can benefit from restoration treatments, particularly if aspects of their evolutionary environment (e.g., large trees, snags, woody debris) are restored. The occupancy modeling approach we used resulted in precise species-level estimates of occupancy in response to habitat attributes for a greater number of small mammal species than in other comparable studies. We recommend our approach for other studies faced with high variability and broad spatial and temporal scales in assessing impacts of treatments or habitat alteration on wildlife species. Moreover, since forest planning efforts are increasingly focusing on progressively larger treatment implementation, better and more efficiently obtained ecological information is needed to inform these efforts.     

鼎湖山马尾松林凋落物分解对凋落物输入变化的响应

为了了解我国南方森林常见的人为干扰（凋落物收取）活动对生态系统养分循环的影响,研究了鼎湖山马尾松林3种主要树种凋落物分解及其养分释放对凋落物输入量变化的响应。这3种树种分别为马尾松（Pinus massoniana）、荷木（Schimasuperba）和锥栗（Castanopsis chinensis）。凋落物输入量变化分别为凋落物去除（L-）、加倍（L＋）和对照（L）3种处理,每种处理25个重复。经过18个月的处理试验,凋落物分解速率及其养分释放随树种、分解阶段和凋落物处理不同而异。荷木、马尾松和锥栗分解物平均残留率分别为0.46±0.01、0.42±0.01、0.40±0.02,其中,荷木与锥栗、马尾松差异性显著。不同处理间的凋落物分解速率差异显著,加倍、对照和去除处理样地凋落物的平均残留率分别为0.51±0.08、0.53±0.09和0.55±0.08。凋落物加倍处理促进了凋落物分解过程中C的释放,而去除凋落物处理则抑制了N、P的释放。以上结果表明,凋落物收取活动不仅直接带走凋落物中的大量养分,而且抑制了凋落物分解及其养分释放。     

Drought induced changes in growth, leaf gas exchange and biomass production in Albizia lebbeck and Cassia siamea seedlings

Diurnal trends in net photosynthesis rate (P(N)), stomatal conductance (g(s)), water use efficiency (WUE) and biomass were compared in six-month-old seedlings of Albizia lebbeck and Cassia siamea, under different levels of drought stress. The potted plants were subjected to four varying drought treatment by withholding watering for 7 (D1), 14(D2) and 25 (D3) days. The fourth group (C) was watered daily and treated as unstressed (control). Species differed significantly (p < 0.001) in their physiological performance under varying stress conditions. Higher P(N) of 11.6 +/- 0.05 in control followed by 4.35 +/- 0.4 in D1 and 2.83 +/- 0.18 micromol m(-2) s(-1) in D2 was observed in A. lebbeck. A significant (p < 0.001) reduction in P(N) was observed in C. siamea (C 7.65 +/- 0.5 micromol m(-2) s(-1), D1, 2.56 +/- 0.33 micromol m(-2) s(-1) and D2, 1.4 +/- 0.01 micromol m(-2) s(-1)) at 9 hr. A positive correlation was seen between P(N) and g(s) (A. lebbeck, r2 = 0.84; C. siamea, r2 = 0.82). Higher WUE was observed in C. siamea (D2, 7.1 +/- 0.18 micromol m(-2) s(-1); D3, 8.39 +/- 0.11 micromol m(-2) s(-1)) than A. lebbeck, (control, 7.58 +/- 0.3 micromol m(-2) s(-1) and D3, 8.12 +/- 0.15 micromol m(-2) s(-1)). The chlorophyll and relative water content (RWC) was more in A. lebbeck than C. siamea. Maximum biomass was produced by A. lebbeckthan C. siamea. From the study, one could conclude that A. lebbeckis better than C. siamea in adopting suitable resource management strategy and be best suited for the plantation programs in the semi-arid dry lands.     

Assessing the past and future distribution and productivity of ponderosa pine in the Pacific Northwest using a process model, 3-PG

Climate variability at decadal scales influences not only the growth of widely distributed species such as Pinus ponderosa, but also can have an effect on the timing and severity of fire and insect outbreaks that may alter species distributions. In this paper, we present a spatial modelling technique to assess the influence of climatic variability on the annual productivity of P. ponderosa in the Pacific Northwest (PNW) of North America over the past 100 years and infer how a sustained change in climate might alter the geographic distribution of this species across defined ecotones. Field observations were used to establish criteria for P. ponderosa dominance including: (1) maximum summer leaf area index (LAI), ranges between 1.5 and 2.5; (2) 80% of available soil water is depleted during summer months; and (3) soil water will return to full capacity at least once during the year. Where these three criteria were not met, eventual replacement of P. ponderosa would be predicted. We utilized a simple physiological model, Physiological Principles for Predicting Growth (3-PG) to predict annual variation in LAI from climatic data provided by the Oregon Climate Service over the period from 1900 to 2000 and from broad scale 0.5°-spatial resolution future climate projections produced by the Hadley Climate Center, UK. From these simulations we produced a series of maps that display predicted shifts of zones where ponderosa pine might be expected to contract or expand its range if modeled climatic conditions at annual and decadal intervals were sustained. From the historical simulations, the most favorable year for pine dominance was 1958 and the least favorable, 1924. The most favorable decade was in the 1900s and the least favorable in the 1930s. The future predictions indicate a reduction in the current range of the P. ponderosa type along the western Cascade Range however, an increase along the east side and inland PNW. The model predicts that pine dominance should increase between 5 and 10% over the next century, mainly in inland Oregon, Idaho, and Washington.     

The plankton tower. II. Release of nutrients from sediments due to changes in the density of bottom water

Results of an enclosure experiment carried out in Kiel Bight are presented. A water column of about 30 m³, extending from the surface to the bottom over a 3 m² patch of sediment (coarse sand), was isolated and observed over a period of 33 days. Considerable water exchange took place with the surroundings partly because of near-surface openings that appeared in the enclosure but mainly because of density changes due to salinity fluctuations in the surrounding water. This denser water entered the enclosure through the sediments, displacing the lighter water through the near-surface openings. Very high nutrient and low oxygen values were measured in the bottom water immediately following this higher-density water influx. The same effect was observed outside the enclosure, but this phenomenon was not marked here, probably due to greater turbulent mixing obscuring this effect. It is postulated that interstitial, water is flushed out of coarse-grained sediments by gravity displacement due to changes in the density of bottom water. In certain areas this mechanism of nutrient release from the sediments is presumably of great ecological importance, both for the phytoplankton and the benthos.Publication no. 83 of the Joint Research Program at Kiel University (Sonderforschungsbereich 95 der Deutschen Forschungsgemeinschaft).     

A century of increasing pine density and associated shifts in understory plant strategies

We analyzed one of the longest-term ecological data sets to evaluate how forest overstory structure is related to herbaceous understory plant strategies in a ponderosa pine forest. Eighty-two permanent 1-m2 chart quadrats that were established as early as 1912 were remeasured in 2007. We reconstructed historical forest structure using dendrochronological techniques. Ponderosa pine basal area increased from an average of 4 m2/ha in the early 1900s to 29 m2/ha in 2007. Understory plant foliar cover declined by 21%, species richness declined by two species per square meter, and functional diversity also declined. The relative cover of C4 graminoids decreased by 18% and C3 graminoids increased by 19%. Herbaceous plant species with low leaf and fine root nitrogen concentrations, low specific leaf area, high leaf dry matter content, large seed mass, low specific root length, short maximum height, and early flowering date increased in relative abundance in sites where pine basal area increased the most. Overall, we observed a long-term shift in composition toward more conservative shade- and stress-tolerant herbaceous species. Our analysis of temporal changes in plant strategies provides a general framework for evaluating compositional and functional changes in terrestrial plant communities.     

Vole response to unintentional changes in the chemistry of GM poplars

Summary.  There is an increased interest for the use of GM trees in forestry and several commercially promising lines are now available. However, the ecological implications of the use of GM trees, e.g. effects on non-target natural enemies, have rarely been explored. The aim of this study was to determine if modification of non-defensive traits in GM poplars unintentionally can influence plant chemistry in a way that has consequences for palatability to voles. In a greenhouse experiment, we used two lines, SPS33A and SPS26, of GM hybrid poplars (Populus tremula x tremuloides) with 1.5 and 4 times, respectively, over-expression sucrose-phosphate synthase (SPS). This enzyme plays a central role in sucrose synthesis, affecting cold acclimation, mesophyll sucrose content and biomass production. As a control we used the isogenic unmodified wild type. Stems of these poplars were presented to bank voles and field voles in cafeteria experiments. The concentration of condensed tannins was higher in leaves of lines SPS33A and SPS26 than in the isogenic wild-type and the concentration of nitrogen was higher in line SPS33A than in both the wild-type and line SPS26. Although the bank voles consumed slightly less bark from SPS33A, there were no significant differences in the preference of bank vole or field vole for the different poplar lines. This indicates that the changes in plant chemistry were insufficient to produce any strong herbivore response or that alteration in tannins and nitrogen counteracted each other. Still, changes in the interactions between mammalian herbivores and GM trees are important to consider in future cost-benefits analyses of GM trees.     

Prediction of response of zooplankton biomass to climatic and oceanic changes

This paper examines the long-term variation in zooplankton biomass in response to climatic and oceanic changes, using a neural network as a nonlinear multivariate analysis method. Zooplankton data collected from 1951 to 1990 off the shore of northeastern Japan were analyzed. We considered patterns of the Kuroshio and the Oyashio, sea surface temperature, and meteorological parameters as environmental factors that affect zooplankton biomass. Back propagation neural networks were trained to generate mapping functions between environmental variables and zooplankton biomass. The performance of the network models was tested by varying the numbers of input and hidden units. Changes in zooplankton biomass could be predicted from environmental conditions. The neural network yielded predictions with smaller errors than those of predictions determined by linear multiple regression. The sensitivity analysis of networks was used to extract predictive knowledge. The air pressure, sea surface temperature, and some indices of atmospheric circulation were the primary factors for predictions. The patterns of the Kuroshio and the Oyashio demonstrated different effects among sea areas.     

Modeling seasonal course of carbon fluxes and evapotranspiration in response to low temperature and moisture in a boreal Scots pine ecosystem

Environmental conditions act above and below ground, and regulate carbon fluxes and evapotranspiration. The productivity of boreal forest ecosystems is strongly governed by low temperature and moisture conditions, but the understanding of various feedbacks between vegetation and environmental conditions is still unclear. In order to quantify the seasonal responses of vegetation to environmental factors, the seasonality of carbon and heat fluxes and the corresponding responses for temperature and moisture in air and soil were simulated by merging a process-based model (CoupModel) with detailed measurements representing various components of a forest ecosystem in Hyytiälä, southern Finland. The uncertainties in parameters, model assumptions, and measurements were identified by generalized likelihood uncertainty estimation (GLUE). Seasonal and diurnal courses of sensible and latent heat fluxes and net ecosystem exchange (NEE) of CO₂ were successfully simulated for two contrasting years. Moreover, systematic increases in efficiency of photosynthesis, water uptake, and decomposition occurred from spring to summer, demonstrating the strong coupling between processes. Evapotranspiration and NEE flux both showed a strong response to soil temperature conditions via different direct and indirect ecosystem mechanisms. The rate of photosynthesis was strongly correlated with the corresponding water uptake response and the light use efficiency. With the present data and model assumptions, it was not possible to precisely distinguish the various regulating ecosystem mechanisms. Our approach proved robust for modeling the seasonal course of carbon fluxes and evapotranspiration by combining different independent measurements. It will be highly interesting to continue using long-term series data and to make additional tests of optional stomatal conductance models in order to improve our understanding of the boreal forest ecosystem in response to climate variability and environmental conditions.     

Ovarian changes in response to heavy metal exposure to the fish,Notopterus notopterus (Pallas)

The ovarian and hepatic protein, lipid and cholesterol content were estimated in the fish, N. notopterus after exposing it to heavy metals at sublethal concentrations. The protein, lipid and cholesterol content of ovary and liver got reduced and amongst the three exposures (mercuric chloride, cadmium chloride and their combination) it was significantly reduced in the order of (HgCl2>HgCl2>+CdCl2>CdCl2>control). The above results indicated that HgCl2 is highly toxic and its toxicity gets reduced in combination.     

Skeletal changes in the test and jaws of the sea urchinDiadema antillarum in response to food limitation

When food-limited,Diadema antillarum increases the relative size of Aristotle's lantern compared to the size of the test. This is accomplished primarily by decreasing the size of the test: the demipyramid grows at a reduced rate as the test shrinks. These results suggest that an altered relationship between demipyramid and test size can provide evidence for food-limitation in field populations. The sizes of the lantern and test of museum and fossil specimens can be investigated for the presence of food limitation and biotic interactions in recent and past times.     

Adaptive behaviour of chironomid larvae (Chironomus riparius) in response to chemical stimuli from predators and resource density

Chemical-mediated effects of predatory fish on chironomid larvae behaviour have been ignored so far. Sediment-dwelling chironomid larvae inhabit protective burrows from which they extend their bodies only to feed on deposited detritus and microalgae from the surrounding sediment. Here, we performed factorial laboratory experiments to study whether fish-borne chemical cues (kairomones) are responsible for behavioural trait changes of chironomid larvae, and whether chironomid larvae are able to assess the densities of fish predators and food resources and the trade-off between them. We exposed naïve Chironomus riparius larvae to the chemical presence of zero, one, and ten predator fish (Rutilus rutilus) and offered two resource levels (low food, high food) for each treatment. Kairomones induced significant inherent behavioural trait changes in chironomid larvae. During the first 120 min after exposing chironomids to fish-conditioned water, we found a significant increase in digging activity with increasing predator density. After 3 days of exposure, the deepest chironomid burrows were found in treatments with the highest predator density. Chironomid larvae were significantly able to adjust their foraging behaviour to different predator densities and food concentrations and trade off between them; that is, when fish predators were more abundant or when more food resources were available, the foraging activities of larvae were significantly reduced. Our data suggest that chemically mediated trait changes (burrowing and foraging behaviour) may cascade through the littoral food web.