Automated escape system: identifying prey's kinematic and behavioral features critical for predator evasion.

Identifying the kinematic and behavioral variables of prey that influence evasion from predator attacks remains challenging. To address this challenge, we have developed an automated escape system that responds quickly to an approaching predator and pulls the prey away from the predator rapidly, similar to real prey. Reaction distance, response latency, escape speed and other variables can be adjusted in the system. By repeatedly measuring the response latency and escape speed of the system, we demonstrated the system's ability to exhibit fast and rapid responses while maintaining consistency across successive trials. Using the live predatory fish species Coreoperca kawamebari, we show that escape speed and reaction distance significantly affect the outcome of predator-prey interactions. These findings indicate that the developed escape system is useful for identifying kinematic and behavioral features of prey that are critical for predator evasion, as well as for measuring the performance of predators.

When things get MESI: The Manipulation Experiments Synthesis Initiative-A coordinated effort to synthesize terrestrial global change experiments.

Responses of the terrestrial biosphere to rapidly changing environmental conditions are a major source of uncertainty in climate projections. In an effort to reduce this uncertainty, a wide range of global change experiments have been conducted that mimic future conditions in terrestrial ecosystems, manipulating CO2 , temperature, and nutrient and water availability. Syntheses of results across experiments provide a more general sense of ecosystem responses to global change, and help to discern the influence of background conditions such as climate and vegetation type in determining global change responses. Several independent syntheses of published data have yielded distinct databases for specific objectives. Such parallel, uncoordinated initiatives carry the risk of producing redundant data collection efforts and have led to contrasting outcomes without clarifying the underlying reason for divergence. These problems could be avoided by creating a publicly available, updatable, curated database. Here, we report on a global effort to collect and curate 57,089 treatment responses across 3644 manipulation experiments at 1145 sites, simulating elevated CO2 , warming, nutrient addition, and precipitation changes. In the resulting Manipulation Experiments Synthesis Initiative (MESI) database, effects of experimental global change drivers on carbon and nutrient cycles are included, as well as ancillary data such as background climate, vegetation type, treatment magnitude, duration, and, unique to our database, measured soil properties. Our analysis of the database indicates that most experiments are short term (one or few growing seasons), conducted in the USA, Europe, or China, and that the most abundantly reported variable is aboveground biomass. We provide the most comprehensive multifactor global change database to date, enabling the research community to tackle open research questions, vital to global policymaking. The MESI database, freely accessible at doi.org/10.5281/zenodo.7153253, opens new avenues for model evaluation and synthesis-based understanding of how global change affects terrestrial biomes. We welcome contributions to the database on GitHub.

Ambient precipitation determines the sensitivity of soil respiration to precipitation treatments in a marsh.

The effects in field manipulation experiments are strongly influenced by amplified interannual variation in ambient climate as the experimental duration increases. Soil respiration (SR), as an important part of the carbon cycle in terrestrial ecosystems, is sensitive to climate changes such as temperature and precipitation changes. A growing body of evidence has indicated that ambient climate affects the temperature sensitivity of SR, which benchmarks the strength of terrestrial soil carbon-climate feedbacks. However, whether SR sensitivity to precipitation changes is influenced by ambient climate is still not clear. In addition, the mechanism driving the above phenomenon is still poorly understood. Here, a long-term field manipulation experiment with five precipitation treatments (-60%, -40%, +0%, +40%, and +60% of annual precipitation) was conducted in a marsh in the Yellow River Delta, China, which is sensitive to soil drying-wetting cycle caused by precipitation changes. Results showed that SR increased exponentially along the experimental precipitation gradient each year and the sensitivity of SR (standardized by per 100 mm change in precipitation under precipitation treatments) exhibited significant interannual variation from 2016 to 2021. In addition, temperature, net radiation, and ambient precipitation all exhibited dramatic interannual variability; however, only ambient precipitation had a significant negative correlation with SR sensitivity. Moreover, the sensitivity of SR was significantly positively related to the sensitivity of belowground biomass (BGB) across 6 years. Structural equation modeling and regression analysis also showed that precipitation treatments significantly affected SR and its autotrophic and heterotrophic components by altering BGB. Our study demonstrated that ambient precipitation determines the sensitivity of SR to precipitation treatments in marshes. The findings underscore the importance of ambient climate in regulating ecosystem responses in long-term field manipulation experiments.

Lowering water table reduces carbon sink strength and carbon stocks in northern peatlands.

Peatlands at high latitudes have accumulated >400 Pg carbon (C) because saturated soil and cold temperatures suppress C decomposition. This substantial amount of C in Arctic and Boreal peatlands is potentially subject to increased decomposition if the water table (WT) decreases due to climate change, including permafrost thaw-related drying. Here, we optimize a version of the Organizing Carbon and Hydrology In Dynamic Ecosystems model (ORCHIDEE-PCH4) using site-specific observations to investigate changes in CO2 and CH4 fluxes as well as C stock responses to an experimentally manipulated decrease of WT at six northern peatlands. The unmanipulated control peatlands, with the WT <20 cm on average (seasonal max up to 45 cm) below the surface, currently act as C sinks in most years (58 ± 34 g C m-2  year-1 ; including 6 ± 7 g C-CH4 m-2  year-1 emission). We found, however, that lowering the WT by 10 cm reduced the CO2 sink by 13 ± 15 g C m-2  year-1 and decreased CH4 emission by 4 ± 4 g CH4 m-2  year-1 , thus accumulating less C over 100 years (0.2 ± 0.2 kg C m-2 ). Yet, the reduced emission of CH4 , which has a larger greenhouse warming potential, resulted in a net decrease in greenhouse gas balance by 310 ± 360 g CO2-eq  m-2  year-1 . Peatlands with the initial WT close to the soil surface were more vulnerable to C loss: Non-permafrost peatlands lost >2 kg C m-2 over 100 years when WT is lowered by 50 cm, while permafrost peatlands temporally switched from C sinks to sources. These results highlight that reductions in C storage capacity in response to drying of northern peatlands are offset in part by reduced CH4 emissions, thus slightly reducing the positive carbon climate feedbacks of peatlands under a warmer and drier future climate scenario.

Rainfall manipulation experiments as simulated by terrestrial biosphere models: Where do we stand?

Changes in rainfall amounts and patterns have been observed and are expected to continue in the near future with potentially significant ecological and societal consequences. Modelling vegetation responses to changes in rainfall is thus crucial to project water and carbon cycles in the future. In this study, we present the results of a new model-data intercomparison project, where we tested the ability of 10 terrestrial biosphere models to reproduce the observed sensitivity of ecosystem productivity to rainfall changes at 10 sites across the globe, in nine of which, rainfall exclusion and/or irrigation experiments had been performed. The key results are as follows: (a) Inter-model variation is generally large and model agreement varies with timescales. In severely water-limited sites, models only agree on the interannual variability of evapotranspiration and to a smaller extent on gross primary productivity. In more mesic sites, model agreement for both water and carbon fluxes is typically higher on fine (daily-monthly) timescales and reduces on longer (seasonal-annual) scales. (b) Models on average overestimate the relationship between ecosystem productivity and mean rainfall amounts across sites (in space) and have a low capacity in reproducing the temporal (interannual) sensitivity of vegetation productivity to annual rainfall at a given site, even though observation uncertainty is comparable to inter-model variability. (c) Most models reproduced the sign of the observed patterns in productivity changes in rainfall manipulation experiments but had a low capacity in reproducing the observed magnitude of productivity changes. Models better reproduced the observed productivity responses due to rainfall exclusion than addition. (d) All models attribute ecosystem productivity changes to the intensity of vegetation stress and peak leaf area, whereas the impact of the change in growing season length is negligible. The relative contribution of the peak leaf area and vegetation stress intensity was highly variable among models.

Growth rates of marine prokaryotes are extremely diverse, even among closely related taxa.

Marine prokaryotes play crucial roles in ocean biogeochemical cycles, being their contribution strongly influenced by their growth rates. Hence, elucidating the variability and phylogenetic imprint of marine prokaryotes' growth rates are crucial for better determining the role of individual taxa in biogeochemical cycles. Here, we estimated prokaryotic growth rates at high phylogenetic resolution in manipulation experiments using water from the northwestern Mediterranean Sea. Experiments were run in the four seasons with different treatments that reduced growth limiting factors: predators, nutrient availability, viruses, and light. Single-amplicon sequence variants (ASVs)-based growth rates were calculated from changes in estimated absolute abundances using total prokaryotic abundance and the proportion of each individual ASV. The trends obtained for growth rates in the different experiments were consistent with other estimates based on total cell-counts, catalyzed reporter deposition fluorescence in situ hybridization subcommunity cell-counts or metagenomic-operational taxonomic units (OTUs). Our calculations unveil a broad range of growth rates (0.3-10 d-1) with significant variability even within closely related ASVs. Likewise, the impact of growth limiting factors changed over the year for individual ASVs. High numbers of responsive ASVs were shared between winter and spring seasons, as well as throughout the year in the treatments with reduced nutrient limitation and viral pressure. The most responsive ASVs were rare in the in situ communities, comprising a large pool of taxa with the potential to rapidly respond to environmental changes. Essentially, our results highlight the lack of phylogenetic coherence in the range of growth rates observed, and differential responses to the various limiting factors, even for closely related taxa.

Nitrogen and potassium limit fine root growth in a humid Afrotropical forest.

Nutrient limitations play a key regulatory role in plant growth, thereby affecting ecosystem productivity and carbon uptake. Experimental observations identifying the most limiting nutrients are lacking, particularly in Afrotropical forests. We conducted an ecosystem-scale, full factorial nitrogen (N)-phosphorus (P)-potassium (K) addition experiment consisting 32 40 × 40 m plots (eight treatments × four replicates) in Uganda to investigate which (if any) nutrient limits fine root growth. After two years of observations, added N rapidly decreased fine root biomass by up to 36% in the first and second years of the experiment. Added K decreased fine root biomass by 27% and fine root production by 30% in the second year. These rapid reductions in fine root growth highlight a scaled-back carbon investment in the costly maintenance of large fine root network as N and K limitations become alleviated. No fine root growth response to P addition was observed. Fine root turnover rate was not significantly affected by nutrient additions but tended to be higher in N added than non-N added treatments. These results suggest that N and K availability may restrict the ecosystem's capacity for CO2 assimilation, with implications for ecosystem productivity and resilience to climate change.

Microbial impact on initial soil formation in arid and semiarid environments under simulated climate change.

The microbiota is attributed to be important for initial soil formation under extreme climate conditions, but experimental evidence for its relevance is scarce. To fill this gap, we investigated the impact of in situ microbial communities and their interrelationship with biocrust and plants compared to abiotic controls on soil formation in initial arid and semiarid soils. Additionally, we assessed the response of bacterial communities to climate change. Topsoil and subsoil samples from arid and semiarid sites in the Chilean Coastal Cordillera were incubated for 16 weeks under diurnal temperature and moisture variations to simulate humid climate conditions as part of a climate change scenario. Our findings indicate that microorganism-plant interaction intensified aggregate formation and stabilized soil structure, facilitating initial soil formation. Interestingly, microorganisms alone or in conjunction with biocrust showed no discernible patterns compared to abiotic controls, potentially due to water-masking effects. Arid soils displayed reduced bacterial diversity and developed a new community structure dominated by Proteobacteria, Actinobacteriota, and Planctomycetota, while semiarid soils maintained a consistently dominant community of Acidobacteriota and Proteobacteria. This highlighted a sensitive and specialized bacterial community in arid soils, while semiarid soils exhibited a more complex and stable community. We conclude that microorganism-plant interaction has measurable impacts on initial soil formation in arid and semiarid regions on short time scales under climate change. Additionally, we propose that soil and climate legacies are decisive for the present soil microbial community structure and interactions, future soil development, and microbial responses.

Woody plant growth increases with precipitation intensity in a cold semiarid system.

As the atmosphere warms, precipitation events become larger, but less frequent. Yet, there is fundamental disagreement about how increased precipitation intensity will affect vegetation. Walter's two-layer hypothesis and experiments testing it have demonstrated that precipitation intensity can increase woody plant growth. Observational studies have found the opposite pattern. Not only are the patterns contradictory, but inference is largely limited to grasslands and savannas. We tested the effects of increased precipitation intensity in a shrub-steppe ecosystem that receives >30% of its precipitation as snow. We used 11 (8 × 8 m) shelters to collect and redeposit rain and snow as larger, more intense events. Total annual precipitation was the same in all plots, but each plot received different precipitation event sizes ranging from 1 to 18 mm. Over three growing seasons, larger precipitation event sizes increased soil water availability, sagebrush (Artemisia tridentata) stem radius, and canopy greenness, decreased new root growth in shallow soils, and had no effect on herbaceous plant cover. Thus, we found that increased precipitation intensity can increase soil water availability and woody plant growth in a cold semiarid system. Assuming that stem growth is positively correlated with shrub reproduction, establishment and spread, results suggest that increasing precipitation intensity may have contributed to the woody plant encroachment observed around the world in the past 50 yr. Further, continuing increases in precipitation intensity caused by atmospheric warming are likely to continue to contribute to shrub encroachment in the future.

Lineage-Specific Growth Curves Document Large Differences in Response of Individual Groups of Marine Bacteria to the Top-Down and Bottom-Up Controls.

Marine bacterioplankton represent a diverse assembly of species differing largely in their abundance, physiology, metabolic activity, and role in microbial food webs. To analyze their sensitivity to bottom-up and top-down controls, we performed a manipulation experiment where grazers were removed, with or without the addition of phosphate. Using amplicon-reads normalization by internal standard (ARNIS), we reconstructed growth curves for almost 300 individual phylotypes. Grazer removal caused a rapid growth of most bacterial groups, which grew at rates of 0.6 to 3.5 day-1, with the highest rates (>4 day-1) recorded among Rhodobacteraceae, Oceanospirillales, Alteromonadaceae, and Arcobacteraceae. Based on their growth response, the phylotypes were divided into three basic groups. Most of the phylotypes responded positively to both grazer removal as well as phosphate addition. The second group (containing, e.g., Rhodobacterales and Rhizobiales) responded to the grazer removal but not to the phosphate addition. Finally, some clades, such as SAR11 and Flavobacteriaceae, responded only to phosphate amendment but not to grazer removal. Our results show large differences in bacterial responses to experimental manipulations at the phylotype level and document different life strategies of marine bacterioplankton. In addition, growth curves of 130 phylogroups of aerobic anoxygenic phototrophs were reconstructed based on changes of the functional pufM gene. The use of functional genes together with rRNA genes may significantly expand the scientific potential of the ARNIS technique. IMPORTANCE Growth is one of the main manifestations of life. It is assumed generally that bacterial growth is constrained mostly by nutrient availability (bottom-up control) and grazing (top-down control). Since marine bacteria represent a very diverse assembly of species with different metabolic properties, their growth characteristics also largely differ accordingly. Currently, the growth of marine microorganisms is typically evaluated using microscopy in combination with fluorescence in situ hybridization (FISH). However, these laborious techniques are limited in their throughput and taxonomical resolution. Therefore, we combined a classical manipulation experiment with next-generation sequencing to resolve the growth dynamics of almost 300 bacterial phylogroups in the coastal Adriatic Sea. The analysis documented that most of the phylogroups responded positively to both grazer removal and phosphate addition. We observed significant differences in growth kinetics among closely related species, which could not be distinguished by the classical FISH technique.

Tree Growth Under Climate Change: Evidence From Xylogenesis Timings and Kinetics.

Tree growth is one of the most studied aspects of tree biology, particularly secondary growth. In the Mediterranean region, cambial activity is mostly determined by water availability. Climatic projections for the Mediterranean region predict more frequent and intense droughts, and longer periods without precipitation. To investigate tree growth under the predicted scenarios of climate change, a water manipulation experiment was conducted in a maritime pine stand (Pinus pinaster Aiton). In 2017, fifteen trees were divided into three groups: control, rain exclusion, and irrigation. Drought conditions were simulated by installing a continuous plastic sheet on the forest floor from March to September. Trees under irrigation treatment were watered twice a week in September. Cambial activity and xylem formation was monitored every 10 days from February 2017 until March 2018. Cell production was maximal around the spring equinox in all treatments. Trees under rain exclusion decreased cell production rates, xylogenesis duration, and latewood cell wall thickness. The extra irrigation in September did not produce noticeable differences in xylogenesis compared to trees in the control treatment. The synchronization of maximum cambial division rates around the vernal equinox (spring) could allow Mediterranean trees to mitigate the impact of summer drought. With the predicted increase in drought intensity and frequency, lower tree productivity, carbon sequestration, and wood biomass are expected.

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Changes in snowpack induced by climate change can profoundly affect forest litter decomposition. A snow depth manipulation experiment with three treatments (i.e.,control,snow addition, and snow removal) was conducted to assess the effects of snow depth changes on leaf litter decomposition of two temperate tree species [Manchurian ash (Fraxinus mandshurica) and Dahurian larch (Larix gmelinii)]. The annual loss of the litter mass after one year decomposition varied between 51.3% and 57.4% for the ash and between 21.7% and 31.4% for the larch. The decomposition constants (k) ranged from 0.048 to 0.057 and from 0.022 to 0.030 for these two species respectively.The greatest k value occurred under the snow addition treatment, while the least occurred under the snow removal treatment.Snow addition treatment shortened the 50% and 95% decomposition time by 1.1 months and 4.2 months for the ash, respectively, and by 3.7 months and 15.5 months for the larch, respectively. The snow removal treatment lengthened those decomposition time by 1.8 months and 6.4 months for the ash,and by 5.0 months and 21.1 months for the larch, respectively. Litter decomposition rate was significantly correlated with tree species, snow depth, decomposition time, and soil temperature, but its major influencing factors varied with decomposition stage. Soil temperature and the initial litter quality were the major factors affecting decomposition rates during the snow covered and following snow free periods, respectively. Our findings highlight that changes in snow depth exert significantly instantaneous and prolonged effects on forest litter decomposition.

Distinct seasonal dynamics of responses to elevated CO2 in two understorey grass species differing in shade-tolerance.

Understorey plant communities are crucial to maintain species diversity and ecosystem processes including nutrient cycling and regeneration of overstorey trees. Most studies exploring effects of elevated CO2 concentration ([CO2]) in forests have, however, been done on overstorey trees, while understorey communities received only limited attention.The hypothesis that understorey grass species differ in shade-tolerance and development dynamics, and temporally exploit different niches under elevated [CO2], was tested during the fourth year of [CO2] treatment. We assumed stimulated carbon gain by elevated [CO2] even at low light conditions in strongly shade-tolerant Luzula sylvatica, while its stimulation under elevated [CO2] in less shade-tolerant Calamagrostis arundinacea was expected only in early spring when the tree canopy is not fully developed.We found evidence supporting this hypothesis. While elevated [CO2] stimulated photosynthesis in L. sylvatica mainly in the peak of the growing season (by 55%-57% in July and August), even at low light intensities (50 µmol m-2 s-1), stimulatory effect of [CO2] in C. arundinacea was found mainly under high light intensities (200 µmol m-2 s-1) at the beginning of the growing season (increase by 171% in May) and gradually declined during the season. Elevated [CO2] also substantially stimulated leaf mass area and root-to-shoot ratio in L. sylvatica, while only insignificant increases were observed in C. arundinacea.Our physiological and morphological analyses indicate that understorey species, differing in shade-tolerance, under elevated [CO2] exploit distinct niches in light environment given by the dynamics of the tree canopy.

Long-term interactive effects of N addition with P and K availability on N status of Sphagnum.

Little information exists concerning the long-term interactive effect of nitrogen (N) addition with phosphorus (P) and potassium (K) on Sphagnum N status. This study was conducted as part of a long-term N manipulation on Whim bog in south Scotland to evaluate the long-term alleviation effects of phosphorus (P) and potassium (K) on N saturation of Sphagnum (S. capillifolium). On this ombrotrophic peatland, where ambient deposition was 8 kg N ha-1 yr-1, 56 kg N ha-1 yr-1 of either ammonium (NH4+, Nred) or nitrate (NO3-, Nox) with and without P and K, were added over 11 years. Nutrient concentrations of Sphagnum stem and capitulum, and pore water quality of the Sphagnum layer were assessed. The N-saturated Sphagnum caused by long-term (11 years) and high doses (56 kg N ha-1 yr-1) of reduced N was not completely ameliorated by P and K addition; N concentrations in Sphagnum capitula for Nred 56 PK were comparable with those for Nred 56, although N concentrations in Sphagnum stems for Nred 56 PK were lower than those for Nred 56. While dissolved inorganic nitrogen (DIN) concentrations in pore water for Nred 56 PK were not different from Nred 56, they were lower for Nox 56 PK than for Nox 56 whose stage of N saturation had not advanced compared to Nred 56. These results indicate that increasing P and K availability has only a limited amelioration effect on the N assimilation of Sphagnum at an advanced stage of N saturation. This study concluded that over the long-term P and K additions will not offset the N saturation of Sphagnum.

Sphagnum can 'filter' N deposition, but effects on the plant and pore water depend on the N form.

The ability of Sphagnum moss to efficiently intercept atmospheric nitrogen (N) has been assumed to be vulnerable to increased N deposition. However, the proposed critical load (20kgNha(-1)yr(-1)) to exceed the capacity of the Sphagnum N filter has not been confirmed. A long-term (11years) and realistic N manipulation on Whim bog was used to study the N filter function of Sphagnum (Sphagnum capillifolium) in response to increased wet N deposition. On this ombrotrophic peatland where ambient deposition was 8kgNha(-1)yr(-1), an additional 8, 24, and 56kgNha(-1)yr(-1) of either ammonium (NH4(+)) or nitrate (NO3(-)) has been applied for 11years. Nutrient status of Sphagnum and pore water quality from the Sphagnum layer were assessed. The N filter function of Sphagnum was still active up to 32kgNha(-1)yr(-1) even after 11years. N saturation of Sphagnum and subsequent increases in dissolved inorganic N (DIN) concentration in pore water occurred only for 56kgNha(-1)yr(-1) of NH4(+) addition. These results indicate that the Sphagnum N filter is more resilient to wet N deposition than previously inferred. However, functionality will be more compromised when NH4(+) dominates wet deposition for high inputs (56kgNha(-1)yr(-1)). The N filter function in response to NO3(-) uptake increased the concentration of dissolved organic N (DON) and associated organic anions in pore water. NH4(+) uptake increased the concentration of base cations and hydrogen ions in pore water though ion exchange. The resilience of the Sphagnum N filter can explain the reported small magnitude of species change in the Whim bog ecosystem exposed to wet N deposition. However, changes in the leaching substances, arising from the assimilation of NO3(-) and NH4(+), may lead to species change.

Effects of temperature and drought manipulations on seedlings of Scots pine provenances.

Rising temperatures and more frequent and severe climatic extremes as a consequence of climate change are expected to affect growth and distribution of tree species that are adapted to current local conditions. Species distribution models predict a considerable loss of habitats for Pinus sylvestris. These models do not consider possible intraspecific differences in response to drought and warming that could buffer those impacts. We tested 10 European provenances of P. sylvestris, from the southwestern to the central European part of the species distribution, for their response to warming and to drought using a factorial design. In this common-garden experiment the air surrounding plants was heated directly to prevent excessive soil heating, and drought manipulation, using a rain-out shelter, permitted almost natural radiation, including high light stress. Plant responses were assessed as changes in phenology, growth increment and biomass allocation. Seedlings of P. sylvestris revealed a plastic response to drought by increased taproot length and root-shoot ratios. Strongest phenotypic plasticity of root growth was found for southwestern provenances, indicating a specific drought adaptation at the cost of overall low growth of aboveground structures even under non-drought conditions. Warming had a minor effect on growth but advanced phenological development and had a contrasting effect on bud biomass and diameter increment, depending on water availability. The intraspecific variation of P. sylvestris provenances could buffer climate change impacts, although additional factors such as the adaptation to other climatic extremes have to be considered before assisted migration could become a management option.

Populations, pools, and peccaries: simulating the impact of ecosystem engineers on rainforest frogs.

"Ecosystem engineering" describes habitat alteration by an organism that affects another organism; such nontrophic interactions between organisms are a current focus in ecological research. Our study quantifies the actual impact an ecosystem engineer can have on another species by using a previously identified model system-peccaries and rainforest frogs. In a 4-year experiment, we simulated the impact of peccaries on a population of Allobates femoralis (Dendrobatidae) by installing an array of artificial pools to mimic a forest patch modified by peccaries. The data were analyzed using a gradual before-after control-impact (gBACI) model. Following the supplementation, population size almost doubled as a result of increased autochthonous recruitment driven by a higher per-capita reproduction of males and a higher proportion of reproducing females. The effect was evenly distributed across the population. The differential response of males and females reflects the reproductive behavior of A. femoralis, as only the males use the aquatic sites for tadpole deposition. Our study shows that management and conservation must consider nontrophic relationships and that human "ecosystem engineering" can play a vital role in efforts against the "global amphibian decline."