Revisiting the classics: considering nonconsumptive effects in textbook examples of predator-prey interactions

Predator effects on prey dynamics are conventionally studied by measuring changes in prey abundance attributed to consumption by predators. We revisit four classic examples of predator-prey systems often cited in textbooks and incorporate subsequent studies of nonconsumptive effects of predators (NCE), defined as changes in prey traits (e.g., behavior, growth, development) measured on an ecological time scale. Our review revealed that NCE were integral to explaining lynx-hare population dynamics in boreal forests, cascading effects of top predators in Wisconsin lakes, and cascading effects of killer whales and sea otters on kelp forests in nearshore marine habitats. The relative roles of consumption and NCE of wolves on moose and consequent indirect effects on plant communities of Isle Royale depended on climate oscillations. Nonconsumptive effects have not been explicitly tested to explain the link between planktonic alewives and the size structure of the zooplankton, nor have they been invoked to attribute keystone predator status in intertidal communities or elsewhere. We argue that both consumption and intimidation contribute to the total effects of keystone predators, and that characteristics of keystone consumers may differ from those of predators having predominantly NCE. Nonconsumptive effects are often considered as an afterthought to explain observations inconsistent with consumption-based theory. Consequently, NCE with the same sign as consumptive effects may be overlooked, even though they can affect the magnitude, rate, or scale of a prey response to predation and can have important management or conservation implications. Nonconsumptive effects may underlie other classic paradigms in ecology, such as delayed density dependence and predator-mediated prey coexistence. Revisiting classic studies enriches our understanding of predator-prey dynamics and provides compelling rationale for ramping up efforts to consider how NCE affect traditional predator-prey models based on consumption, and to compare the relative magnitude of consumptive and NCE of predators.     

Trophic levels and trophic tangles: the prevalence of omnivory in real food webs

The concept of trophic levels is one of the oldest in ecology and informs our understanding of energy flow and top-down control within food webs, but it has been criticized for ignoring omnivory. We tested whether trophic levels were apparent in 58 real food webs in four habitat types by examining patterns of trophic position. A large proportion of taxa (64.4%) occupied integer trophic positions, suggesting that discrete trophic levels do exist. Importantly however, the majority of those trophic positions were aggregated around integer values of 0 and 1, representing plants and herbivores. For the majority of the real food webs considered here, secondary consumers were no more likely to occupy an integer trophic position than in randomized food webs. This means that, above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores. Omnivory was most common in marine systems, rarest in streams, and intermediate in lakes and terrestrial food webs. Trophic-level-based concepts such as trophic cascades may apply to systems with short food chains, but they become less valid as food chains lengthen.     

Controls on vegetation structure in Southwestern ponderosa pine forests, 1941 and 2004

Long-term studies can broaden our ecological understanding and are particularly important when examining contingent effects that involve changes to dominance by long-lived species. Such a change occurred during the last century in Southwestern (USA) ponderosa pine (Pinus ponderosa) forests. We used five livestock grazing exclosures established in 1912 to quantify vegetation structure in 1941 and 2004. Our objectives were to (1) assess the effects of historical livestock grazing on overstory structure and age distribution, (2) assess the effects of recent livestock grazing and overstory on understory vegetation, and (3) quantify and explain changes in understory vegetation between 1941 and 2004. In 1941, canopy cover of tree regeneration was significantly higher inside exclosures. In 2004, total tree canopy cover was twice as high, density was three times higher, trees were smaller, and total basal area was 40% higher inside exclosures. Understory species density, herbaceous plant density, and herbaceous cover were negatively correlated with overstory vegetation in both years. Most understory variables did not differ between grazing treatments in 1941 but were lower inside exclosures in 2004. Differences between grazing treatments disappeared once overstory effects were accounted for, indicating that they were due to the differential overstory response to historical livestock grazing practices. Between 1941 and 2004, species density declined by 34%, herbaceous plant density by 37%, shrub cover by 69%, total herbaceous cover by 59%, graminoid cover by 39%, and forb cover by 82%. However, these variables did not differ between grazing treatments or years once overstory effects were accounted for, indicating that the declines were driven by the increased dominance of the overstory during this period. Our results demonstrate that historical livestock grazing practices are an aspect of land-use history that can affect ecosystem development. Grazing history must be considered when extrapolating results from one site to another. In addition, the understory vegetation was more strongly controlled by the ponderosa pine overstory than by recent livestock grazing or by temporal dynamics, indicating that overstory effects must be accounted for when examining understory responses in this ecosystem.     

Independent gradients of producer, consumer, and microbial diversity in lake plankton

Interactions between trophic levels during food web assembly can drive positive correlations in diversity between producers, consumers, and decomposers. However, the contribution of trophic interactions relative to local environmental factors in promoting species diversity is poorly understood, with many studies only considering two trophic levels. Here we examine correlations in diversity among zooplankton, phytoplankton, and bacteria in the pelagic zone of 31 lakes in British Columbia, Canada. We sampled species diversity of zooplankton and phytoplankton through morphological identification, and bacterial genetic diversity was estimated by denaturing gradient gel electrophoresis (DGGE) of 16S rDNA polymorphisms. We looked for correlations in diversity that were independent of the abiotic environment by statistically controlling for 18 limnological variables. No significant correlations were found between the diversity of zooplankton, phytoplankton, and bacteria. In addition, the physical factors that were associated with species composition in one trophic level were independent of those that were important for another. Our results provide no support for the importance of direct feedbacks between producers, consumers, and decomposers in maintaining diversity. Zooplankton, phytoplankton, and bacterial diversity and composition are regulated independently from one another and respond to different environmental variables. These results suggest that species of lake plankton show loose trophic associations with one another due to broad diets in consumers and decomposers.     

Quantitative threat analysis for management of an imperiled species: Chinook salmon (Oncorhynchus tshawytscha).

Chinook salmon (Oncorhynchus tshawytscha) have declined dramatically across the Pacific Northwest because of multiple human impacts colloquially characterized as the four "H's": habitat degradation, harvest, hydroelectric and other dams, and hatchery production. We use this conceptual framework to quantify the relative importance of major threats to the current status of 201 Chinook populations. Current status is characterized by two demographic indices: population density and trend. We employ path analytic models and information theoretic methods for multi-model inference. Our results indicate that dams most strongly affect variation in population density, while harvest and hatchery production most strongly affect variation in population trend. Comparable results arise when the sample size of the analysis is reduced to 22 Chinook populations within a smaller region typical of the scale at which salmon recovery planning is conducted. Results from these threat analyses suggest that recovery strategies targeting specific demographic indices, and those considering natural and human-mediated interdependencies of major threats, are most likely to succeed.     

Inorganic and organic contaminants in sediments from an urban playa and associated toxicity among Hyalella azteca

Playa wetlands are important components of the Southern High Plains (USA) landscape as they are the major aquatic surface feature. Chemical contaminants associated with playas have been documented, particularly for grassland and agricultural watersheds, but not for playas in urban settings. The objectives of this study were to determine concentrations of inorganic and organic contaminants in sediments from an urban playa within the I-20 Wildlife Preserve and Jenna Welch Nature Study Center in Midland, TX, and evaluate toxicity of these sediments to Hyalella azteca. Concentrations of most trace elements were below sediment quality guidelines with exceptions of lead, cadmium, and arsenic. Concentrations of organic contaminants, particularly PAHs, DDT, DDE, and malathion, were above sediment quality guidelines at various locations within the playa. Decreased survival was observed among H. azteca exposed to sediment from a single location when compared those exposed to reference sediments. This location also produced maximum observed concentrations for five of seven trace elements, potentially due to its location at the lowest elevation within the playa. This study documented concentrations of contaminants in sediments of an urban playa associated with past and present land uses in its urban setting, including those from automotive emissions and historical pesticide use.     

Vivid birds respond more to acoustic signals of predators

Because conspicuous morphology such as colorful plumage may increase predation risk, we aimed to see if variation in plumage coloration could explain variation in avian anti-predator behavior. We included several measures of plumage coloration: human perception of vividness from images in field guides, total intensity from reflectance spectra of museum skins, contrasts calculated from physiological models of these spectra parameterized for both raptors and humans, chroma, and spectral saturation. We investigated how well these measurements predicted risk assessment in ten species of birds in St. John, U.S. Virgin Islands. We quantified how each species responded to playbacks of a predator’s calls and compared this response to that elicited by songs from a non-predatory, sympatric bird. We found that human-determined measures of vividness best predicted anti-predator responses of birds—more vividly colored species responded more to predators than duller species. No spectrophotometric variable explained variation in species reactions to a predator call. Our results suggest that vivid birds may compensate for their conspicuousness by being more responsive to the sound of predators and that more work is needed to better evaluate how animal coloration is quantified in comparative studies.     

Lipid components as a measure of nutritional condition in fish larvae (Pleuragramma antarcticum) in East Antarctica

Pleuragramma antarcticum is a key component of the neritic assemblages in the Antarctic coastal waters. Larvae of this species were sampled from 2008 to 2011 in the Dumont d’Urville Sea (East Antarctica). The lipid class composition [triacylglycerols (TAG), cholesterol (Chol) and polar lipids (PL)] of larvae was measured to assess the larval condition. The total amount of lipids was linearly related to the quantity of structural polar lipids, suggesting that growth is favored over lipid storage. The TAG:Chol ratio showed interannual variability in the condition of fish larvae, probably related to prey availability. Nevertheless, the essential fatty acids composition of polar lipids illustrates that larvae with low levels of TAG:Chol could be either growing or under starvation. Only the combination of a low TAG:Chol ratio and low polar lipids content, which can also be mobilized during starvation periods, allowed identification of larvae in poor condition. This lipid condition index should be of great assistance to evaluate the probability of survival of P. antarcticum larvae in long-term monitoring. It has widespread applicability and should also be useful in the diagnosis of nutritional condition in other species.     

Stingray life history trade-offs associated with nursery habitat use inferred from a bioenergetics model

Consumption rates of marine predators are vital to assessing their trophic impacts and potential consequences of fisheries removal and habitat alteration, yet are rarely estimated. Standard metabolic rates were estimated for juvenile brown stingrays, Dasyatis lata, and used as input parameters for a bioenergetics model to predict consumption rates. Temperature and mass had significant effects on metabolic rates. The energy budget of juvenile brown stingrays was heavily weighted toward metabolism, accounting for 66 % of consumed energy. Growth accounted for 7 % of the energy budget indicating very slow growth potentially due to limited food resources. Population consumption rates suggest potential for strong top-down effects on prey populations due to stingray predation. This study suggests the use of Kāne‘ohe Bay as a nursery habitat for juvenile brown stingrays is a trade-off between increased juvenile survival through predator avoidance and a late age at first maturity due to slow growth rates resulting from low prey availability.     

Health care system hazard vulnerability analysis: an assessment of all public hospitals in Abu Dhabi

Hazard vulnerability analysis (HVA) is used to risk‐stratify potential threats, measure the probability of those threats, and guide disaster preparedness. The primary objective of this project was to analyse the level of disaster preparedness in public hospitals in the Emirate of Abu Dhabi, utilising the HVA tool in collaboration with the Disaster Medicine Section at Harvard Medical School. The secondary objective was to review each facility's disaster plan and make recommendations based on the HVA findings. Based on the review, this article makes eight observations, including on the need for more accurate data; better hazard assessment capabilities; enhanced decontamination capacities; and the development of hospital‐specific emergency management programmes, a hospital incident command system, and a centralised, dedicated regional disaster coordination centre. With this project, HVAs were conducted successfully for the first time in health care facilities in Abu Dhabi. This study thus serves as another successful example of multidisciplinary emergency preparedness processes.