Current food web models cannot explain the overall topological structure of observed food webs

Topological food webs illustrating “who eats whom” in different systems exhibit similar, non‐random, structures suggesting that general rules govern food web structure. Current food web models correctly predict many measures of food web topology from knowledge of species richness and connectance (fraction of possible predator–prey links that actually occur), together with assumptions about the ecological rules governing “who eats whom”. However, current measures are relatively insensitive to small changes in topology. Improvement of, and discrimination among, current models requires development of new measures of food web structure. Here I examine whether current food web models (cascade, niche, and nested hierarchy models, plus a random null model) can predict a new measure of food web structure, structural stability. Structural stability complements other measures of food web topology because it is sensitive to changes in topology that other measures often miss. The cascade and null models respectively over‐ and underpredict structural stability for a set of 17 high‐quality food webs. While the niche and nested hierarchy models provide unbiased predictions on average, their 95% confidence intervals frequently fail to include the observed data. Observed structural stabilities for all models are overdispersed compared to model predictions, and predicted and observed structural stabilities are uncorrelated, indicating that important sources of variation in structural stability are not captured by the models. Crucially, poor model performance arises because observed variation in structural stability is unrelated to variation in species richness and connectance. In contrast, almost all other measures of food web topology vary with species richness and connectance in natural webs. No model that takes species richness and connectance as the only input parameters can reproduce observed variation in structural stability. Further progress in predicting and explaining food web topology will require fundamentally new models based on different input parameters.     

Prey preference of top predators manipulates the functioning and stability of multi-trophic ecosystems

The relationship between biodiversity and ecosystem functioning, and the mechanisms underpinning the food web stability, have been intensively investigated in ecological research. The ubiquities of generalists in natural food webs and its important role in dictating these ecosystem properties have been generally recognized. However, how competition between multiple top predators shape these ecosystem properties and determine the success of invasive predators remain largely unexplored. Here, we use a well-developed food web model to investigate the effects of prey preference of top predators on ecosystem functioning and food web stability in both local and invasive conditions. We design several modeling scenarios to mimic combinations of different types of top predators (specialist/generalist) and their origins (local/invasive). Our model theoretically shows that lower exploitation competition for prey between top predators (with distinct prey preferences featured by higher attack rates) would be beneficial for the ecosystem functioning and food web stability. We also demonstrate that the success of top predator invasion depends on the prey preference of both local and invasive top predators. Sensitivity analysis on the model further supports our findings. Our results highlight the importance of prey preference of multiple top predators in manipulating the properties of multi-trophic ecosystems. Our findings may have important implications because the current ongoing global changes profoundly change the phenology of many biological systems and create trophic mismatch, which may manipulate prey preference of top predators and in turn deteriorate ecosystem functioning and food web stability.     

Spatiotemporal food web dynamics along a desert riparian–upland transition

Increased awareness of spatiotemporal variation in species interactions has motivated the study of temporally-resolved food web dynamics at the landscape level. Empiricists have focused attention on cross-habitat flows of materials, nutrients, and prey, largely ignoring the movement of predators between habitats that differ in productivity (and how predators integrate pulses in resource availability over time). We set out to study seasonal variation in food web interactions between mammalian carnivores and their rodent prey along a riparian–upland gradient in semi-arid southeastern Arizona which features both spatial and temporal heterogeneity in resource availability. Specifically, we tested whether mammalian carnivores spill over from productive, near-river habitats into adjacent, desert-scrub habitats; and if they do, to document the effects of this spillover on rodent communities. Furthermore, we examined seasonal variation in top-down effects by measuring changes in carnivore diet and distribution patterns and rodent populations over time. The results indicate that carnivores track seasonally-abundant resources across the landscape, varying both their diet and movement patterns. In turn, desert-scrub rodent population dynamics track seasonal shifts in carnivore habitat use but not resource availability, suggesting that predation plays a role in structuring rodent communities along the San Pedro River. Further evidence comes from data on rodent community composition, which differs between desert-scrub habitats near and far from the river, despite similarities in resource availability. Our data also suggest that seasonal omnivory helps predators survive lean times, increasing their effects on prey populations. Taken together, these results underscore the importance of spatiotemporal variation in species interactions, highlighting the complexity of natural systems and the need for further detailed studies of food web dynamics.     

Effects of evolutionary changes in prey use on the relationship between food web complexity and stability

The relationship between food web complexity and stability has been the subject of a long-standing debate in ecology. Although rapid changes in the food web structure through adaptive foraging behavior can confer stability to complex food webs, as reported by Kondoh (Science 299:1388–1391, 2003), the exact mechanisms behind this adaptation have not been specified in previous studies; thus, the applicability of such predictions to real ecosystems remains unclear. One mechanism of adaptive foraging is evolutionary change in genetically determined prey use. We constructed individual-based models of evolution of prey use by predators assuming explicit population genetics processes, and examined how this evolution affects the stability (i.e., the proportion of species that persist) of the food web and whether the complexity of the food web increased the stability of the prey–predator system. The analysis showed that the stability of food webs decreased with increasing complexity regardless of evolution of prey use by predators. The effects of evolution on stability differed depending on the assumptions made regarding genetic control of prey use. The probabilities of species extinctions were associated with the establishment or loss of trophic interactions via evolution of the predator, indicating a clear link between structural changes in the food web and community stability.     

Comparative analysis of food-finding behavior of an herbivorous and a carnivorous land snail

Abstract. Although the olfactory capabilities of land snail tentacles have been tested by lesion studies and unilateral exposure of tentacles to specific odors, studies of a carnivorous species suggest that the anatomical similarities of herbivorous and carnivorous land snails may belie a fundamental difference in the way these structures are used to find food. Therefore, we challenged the herbivore, Anguispira alternata , and the carnivore, Haplotrema concavum , to find a stationary food source (carrot and caged young prey snail, respectively) under identical still air conditions. The herbivore traveled a significantly shorter distance to the food, even negotiating a barrier placed halfway between the snail and its food. The carnivore, on the other hand, followed a circuitous, apparently random, path to the food. Subsequent tests revealed that H. concavum readily follows prey slime trails while A. alternata seldom follows conspecific slime trails when a distant food source is available. These results are consistent with what might be expected as adaptations to the usually mobile nature of carnivore prey and the stationary nature of herbivore food plants. The ability of A. alternata to exhibit typical detour behavior is noted.     

The role of exotic mammals in the diet of native carnivores from South America

相似文献   

Seasonal subsidy stabilizes food web dynamics: Balance in a heterogeneous landscape

Resource subsidies from external habitats can substantially affect the food web dynamics of local habitats. In this paper, we explore a mathematical model that is tailored for a stream food web, studied by Nakano and colleagues, in which consumers, in situ prey and subsidies all show seasonal fluctuation. The model reveals that the food web dynamics are stabilized if subsidies increase in summer when in situ productivity is low. Consumer dynamics are stabilized because subsidies complement seasonal resource deficiency. In situ prey dynamics are stabilized because subsidies indirectly balance the predation pressure by consumers, with seasonal change in prey carrying capacity. In summer when prey carrying capacity is low, seasonally abundant subsidies indirectly decrease predation pressure, whereas in winter, with high prey carrying capacity, scarce subsidies increase the predation pressure. Our results suggest that temporal productivity differences between spatially linked habitats are important to promote the stability of food web dynamics in a landscape context.     

Rethinking trophic niches: Speed and body mass colimit prey space of mammalian predators

1. Realized trophic niches of predators are often characterized along a one‐dimensional range in predator–prey body mass ratios. This prey range is constrained by an “energy limit” and a “subdue limit” toward small and large prey, respectively. Besides these body mass ratios, maximum speed is an additional key component in most predator–prey interactions.
2. Here, we extend the concept of a one‐dimensional prey range to a two‐dimensional prey space by incorporating a hump‐shaped speed‐body mass relation. This new “speed limit” additionally constrains trophic niches of predators toward fast prey.
3. To test this concept of two‐dimensional prey spaces for different hunting strategies (pursuit, group, and ambush predation), we synthesized data on 63 terrestrial mammalian predator–prey interactions, their body masses, and maximum speeds.
4. We found that pursuit predators hunt smaller and slower prey, whereas group hunters focus on larger but mostly slower prey and ambushers are more flexible. Group hunters and ambushers have evolved different strategies to occupy a similar trophic niche that avoids competition with pursuit predators. Moreover, our concept suggests energetic optima of these hunting strategies along a body mass axis and thereby provides mechanistic explanations for why there are no small group hunters (referred to as “micro‐lions”) or mega‐carnivores (referred to as “mega‐cheetahs”).
5. Our results demonstrate that advancing the concept of prey ranges to prey spaces by adding the new dimension of speed will foster a new and mechanistic understanding of predator trophic niches and improve our predictions of predator–prey interactions, food web structure, and ecosystem functions.

     

Predator-prey size relationships in an African large-mammal food web

1. Size relationships are central in structuring trophic linkages within food webs, leading to suggestions that the dietary niche of smaller carnivores is nested within that of larger species. However, past analyses have not taken into account the differing selection shown by carnivores for specific size ranges of prey, nor the extent to which the greater carcass mass of larger prey outweighs the greater numerical representation of smaller prey species in the predator diet. Furthermore, the top-down impact that predation has on prey abundance cannot be assessed simply in terms of the number of predator species involved. 2. Records of found carcasses and cause of death assembled over 46 years in the Kruger National Park, South Africa, corrected for under-recording of smaller species, enabled a definitive assessment of size relationships between large mammalian carnivores and their ungulate prey. Five carnivore species were considered, including lion (Panthera leo), leopard (Panthera pardus), cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus) and spotted hyena (Crocuta crocuta), and 22 herbivore prey species larger than 10 kg in adult body mass. 3. These carnivores selectively favoured prey species approximately half to twice their mass, within a total prey size range from an order of magnitude below to an order of magnitude above the body mass of the predator. The three smallest carnivores, i.e. leopard, cheetah and wild dog, showed high similarity in prey species favoured. Despite overlap in prey size range, each carnivore showed a distinct dietary preference. 4. Almost all mortality was through the agency of a predator for ungulate species up to the size of a giraffe (800-1200 kg). Ungulates larger than twice the mass of the predator contributed substantially to the dietary intake of lions, despite the low proportional mortality inflicted by predation on these species. Only for megaherbivores substantially exceeding 1000 kg in adult body mass did predation become a negligible cause of mortality. 5. Hence, the relative size of predators and prey had a pervasive structuring influence on biomass fluxes within this large-mammal food web. Nevertheless, the large carnivore assemblage was dominated overwhelmingly by the largest predator, which contributed the major share of animals killed across a wide size range.     

The Coupling Between Grazing and Detritus Food Chains and the Strength of Trophic Cascades Across a Gradient of Nutrient Enrichment

A minimal food web model was constructed comprising one grazing and one detritus food chain coupled by nutrient cycling and generalist carnivores to investigate how prey preference by carnivores may affect the strength of trophic cascades across a gradient of nutrient enrichment. The equilibrium or mean abundance of each food web component and the magnitude of the carnivore effect on lower trophic levels were calculated for different values of the prey preference and nutrient input parameters. Our model predicts that nutrient enrichment increases the mean abundances of carnivores, autotrophs and detritus, but the magnitude of this effect is dependent on the prey preference term. On the other hand, herbivores and detritivores are relatively unaffected by enrichment but are strongly affected by carnivore preference. Carnivores have a negative effect on herbivores and a positive effect on autotrophs and detritus, whereas the effect on detritivores can be both positive and negative. At high preference for herbivores, carnivores have a positive effect on detritivores, because the positive effect of increased detritus availability due to reduced herbivore grazing outweighs the negative effect of predation. At high preference for detritivores, the balance is changed in the other direction. We argue that in systems where authochtonous primary production is the major source of detritus, herbivores can control the rates of detritus production and have indirect effects on detritivores, which may feed back into effects on herbivores through their shared enemies. This positive feedback is probably one mechanism affecting the resilience of alternative stable states in shallow lakes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  J.L.A. conceived the model and the study, J.R. wrote the Matlab programs and ran the simulations and J.L.A. wrote most of the paper.     

海州湾5种关键饵料生物的主要捕食者及其捕食压力

为了解海州湾关键饵料生物在食物网中的被捕食压力及自然死亡率波动情况,研究基于2011和2013—2020年在海州湾及其邻近海域进行的渔业资源底拖网调查资料和胃含物分析数据,以物种间的营养联系为基础,选择细螯虾(Leptochela gracilis)、日本鼓虾(Alpheus japonicus)、枪乌贼(Loligo sp.)、小黄鱼(Larimichthys polyactis)和口虾蛄(Oratosquilla oratoria)5种关键饵料生物为研究对象,通过计算捕食压力指数(Predation pressure index, PPI),分析它们在海州湾食物网中的主要捕食者及其捕食压力,并计算了纳入捕食压力指数的自然死亡系数。结果表明,细螯虾的捕食者对其年平均捕食压力指数最高,小黄鱼的捕食者对其年平均捕食压力指数最低。小眼绿鳍鱼(Chelidonichthys kumu)对细螯虾和枪乌贼的年平均捕食压力指数最高,分别为168.89和75.77;(Miichthys miiuy)对日本鼓虾和口虾蛄的年平均捕食压力指数最高,分别为39.41和9.85;海鳗(Muraenesox c...     

Relations between bacterioplankton,heterotrophic nanoflagellates,and virioplankton in the littoral zone of a large plain reservoir: Impact of bird colonies

Interactions of the main components of microbial planktonic food web (bacteria, heterotrophic nanoflagellates, and viruses) were studied in a protected overgrown littoral zone of the Rybinsk Reservoir (Upper Volga). The effect of colonial bird settlements (the Laridae family) on these processes was determined. The following systems exhibited significant negative correlations: “heterotrophic nanoflagellates–large rod-shaped bacteria” (“predator–prey”), “viruses-bacteriophages–bacterial products” (“parasite–host”) and “heterotrophic nanoflagellates–viruses-bacteriophages”. Relations between biotic factors controlling bacterial development were more pronounced outside the zone affected by colonial bird settlements. Near the bird colony the role of viruses in mortality of planktonic bacteria increased. Reproduction of bacterial cells accelerated in response to the increase in feeding activity of heterotrophic nanoflagellates. Virusesbacteriophages and heterotrophic nanoflagellates probably eliminate different targets until medium-sized cells become predominant in the bacterial community. Then heterotrophic nanoflagellates consume bacterial cells infected with viruses.     

Inducible defenses,competition and shared predation in planktonic food chains

Ecologists have long debated the role of predation in mediating the coexistence of prey species. Theory has mainly taken a bitrophic perspective that excludes the effects of inducible defenses at different trophic levels. However, inducible defenses could either limit or enhance the effects of predation on coexistence, by means of effects on bottom-up control and population stability. Our aim was to investigate how inducible defenses at different trophic levels affect the possibilities for predator-mediated coexistence, as opposed to competitive exclusion, in replicated experimental plankton communities. In particular, we analyzed how the presence or absence of inducible defenses in algal basal prey affected the outcome of competition between an inducible defended and an undefended herbivore, in the presence or absence of a carnivore. We found the undefended herbivore to be a superior competitor in the absence of predation. This outcome was reversed in the presence of a shared carnivore: populations of the undefended herbivore then strongly declined. The extent of this population decline differed between food webs based on undefended as opposed to inducible defended algal prey. In the former the undefended herbivore became undetectable for most of the duration of the experiment. In the latter the undefended herbivore also crashed to low densities, but it could still be detected during most of the experiment. In food webs based on inducible defended algae, the carnivore failed to reach high densities and exerted weaker top-down control on the two competing herbivores. We conclude that the inducible defense in one of our two competing herbivores allowed the outcome of competition to be reversed when a shared carnivore was added. Inducible defenses in algae did not change this outcome, but they significantly delayed extinction of the undefended herbivore. Predation itself did not promote coexistence in these experimental plankton communities.     

基于SURF指数识别海州湾食物网的关键饵料生物

关键种(keystone species)在生态系统中发挥着不可替代的重要作用,对于群落结构的稳定与演替起着决定性的作用。基于2011年3—12月在海州湾及其邻近海域进行的渔业资源底拖网调查资料以及胃含物分析数据和参考历史文献数据,以物种间的摄食关系作为基础,采用SURF(Supportive Role to Fishery ecosystems)指数识别海州湾食物网中的关键饵料生物。在矩阵中分析每个物种作为饵料生物为捕食者提供的摄食比例及捕食者数量,计算每个物种的关键指标(SURFi)的值,结果表明前五位数值较高的物种是细鳌虾(Leptochela gracilis)、毛虾(Acetessp.)、疣背宽额虾(Latreutes planirostris)、日本鼓虾(Alpheus japonicus)和鳀鱼(Engraulis japonicus),它们是海州湾的关键饵料生物,在整个食物网中起到关键的控制作用,它们的数量波动会对海州湾其他物种产生直接或间接的影响。加强关键饵料生物的保护,对于维持海州湾生态系统的健康和稳定至关重要。     

Temperature alters food web body-size structure

The increased temperature associated with climate change may have important effects on body size and predator–prey interactions. The consequences of these effects for food web structure are unclear because the relationships between temperature and aspects of food web structure such as predator–prey body-size relationships are unknown. Here, we use the largest reported dataset for marine predator–prey interactions to assess how temperature affects predator–prey body-size relationships among different habitats ranging from the tropics to the poles. We found that prey size selection depends on predator body size, temperature and the interaction between the two. Our results indicate that (i) predator–prey body-size ratios decrease with predator size at below-average temperatures and increase with predator size at above-average temperatures, and (ii) that the effect of temperature on predator–prey body-size structure will be stronger at small and large body sizes and relatively weak at intermediate sizes. This systematic interaction may help to simplify forecasting the potentially complex consequences of warming on interaction strengths and food web stability.     

Food web complexity and stability across habitat connectivity gradients

The effects of habitat connectivity on food webs have been studied both empirically and theoretically, yet the question of whether empirical results support theoretical predictions for any food web metric other than species richness has received little attention. Our synthesis brings together theory and empirical evidence for how habitat connectivity affects both food web stability and complexity. Food web stability is often predicted to be greatest at intermediate levels of connectivity, representing a compromise between the stabilizing effects of dispersal via rescue effects and prey switching, and the destabilizing effects of dispersal via regional synchronization of population dynamics. Empirical studies of food web stability generally support both this pattern and underlying mechanisms. Food chain length has been predicted to have both increasing and unimodal relationships with connectivity as a result of predators being constrained by the patch occupancy of their prey. Although both patterns have been documented empirically, the underlying mechanisms may differ from those predicted by models. In terms of other measures of food web complexity, habitat connectivity has been empirically found to generally increase link density but either reduce or have no effect on connectance, whereas a unimodal relationship is expected. In general, there is growing concordance between empirical patterns and theoretical predictions for some effects of habitat connectivity on food webs, but many predictions remain to be tested over a full connectivity gradient, and empirical metrics of complexity are rarely modeled. Closing these gaps will allow a deeper understanding of how natural and anthropogenic changes in connectivity can affect real food webs.     

Evidence of two subaggregations of humpback whales on the Kodiak,Alaska, feeding ground revealed from stable isotope analysis

Knowledge of humpback whale (Megaptera novaeangliae) foraging on feeding grounds is becoming increasingly important as the growing North Pacific population recovers from commercial whaling and consumes more prey, including economically important fishes. We explored spatial and temporal (interannual, within‐season) variability in summer foraging by humpback whales along the eastern side of the Kodiak Archipelago as described by stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of humpback whale skin (n = 118; 2004–2013). The trophic level (TL) of individual whales was calculated using basal food web δ¹⁵N values collected within the study area. We found evidence for the existence of two subaggregations of humpback whales (“North,” “South”) on the feeding ground that fed at different TLs throughout the study period. Linear mixed models suggest that within an average year, Kodiak humpback whales forage at a consistent TL during the feeding season. TL estimates support mixed consumption of fish and zooplankton species in the “North” (mean ± SE; 3.3 ± 0.1) and predominant foraging on zooplankton in the “South” (3.0 ± 0.1). This trend appears to reflect spatial differences in prey availability, and thus, our results suggest North Pacific humpback whales may segregate on feeding aggregations and target discrete prey species.     

Aboriginal overkill

Prior to European influence, predation by Native Americans was the major factor limiting the numbers and distribution of ungulates in the Intermountain West. This hypothesis is based on analyses of (1) the efficiency of Native American predation, including cooperative hunting, use of dogs, food storage, use of nonungulate foods, and hunting methods; (2) optimal-foraging studies; (3) tribal territory boundary zones as prey reservoirs; (4) species ratios, and sex and age of aboriginal ungulate kills; (5) impact of European diseases on aboriginal populations; and (6) synergism between aboriginal and carnivore predation. Native Americans had no effective conservation practices, and the manner in which they harvested ungulates was the exact opposite of any predicted conservation strategy. Native Americans acted in ways that maximized their individual fitness regardless of the impact on the environment. For humans, conservation is seldom an evolutionarily stable strategy. By limiting ungulate numbers and purposefully modifying the vegetation with fire, Native Americans structured entire plant and animal communities. Because ecosystems with native peoples are entirely different than those lacking aboriginal populations, a “hands-off” or “natural regulation” approach by today’s land managers will not duplicate the ecological conditions under which those ecosystems developed. The modern concept of wilderness as areas without human influence is a myth. North America was not a “wilderness” waiting to be discovered, instead it was home to tens of millions of aboriginal peoples before European-introduced diseases decimated their numbers.     

Community Response of Mammalian Predators and Their Prey to Motorways: Implications for Predator–Prey Dynamics

Understanding the interactions between predators and prey is essential for predicting the effects of disturbances to ecosystems. Motorways produce changes in the surrounding biotic and abiotic environment and hence have multiple impacts on wildlife. Some species are known to change their activity patterns in the proximity of motorways but the implications for the structure of food webs are unknown. This study analyzes the activity patterns of both mammalian predators and their prey species near nine motorways in attempt to clarify how motorways affect the mammalian community. Habitat structural variables were also sampled to control the effects of microhabitat on relative prey abundance. Our results revealed different activity patterns of both predators and prey near motorways that are independent of structural differences in microhabitat. Both the red fox and small mammals were found to use the zone close to the motorways more frequently, whereas lagomorphs and mustelids were less active there. These differences suggest that motorways favor the population of the predator that is most tolerant of human activity, the red fox, whose activity could have both direct and indirect effects on that of other members of the predator and prey community. On the one hand, the red fox seems to act as “top predator” and mustelids to follow a “safety match” strategy avoiding the area close to the motorway where fox is more active. On the other hand, abundances of prey species are negatively associated with the activity of their most frequent predators. This study is the first to assess how the proximity to motorways affects the activity of mammals in two levels of the food web and opens the field for research to understand the processes driving the detected patterns. Moreover, such effects at the community scale should be taken into account when evaluating the impacts of motorways on the surrounding ecosystems.     

Evaluating the effects of trophic complexity on a keystone predator by disassembling a partial intraguild predation food web

1. Many taxa can be found in food webs that differ in trophic complexity, but it is unclear how trophic complexity affects the performance of particular taxa. In pond food webs, larvae of the salamander Ambystoma opacum occupy the intermediate predator trophic position in a partial intraguild predation (IGP) food web and can function as keystone predators. Larval A. opacum are also found in simpler food webs lacking either top predators or shared prey. 2. We conducted an experiment where a partial IGP food web was simplified, and we measured the growth and survival of larval A. opacum in each set of food webs. Partial IGP food webs that had either a low abundance or high abundance of total prey were also simplified by independently removing top predators and/or shared prey. 3. Removing top predators always increased A. opacum survival, but removal of shared prey had no effect on A. opacum survival, regardless of total prey abundance. 4. Surprisingly, food web simplification had no effect on the growth of A. opacum when present in food webs with a low abundance of prey but had important effects on A. opacum growth in food webs with a high abundance of prey. Simplifying a partial IGP food web with a high abundance of prey reduced A. opacum growth when either top predators or shared prey were removed from the food web and the loss of top predators and shared prey influenced A. opacum growth in a non-additive fashion. 5. The non-additive response in A. opacum growth appears to be the result of supplemental prey availability augmenting the beneficial effects of top predators. Top predators had a beneficial effect on A. opacum populations by reducing the abundance of A. opacum present and thereby reducing the intensity of intraspecific competition. 6. Our study indicates that the effects of food web simplification on the performance of A. opacum are complex and depend on both how a partial IGP food web is simplified and how abundant prey are in the food web. These findings are important because they demonstrate how trophic complexity can create variation in the performance of intermediate predators that play important roles in temporary pond food webs.