Partitioning mechanisms of Predator Interference in different Habitats

Prey are often consumed by multiple predator species. Predation rates on shared prey species measured in isolation often do not combine additively due to interference or facilitation among the predator species. Furthermore, the strength of predator interactions and resulting prey mortality may change with habitat type. We experimentally examined predation on amphipods in rock and algal habitats by two species of intertidal crabs, Hemigrapsus sanguineus (top predators) and Carcinus maenas (intermediate predators). Algae provided a safer habitat for amphipods when they were exposed to only a single predator species. When both predator species were present, mortality of amphipods was less than additive in both habitats. However, amphipod mortality was reduced more in rock than algal habitat because intermediate predators were less protected in rock habitat and were increasingly targeted by omnivorous top predators. We found that prey mortality in general was reduced by (1) altered foraging behavior of intermediate predators in the presence of top predators, (2) top predators switching to foraging on intermediate predators rather than shared prey, and (3) density reduction of intermediate predators. The relative importance of these three mechanisms was the same in both habitats; however, the magnitude of each was greater in rock habitat. Our study demonstrates that the strength of specific mechanisms of interference between top and intermediate predators can be quantified but cautions that these results may be habitat specific. An erratum to this article can be found at     

Estimating the prevalence and strength of non-independent predator effects

Understanding whether multiple predator species have independent effects on shared prey is critical for understanding community dynamics. We describe the prevalence and strength of non-independence between predators by quantifying the prey’s risk of predation and the degree to which it deviates from the risk predicted from a null model of independent predator effects. Specifically, we document how frequently non-independent effects occur among ten different multiple predator combinations with mayfly larvae as prey. These predator combinations vary both predator density and predator species richness. Overall, the predator effects were non-independent and translated to an average of 27% fewer prey being consumed compared to independent predator effects. Non-independence of this magnitude is likely to have population level consequences for the prey and influence the distribution or prey preference of predators. Closer inspection shows that much of the risk reduction in this system is weak, to the point of being indistinguishable from independent predator effects, while few effects are strong. This pattern of many weak interactions and few strong ones parallels the pattern of interaction strengths documented previously in intertidal communities. Consequently, understanding strong interactors in multiple predator systems may help us understand the importance of a species.     

Invasive species cause large-scale loss of native California oyster habitat by disrupting trophic cascades

Although invasive species often resemble their native counterparts, differences in their foraging and anti-predator strategies may disrupt native food webs. In a California estuary, we showed that regions dominated by native crabs and native whelks have low mortality of native oysters (the basal prey), while regions dominated by invasive crabs and invasive whelks have high oyster mortality and are consequently losing a biologically diverse habitat. Using field experiments, we demonstrated that the invasive whelk’s distribution is causally related to a large-scale pattern of oyster mortality. To determine whether predator–prey interactions between crabs (top predators) and whelks (intermediate consumers) indirectly control the pattern of oyster mortality, we manipulated the presence and invasion status of the intermediate and top trophic levels in laboratory mesocosms. Our results show that native crabs indirectly maintain a portion of the estuary’s oyster habitat by both consuming native whelks (density-mediated trophic cascade) and altering their foraging behavior (trait-mediated trophic cascade). In contrast, invasive whelks are naive to crab predators and fail to avoid them, thereby inhibiting trait-mediated cascades and their invasion into areas with native crabs. Similarly, when native crabs are replaced with invasive crabs, the naive foraging strategy and smaller size of invasive crabs prevents them from efficiently consuming adult whelks, thereby inhibiting strong density-mediated cascades. Thus, while trophic cascades allow native crabs, whelks, and oysters to locally co-exist, the replacement of native crabs and whelks by functionally similar invasive species results in severe depletion of native oysters. As coastal systems become increasingly invaded, the mismatch of evolutionarily based strategies among predators and prey may lead to further losses of critical habitat that support marine biodiversity and ecosystem function. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Predator diversity and density affect levels of predation upon strongly interactive species in temperate rocky reefs

Indirect effects of predators in the classic trophic cascade theory involve the effects of basal species (e.g. primary producers) mediated by predation upon strongly interactive consumers (e.g. grazers). The diversity and density of predators, and the way in which they interact, determine whether and how the effects of different predators on prey combine. Intraguild predation, for instance, was observed to dampen the effects of predators on prey in many ecosystems. In marine systems, species at high trophic levels are particularly susceptible to extinction (at least functionally). The loss of such species, which is mainly attributed to human activities (mostly fishing), is presently decreasing the diversity of marine predators in many areas of the world. Experimental studies that manipulate predator diversity and investigate the effects of this on strongly interactive consumers (i.e. those potentially capable of causing community-wide effects) in marine systems are scant, especially in the rocky sublittoral. I established an experiment that utilised cage enclosures to test whether the diversity and density of fish predators (two sea breams and two wrasses) would affect predation upon juvenile and adult sea urchins, the most important grazers in Mediterranean sublittoral rocky reefs. Changes in species identity (with sea breams producing major effects) and density of predators affected predation upon sea urchins more than changes in species richness per se. Predation upon adult sea urchins decreased in the presence of multiple predators, probably due to interference competition between sea breams and wrasses. This study suggests that factors that influence both fish predator diversity and density in Mediterranean rocky reefs (e.g. fishing and climate change) may have the potential to affect the predators' ability to control sea urchin population density, with possible repercussions for the whole benthic community structure.     

Food quality, nutrient limitation of secondary production, and the strength of trophic cascades

Recent meta‐analyses confirm that the strength of trophic cascades (indirect positive effects of predators on plant biomass through control of herbivores) varies among ecosystem types. In particular, most terrestrial systems show smaller cascades than most aquatic ones. Ecologists still remain challenged to explain this variation. Here, we examine a food quality hypothesis which states that higher quality plants should promote stronger trophic cascades. Food quality involves two components: digestion resistance of plants and magnitude of stoichiometric imbalance between plants and herbivores (where stoichiometry involves ratios of nutrient:carbon ratio of tissues). Both factors vary among ecosystems and could mediate conversion efficiency of plants into new herbivores (and hence control of plants by herbivores). We explored the food quality hypothesis using two models, one assuming that plant stoichiometry is a fixed trait, the other one allowing this trait to vary dynamically (but with a minimal nutrient:carbon ratio of structural mass). Both models produce the same suite of results. First, as expected, systems with more easily digested plants promote stronger cascades. Second, contrary to expectations, higher (fixed or minimal) nutrient:carbon ratio of plants do not promote stronger cascades, largely because of the net result of ecosystem feedbacks. Still, the model with dynamic stoichiometry permits positive correlations of realized plant nutrient:carbon ratio and cascade strength (as predicted), mediated through digestion resistance. Third, lower nutrient:carbon ratio of herbivores promotes stronger cascades. However, this result likely cannot explain variation in cascade strength because nutrient:carbon stoichiometry of herbivores does not vary greatly between terrestrial and aquatic ecosystems. Finally, we found that predation promotes nutrient limitation of herbivores. This finding highlights that food web processes, such as predation, can influence stoichiometry‐mediated interactions of plants and herbivores.     

Community impacts of two invasive crabs: the interactive roles of density, prey recruitment, and indirect effects

Assessing the implications of species invasion for native communities requires determining whether effects of invaders are novel, or are redundant with effects of species that are already present. Using a pair of field experiments conducted over two successive years, we examined factors that influence community impacts of a recent predatory crab invader (Hemigrapsus sanguineus) and a previously established invasive crab (Carcinus maenas) on New England coasts. We demonstrate that effects of these species differ temporally with changes in the ambient prey community, and are influenced by density differences between the two species and by different strengths and types of indirect effects that each elicits. Our study highlights the importance of including bottom-up processes (i.e., prey recruitment) when examining the redundancy of consumers.     

Energy gradients and the geographic distribution of local ant diversity

Geographical diversity gradients, even among local communities, can ultimately arise from geographical differences in speciation and extinction rates. We evaluated three models—energy-speciation, energy-abundance, and area—that predict how geographic trends in net diversification rates generate trends in diversity. We sampled 96 litter ant communities from four provinces: Australia, Madagascar, North America, and South America. The energy-speciation hypothesis best predicted ant species richness by accurately predicting the slope of the temperature diversity curve, and accounting for most of the variation in diversity. The communities showed a strong latitudinal gradient in species richness as well as inter-province differences in diversity. The former vanished in the temperature-diversity residuals, suggesting that the latitudinal gradient arises primarily from higher diversification rates in the tropics. However, inter-province differences in diversity persisted in those residuals—South American communities remained more diverse than those in North America and Australia even after the effects of temperature were removed.     

Vascular plant diversity and climate change in the alpine zone of the Lefka Ori,Crete

The aim of this study is to analyse the vascular flora and the local climate along an altitudinal gradient in the Lefka Ori massif Crete and to evaluate the potential effects of climate change on the plant diversity of the sub-alpine and alpine zones. It provides a quantitative/qualitative analysis of vegetation-environment relationships for four summits along an altitude gradient on the Lefka Ori massif Crete (1664–2339 m). The GLORIA multi-summit approach was used to provide vegetation and floristic data together with temperature records for every summit. Species richness and species turnover was calculated together with floristic similarity between the summits. 70 species were recorded, 20 of which were endemic, belonging to 23 different families. Cretan endemics dominate at these high altitudes. Species richness and turnover decreased with altitude. The two highest summits showed greater floristic similarity. Only 20% of the total flora recorded reaches the highest summit while 10% is common among summits. Overall there was a 4.96°C decrease in temperature along the 675 m gradient. Given a scenario of temperature increase the ecotone between the sub-alpine and alpine zone would be likely to have the greatest species turnover. Southern exposures are likely to be invaded first by thermophilous species while northern exposures are likely to be more resistant to changes. Species distribution shifts will also depend on habitat availability. Many, already threatened, local endemic species will be affected first.     

Positive effects of plant diversity on soil microbial biomass and activity are associated with more root biomass production

This study aims to explore relationships between plant diversity and soil microbial function and the factors that mediate the relationships. Artificial plant communities (1, 2, 4 and 8 species) were established filled with natural and mine tailing soils, respectively. After 12 months, the plant species richness positively affected the soil microbial functional diversity in both soil environments but negatively affected microbial biomass and soil basal respiration in the natural soil. The root biomass positively correlated with the microbial biomass, cultural bacterial activity and soil basal respiration in both soil environments. Moreover, the D_i (deviations between observed performances and expected performances from the monoculture performance of each species of mixture) of microbial biomass, cultural bacterial activity and soil basal respiration positively correlated with the D_i of root biomass in both soil environments. Consistent with stress-gradient hypothesis, the D_mix (over-function index) of aboveground biomass positively correlated plant species richness in the mine tailing soil. Results suggest that the root biomass production is an important mechanism that affects the effects of plant diversity on soil microbial functions. Different responses of soil microbial function to increasing plant diversity may be due to root biomass production mediated by other factors.     

Parasites,info-disruption,and the ecology of fear

There is growing interest in the ecological consequences of fear, as evidenced by the numerous studies on the nonconsumptive, trait-mediated effects of predators. Parasitism, however, has yet to be fully integrated into research on the ecology of fear, despite it having direct negative and often lethal effects on hosts and being the most common life history strategy on the planet. This might at least be partly due to the traditional, but untested, assumption that anti-parasite responses are weak relative to anti-predator responses. To test this hypothesis, we quantified the activity and location responses of Bufo americanus tadpoles to one of six chemical cues: water; cercariae of Echinostoma trivolvis, a trematode which infects and can kill amphibians; a snail releasing E. trivolvis cercariae; an uninfected snail; food; or conspecific alarm chemicals signaling predation. There is also literature encouraging research on the context dependency and pollution-induced disruption of fear responses. Consequently, before quantifying responses to the chemical cues, half of the B. americanus were exposed to the herbicide atrazine (201 μg/l for 4 days), a reported inhibitor of fear responses in fish. Tadpoles were attracted to food, were indifferent to an uninfected snail, avoided alarm chemicals, and exhibited avoidance and elevated activity in response to a snail shedding cercariae and cercariae alone. Atrazine had no detectable effects on B. americanus’ responses to the tested cues despite the use of a higher concentration and longer exposure duration than has been repeatedly shown to inhibit chemical cue detection in fish. The magnitude of anti-parasite and anti-predator responses were qualitatively similar, suggesting that the fear of disease and its ecological consequences could be comparable to that of predation. Consequently, we call for a greater integration of parasites into research on the ecology of fear and trait-mediated indirect effects.     

Plant species diversity is correlated with climatic factors differently at the community and the functional group levels: A case study of desert steppe in Inner Mongolia,China

In the desert steppe of Inner Mongolia, mean annual precipitation was found to be the dominant factor that correlated positively with species richness at the community level and for the perennial forb functional group. Shrub and semi-shrub functional group exhibited positive correlation with mean temperature of the coldest month.     

Potential impacts of climate change on the distributions and diversity patterns of European mammals

The Intergovernmental Panel on Climate Change (IPCC) predicts an increase in global temperatures of between 1.4°C and 5.8°C during the 21st century, as a result of elevated CO₂ levels. Using bioclimatic envelope models, we evaluate the potential impact of climate change on the distributions and species richness of 120 native terrestrial non-volant European mammals under two of IPCC’s future climatic scenarios. Assuming unlimited and no migration, respectively, our model predicts that 1% or 5–9% of European mammals risk extinction, while 32–46% or 70–78% may be severely threatened (lose > 30% of their current distribution) under the two scenarios. Under the no migration assumption endemic species were predicted to be strongly negatively affected by future climatic changes, while widely distributed species would be more mildly affected. Finally, potential mammalian species richness is predicted to become dramatically reduced in the Mediterranean region but increase towards the northeast and for higher elevations. Bioclimatic envelope models do not account for non-climatic factors such as land-use, biotic interactions, human interference, dispersal or history, and our results should therefore be seen as first approximations of the potential magnitude of future climatic changes.     

Peracarid composition,diversity and species richness in the area of the Northeast Water polynya,East Greenland (Crustacea,Malacostraca)

During the ARK XI-2 expedition with RV Polarstern in September/October 1995, a transect of epibenthic sledge (EBS) samples was taken in the area of the Northeast Water Polynya off the Greenland coast, from the shelf down into the deep sea. A total of 85,304 specimens of Peracarida were collected at seven stations. These individuals comprised 45 families, 103 genera and 180 species. With regard to abundance, Cumacea occurred with highest numbers, 31,269, followed by Isopoda, Amphipoda, Mysidacea and Tanaidacea. Species richness was highest in Amphipoda with 94 species, followed by Isopoda with 43 species, Cumacea with 20 species, Tanaidacea with 15 and Mysidacea with 8 species. Species richness was highest at the shallowest station 37-016 and lowest at the deepest station 37-021, whereas the opposite pattern was found for abundance. Diversity and eveness were highest at the southernmost station and lowest at the deepest station. Amphipoda occurred more frequently at the shallower stations, while Cumacea were very frequent at the deepest station. Numbers of species were lowest for Cumacea at the deepest station, while Amphipoda and Isopoda generally occurred with high species richness at all stations. On the basis of the species composition stations were compared.     

Different effects of coarse woody material on the species diversity of three saproxylic beetle families (Cerambycidae, Melandryidae, and Curculionidae)

The relationships between coarse woody material (CWM) and the diversity of three saproxylic beetle families (Cerambycidae, Melandryidae, and Curculionidae) were investigated. These three families responded differently toward the quality and quantity of CWM. The species richness of cerambycid beetles increased in forests with more CWM in the early stage of decomposition. The richness of saproxylic curculionids increased in stands with more tree species. No clear trend was evident for melandryid diversity. The CWM differed in quality and quantity among three forest types [middle-aged larch (Larix kaempferi [Lamb.] Carrière) plantation, secondary forest, and old-growth forest] and between two forest-management practices (thinning and long rotation). Forest type and management practices affected CWM and the number of tree species, and CWM and the number of tree species influenced the diversity of saproxylic families differently. Because each decomposition stage is considered to be important for saproxylic beetles at the species level, a constant supply of CWM is necessary to maintain a suitable balance of CWM through these decomposition stages in forests to maintain the diversity of saproxylic beetles.     

Littoral species diversity and biomass: concordance among organismal groups and the effects of environmental variables

Biomass and species diversity (richness and evenness) of littoral organisms were explored in 27 sites in three basins of the large Lake Saimaa system in eastern Finland. The basins differed in degree of nutrient loading and trophic status. Six organismal groups, i.e., phytoplankton, periphyton, macrophytes, crustacean zooplankton, benthic macroinvertebrates and fish were studied. Factors affecting the biomass and diversity of these groups were explored by multiple stepwise regression analysis. The biomass of different groups was explained by the same variables, mainly nutrients, while diversity was associated with different environmental factors among the studied groups. The biomass of periphyton, phytoplankton, zooplankton, and fish correlated significantly with each other. There was also an apparent association between the biomass of macrophytes and that of benthic invertebrates. However, no significant correlations were found among the diversity of the studied groups. In accordance with previous studies, our results did not support the existence of species-rich hotspots or the possibility of using any surrogate taxon to reveal overall biodiversity. Thus, for conservation planning, biological surveys should include extensive collection of taxonomic groups and organisms at all trophic levels.     

Habitat structure,resources and diversity: the separate effects of surface roughness and macroalgae on stream invertebrates

Habitat structure has pervasive effects on community composition and diversity, with physically complex habitats often containing more species than physically simple ones. What factors or mechanism drive this pattern is little understood, but a complicating problem is that different sources of habitat structure can be confounded in both surveys and experiments. In this study, we carried out an experiment in which two sources of habitat structure, attached macroalgae and substrate surface texture, were separately manipulated to discern their joint and separate effects upon the diversity and composition of colonizing macroinvertebrates in a stony, upland stream. Because stream algae vary markedly in abundance in both space and time, we also sampled the epilithon of stream stones at two spatial scales on eight dates over 2 years to gain some preliminary data on how stream algae vary between individual substrata over time. Experimental substrata had either a smooth (siltstones, sandstones, crystal-poor felsic volcanics, plain paving bricks) or rough (granodiorites, crystal-rich felsic volcanics, sand-blasted paving bricks) surface. We allowed these substrata to be colonized naturally by macroalgae, mostly the filamentous red alga Audouinella hermannii. Half of each of the rough and smooth substrata were selected at random and the macroalgae gently sheared off. All substrata were defaunated with a household insecticide with little field persistence, set out randomly through the study riffle, and invertebrates allowed to colonize them for 14 days. Some substrata were sampled immediately to check the efficacy of faunal and algal removals, which proved to be successful. Experimental results showed that both surface texture and macroalgae increase species richness independently of each other. Surface texture had no effect on densities, while macroalgae increased colonization densities, but rarefaction showed that both sources of habitat structure increased species richness above values expected simply on the basis of the numbers of colonists. However, reference stones with high macroalgal cover had the same species richness as those with low cover, suggesting that the effects of macroalgae on species richness are transient relative to those associated with surface texture. Epilithon samples taken at different times suggest that the magnitude of spatial variation in plant growth alters with time. If plants generally recolonize rough surfaces more quickly than smooth, then the effects of habitat structure on macroinvertebrates ought to be strongest after major disturbances during growing seasons of plants. Received: 1 September 1999 / Accepted: 10 January 2000     

Studies on the diversity of arbuscular mycorrhizal fungi and the efficacy of two native isolates in a highly alkaline anthropogenic sediment

A field survey of the arbuscular mycorrhizal status of herbaceous plant species was conducted in a highly alkaline anthropogenic sediment resulting from the disposal of waste from an acetylene and polyvinyl chloride factory. Most plant species found at the site were mycorrhizal and the dominant mycotrophic plant species was Conyza bilbaoana. Fungal species richness was assessed by identification of spores extracted from the sediment and from continuously propagated trap pot cultures. All of the six species of arbuscular mycorrhizal fungi (AMF) found were from the genus Glomus. Glomus intraradices and G. mosseae were found in field-collected sediment samples and also occurred most frequently in trap cultures. To test the symbiotic effectiveness of these two fungi, seedlings of C. bilbaoana were inoculated with either native G. intraradices BEG163 or G. mosseae BEG198 and non-native G. intraradices BEG75 or G. mosseae BEG25 isolates in sterile and non-sterile sediment collected from the study site. All four isolates were able to colonise C. bilbaoana. However, AMF native to the target sediments were generally more effective than the non-native fungi in promoting plant establishment and growth under highly alkaline conditions. The non-native G. intraradices was, however, more effective than the non-native G. mosseae. The results of this study suggest the use of adapted AMF as inoculants for phytorestoration of alkaline anthropogenic-stressed sediments.     

Elevational gradients of diversity for lizards and snakes in the Hengduan Mountains,China

Comparing elevational gradients across a wide spectrum of climatic zones offers an ideal system for testing hypotheses explaining the altitudinal gradients of biodiversity. We document elevational patterns of lizard and snake species richness, and explore how land area and climatic factors may affect species distributions of lizards and snakes. Our synthesis found 42 lizard species and 94 snake species known from the Hengduan Mountains. The lizards are distributed between 500 and 3500 m, and the snakes are distributed between 500 and 4320 m. The relationship between species richness and elevation for lizards and snakes is unimodal. Land area explains a significant amount of the variation in lizard and snake species richness. The cluster analysis reveals pronounced distinct assemblages for lizards and snakes to better reflect the vertical profiles of climate in the mountains. Climatic variables are strongly associated with lizard and snake richness along the elevational gradient. The data strongly implicate water availability as a key constraint on lizard species richness, and annual potential evapotranspiration is the best predictor of snake species richness along the elevational gradient in the Hengduan Mountains.     

Indirect and direct controls of macroinvertebrates and small fish by abiotic factors and trophic interactions in the Florida Everglades

1. The roles of nutrients, disturbance and predation in regulating consumer densities have long been of interest, but their indirect effects have rarely been quantified in wetland ecosystems. The Florida Everglades contains gradients of hydrological disturbance (marsh drying) and nutrient enrichment (phosphorus), often correlated with densities of macroinvertebrate infauna (macroinvertebrates inhabiting periphyton), small fish and larger invertebrates, such as snails, grass shrimp, insects and crayfish. However, most causal relationships have yet to be quantified. 2. We sampled periphyton (content and community structure) and consumer (small omnivores, carnivores and herbivores, and infaunal macroinvertebrates inhabiting periphyton) density at 28 sites spanning a range of hydrological and nutrient conditions and compared our data to seven a priori structural equation models. 3. The best model included bottom‐up and top‐down effects among trophic groups and supported top‐down control of infauna by omnivores and predators that cascaded to periphyton biomass. The next best model included bottom‐up paths only and allowed direct effects of periphyton on omnivore density. Both models suggested a positive relationship between small herbivores and small omnivores, indicating that predation was unable to limit herbivore numbers. Total effects of time following flooding were negative for all three consumer groups even when both preferred models suggested positive direct effects for some groups. Total effects of nutrient levels (phosphorus) were positive for consumers and generally larger than those of hydrological disturbance and were mediated by changes in periphyton content. 4. Our findings provide quantitative support for indirect effects of nutrient enrichment on consumers, and the importance of both algal community structure and periphyton biomass to Everglades food webs. Evidence for top‐down control of infauna by omnivores was noted, though without substantially greater support than a competing bottom‐up‐only model.