The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

Types of larval development in marine bottom invertebrates,their distribution and ecological significance: a re-evaluation

The distribution of various types of larval development among marine bottom invertebrates has been discussed on the basis of ecological evidence by Thorson (1936, 1946, 1950, 1952) and Mileikovsky (1961b, 1965). The information at hand is reviewed anew in this paper and is re-evaluated in the light of modern pertinent literature. The interrelationships between certain larval types and their distribution are not as rigid and direct as originally assumed. This can be proved even by the copy book example of the distribution of the various forms of development among species of the coastal gastropod genus Littorina. Especially among species with wide distributional areas, local populations may exhibit greater diversity in larval types than has previously been thought. Different types of larval development have now become known to exist in different populations of opisthobranch gastropods and lamellibranchs, i.e., in invertebrate groups in which such variability had been ruled out by Thorson. Variability in the type of larval development within given species — as a function of geographical, seasonal and other environmental parameters —is also more common in other marine bottom invertebrates than formerly considered. Marine bottom invertebrates are characterized not only by the 3 main different types of larval development proposed by Thorson (pelagic, direct, viviparous), but also by a fourth type: demersal (free non-pelagic) development. This fourth type occurs at all water depths and in all geographic zones of the oceans. The most important of the 4 types is pelagic (planktotrophic) development. Thorson's rule (decrease in numbers of species possessing pelagic development from the Equator towards the Poles, and from shallow-shelf waters to greater oceanic depths) is well substantiated by new data. However, one correction is necessary: pelagic development is not completely absent in the abyssal zone, as was proposed by Thorson (1950, and later), but is represented in it by at least several species belonging to various groups of invertebrates, and is also fairly common in the bathyal zone. A detailed analysis of the distributional pattern of the different types of development of marine bottom invertebrates must further take into consideration asexual reproduction with all its different modifications. Asexual reproduction in benthonic invertebrates is ecologically significant because of its common occurrence in nature; in numerous species it is also important as a biological supplement to sexual reproduction. The vast majority of species inhabiting the shallow-shelf zone and, partly, the higher levels of the slope zone of ocean areas located roughly between the polar circles, reveals development by means of planktotrophic larval stages. In the highest latitudes and on the slopes to abyssal depths—characterized by low water temperatures, scarcity of food, increasing hydrostatic pressure and other environmental peculiarities—other types of larval development prevail and, progressively, replace pelagic development with increasing latitude or depth. The distributional patterns of the various types of development among marine bottom invertebrates form one of the most important factors determining the basic distributional dynamics of the whole benthos in all oceans, both in the geological past and at the present time.Dedicated to the memory of Professor G. Thorson —founder of modern reproductive and larval ecology of marine bottom invertebrates.     

Palatability of invertebrate larvae to corals and sea anemones

Risk of larval mortality is an underlying theme in debates and models concerning the ecology and evolution of the differing reproductive characteristics among marine benthic invertebrates. In these discussions, predation is often assumed to be a major source of larval mortality. Previous studies, focused primarily on planktotrophic larvae, suggested that marine larvae generally were susceptible to, and poorly defended against, planktivorous fishes and invertebrates. Larval-planktivore interactions involving larger and more conspicuous lecithotrophic larvae that are typical of many brooding sessile invertebrates have not been well studied. This lack of data for diverse larval types has hindered testing broad generalities about marine larvae and planktivore prey-preferences. This study demonstrates that lecithotrophic larvae of many Caribbean and temperate western Atlantic invertebrates are distasteful to co-occurring corals and anemones. These larval predators frequently rejected larvae of sponges (6 of 9 species), gorgonians (7 of 9 species), corals (3 of 3 species), hydroids (2 of 2 species) and a bryozoan. Larvae of three temperate colonial ascidians were readily consumed. Frequencies of survivorship for larvae captured but rejected by corals and anemones were generally high and, in 20 of 24 assays, were not statistically different from those of unattacked control larvae. Levels of metamorphosis (when it occurred) of rejected larvae also rarely differed significantly from those of unattacked controls. These results provide further evidence that larval palatability to predators may not be as high as once thought, particularly for brooded larvae of sessile colonial invertebrates. The means by which larvae may avoid or deter predators, and the demographic consequences for marine invertebrates and for the evolution of invertebrate life-history patterns, need to be assessed.     

Underwater and above-water search patterns of an Arctic seabird: reduced searching at small spatiotemporal scales

How predators vary search patterns in response to prey predictability is poorly known. For example, marine invertebrates may be predictable but of low energy value, while fish may be of higher energy value but unpredictable at large (pelagic schools) or small (solitary benthics) spatial scales. We investigated the search patterns of the thick-billed murre (Uria lomvia), an Arctic seabird feeding on invertebrates, pelagic fish, or benthic fish. Foraging ranges at the Coats Island colony are generally smaller (<240 min per trip) than at larger colonies, and many birds specialize in foraging tactics and diet. Underwater search times for benthic fish were higher than for pelagic fish or invertebrates while above-water search times for pelagic fish were higher than for benthic fish or invertebrates. There were few stops during trips. Total trip time, flying time, number of flights, and number of dives were intercorrelated and increased with prey energy content, suggesting that longer trips involved fewer prey encounters due to selection of higher-quality, but rarer, prey items. Flight times were not Lévy-distributed and seabirds may have used area-restricted searches. The high degree of specialization, apparent absence of information center effects, and reduced above-water searching times may be linked to the relatively small colony size and the resulting short commuting distances to feeding areas, leading to greater prey predictability. We concluded that prey predictability over various scales affected predator search patterns.     

Stronger predation in the tropics shapes species richness patterns in marine communities

Species interactions are widely assumed to be stronger at lower latitudes, but surprisingly few experimental studies test this hypothesis, and none ties these processes to observed patterns of species richness across latitude. We report here the first experimental field test that predation is both stronger and has a disproportionate effect on species richness in the tropics relative to the temperate zone. We conducted predator-exclusion experiments on communities of sessile marine invertebrates in four regions, which span 32 degrees latitude, in the western Atlantic Ocean and Caribbean Sea. Over a three-month timescale, predation had no effect on species richness in the temperate zone. In the tropics, however, communities were from two to over ten times more species-rich in the absence of predators than when predators were present. While micro-and macro-predators likely compete for the limited prey resource in the tropics, micropredators alone were able to exert as much pressure on the invertebrate communities as the full predator community. This result highlights the extent to which exposure to even a subset of the predator guild can significantly impact species richness in the tropics. Patterns were consistent in analyses that included relative and total species abundances. Higher species richness in the absence of predators in the tropics was also observed when species occurrences were pooled across two larger spatial scales, site and region, demonstrating a consistent scaling relationship. These experimental results show that predation can both limit local species abundances and shape patterns of regional coexistence in the tropics. When preestablished diverse tropical communities were then exposed to predation for different durations, ranging from one to several days, species richness was always reduced. These findings confirmed that impacts of predation in the tropics are strong and consistent, even in more established communities. Our results offer empirical support for the long-held prediction that predation pressure is stronger at lower latitudes. Furthermore, we demonstrate the magnitude to which variation in predation pressure can contribute to the maintenance of tropical species diversity.     

Patterns of selective predation by juvenile,benthivorous fish on estuarine macrofauna

Benthic feeding on macrofauna was studied in juveniles of the sparids Lithognathus lithognathus and Rhabdosargus holubi in the upper reaches of the Gamtoos Estuary, South Africa. Fish and benthic macrofauna were sampled simultaneously, and the selection of invertebrate prey assessed. Both fish species strongly selected for corophioid amphipods and consumed other benthic taxa in low numbers. R. holubi also exploited aquatic autotrophs, while L. lithognathus had a narrow prey-spectrum, feeding almost exclusively on the tube-dwelling amphipod Grandidierella lignorum. G. lignorum was the most abundant prey species, both in the benthos and the fish's diet. This species also showed prominent behavioural differences between the sexes; males were markedly more active on the sediment surface than females, who rarely left their tubes during the day. Males switched from an infaunal to epifaunal microhabitat in search of receptive females, concurrently increasing their exposure to fish predators. Consequently, L. lithognathus selected significantly more males than female amphipods, causing a marked bias towards females in the sex ratio and age-structure of the amphipod population. Juvenile amphipods were less preyed upon, presumably as a result of lower prey-detection or capture efficiency by the predators. Accepting current notions about predation as an important structuring element for benthic communities, our data also stress the prominence of size-and sex-selective predation in structuring individual prey populations.     

Post-settlement mortality of juvenile lagoonal cockles (Cerastoderma glaucum : Mollusca: Bivalvia)

Like many benthic marine invertebrates the lagoonal cockle Cerastoderma glaucum Bruguière (Mollusca: Bivalvia) suffers extensive mortality between settling out of the plankton and reaching adolescence. It is thought that predation could be a major cause of this mortality. Experiments were conducted in aquaria to assess the influence of predation on the survival of juvenile lagoonal cockles. Newly settled spat (collected from Holkham Salts Hole, a lagoon in North Norfolk) were enclosed with potential predators at field densities. Predators used were fish, prawns and polychaetes (singly or combination). The numbers and sizes of the surviving cockles were measured after a 4-week period to establish the extent of predation, and the results were consistent with predation by the fish (Pomatoschistus microps) being a major causative agent of the mortality seen in the field. This reinforces the idea of epibenthic predation as an important structuring element for marine benthic communities. The effect of post-settlement juvenile mortality on cockle population demographics is considered, and other possible causes of mortality in the field are discussed. Received: 1 February 1998 / Accepted: 2 April 1998     

Larvae of a colonial ascidian use a non-contact mode of substratum selection on a coral reef

The rate at which larvae successfully recruit into communities of marine benthic invertebrates is partially dependent upon how well larvae avoid benthic predators and settle on appropriate substrata. Therefore, to be able to predict recruitment success, information is needed on how larvae search for settlement sites, whether larvae preferentially settle on certain substrata, and the extent to which there are adequate cues for larvae to find these substrata. This article describes how larvae of the colonial ascidian Diplosoma similis find settlement sites on a coral reef. Direct field observations of larval settlement were made on a fringing reef in Kaneohe Bay, Oahu, Hawaii, between September 1985 and April 1986. A comparison of the substrata that larvae contacted prior to settlement relative to the percentage cover of these substrata on the study reef suggests that larvae are using a non-contact mode of substratum identification to locate suitable settlement sites. This mode of substratum identification allowed 74% of larvae to evade predation by benthic organisms who would otherwise have eaten larvae if they had been contacted. Of those larvae that evaded predation, 88% subsequently settled on the same two substrata upon which most adults are found (dead coral or the green alga Dictyosphaeria cavernosa). This pattern of settlement was probably a result of active selection, since the two substrata cover only 14.4% of the reef's surface and currents had little effect on the direction in which larvae swam. An important contributing factor to the high success rate of larval settlement on suitable substrata was the lack of any temporal decay in substratum preference. It is concluded that for Diplosoma similis larval supply is a sufficient predictor of larval settlement rate. However, for marine invertebrates whose larvae are passively dispersed and exhibit a greater temporal decay in substratum preference, larval settlement should generally have a greater dependency on spatial variation in the abundance of benthic predators and suitable substrata.     

The influence of pollution on pelagic larvae of bottom invertebrates in marine nearshore and estuarine waters

At present, human activities on marine shores, nearshore waters and estuaries, as well as in rivers which discharge into the seas, are influencing nearshore and estuarine waters so strongly that the resultant changes affect the breeding and spawning of bottom invertebrates which inhabit these waters. The majority of bottom invertebrate species from the shallow shelf of all seas and oceans, and from all except the highest latitudes, pass through a pelagic larval phase during development. Thus, survival and maintenance of such species are determined by survival and distribution of larvae during their planktonic phase and their settling on the substrate. Consequently, an attempt to summarize all available data on the influence of pollution in marine and estuarine waters, not only on adult bottom invertebrates, but also on their pelagic larvae, will be of practical interest from both scientific and economic points of view. This paper represents a first step towards that goal. Despite the fact that all aspects of pollution-industrial wastes (including heated effluents producing thermal pollution), domestic sewage, oil and oil products, oil-spill removers and oil-emulsifiers, various pesticides, synthetic surfactants, etc.—exert harmful effects on pelagic larvae of bottom invertebrates under experimental conditions, in natural environments, free-swimming larvae are influenced only slightly by these pollutants. Pollution becomes a grave danger for larvae when they are settling on the substrate, as much higher concentrations of different pollutants may be present on the substrate than in the water mass above it (the pollution of every nearshore water mass usually varies greatly from place to place). Pelagic larvae of bottom invertebrates inhabiting comparatively clean parts of polluted regions are dispersed throughout all such regions by local currents, tidal oscillations, eddies and other small-scale water movements. They represent, therefore, the potential source for re-establishment of the normal composition of the bottom communities in these regions, after the abatement of pollution by means of natural causes or by man's own improvements.     

Seasonal and daily dynamics in pelagic larvae of marine shelf bottom invertebrates in nearshore waters of Kandalaksha Bay (White Sea)

Seasonal and daily population dynamics have been studied in pelagic larvae of littoral and upper-sublittoral bottom invertebrates in the plankton of the shallow, narrow Velikaya Salma Sound, which connects the inner and outer areas of the Kandalaksha Bay in the western part of the White Sea. Hydrologically, this Sound is characterised by a clearly defined cycle of great seasonal variations in water temperature coupled with more or less stable salinities and regular, pronounced semi-diurnal tides corresponding to daily and lunar monthly tidal cycles. The seasonal dynamics of larvae in the Sound reflect differences in occurrence of spawning periods in local waters of various species and systematic groups of bottom invertebrates. These differences are caused by the correlation of spawning periods of local species of different zoogeographical origin with the different water temperatures. They reflect, also, lunar periodicities of spawning and larval hatchings. The daily dynamics of larval abundancies are related to the daily spawning rhythms of many species with pelagic development affected by the daily tidal cycles of the Velikaya Salma Sound. A daily invasion of the Sound by pelagic larvae of bottom invertebrates from the inner and the outer parts of the Kandalaksha Bay occurs at ebb tide, and also at flood tide; the rhythms of the invasions coincide with the daily spawning rhythms of the Sound's invertebrates. From literature data summarized by Mileikovsky (1958a, b, 1960a, b, c, 1961, 1965, 1968, 1970), it is concluded that seasonal, lunar and daily (tidal) reproductive periodicities for the marine shallowshelf bottom invertebrates concerned, follow world-wide ecological patterns. It is evident that the effects of these rhythms upon the population dynamics of pelagic invertebrate larvae, as demonstrated by the present data on the Velikaya Salma Sound (White Sea), must also follow world-wide regularities.     

Feeding by larvae of intertidal invertebrates: assessing their position in pelagic food webs

One of the leading determinants of the structure and dynamics of marine populations is the rate of arrival of new individuals to local sites. While physical transport processes play major roles in delivering larvae to the shore, these processes become most important after larvae have survived the perils of life in the plankton, where they usually suffer great mortality. The lack of information regarding larval feeding makes it difficult to assess the effects of food supply on larval survival, or the role larvae may play in nearshore food webs. Here, we examine the spectrum of food sizes and food types consumed by the larvae of two intertidal barnacle species and of the predatory gastropod Concholepas concholepas. We conducted replicated experiments in which larvae were exposed to the food size spectrum (phytoplankton, microprotozoan and autotrophic picoplankton) found in nearshore waters in central Chile. Results show that barnacle nauplii and gastropod veligers are omnivorous grazers, incorporating significant fractions of heterotrophs in their diets. In accordance with their feeding mechanisms and body size, barnacle nauplii were able to feed on autotrophic picoplankton (<5 microm) and did not consume the largest phytoplankton cells, which made the bulk of phytoplankton biomass in spring-summer blooms. Balanoid nauplii exhibited higher ingestion rates than the smaller-bodied chthamaloid larvae. Newly hatched C. concholepas larvae also consumed picoplankton cells, while competent larvae of this species ingested mostly the largest phytoplankton cells and heterotrophic protozoans. Results suggest that persistent changes in the structure of pelagic food webs can have important effects on the species-specific food availability for invertebrate larvae, which can result in large-scale differences in recruitment rates of a given species, and in the relative recruitment success of the different species that make up benthic communities.     

Importance of jumbo squid Dosidicus gigas (Orbigny, 1835) in the pelagic ecosystem of the central Gulf of California

The Humboldt squid is an important predator in the pelagic ecosystem of the central Gulf of California and the commercial catch of this species has increased over the past decade, probable due to a decrease of several top predators (sharks, large pelagic fish and the marine mammals) and the optimal feeding conditions in this area. Its high abundance and important position in the pelagic food web was quantified through two trophic models of the pelagic ecosystem of the central Gulf of California. Models represented conditions in 1980 and 2002, to document the decadal changes in ecosystem structure and function. The models were composed of 18 functional groups, including marine mammals, birds, fish, mollusks, crustaceans, and primary producers. Model results show direct negative effects on principal prey groups such as myctophids and pelagic red crab and positive effects on sharks, marine mammals and specifically sperm whales. It thus appears that the jumbo squid has an important role in the ecosystem and plays a central part in the overall energy flow as main food item for most top predators, and due to its predation of organisms on lower tropic levels.     

A review of predator diet effects on prey defensive responses

Numerous studies have examined how predator diets influence prey responses to predation risk, but the role predator diet plays in modulating prey responses remains equivocal. We reviewed 405 predator–prey studies in 109 published articles that investigated changes in prey responses when predators consumed different prey items. In 54 % of reviewed studies, prey responses were influenced by predator diet. The value of responding based on a predator’s recent diet increased when predators specialized more strongly on particular prey species, which may create patterns in diet cue use among prey depending upon whether they are preyed upon by generalist or specialist predators. Further, prey can alleviate costs or accrue greater benefits using diet cues as secondary sources of information to fine tune responses to predators and to learn novel risk cues from exotic predators or alarm cues from sympatric prey species. However, the ability to draw broad conclusions regarding use of predator diet cues by prey was limited by a lack of research identifying molecular structures of the chemicals that mediate these interactions. Conclusions are also limited by a narrow research focus. Seventy percent of reviewed studies were performed in freshwater systems, with a limited range of model predator–prey systems, and 98 % of reviewed studies were performed in laboratory settings. Besides identifying the molecules prey use to detect predators, future studies should strive to manipulate different aspects of prey responses to predator diet across a broader range of predator–prey species, particularly in marine and terrestrial systems, and to expand studies into the field.     

Feeding strategies of some demersal fishes of the continental slope and rise off the Mid-Atlantic Coast of the USA

Stomach contents of 729 fishes comprising 16 species were examined from the continental slope and rise off the Middle Atlantic States of the USA. Two main feeding modes among demersal deep-sea fishes were evident: those feeding primarily on pelagic food items, and those feeding on benthic invertebrates. Both pelagic and benthic predators were euryphagous. Most pelagic predators also fed on the epibenthos. These findings support Dayton and Hessler's (1972) suggestion that benthic predators should have a generalized diet which may be responsible for the high diversity found in the deep-sea infauna. The mesopelagic fauna is an important food source for some demersal fishes on the continental slope. Pelagic prey, which are also important to ecologically dominant demersal fishes on the lower slope and continental rise, may be nutritionally supported by suspended particulate organic matter in a nepheloid layer close to the bottom, and they may exist in much higher concentrations than in the bathypelagic zone above.Virginia Institute of Marine Science Contribution No. 835.     

The “pelagic larvaton” and its role in the biology of the World Ocean,with special reference to pelagic larvae of marine bottom invertebrates

Ecological subdivision of marine organisms is often based on two characteristics: presence in a defined environment, and types of locomotion (degree of free active movement) in such an environment. The use of these characteristics results in a simple scheme: (1) Inhabitants of the boundary surface “ocean-atmosphere” (a zone including not only the surface film but also the thin subsurface water layer below it and the air layer just above it, i.e., pleuston and neuston). (2) Inhabitants of the deeper water layers of the ocean i.e., excluding the zone mentioned under (1): (a) passively drifting forms with very limited locomotory capacity, moving practically in the vertical plane only (plankton); (b) actively moving forms which migrate both vertically and horizontally (nekton). (3) Inhabitants of the “bottom”-benthos (level-bottom of oceans and coastal waters, tidal zones up to the upper supralittoral, different types of drifting and floating substrata, e.g. ship bottoms, harbour structures, buoys, driftwood, sargassum, whales, etc.). This simple scheme is essentially based on characteristics of adults. If developmental stages are considered, pelagic larvae of bottom invertebrates, eggs and larvae of fishes and other forms, usually present only temporarily in the plankton, neuston, and pleuston, can be distinguished as “mero-plankton”, “mero-neuston” and “mero-pleuston”, from the permanent “holo”-components of these groups. Division into “mero”-subgroups opposes all these larvae to those of planktonic, neustonic and pleustonic forms developing within the “parental” groups and their environments. However, the last category of larvae in the light of world-wide distribution of the seasonal reproductive pattern of marine invertebrates and some other organisms — especially in temperate and high latitudes — can also be rated to some degree as “mero”-(not “holo”-) components. The present paper proposes to unite all larvae of marine invertebrates (and of other organisms) undergoing pelagic development into one biological group, the “pelagic larvaton”. The main characteristic for all forms of this group is the presence of one and the same life-cycle stage in one and the same environment. All forms of the “pelagic larvaton” are, to various degrees, biologically different from their respective adult forms. Even the pelagic larvae of the holoplanktonic species exhibit some differences. Within the “pelagic larvaton”, 3 subgroups can be distinguished on the basis of their ecological peculiarities;

1. Larvae undergoing their whole development in an environment different from that inhabited by their parents and belonging to a group different from that of their parental forms; e.g. the pelagic larvae of bottom invertebrates which develop in the plankton, neuston or pleuston.
2. Larvae undergoing development in the same general pelagic environment, but in “non-parental” ecological groups; e.g. larvae of nektonic species developing in the plankton, neuston or pleuston; larvae of planktonic species in the neuston or pleuston; larvae of neustonic and pleustonic species in the plankton.
3. Larvae undergoing development in the “parental” groups; e.g. larvae of planktonic species in the plankton, of neustonic species in the neuston, or of pleustonic species in the pleuston.

In contrast to the 5 ecological groups: benthos, plankton, nekton, neuston and pleuston, the “pelagic larvaton” represents rather a biological than an ecological group. The “pelagic larvaton” comprises the 5 ecological groups and maintains the permanent turnover of organic substances between water and bottom. This group short-circuits the interrelations between the 5 ecological groups in all possible combinations. The existence of the “pelagic larvaton” presents another illustration of the unity of the biological nature of the oceans. The present paper also discusses the specific distributional patterns of the pelagic larvae of bottom invertebrates and their biological role in the seas.     

Predator identity and additive effects in a treehole community

Multiple predator species can interact as well as strongly affect lower trophic levels, resulting in complex, nonadditive effects on prey populations and community structure. Studies of aquatic systems have shown that interactive effects of predators on prey are not necessarily predictable from the direct effects of each species alone. To test for complex interactions, the individual and combined effects of a top and intermediate predator on larvae of native and invasive mosquito prey were examined in artificial analogues of water-filled treeholes. The combined effects of the two predators were accurately predicted from single predator treatments by a multiplicative risk model, indicating additivity. Overall survivorship of both prey species decreased greatly in the presence of the top predator Toxorhynchites rutilus. By itself, the intermediate predator Corethrella appendiculata increased survivorship of the native prey species Ochlerotatus triseriatus and decreased survivorship of the invasive prey species Aedes albopictus relative to treatments without predators. Intraguild predation did not occur until alternative prey numbers had been reduced by approximately one-half. Owing to changes in size structure accompanying its growth, T. rutilus consumed more prey as time progressed, whereas C. appendiculata consumed less. The intermediate predator, C. appendiculata, changed species composition by preferentially consuming A. albopictus, while the top predator, T. rutilus, reduced prey density, regardless of species. Although species interactions were in most cases predicted from pairwise interactions, risk reduction from predator interference occurred when C. appendiculata densities were increased and when the predators were similarly sized.     

Predator-induced morphological defenses in marine zooplankton: a larval case study

While there are numerous reports of predator-induced morphological defenses for freshwater zooplankton, freshwater larvae, and benthic marine animals, a literature search revealed no reports of predator-induced morphological defenses for marine zooplankton. Rarity of predator-induced morphological defenses in marine zooplankton would imply a difference in predation risks compared to those experienced by freshwater organisms and benthic marine adults, whereas the presence of such plasticity in defenses would imply that risks are modified by developmental responses. This study reports a predator-induced change in defenses and vulnerability of a marine planktonic larva. Specifically, when reared in the presence of zoea larvae of Cancer spp., veliger larvae of the intertidal snail Littorina scutulata developed significantly smaller shell apertures and rounder shells than did cohort veligers reared in the absence of predator cues. Pairwise predation trials demonstrated that veligers reared with caged zoeas throughout development had greater survival than predator-naive veligers during short-term exposure to zoeas. The development of predator-induced morphological defenses by some marine larvae introduces a range of testable hypotheses on developmental plasticity that reduces vulnerability of planktonic larvae and other marine zooplankton to predators.     

Comparison of laboratory rates of predation of five species of marine fish larvae by three planktonic invertebrates: effects of larval size on vulnerability

Rates of predation by the invertebrates Aurelia aurita, Thysanoessa raschi and Euchaeta norvegica on larval stages of cod (Gadus morhua L.), flunder (Platichthys flesus L.), plaice (Pleuronectes platessa L.), herring (Clupea harengus L.), and turbot (Scophthalmus maximus L.) were determined. Experiments were conducted in late winter and early spring 1982 with predators collected in Loch Etive, Scotland and prey obtained from several locations in Great Britain. Early stages of the smallest species, cod, flounder and turbot, tended to be most vulnerable to all three predators, while the early stages of the larger species, plaice and herring, and older stages of all species, were less vulnerable. For all stages and species of larvae, predation rates by the three predators were most closely related to larval length and escape swimming speed. Larval length itself was closely correlated to indices of larval escape ability. Low predation rates on large larvae by E. norvegica could be due to handling difficulties, whereas for A. aurita and T. raschi these low rates were due to escape abilities of the larger larvae. Prey movement is an important stimulus eliciting predation in E. norvegica but not in A. aurita or T. raschi.     

Predators, prey, and transient states in the assembly of spatially structured communities

Ecological theory suggests that both dispersal limitation and resource limitation can exert strong effects on community assembly. However, empirical studies of community assembly have focused almost exclusively on communities with a single trophic level. Thus, little is known about the combined effects of dispersal and resource limitation on assembly of communities with multiple trophic levels. We performed a landscape-scale experiment using spatially arranged mesocosms to study effects of dispersal and resource limitation on the assembly dynamics of aquatic invertebrate communities with two trophic levels. We found that interplay between dispersal and resource limitation regulated the assembly of predator and prey trophic levels in these pond communities. Early in assembly, predators and prey were strongly dispersal limited, and resource (i.e., prey) availability did not influence predator colonization. Later in assembly, after predators colonized, resource limitation was the strongest driver of predator abundance, and dispersal limitation played a negligible role. Thus, habitat isolation affected predators directly by reducing predator colonization rate, and indirectly through the effect of distance on prey availability. Dispersal and resource limitation of predators resulted in a transient period in which predators were absent or rare in isolated habitats. This period may be important for understanding population dynamics of vulnerable prey species. Our findings demonstrate that dispersal and resource limitation can jointly regulate assembly dynamics in multi-trophic systems. They also highlight the need to develop a temporal picture of the assembly process in multi-trophic communities because the availability and spatial distribution of limiting resources (i.e., prey) and the distribution of predators can shift radically over time.     

Predator hunting mode and habitat domain alter nonconsumptive effects in predator-prey interactions

Predators can affect prey populations through changes in traits that reduce predation risk. These trait changes (nonconsumptive effects, NCEs) can be energetically costly and cause reduced prey activity, growth, fecundity, and survival. The strength of nonconsumptive effects may vary with two functional characteristics of predators: hunting mode (actively hunting, sit-and-pursue, sit-and-wait) and habitat domain (the ability to pursue prey via relocation in space; can be narrow or broad). Specifically, cues from fairly stationary sit-and-wait and sit-and-pursue predators should be more indicative of imminent predation risk, and thereby evoke stronger NCEs, compared to cues from widely ranging actively hunting predators. Using a meta-analysis of 193 published papers, we found that cues from sit-and-pursue predators evoked stronger NCEs than cues from actively hunting predators. Predator habitat domain was less indicative of NCE strength, perhaps because habitat domain provides less reliable information regarding imminent risk to prey than does predator hunting mode. Given the importance of NCEs in determining the dynamics of prey communities, our findings suggest that predator characteristics may be used to predict how changing predator communities translate into changes in prey. Such knowledge may prove particularly useful given rates of local predator change due to habitat fragmentation and the introduction of novel predators.