Predator (<Emphasis Type="Italic">Carcinus maenas</Emphasis>) nonconsumptive limitation of prey (<Emphasis Type="Italic">Nucella lapillus</Emphasis>) feeding depends on prey density and predator cue type

Predators often have nonconsumptive effects (NCEs) on prey. For example, upon detection of predator cues, prey can reduce feeding activities to hamper being detected by predators. Previous research showed that waterborne chemical cues from green crabs (Carcinus maenas, predator) limit the dogwhelk (Nucella lapillus, prey) consumption of barnacles regardless of dogwhelk density, even though individual predation risk for dogwhelks decreases with conspecific density. Such NCEs might disappear with dogwhelk density if dogwhelks feed on mussels, as mussel stands constitute better antipredator refuges than barnacle stands. Through a laboratory experiment, we effectively found that crab chemical cues limit the per-capita consumption of mussels by dogwhelks at low dogwhelk density but not at high density. The combination of tactile and chemical cues from crabs, however, limited the dogwhelk consumption of mussels at both dogwhelk densities. The occurrence of such NCEs at both dogwhelk densities could have resulted from tactile cues indicating a stronger predation risk than chemical cues alone. Overall, the present study reinforces the notions that prey evaluate conspecific density when assessing predation risk and that predator cue type affects their perception of risk.     

Behavioural components of prey-selection by dogwhelks, Nucella lapillus (L.), feeding on barnacles, Semibalanusbalanoides (L.), in the laboratory

Adult dogwhelks, maintained on mussels for 60 days before experimentation to reduce prior effects of ingestive conditioning and handling skills appropriate to barnacles, and thence deemed “inexperienced”, preferred the largest barnacles presented to them. Juvenile and adult dogwhelks, maintained on barnacles and deemed experienced, preferred barnacles of intermediate sizes that were correlated with the sizes of the dogwhelks. “Inexperienced” dogwhelks penetrated barnacles significantly more often by drilling than by prising, whereas experienced dogwhelks did the reverse. A predominant tendency to prise open barnacles was developed by previously “inexperienced” dogwhelks after they had eaten six to eight consecutive prey. Larger dogwhelks prised open greater proportions of larger barnacles than did smaller dogwhelks. Experienced adult dogwhelks prised open all barnacles of ≈2-mm opercular diameter but only 20% of those ≈ 6 mm in diameter. The preferred barnacles, 4–5 mm in opercular diameter, were of a size that the experienced adult dogwhelks could prise open in 50% of attacks.Dogwhelks inspected barnacles by crawling over them for ≈20 min when the prey were subsequently rejected and ≈30 min when subsequently eaten. Penetration time and ingestion time were linear functions of barnacle diameter, total handling time ranging from 4–26 h for barnacles of 2.5–2.7 mm opercular diameter. Experienced dogwhelks handled barnacles faster than “inexperienced” dogwhelks, largely because they used the quicker method of penetration by prising whenever possible. The yield of flesh per unit handling time was an accelerating function of barnacle size and, with experience, could be increased by a factor of ≈1.2 for barnacles of 7 mm opercular diameter to ≈2.0 for barnacles of 2 mm opercular diameter. The preference of experienced dogwhelks for barnacles somewhat smaller than the largest available, may have reflected the greater frequency with which these prey could be penetrated by the quicker method of prising. Minimizing handling time could be important in nature if there is a risk to feeding dogwhelks of being displaced by competitors. In the laboratory, 12% of dogwhelks where thus displaced.Barnacles of ≈5 mm opercular diameter were estimated to be slightly more profitable than mussels of ≈17.5-mm shell length to a dogwhelk experienced with barnacles, but less profitable to a dogwhelk that has recently fed extensively on mussels. Dogwhelks, therefore, might switch between diets predominately of barnacles or of mussels if prey of comparable profitabilities change drastically in their relative abundance on the shore.     

Using polyclonal antibodies to detect predation by dogwhelks Nucella lapillus (L.)

Interactions between predators and their multiple prey species can vary greatly among locations where they coexist. As a method to assess spatial variation in predation by intertidal dogwhelks on their dominant prey, immunoassays of dogwhelk gut contents from experimental populations and field collected individuals were evaluated using polyclonal antibodies raised separately to soluble proteins from Mytilus edulis L. mussels and Semibalanus balanoides (L.) barnacles. Both antisera produced strong reactions against their homologous antigens but no cross reactions between prey species. Experimental trials tested the critical hypothesis that prey species had equal detection intervals in dogwhelk guts. Two groups of 225 dogwhelks were starved for 14 days, provided with either mussels or barnacles for five days, and then sampled over 22 days. Independent immunoassays of dogwhelk gut contents against each antibody revealed a consistent, weak cross reaction between the anti-mussel antibody and dogwhelk gut tissues. After accounting for this cross reaction, the strength of immunoassays against both prey species declined exponentially and at similar rates. The proportions of dogwhelks that tested positive for their provided prey species declined linearly through time and were not significantly influenced by prey type. Prey were detectable throughout the sampled post-feeding period and were projected to have detection limits of 24.4 days (barnacles) and 26.5 days (mussels), demonstrating that immunoassay results are not biased by dissimilar prey detection intervals. Reactions against the antibody from the non-provided prey were time invariant and occurred at relatively low frequencies. Immunoassays of dogwhelks collected from five intertidal sites on Swans Island, Maine, USA revealed patterns similar to field observations, though immunoassays classified far fewer individuals as non-feeders and more as barnacle feeders than indicated by direct field observations. Unlike single observations, immunoassays also revealed the presence of both prey in dogwhelks from four sites, though most individuals tested positive for only a single prey type. Immunoassays facilitate concurrent collections of predation data from many individuals and will enable further local- to regional-scale assessments of dogwhelk predation at additional sites around the Gulf of Maine.     

Variation in barnacle recruitment over small scales: larval predation by adults and maintenance of community pattern

Over small spatial scales, variation in the density of settlers of benthic sessile species is the result of interactions among larval behavior, local hydrodynamic conditions, and the physical, chemical and biological characteristics of the benthic habitat. It has been shown repeatedly that adult benthic filter-feeders can consume larvae of their own and other species, but their effects on the distribution and abundance of recruits have rarely been demonstrated under natural conditions in the field, particularly on hard substrata. Here we experimentally quantified the effect of the large intertidal barnacle, Semibalanus cariosus (Pallas), on the density of recruits of three common barnacle species. The experiments were conducted at the peak of the barnacle recruitment season over three successive years, on the west coast of San Juan Island, Washington. A persistent and well documented community pattern in the mid intertidal zone of the study site is a sparse bed of adult S. cariosus with bare rock spaces essentially devoid of small barnacles among the large individuals. Field experiments consisted of small areas from which either all adult S. cariosus were killed leaving the shells attached to the rock, or live adult barnacles were left intact. Our results showed that over small spatial scales of a few to tens of centimeters, the large barnacle S. cariosus can interfere and significantly reduce net settlement and recruitment of conspecific as well as other barnacle species. Between 65 and 100% reduction in settlement could be attributed to larval predation by adults, as implied by barnacle settlement patterns on different treatments and by the presence of nauplius larvae in cirri and stomach contents of S. cariosus. The negative effect on barnacle settlement was consistent between years of relatively low barnacle recruitment, which appears to be the most common situation at the study site, but it disappeared on a year of unusually high recruitment, when settling larvae seem to have swamped the filtration ability of adult S. cariosus. The different barnacle species displayed contrasting settlement patterns on bare rock and on the lateral shells of the large barnacles, which appear to be a result of differences in larval behavior. Comparisons against the relative availability of these substrata in the experimental plots suggested that larvae of different species sample the benthic microhabitat in very different ways.     

Effects of early recruits on temperate sessile marine community composition depend on other species recruiting at the time

In many environments recruitment of dispersive propagules (e.g. seeds, spores and larvae) can vary from situations when particular taxa recruit in relative isolation to times when they recruit simultaneously with other, functionally quite different taxa. Differences in the identity and density of recruiting taxa can have important consequences on community structure, but it is still not clear how the effects of individual taxa on communities are modified when they recruit together with other species. Using an experimental approach we compared early development of a temperate marine sessile community after the recruitment of mixtures of botryllid ascidians and barnacles to that when barnacles or botryllid ascidians recruited alone. Communities exposed to recruitment of botryllid ascidians in isolation differed from those that received barnacles, a mixture of botryllids and barnacles or no recruitment in 2-week-old communities. These early differences were driven by higher abundances of the species that were present as initial recruits in experimental treatments. After 2 months communities also differed between barnacle and mixed recruitment treatments but not mixed and botryllid or botryllid and barnacle treatments. These differences were not directly due to differences in the abundances of our manipulated taxa but occurred because of two abundant arborescent bryozoans, Bugula dentata, which occupied more space in communities that initially received mixed recruitment than in those that received barnacle or no recruitment, and Zoobotryon verticillatum, which occupied more space in communities that initially received only barnacle recruitment than those that initially received botryllid or mixed recruitment. These effects did not persist, and communities did not differ after 6 months. These results suggest that, more generally, species may influence community dynamics differently when they recruit alongside other species than when they recruit in relative isolation.     

Generalizing from experiments: is predation strong or weak in the New England rocky intertidal?

Summary Petraitis (1990) recently critized previous generalizations regarding the effects of predation in the New England rocky intertidal region (e.g., Menge 1976; Lubchenco and Menge 1978). Contrary to Lubchenco's and my conclusions, Petraitis concluded that (1) barnacles and not mussels are the favored prey of dogwhelks and (2) barnacles and not dogwhelks control mussel abundances in the mid and low rocky intertidal region. I provide evidence that these criticisms are unwarranted. First, Lubchenco and I never claimed that diet composition reflected prey preference. Moreover, predators can influence prey abundance without preferring the prey. Hence, claims regarding preferences have no bearing on our conclusions. Second, Petraitis' experiments do not invalidate Lubchenco's and my experimental results. Reanalyses of our experimental data support the earlier conclusion that at wave-sheltered sites, whelks reduce the abundance of mussels independently of barnacle abundances. Further, at all but one of Lubchenco's and my study sites, predator densities were higher than at Petraitis' site. Thus, the absence of a predator effect in Petraitis's study was most likely due to low predator density rather than a lack of generality of our earlier results. This reevaluation therefore suggests that within a broader conceptual framework, Petraitis' apparently divergent results are actually consistent with ours.     

Enhanced condition in dogwhelks, Nucella lapillus (L.) living under mussel hummocks

Dogwhelks, Nucella lapillus eat both barnacles and mussels, Mytilus edulis. However, mussels are dangerous prey because they can snare and immobilize dogwhelks with their byssi. Despite this danger, adult dogwhelks are common inside mussel hummocks or stacks found on mature mussel beds. The whelks live in debris-filled chambers within the hummocks, unsnared by byssi. Evidence is presented to show that this is a highly favourable microhabitat for dogwhelks. Condition index (CI) measurements showed that mean CI for “hummock” dogwhelks was 20.99 in early spring, 29.65 in midsummer and 34.86 in early winter. In comparison, CI values for dogwhelks from nearby barnacle zones were 20.13, 25.03 and 24.66 respectively. The summer and winter CI values of “hummock” dogwhelks were enhanced to a highly significant extent. Data are also presented to show that shore elevation or dogwhelk shell height (over the size range of animals studied) had no confounding effect on this result. Predicted flesh and shell mass values for a standard 25 mm high whelk showed that occupation of hummocks was associated with a thin-walled shell morphology, and that the shell mass became proportionately lower in summer and early winter. A feeding experiment showed that CI rises slowly, even in dogwhelks fed to satiation on mussels. Dogwhelks were taken from the barnacle zone in March 1997 and either fed to satiation on Mytilus or starved, whilst being held in seawater at ambient seawater temperature. The CI of fed dogwhelks rose from 19.7 to 27.4 in 3 months, while that of starved Nucella fell from 19.7 to 16.3. In the meantime, the CI of dogwhelks in situ in the barnacle zone had risen from 19.7 to 23.8. Hence, 3 months' satiation ration had led to a nett 3.6 unit rise in CI; starvation to a nett 7.5 unit fall. All of the changes in CI were due to changes in flesh mass; shell mass was stable. Since “hummock” dogwhelks collected in early winter have a CI value about 10.2 units greater than barnacle zone dogwhelks, this indicates that “hummock” dogwhelks are long-term inhabitants of this microhabitat.     

Response of predatory snails to a novel prey following the geographic range expansion of an intertidal barnacle

During a geographic range expansion, the success of a species colonizing a new region may be influenced by how resident predators respond to this novel prey. The volcano barnacle Tetraclita rubescens has recently expanded its range > 300 km northward along the coast of California, USA. In the historic portion of its range, Tetraclita is preyed upon by a southern dogwhelk, Nucella emarginata. In newly colonized northern habitats, Tetraclita has joined an assemblage of mid-intertidal barnacles preyed upon by two species of northern dogwhelks (N. ostrina and N. canaliculata). Using a combination of field enclosures and laboratory experiments, we tested the impacts of northern whelks on volcano barnacles relative to other barnacle species. In the field, juvenile Tetraclita were rarely consumed. In laboratory trials with larger barnacles, Tetraclita were nearly immune to drilling by northern whelks, perhaps due to superior morphological defenses. In all experiments, prior encounters with Tetraclita did not appear to increase the frequency of predation on this barnacle; northern whelk species rarely preyed on Tetraclita even when collected from a site where this barnacle was abundant. In contrast, the southern whelk N. emarginata readily preyed on Tetraclita in laboratory trials, perhaps reflecting a longer shared evolutionary history between these species. Overall, our results suggest that the success of a population established beyond a species' historical range boundary may be influenced by the functional traits of that species in relation to the composition and evolutionary history of the surrounding community.     

Prey risk assessment depends on conspecific density

In many systems, the number of prey killed by predators increases with prey density. This in turn generates higher levels of the indirect signals that prey use to assess predation risk. A model developed by Peacor (2003) showed that prey that respond to predator cues without accounting for conspecific density will consistently over‐ or under‐estimate risk and therefore invest improperly in anti‐predator defense. We tested this model using Rana temporaria tadpoles as prey and Aeshna cyanea dragonfly larvae as predators. As assumed by the model, prey reduced risky activity with increasing concentrations of predator kairomones and increased activity at high prey density. However, prey did not react to changes in cue or density if the ratio of cue‐to‐density remained constant. Prey therefore monitored their per capita risk, strongly supporting Peacor's model.     

Desiccation,predation, and mussel-barnacle interactions in the northern Gulf of California

Summary Field experiments were conducted in order to determine the potential for desiccation and predation to mediate the effect of mussels (Brachidontes semilaevis) on barnacles (Chthamalus anisopoma) in the highly seasonal northern Gulf of California. We did this by removing both mussels and a common mussel predator (Morula ferruginosa: Gastropoda) and by spraying selected sites with sea water during summertime spring low tides. We also determined the effect of crowding on resistance to desiccation in barnacles, and the effect of barnacles on colonization by mussels. The mussel-barnacle community was not affected by keeping experimental quadrats damp during daytime low tides throughout the summer. Exposure to summertime low tides, however, did affect the survivorship of isolated, but not crowded, barnacles; and barnacle clumps enhanced the recruitment of mussels. Hence crowding in barnacles had a positive effect on both barnacle survivorship and mussel recruitment. Morula had a negative effect on mussel density, and mussels had a negative effect on barnacle density. The effect of Morula on barnacle density was positive, presumably due to its selective removal of mussels. These results suggest an indirect mutualism between barnacles and the gastropod predator, because barnacles attract settlement or enhance the survival of mussels, and the predator reduces the competitive effect of mussels on barnacles.     

Prey perception of predation risk: volatile chemical cues mediate non-consumptive effects of a predator on a herbivorous insect

Predators can affect prey in two ways—by reducing their density (consumptive effects) or by changing their behavior, physiology or other phenotypic traits (non-consumptive effects). Understanding the cues and sensory modalities prey use to detect predators is critical for predicting the strength of non-consumptive effects and the outcome of predator–prey encounters. While predator-associated cues have been well studied in aquatic systems, less is known about how terrestrial prey, particularly insect larvae, detect their predators. We evaluated how Colorado potato beetle, Leptinotarsa decemlineata, larvae perceive predation risk by isolating cues from its stink bug predator, the spined soldier bug, Podisus maculiventris. When exposed to male “risk” predators that were surgically manipulated so they could hunt but not kill, beetles reduced feeding 29 % compared to controls. Exposure to risk females caused an intermediate response. Beetles ate 24 % less on leaves pre-exposed to predators compared to leaves never exposed to predators, indicating that tactile and visual cues are not required for the prey’s response. Volatile odor cues from predators reduced beetle feeding by 10 % overall, although male predators caused a stronger reduction than females. Finally, visual cues from the predator had a weak effect on beetle feeding. Because multiple cues appear to be involved in prey perception of risk, and because male and female predators have differential effects, beetle larvae likely experience tremendous variation in the information about risk from their local environment.     

Effects of prey density,prey size and predator size on rates of feeding by an intertidal predatory gastropod Morula marginalba Blainville (Muricidae), on several species of prey

As a prerequisite for models of foraging behaviour of the whelk, Morula marginalba Blainville (Muricidae), the effects of variation in density of prey on the rate of feeding of the predator were examined in field conditions for three coexisting species of prey. Densities of prey used were those at which the prey, two limpets and a barnacle, occurred naturally in the rocky intertidal habitat.Large limpets, Cellana tramoserica (Sowerby) can resist attacks by predatory gastropods by raising the mantle over the outside of the shell. These experiments showed that no C. tramoserica were killed by Morula marginalba even at very great densities and with no alternative prey present. For the small limpet Patelloida latistrigata (Angas), one of the whelk's most highly preferred prey, juveniles were eaten 1.4 times as fast as adults. Fitting the random predator equation gave greater attack coefficients and shorter handling times for juvenile than adult limpets.Sizes of both predator and prey affected rates of eating barnacles, Tesseropora rosea (Krauss), but not in a simple way. Whelks of 15-mm aperture length ate adult barnacles 4.2 times faster than did 12-mm whelks, but there was no significant difference in the rates at which the two sizes of snail ate juvenile barnacles.Rates of feeding on T. rosea and Patelloida latistrigata increased significantly with prey density. These results form a basis for including the density of prey in models of spatial dispersion of the predatory gastropod Morula marginalba.     

Keystone nonconsumptive effects within a diverse predator community

The number of prey killed by diverse predator communities is determined by complementarity and interference among predators, and by traits of particular predator species. However, it is less clear how predators' nonconsumptive effects (NCEs) scale with increasing predator biodiversity. We examined NCEs exerted on Culex mosquitoes by a diverse community of aquatic predators. In the field, mosquito larvae co‐occurred with differing densities and species compositions of mesopredator insects; top predator dragonfly naiads were present in roughly half of surveyed water bodies. We reproduced these predator community features in artificial ponds, exposing mosquito larvae to predator cues and measuring resulting effects on mosquito traits throughout development. Nonconsumptive effects of various combinations of mesopredator species reduced the survival of mosquito larvae to pupation, and reduced the size and longevity of adult mosquitoes that later emerged from the water. Intriguingly, adding single dragonfly naiads to ponds restored survivorship of larval mosquitoes to levels seen in the absence of predators, and further decreased adult mosquito longevity compared with mosquitoes emerging from mesopredator treatments. Behavioral observations revealed that mosquito larvae regularly deployed “diving” escape behavior in the presence of the mesopredators, but not when a dragonfly naiad was also present. This suggests that dragonflies may have relaxed NCEs of the mesopredators by causing mosquitoes to abandon energetically costly diving. Our study demonstrates that adding one individual of a functionally unique species can substantially alter community‐wide NCEs of predators on prey. For pathogen vectors like mosquitoes, this could in turn influence disease dynamics.     

Non-consumptive effects of larval Salamandra on crustacean prey: can eggs detect predators?

Predators affect prey populations not only by prey consumption but also in nonconsumptive ways including modifying prey behavior. I tested the effects of fire salamander larvae (Salamandra infraimmaculata) on populations of co-occurring crustacean species in artificial outdoor pools. I also tested whether these effects were due entirely to prey consumption by Salamandra larvae or alternatively to some nonconsumptive effect. The soil (containing crustacean eggs) added to the artificial pools was collected from a dried-out temporary pool that is inhabited by Salamandra during the early part of the hydroperiod. I randomly assigned the pools to one of three treatments: control, free Salamandra, or caged Salamandra. Free salamander larvae could roam the entire pool and prey upon crustaceans. Caged salamander larvae were placed within a cage with having 250-μm mesh windows. They could not prey upon the crustaceans but could, for example, influence them by chemical cues. Densities of the three dominant crustacean species (Arctodiaptomus similis, Ceriodaphnia quadrangula and Cyzicus sp.) were drastically reduced in both salamander treatments compared to the control. Crustacean densities, however, were not significantly different between the two salamander treatments. One plausible explanation is that crustacean eggs can detect the presence of this predator via chemical cues and delay hatching. Received: 4 March 1996 / Accepted, 23 October 1996     

Female blue tits with brighter yellow chests transfer more carotenoids to their eggs after an immune challenge

Predicting the consequences of predator biodiversity loss on prey requires an understanding of multiple predator interactions. Predators are often assumed to have independent and additive effects on shared prey survival; however, multiple predator effects can be non-additive if predators foraging together reduce prey survival (risk enhancement) or increase prey survival through interference (risk reduction). In marine communities, juvenile reef fish experience very high mortality from two predator guilds with very different hunting modes and foraging domains—benthic and pelagic predator guilds. The few previous predator manipulation studies have found or assumed that mortality is independent and additive. We tested whether interacting predator guilds result in non-additive prey mortality and whether the detection of such effects change over time as prey are depleted. To do so, we examined the roles of benthic and pelagic predators on the survival of a juvenile shoaling zooplanktivorous temperate reef fish, Trachinops caudimaculatus, on artificial patch reefs over 2 months in Port Phillip Bay, Australia. We observed risk enhancement in the first 7 days, as shoaling behaviour placed prey between predator foraging domains with no effective refuge. At day 14 we observed additive mortality, and risk enhancement was no longer detectable. By days 28 and 62, pelagic predators were no longer significant sources of mortality and additivity was trivial. We hypothesize that declines in prey density led to reduced shoaling behaviour that brought prey more often into the domain of benthic predators, resulting in limited mortality from pelagic predators. Furthermore, pelagic predators may have spent less time patrolling reefs in response to declines in prey numbers. Our observation of the changing interaction between predators and prey has important implications for assessing the role of predation in regulating populations in complex communities.     

Attraction of Flatworms at Various Hunger Levels Toward Cues from an Odonate Predator

Different hunger levels can modify a prey's antipredator behavior in the presence or absence of food. Satiated animals often forego foraging if a predator is nearby, whereas starved animals may risk a predator encounter to search for food. This study evaluated the influence of nutritional state on the behavior of the flatworm Dugesia dorotocephala in the presence of food, predator, and crushed conspecific cues. We found that flatworms are attracted to cues originating from a food source, crushed conspecifics, and a predator (dragonfly larva) compared with control cues. Among the different hunger level treatment groups, levels of satiation had no influence on activity levels but significantly influenced time spent close to and distance from the cue source. Flatworm movement toward predator cues emitted from dragonfly larvae is contrary to our expectations. These results suggest either a unique case of chemical mimicry from the dragonfly larvae or an inherent attraction of planarians to odonate predators that allows them to scavenge the remains of other odonate prey items.     

Prey stage preference and feeding behaviour ofOligota kashmirica benefica (Coleoptera: Staphylinidae), an insect predator of the spider miteTetranychus urticae (Acari: Tetranychidae)

We studied the prey stage preference and feeding behaviour of the first to third instar larvae and adult females ofOligota kashmirica benefica Naomi (Coleoptera: Staphylinidae), a predator of the spider miteTetranychus urticae Koch (red form) (Acari: Tetranychidae), on leaves of the kudzu vine (Pueraria lobata (Wild.) Ohwi (Leguminosae)) under laboratory conditions. The number of mites eaten increased with the growth of predator larvae. Third instar larvae preyed on all stages of spider mite, whereas first instar larvae preyed mainly on immobile stages (eggs and quiescent stages). The predator larvae showed two types of foraging behaviour (active searching and ambush behaviour) when targeting the mobile stages (larval nymph and adult stages of prey). Although no difference was found in the number of prey consumed by adult females and third instar larvae of the predator, the adult females mainly attacked and consumed the immobile stages.     

Settlement in different-sized patches by the gregarious intertidal barnacle Chamaesipho tasmanica Foster and Anderson in New South Wales

Most models on settlement of open marine invertebrate populations are based on space-limitation. These models, however, do not recognise that free space may not drive the demography of populations when larval numbers are small or when larval supply varies along a gradient in the habitat. They also do not incorporate the effects of larval choice when settling. It has been hypothesised that, in gregarious barnacles, the effects of adult conspecifics, rather than available free space, may play a primary role in settlement. That is, cues from adults along perimeters of patches, rather than space available, may enhance colonisation. This study therefore aimed to distinguish between these separate influences on populations of Chamaesipho tasmanica, a gregarious barnacle characterised by relatively few larvae arriving to settle each year. Patches of 6, 3 and 1.5 cm diameter were cleared within aggregations of barnacles at three heights (Low, Mid, Upper) of Chamaesipho's distribution at two sites and during 2 years of settlement. Total numbers of settlers in each year were manipulated to determine the separate influences on settlement due to availability of substratum or the effects of conspecific adults. To test for the effects of available free space, numbers of settlers per unit area were analysed. To test for gregarious effects due to the presence of adults, numbers of settlers per unit perimeter were analysed. While available substratum was found not to affect settlement of this barnacle, gregarious settlement in response to adults at perimeters of patches was thought to be confounded by differential larval supply and differential conspecific cues among heights on the shore. Results from this study therefore have important implications for survival of gregarious populations following disturbances, especially in species where larval supply is poor.     

Attracted to the enemy: Aedes aegypti prefers oviposition sites with predator-killed conspecifics

Oviposition habitat choices of species with aquatic larvae are expected to be influenced by both offspring risk of mortality due to predation, and offspring growth potential. Aquatic predators may indirectly influence growth potential for prey by reducing prey density and, for filter-feeding prey, by increasing bacterial food for prey via added organic matter (feces, partially eaten victims), creating the potential for interactive effects on oviposition choices. We tested the hypothesis that the mosquito Aedes aegypti preferentially oviposits in habitats with predatory Toxorhynchites larvae because of indirect effects of predation on chemical cues indicating bacterial abundance. We predicted that A. aegypti would avoid oviposition in sites with Toxorhynchites, but prefer to oviposit where bacterial food for larvae is abundant, and that predation by Toxorhynchites would increase bacterial abundances. Gravid A. aegypti were offered paired oviposition sites representing choices among: predator presence; the act of predation; conspecific density; dead conspecific larvae; and bacterial activity. A. aegypti preferentially oviposited in sites with Toxorhynchites theobaldi predation, and with killed conspecific larvae, but failed to detect preferences for other treatments. The antibiotic tetracycline eliminated the strongest oviposition preference. Both predation by Toxorhynchites and killed larvae increased bacterial abundances, suggesting that oviposition attraction is cued by bacteria. Our results show the potential for indirect effects, like trophic cascades, to influence oviposition choices and community composition in aquatic systems. Our results suggest that predators like Toxorhynchites may be doubly beneficial as biocontrol agents because of the attraction of ovipositing mosquitoes to bacterial by-products of Toxorhynchites feeding.     

Determining sensitive stages for learning to detect predators in larval bronzed frogs: Importance of alarm cues in learning

Successful survival and reproduction of prey organisms depend on their ability to detect their potential predators accurately and respond effectively with suitable defences. Predator detection can be innate or can be acquired through learning. We studied prey–predator interactions in the larval bronzed frogs (Sylvirana temporalis), which have the innate ability to detect certain predators. We conducted a series of experiments to determine if the larval S. temporalis rely solely on innate predator detection mechanisms or can also learn to use more specific cues such as conspecific alarm cues for the purpose. The results of our study clearly indicate that larval S. temporalis use both innate and learned mechanisms for predator detection. Predator-naïve tadpoles could detect kairomones alone as a potential threat and responded by reducing activity, suggesting an innate predator detection mechanism. Surprisingly, predator-naïve tadpoles failed to detect conspecific alarm cues as a potential threat, but learned to do so through experience. After acquiring the ability to detect conspecific alarm cues, they could associate novel predator cues with conspecific alarm cues. Further, post feeding stages of larval S. temporalis are sensitive for learning to detect conspecific alarm cues to label novel predators.