Threat-sensitive foraging by larval threespine sticklebacks (Gasterosteus aculeatus)

Summary The threat-sensitive predator avoidance hypothesis predicts that prey can assess the relative threat posed by a predator and adjust their behaviour to reflect the magnitude of the threat. We tested the ability of larval threespine sticklebacks to adjust their foraging in the presence of predators by exposing them to conspecific predators of various sizes and recording their foraging and predator avoidance behaviours. Larvae (<30 days post-hatch) displayed predator escape behaviours only towards attacking predators. At 3 weeks post-hatch larvae approached the predator after fleeing, a behaviour which may be the precursor to predator inspection. Larvae reduced foraging and spent less time in the proximity of large and medium-sized predators compared to small predators. The reduction in foraging was negatively correlated to the predator/larva size ratio, indicating that larvae increased their foraging as they increased in size relative to the predator. We conclude that larval sticklebacks can assess the threat of predation early in their ontogeny and adjust their behaviour accordingly.Correspondence to: J.A. Brown     

A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus

Organisms in natural habitats participate in complex ecological interactions that include competition, predation, and foraging. Under natural aquatic environmental conditions, amphibian larvae can simultaneously receive multiple signals from conspecifics, predators, and prey, implying that predator-induced morphological defenses can occur in prey and that prey-induced offensive morphological traits may develop in predators. Although multiple adaptive plasticity, such as inducible defenses and inducible offensive traits, can be expected to have not only ecological but also evolutionary implications, few empirical studies report on species having such plasticity. The broad-headed larval morph of Hynobius retardatus, which is induced by crowding with heterospecific anuran (Rana pirica) larvae, is a representative example of prey-induced polyphenism. The morph is one of two distinct morphs that have been identified in this species; the other is the typical morph. In this paper, we report that typical larval morphs of Hynobius can respond rapidly to a predatory environment and show conspicuous predator-induced plasticity of larval tail depth, but that broad-headed morphs cannot respond similarly to a predation threat. Our findings support the hypothesis that induction or maintenance of adaptive plasticity (e.g., predator-induced polyphenism) trades off against other adaptive plastic responses (e.g., prey-induced polyphenism). For a species to retain both an ability to forage for larger prey and an ability to more effectively resist predation makes sense in light of the range of environments that many salamander larvae experience in nature. Our results suggest that the salamander larvae clearly discriminate between cues from prey and those from predators and accurately respond to each cue; that is, they adjust their phenotype to the current environment.     

Feeding selectivity in larvae of the European hake (<Emphasis Type="Italic">Merluccius merluccius</Emphasis>) in relation to ontogeny and visual capabilities

Feeding ecology was analysed for the first time in the larvae of the European hake (Merluccius merluccius) to determine whether their diet and selectivity were constrained by environmental conditions and how these feeding characteristics were related to ontogeny, prey availability and visual capabilities. Larvae collected during both day and night were analysed, and it was found that feeding incidence was high, regardless of the time of day. Examination of the visual system corroborated the hypothesis that hake larvae should be able to cope with a wide range of photic conditions and to forage even at low light intensity. A clear preference for adult calanoid copepods and, especially, for Clausocalanus spp. was observed in all sizes analysed. Prey number increased with larval size, but prey size did not. This finding indicates that hake larvae behave as selective and specialist predators that consume an increasing number of prey rather than larger prey during larval growth.     

Adaptive behaviour of chironomid larvae (Chironomus riparius) in response to chemical stimuli from predators and resource density

Chemical-mediated effects of predatory fish on chironomid larvae behaviour have been ignored so far. Sediment-dwelling chironomid larvae inhabit protective burrows from which they extend their bodies only to feed on deposited detritus and microalgae from the surrounding sediment. Here, we performed factorial laboratory experiments to study whether fish-borne chemical cues (kairomones) are responsible for behavioural trait changes of chironomid larvae, and whether chironomid larvae are able to assess the densities of fish predators and food resources and the trade-off between them. We exposed naïve Chironomus riparius larvae to the chemical presence of zero, one, and ten predator fish (Rutilus rutilus) and offered two resource levels (low food, high food) for each treatment. Kairomones induced significant inherent behavioural trait changes in chironomid larvae. During the first 120 min after exposing chironomids to fish-conditioned water, we found a significant increase in digging activity with increasing predator density. After 3 days of exposure, the deepest chironomid burrows were found in treatments with the highest predator density. Chironomid larvae were significantly able to adjust their foraging behaviour to different predator densities and food concentrations and trade off between them; that is, when fish predators were more abundant or when more food resources were available, the foraging activities of larvae were significantly reduced. Our data suggest that chemically mediated trait changes (burrowing and foraging behaviour) may cascade through the littoral food web.     

Foraging behavior in early Atlantic cod larvae (Gadus morhua) feeding on a protozoan (Balanionsp.) and a copepod nauplius (Pseudodiaptomussp.)

Food limitation is likely to be a source of mortality for fish larvae in the first few weeks after hatching. In the laboratory, we analyzed all aspects of foraging in cod larvae (Gadus morhua Linnaeus) from 5 to 20 d post-hatching using protozoa (Balanion sp.) and copepod nauplii (Pseudodiaptomus sp.) as prey. A camera acquisition system with two orthogonal cameras and a digital image analysis program was used to observe patterns of foraging. Digitization provided three-dimensional speeds, distances, and angles for each foraging event, and determined prey and fish larval head and tail positions. Larval cod swimming speeds, perception distances, angles, and volumes increased with larval fish size. Larval cod swam in a series of short intense bursts interspersed with slower gliding sequences. In 94% of all foraging events prey items were perceived during glides. Larval cod foraging has three possible outcomes: unsuccessful attacks, aborted attacks, and successful attacks. The percentage of successful attacks increased with fish size. In all larval fish size classes, successful attacks had smaller attack distances and faster attack speeds than unsuccessful attacks. Among prey items slowly swimming protozoans were the preferred food of first-feeding cod larvae; larger larvae had higher swimming speeds and captured larger, faster copepod nauplii. Protozoans may be an important prey item for first-feeding larvae providing essential resources for growth to a size at which copepod nauplii are captured. Received: 20 April 1999 / Accepted: 12 January 2000     

Time stress,predation risk and diurnal–nocturnal foraging trade-offs in larval prey

Insect larvae increase in size with several orders of magnitude throughout development making them more conspicuous to visually hunting predators. This change in predation pressure is likely to impose selection on larval anti-predator behaviour and since the risk of detection is likely to decrease in darkness, the night may offer safer foraging opportunities to large individuals. However, forsaking day foraging reduces development rate and could be extra costly if prey are subjected to seasonal time stress. Here we test if size-dependent risk and time constraints on feeding affect the foraging–predation risk trade-off expressed by the use of the diurnal–nocturnal period. We exposed larvae of one seasonal and one non-seasonal butterfly to different levels of seasonal time stress and time for diurnal–nocturnal feeding by rearing them in two photoperiods. In both species, diurnal foraging ceased at large sizes while nocturnal foraging remained constant or increased, thus larvae showed ontogenetic shifts in behaviour. Short night lengths forced small individuals to take higher risks and forage more during daytime, postponing the shift to strict night foraging to later on in development. In the non-seasonal species, seasonal time stress had a small effect on development and the diurnal–nocturnal foraging mode. In contrast, in the seasonal species, time for pupation and the timing of the foraging shift were strongly affected. We argue that a large part of the observed variation in larval diurnal–nocturnal activity and resulting growth rates is explained by changes in the cost/benefit ratio of foraging mediated by size-dependent predation and time stress.     

Activity and metabolism of larval Atlantic cod (Gadus morhua) from Scotian Shelf and Newfoundland source populations

Patterns of activity and metabolism were investigated in larval Atlantic cod (Gadus morhua L.) between December 1991 and July 1992: (1) throughout larval development; (2) between two genetically discrete populations (Scotian Shelf and Newfoundland) and (3) as a function of two different culture temperatures. During the yolk-sac stage (0 to 5 d post-hatch), changes in swimming speed were not related to mass-specific metabolic rates; no portion of the mass-specific oxygen consumption could be explained by changes in activity. In the mixed feeding stage (6 to 14 d posthatch), there was a tendency for oxygen consumption to be related to changes in swimming speed. In the exogenous feeding stage (>14 d post-hatch), oxygen consumption significantly increased with swimming speed. These ontogenetic patterns of activity and metabolism were the same for larvae from the Scotian Shelf and Newfoundland populations. However, over the entire larval life and among ontogenetic stages, the metabolic cost of activity (mass-specific O₂ consumption/swimming speed) of Scotian Shelf larvae was significantly higher than that of Newfoundland larvae. When cod larvae, that had developed at 5°C, were acutely exposed to 10°C, Scotian Shelf larvae had a higher intrinsic cost of activity than Newfoundland larvae, over the entire larval life. During the exogenous feeding stage, the mean metabolic cost of activity for Newfoundland larvae raised at 10°C and tested at 10°C was significantly higher and more variable than that of larvae raised at lower temperatures. However, the metabolic cost of activity of larvae raised and tested at 10°C was not significantly different between source populations. Together these findings suggest that differences in swimming energetics reflect changing energy requirements for activity among ontogenetic stages, and reflect adaptation to regional environments among genetically discrete populations.     

Active salinity choice and enhanced swimming endurance in 0 to 8-d-old larvae of diadromous gobies,including Sicydium punctatum (Pisces), in Dominica,West Indies

We studied the early life history of diadromous gobies in Dominica, West Indies, from May 1989 to May 1991, emphasising Sicydium punctatum Perugia. The transition of newly hatched larvae from upriver nest sites to the sea was studied in laboratory experiments. Newly hatched larvae are negatively buoyant but avoid settling to the bottom by active swimming during drift to the sea. Laboratory experiments evaluated salinity preferences and effects on swimming endurance. Larvae in haloclines actively selected low to intermediate salinities. Initially (0 to 5-d post-hatch), larvae minimized exposure to salinities >10 ppt, but later (5 to 8-d) occupied increasingly saline water. Larvae in no-choice freshwater or seawater treatments ceased activity at 4 to 5 d, but in haloclines larvae remained active up to 8 d post-hatch. Salinities <10 ppt are important for early survival of sicydiine gobies. Implications for larval survival and transport are discussed.     

Threat-sensitive learning of predators by larval mosquitoes <Emphasis Type="Italic">Culex restuans</Emphasis>

A prerequisite for prey to show adaptive behavioural responses to predators is that the prey has the ability to recognise predators as threats. While predator recognition can be innate in many situations, learning is often essential. For many aquatic species, one common way to learn about predators is through the pairing of a novel predator odour with alarm cues released from injured conspecifics. One study with fish demonstrated that this mode of learning not only allows the prey to recognise the predatory cues as a threat, but also mediates the level of threat associated with the predator cues (i.e. threat-sensitive learning). When the prey is exposed to the novel predator with a high concentration of alarm cues, they subsequently show a high intensity of antipredator response to the predator cues alone. When exposed to the predator with a low concentration of alarm cues, they subsequently show a low-intensity response to the predator cues. Here, we investigated whether larval mosquitoes Culex restuans have the ability to learn to recognise salamanders as a threat through a single pairing of alarm cues and salamander odour and also whether they would learn to respond to salamander cues in a threat-sensitive manner. We conditioned individual mosquitoes with water or a low, medium or high concentration of crushed conspecific cues (alarm cues) paired with salamander odour. Mosquitoes exposed to salamander odour paired with alarm cues and subsequently exposed to salamander odour alone responded to the salamander as a threat. Moreover, the intensity of antipredator response displayed during the conditioning phase matched the response intensity during the testing phase. This is the first demonstration of threat-sensitive learning in an aquatic invertebrate.     

Foraging behaviour of larval cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) at low light intensities

The ability to forage at low light intensities can be of great importance for the survival of fish larvae in a pelagic environment. Three-dimensional silhouette imaging was used to observe larval cod foraging and swimming behaviour at three light intensities (dusk ~1.36 × 10⁻³ W/m², night ~1.38 × 10⁻⁴ W/m² and darkness ~3.67 × 10⁻⁶ W/m²) at 4 different ages from 6 to 53 days post-hatch (dph). At 6 dph, active pursuit of prey was only observed under dusk conditions. Attacks, and frequent orientations, were observed from 26 dph under night conditions. This was consistent with swimming behaviour which suggested that turn angles were the same under dusk and night conditions, but lower in darkness. Cod at 53 dph attacked prey in darkness and turn angles were not different from those under other light conditions. This suggests that larvae are still able to feed at light intensities of 3.67 × 10⁻⁶ W/m². We conclude that larval cod can maintain foraging behaviour under light intensities that correspond to night-time at depths at which they are observed in the field, at least if they encounter high-density patches of prey such as those that they would encounter at thin layers or fronts.     

Threat-sensitive predator avoidance in damselfish-trumpetfish interactions

Summary Predatory threat can vary during a predator-prey interaction as an attack escalates or among predators at different times. A Threat-sensitivity hypothesis is presented which predicts that prey individuals will trade-off predator avoidance against other activities by altering their avoidance responses in a manner that reflects the magnitude of the predatory threat. This hypothesis was tested in the field by presenting prey (threespot damselfish, Stegastes planifrons) with models of foraging predators (Atlantic trumpetfish, Aulostomus maculatus). During a presentation, damselfish displayed progressively stronger avoidance as predator models were brought nearer; response waned rapidly once predator models passed overhead. Larger predator models and those oriented in a strike pose evoked stronger avoidance reactions than smaller and non-attacking models, intermediate responses were evoked by size and orientation combinations that were intermediate in threat, and habituation was more common to weakly-threatening presentations. Smaller damselfish showed stronger avoidance of models than did larger damselfish. Nonavoidance activities, such as feeding and territorial defense, were curtailed during presentations or were more common during weakly threatening presentations. Approaches to the models, equated with mobbing, were more common among large damselfish, again reflecting degrees of vulnerability among different size prey individuals. These initial results indicate that damselfish threatened by predators respond in a graded manner that reflects the degree of threat posed by the predator, in accordance with the Threat-sensitivity hypothesis.     

The interplay between foraging mode, habitat structure, and predator presence in antlions

Antlion larvae are sand-dwelling insect predators, which ambush small arthropod prey while buried in the sand. In some species, the larvae construct conical pits and are considered as sit-and-wait predators which seldom relocate while in other species, they ambush prey without a pit but change their ambush site much more frequently (i.e., sit-and-pursue predators). The ability of antlion larvae to evade some of their predators which hunt them on the sand surface is strongly constrained by the degree of sand stabilization or by sand depth. We studied the effect of predator presence, predator type (active predatory beetle vs. sit-and-pursue wolf spider), and sand depth (shallow vs. deep sand) on the behavioral response of the pit building Myrmeleon hyalinus larvae and the sit-and-pursue Lopezus fedtschenkoi larvae. Predator presence had a negative effect on both antlion species activity. The sit-and-wait M. hyalinus larvae showed reduced pit-building activity, whereas the sit-and-pursue L. fedtschenkoi larvae decreased relocation activity. The proportion of relocating M. hyalinus was negatively affected by sand depth, whereas L. fedtschenkoi was negatively affected also by the predator type. Specifically, the proportion of individual L. fedtschenkoi that relocated in deeper sand was lower when facing the active predator rather than the sit-and-pursue predator. The proportion of M. hyalinus which constructed pits decreased in the presence of a predator, but this pattern was stronger when exposed to the active predator. We suggest that these differences between the two antlion species are strongly linked to their distinct foraging modes and to the foraging mode of their predators. Reut Loria and Inon Scharf contributed equally to the paper.     

Larval ecology of the great barracuda, <Emphasis Type="Italic">Sphyraena barracuda</Emphasis>, and other sphyraenids in the Straits of Florida

The great barracuda (Sphyraena barracuda) is a widespread, ecologically and socioeconomically important coastal fish, yet very little is known about its larvae. We examined spawning and larval ecology of Western Atlantic sphyraenids using monthly ichthyoplankton samples collected over 2 years along a transect spanning the east–west axis of the Straits of Florida (SOF). Samples were dominated by the great barracuda (92.8%) and sennets (Sphyraena borealis and Sphyraena picudilla; 6.6%). While larval sennets and S. barracuda displayed similar vertical distributions (majority in upper 25 m), horizontal and temporal patterns of abundance suggested a spatial and temporal species replacement between larval S. barracuda and sennets that tracks adult ecology. The diet of both taxa consisted largely of copepods, with inclusion of fish larvae at 8 mm SL, and in S. barracuda alone, a switch in the wet season to exclusive piscivory by 12 mm SL (18 days post-hatch). A lack of piscivory in S. barracuda larvae captured in the dry season corresponded to slower larval growth than in the wet season. Larval growth was also related to size-at-hatch and larval age such that larvae that were larger at hatch or larger (older) at capture grew faster at earlier ages, suggesting faster larval growth, and indirectly larger hatch size, conveys a survival advantage. Unlike larval growth, instantaneous mortality rate did not differ with season, and no lunar cyclic patterns in spawning output were identified. Our results provide insight into the pelagic phase of sphyraenids and highlight the importance of both diet and hatch size to the growth and survival of fish larvae in low latitude oceanic environments.     

Why run and hide when you can divide? Evidence for larval cloning and reduced larval size as an adaptive inducible defense

Predator-induced cloning (asexual reproduction), with reduced size as consequence of cloning, suggests a novel adaptation to the threat of predation. Although cloning is a common reproductive strategy of many plants and animals, cloning in response to stimuli from predators has, at present, been documented only in the larvae (plutei) of the sand dollar, Dendraster excentricus. Other studies report larval cloning in echinoderms under optimal conditions of food and temperature. A burst of asexuality should be favored when environmental conditions are conducive to growth, but it is less clear that cloning is advantageous when conditions indicate risk from predators. This study tested the hypothesis that the small size of predator-induced clones reduces vulnerability during encounters with planktivorous fish. Successful cloning was inferred from an increase in larval density, a reduction in larval size and stage, and some direct observations of budding. All clones were smaller than uncloned sibling larvae, suggesting an advantage against visual predators. Pair-wise predation trials demonstrated that planktivorous fish ate more uncloned sibling plutei than small clones. These results offer a new ecological context for asexual reproduction: rapid size reduction as a defense. If the identifiable cues for cloning in echinoderm larvae (food and predators) are linked in nature, then larval cloning may be a response to a single ecological scenario rather than two separate and unrelated conditions.     

Vertical distribution and mortality of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during overwintering in oceanic waters southwest of Iceland

The seasonal vertical distribution and the predatory regime encountered by Calanus finmarchicus were studied along a transect across the Reykjanes Ridge in the oceanic area southwest of Iceland from data collected during four cruises between November 1996 and June 1997. The mortality for the overwintering period was estimated using linear regressions of density estimates from November 1996 to April 1997. In addition, we also estimated the mortality of the oldest population stages (C4 and older) in April and June by applying the vertical table method. During winter (November/December–January/February), the animals mainly resided at a depth of ∼300–1,500 m in the water of Atlantic origin. Ascent to upper layers took place mainly during March and April, and continued until May. During all cruises, continuous deep-scattering layers were observed, mainly within the range of 400–500 m to 700–800 m depth. Based on sampling with a Harstad pelagic trawl in April, the scattering was mainly ascribed to jellyfish (mainly Periphylla periphylla), small mesopelagic oceanic fishes (several species but Benthosema glaciale and Maurolicus muelleri were most abundant), euphausiids (mainly Meganyctiphanes norvegica) and shrimps (mainly Sergestes arcticus). These species may represent a predatory threat to overwintering C. finmarchicus. From November to April, daily per capita mortality rates were estimated to be (mean ± 95% CL) 0.004 ± 0.0028 for the total data set, and 0.004 ± 0.0033 and 0.004 ± 0.0023 (day^-1) for the Iceland Basin and Irminger Basin, respectively. Mortality rates were higher later in life (mean ± 95% CL) for C5/females (0.13 ± 0.044) and C5/males (0.19 ± 0.051) than for C4/C5 (0.00 ± 0.035) when averaged over all samples taken in April and June 1997. We discuss how the observed distribution and mortality rates of overwintering C. finmarchicus might be related to predatory regime.     

Client preferences by Caribbean cleaning gobies: food,safety or something else?

Predation risk is amongst the most pervasive selective pressures influencing behaviour and animals have been repeatedly shown to trade-off foraging success for safety. We examined the nature of this trade-off in cleaning symbioses amongst Caribbean coral reef fishes. We predicted that cleaning gobies (Elacatinus evelynae and Elacatinus prochilos) should prefer fish clients that pose a low risk of predation (e.g. herbivores) over clients that may have more ectoparasites but pose a higher risk (e.g. piscivores). Our field observations revealed that cleaners did clean preferentially client species with more parasites but predatory and non-predatory clients had similar ectoparasite loads. Despite the lack of a foraging advantage for inspecting predators, cleaners did not avoid risky clients. On the contrary, a larger proportion of visiting predators than non-predators was inspected, gobies initiated more interactions with predatory clients, and predators were attended to immediately upon arrival at cleaning stations. This preferential treatment of dangerous clients may allow the rapid identification of cleaners as non-prey item or may be due to the effect of predators on the rest of the cleaners’ clientele, which avoided cleaning stations whilst predators were present. Dealing with potentially risky clients may allow gobies to regain access to their main food source: non-predatory clients.     

Effect of unfavorable trophic scenarios on amylase and protease activity of <Emphasis Type="Italic">Nephrops norvegicus</Emphasis> (L.) larvae during their first vertical migration: a laboratory approach

In Portuguese waters, Nephrops norvegicus larvae hatch at 400–800 m depth and need to perform a vertical migration to food-rich shallower waters to find suitable prey. The effect of suboptimal feeding on digestive enzymes activity of N. norvegicus larvae during this early period of their larval life remains unknown. Protease and amylase activities were investigated ex situ using flurometry in laboratory-hatched larvae exposed to different feeding and/or starving scenarios in the 24 h following hatching, the period during which they typically accomplish their upward vertical migration. Amylase activity was very low in comparison with protease activity, indicating that carbohydrates are not a primary energy reserve. Larvae starved for 12 h and subsequently fed displayed no increase in amylase activity, which suggests that feeding may be required before 12 h post-hatch to trigger amylase activity. Protease activity was high under all feeding conditions, and the increase in protease activity under sustained starvation indicated the catabolism of protein reserves. The ability of first-stage N. norvegicus larvae to metabolize protein reserves may play a decisive role for their survival during their first vertical migration, as it enables them to overcome the deleterious effects of short-term starvation and/or suboptimal feeding.     

Control of larval release in the Caribbean spiny lobster, <Emphasis Type="Italic">Panulirus argus</Emphasis>: role of chemical cues

The current model for larval release in subtidal crustaceans suggests that hatching time is controlled by the embryos, which release a pheromone that stimulates the parent female to undergo behaviors that synchronize larval release. Alternatively, hatching could be controlled by the females. Ovigerous spiny lobsters Panulirus argus (Latreille) exhibit stereotypic behaviors during larval release, including rapid abdominal extensions and pleopod-pumping activity. Ovigerous P. argus were collected from coral reefs in the Florida Keys, USA during the summers of 2005 and 2006. Pleopod-pumping activity was quantified to determine if a female’s pumping activity correlates with the developmental state of the embryos. The role of pheromones released by developing and hatching embryos in controlling pumping behaviors was tested by measuring the pumping response of ovigerous lobsters to (1) hatch water, (2) homogenized embryo water, (3) embryo-conditioned water (unhatched late-stage embryos soaked for 20 h), and (4) water containing homogenized post-hatch embryo cases. Bioassays were conducted under constant conditions (dim-red light) in the laboratory at random times during the day to control for any possible rhythm in pumping activity. Spontaneous pleopod-pumping activity increased significantly with increasing embryo development. Upon exposure to hatch water, ovigerous lobsters with late-stage embryos displayed increased pleopod pumping with increased treatment concentration. Water individually conditioned with homogenized late-stage embryos, intact late-stage embryos, and homogenized post-hatch embryo cases all induced larval release behaviors in females with late-stage embryos. Ovigerous females with early-stage embryos did not respond to water conditioned with homogenized early- or late-stage embryos. These results suggest that active substances are released by embryos at the time of hatching and induce the stereotypical pumping behaviors of the female that synchronizes larval release. The results support the model that larval release in subtidal crustaceans is controlled by pheromones released from hatching embryos.     

Does cannibalism in Spodoptera frugiperda (Lepidoptera: Noctuidae) reduce the risk of predation?

The incidence of cannibalism of larval Spodoptera frugiperda (Lepidoptera: Noctuidae) on maize under field conditions was investigated using field cages. Cannibalism was found to account for approximately 40% mortality when maize plants were infested with two or four fourth-instar larvae over a 3-day period. Field trials examined the effect of larval density on the prevalence of natural enemies of S. frugiperda. The abundance of predators (earwigs, staphylinids, other predatory beetles, and Chrysoperla spp.) was significantly greater on maize plants with higher levels of larval feeding damage, while the relationship between predator abundance and number of S. frugiperda larvae per plant was less clear. As larval damage is probably a more reliable indicator of previous larval density than numbers collected at an evaluation, this indicates that predation risk will be greater for larvae living in large groups. Parasitism accounted for 7.1% mortality of larvae in sorghum, and involved six species of Hymenoptera and Tachinidae. There was no effect of larval density or within-plant distribution on the probability of larval attack by parasitoids. The selective benefits of cannibalism, in relation to the risk of predation and parasitism, are discussed. Received: 23 March 2000 / Accepted: 24 June 2000     

The effect of hunger on the strength and duration of the antipredator behavioral response of green frog (<Emphasis Type="Italic">Rana clamitans</Emphasis>) tadpoles

The activity level of prey reflects a trade-off between predation risk and foraging gain. A number of theoretical and empirical studies have shown that a prey's energetic state or the level of its resource should influence this trade-off (i.e., what the optimal activity level at a level of predation risk is). Here, I show that the energetic state of prey may also influence the duration of their antipredator behavioral response. Green frog tadpoles (Rana clamitans) reduced their activity level for a shorter time during exposure to the chemical cue of predatory larval dragonflies (Anax spp.) as their time since last feeding increased (i.e., as their energetic state decreased). Interestingly, the tadpoles strongly reduced their activity level upon cue exposure in all treatments. Thus, the relative activity level of tadpoles at different energetic states varied over time.