Prey switching in Rhyacophila dorsalis (Trichoptera) alters with larval instar

1. Ontogenetic shifts in predator behaviour can affect the assessment of food‐web structure and the development of predator–prey models. Previous studies have shown that the diel activity pattern and functional response differed between larval instars of the carnivorous caddis, Rhyacophila dorsalis. The present study examines switching by larvae of R. dorsalis presented with different proportions of two prey types; either small (length 2–4 mm) and large (5–8 mm) Chironomus larvae for second, third, fourth and fifth instars of R. dorsalis; or Baetis rhodani (9–12 mm) and large Chironomus larvae for fourth and fifth instars. Experiments were performed in stream tanks with one Rhyacophila larva per tank and 200 prey arranged in nine different combinations of the two prey types (20 : 180, 40 : 160, 60 : 140, 80 : 120, 100 : 100, 120 : 80, 140 : 60, 160 : 40 and 180 : 20). Prey were replaced as they were eaten. A model predicted the functional response in the absence of switching and provided a null hypothesis against which any tendency to switch could be tested. 2. There was no prey switching in the second and third instars, with both instars always showing a preference for small over large Chironomus larvae. Prey switching occurred in the fourth and fifth instars. As the relative abundance of one prey type increased in relation to the alternative, the proportion eaten of the former prey changed from less to more than expected from its availability, the relationship being described by an S‐shaped curve. In the experiments with small and large Chironomus, the two instars switched to large larvae when their percentage of the total available prey exceeded 29% and 37% for fourth and fifth instars, respectively. In the experiments with Baetis and large Chironomus, both instars switched to Baetis larvae when their percentage of the total available prey exceeded 36%. 3. Non‐switching in second and third instars was related to their feeding strategies, both instars preferring smaller prey items. When the fourth and fifth instars foraged actively at night, they preferred larger over small Chironomus larvae, but when they behaved as ambush predators at dusk, they captured the more active Baetis larvae in preference to the more sedentary large Chironomus larvae and only switched to the latter when they were >64% of the available prey.     

Ontogenetic shifts in the functional response and interference interactions of Rhyacophila dorsalis larvae (Trichoptera)

1. Ontogenetic shifts in predator behaviour can affect the assessment of food‐web structure and the development of predator–prey models. Therefore, it is important to establish if the functional response and interference interactions differ between life‐stages. These hypotheses were tested by (i) comparing the functional response of second, third, fourth and fifth larval instars of Rhyacophila dorsalis, using three stream tanks with one Rhyacophila larva per tank and one of 10 prey densities between 20 and 200 larvae of Chironomus sp.; (ii) using other experiments to assess interference within instars (two to five larvae of the same instar per tank), and between pairs of different instars (one, two or three larvae per instar; total predator densities of two, four or six larvae per tank). 2. The first hypothesis was supported. The number of prey eaten by each instar increased with prey density, the relationship being described by a type II model. The curvilinear response was stronger for fourth and fifth instars than for second and third instars. Mean handling time did not change significantly with prey density, and increased with decreasing instar number from 169 s for fifth instars to 200 s for second instars. Attack rate decreased progressively with decreasing instar number. Handling time varied considerably for each predator–prey encounter, but was normally distributed for each predator instar. Variations in attack rate and handling time were related to differences in activity between instars, fourth and fifth instars being more active and aggressive than second and third instars, and having a higher food intake. 3. The second hypothesis was partially supported. In the interference experiments between larvae of the same instar or different instars, mean handling time did not change significantly with increasing predator density, and attack rate did not change for second and third instars but decreased curvilinearly for fourth and fifth instars. Interference between some instars could not be studied because insufficient second instars were available at the same time as fourth and fifth instars, and most third instars were eaten by fourth and fifth instars in the experiments. Prey capture always decreased with decreasing attack rate. Therefore, interference reduced prey consumption in fourth and fifth instars, but not in second and third instars. The varying feeding responses of different instars should be taken into account when assessing their role in predator–prey relationships in the field.     

Contrasting diel activity and feeding patterns of four instars of Rhyacophila dorsalis (Trichoptera)

1. Ontogenetic shifts in prey choice and predator behaviour can affect food‐web structure. Therefore, it is important to establish if the diet and feeding activity differ between life‐stages of the same species. This hypothesis was tested for second, third, fourth and fifth larval instars of Rhyacophila dorsalis by comparing their diel activity and feeding patterns. Second to fifth instars collected from two streams were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks. Food was provided in excess; being organisms living in bryophytes on top of a large stone in each tank, augmented by different‐sized larvae of Ephemeroptera, Simuliidae and Chironomidae. As few first instars for gut analyses were found in the field, the diet of first instars reared in the laboratory was also studied. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 50 larvae per instar for each day or night sample). First and second instars fed on the smaller food items with no significant day‐night differences in diet. Gut contents indicated a progressive trend from feeding chiefly at night in third instars to almost exclusively at night in fifth instars. Fourth and fifth instars fed on the larger food items, whilst the diet of the third instar larvae overlapped with that of both the earlier and later instars. 3. Diel activity patterns of single larvae differed between instars but not within each instar (n = 20 larvae per instar). Second instars were active throughout the 24 h, with peaks at dusk, around midnight, dawn and around midday. A similar pattern was shown by third instars but the peak of activity at midday was less than the other three peaks. Prey were captured only during these peaks for both instars. Fourth and fifth instars were most active, and fed only, at night. They used an ambush strategy to capture more active prey at dusk and dawn (e.g. Baetis, Gammarus), and a searching strategy to capture more sedentary prey during the night (e.g. chironomids, simuliids). These experiments provided support for the hypothesis under test. If competition and/or interference occur between instars, then it could be reduced between earlier and later instars because of differences in their diet and diel pattern of feeding activity.     

Predation and drift of lotic macroinvertebrates during colonization

Summary A field experiment was carried out to determine the effect of an invertebrate predator on the colonization and drift of benthic macroinvertebrates in experimental stream channels. Lotic invertebrates colonized four replicate channels: two controls with no predators, and two channels with low densities (2.8 m^–2) of predatory stonefly nymphs, Doroneuria baumanni (Perlidae). Immigration rates were measured at the inflow of two other channels. Drift rates of invertebrates immigrating to and emigrating from channels were measured daily, and benthic samples were collected every five days. Over a 25-day colonization period, benthic densities of Baetis nymphs and larval Chironomidae were reduced by D. baumanni. Colonization curves were fit with a power function and significantly different colonization rates were indicated for both Baetis and chironomids in predation and control channels. A predator-induced drift response was exhibited by Baetis only and this response was size-dependent. In the presence of D. baumanni, large Baetis drifted more frequently than small nymphs and, correspondingly, small nymphs were more frequent in the benthos. Net predator impacts on invertebrate densities in channel substrates were partitioned into predator-induced drift and prey consumption. These estimates suggest that predator avoidance by Baetis is a prominent mechanism causing density reductions in the presence of predators. Reductions in the density of Chironomidae, however, were attributed to prey consumption only. A rainstorm during the experiment demonstrated that stream flow disruptions can override the influence of predators on benthic invertebrates, at least temporarily, and re-set benthic densities.     

Predation on Mosquito Larvae by Mesocyclops thermocyclopoides (Copepoda: Cyclopoida) in the Presence of Alternate Prey

The cyclopoid copepod Mesocyclops thermocyclopoides, a dominant invertebrate predator in many shallow ponds and temporary water bodies in northern India, feeds on cladocerans, rotifers, ciliates and when present, on mosquito larvae also. We studied in the laboratory the prey consumption rates of the copepod on first and fourth instar larvae of two species of mosquito (Anopheles stephensi and Culex quinquefasciatus) in relation to their density. We also studied its prey selectivity with mosquito larvae in the presence of an alternate prey (the cladocerans‐either Moina macrocopa or Ceriodaphnia cornuta) in different proportions. With either mosquito species, the copepod actively selected Instar‐I larvae, avoiding the Instar‐IV larvae, and with either instar, selected Anopheles stephensi over Culex quinquefasciatus. When prey choice included the cladoceran as an alternate prey, the copepod selected the cladoceran only when the other prey was Instar‐IV mosquito larvae. Our results point to the potential and promise of M. thermocyclopoides as a biological agent for controlling larval populations of vectorially important mosquito species.     

The impact of vertebrate and invertebrate predators on a stream benthic community

I assessed the impact of both vertebrate and invertebrate predators on a lotic benthic community in a 1-month-long experiment, using enclosures containing cobble/gravel bottoms, with large-mesh netting that allowed invertebrates to drift freely. Brown trout (Salmo trutta) and leeches (Erpobdella octoculata) were used as predators and four treatments were tested: a predator-free control, leeches only, trout only, and leeches and trout together. A density of 26.7 leeches/m² (20 leeches/enclosure) and 1.3 trout/m² (one trout per enclosure) was stocked into the enclosures. The total biomass of invertebrate prey was significantly lower in the trout and trout plus leech treatments than in the leech and control treatments, which were due to strong negative effects of trout on Gammarus. On the individual prey taxon level, both trout and leeches affected the abundance of Asellus , Baetis and Ephemerella, whereas the abundance of Gammarus was only affected by trout, and the abundance of Orthocladiinae and Limnephilidae was only affected by leeches. In the treatment with trout and leeches together, the abundance of Ephemerella and Baetis was higher than when trout or leeches were alone, which was probably due to predator interactions. Leeches and trout had no effects on prey immigration but did affect per capita emigration rates. Both trout and leeches indirectly increased periphyton biomass in enclosures, probably due to their strong effects on grazers. Both trout and leeches were size-selective predators, with trout selecting large prey, and leeches selecting small prey. Size-selective predation by trout and leeches affected the size structure of five commonly consumed prey taxa. Trout produced prey populations of small sizes owing to consumption of large prey as well as increased emigration out of enclosures by these large prey. Leech predation produced prey assemblages of larger size owing to consumption and increased emigration of small prey. These results suggest that in lotic habits, predatory invertebrates can be as strong interactors as vertebrate predators. Received: 23 June 1997 / Accepted: 4 May 1998     

The role of amphibian prey in the diet and growth of giant water bug nymphs in Japanese rice fields

Predatory insects that depend upon particular prey animals are commonly regulated by the prey animal’s abundance. Nymphs of the giant water bug Kirkaldyia (=Lethocerus) deyrolli (Heteroptera: Belostomatidae) are predators regarded as specialists in feeding on tadpoles. We studied the ontogenetic diet shift of aquatic nymphs by quantifying instar abundance and by analyzing captured prey and prey relative abundance during the period of rice irrigation in three localities. We also evaluated the contribution of major prey items (tadpoles, frogs, and Odonata nymphs) on specific growth rates of each nymphal stage in a rearing experiment. First to third-instar nymphs of K. deyrolli fed mainly on tadpoles, regardless of differences in prey availability. Nymphs of subsequent fourth and fifth instar stages shifted from tadpoles to other prey animals within each rice field. A rearing experiment demonstrated that giant water bug nymphs provided with tadpoles had greater specific growth rates at all nymphal stages, except for the final stage, than nymphs fed other prey (frogs and Odonata nymphs). The emergence of young K. deyrolli nymphs seemed to coincide with the period during which tadpoles became abundant in the rice fields. Consumption of tadpoles seems important to allow the nymph to complete its larval development in an unstable temporary habitat. An erratum to this article can be found at     

Fitness and predation potential of Macrolophus pygmaeus reared under artificial conditions

Abstract The biological parameters of Macrolophus pygmaeus Rambur after prolonged rearing in the absence of plant materials were compared with those of conventionally plant‐reared predators. When eggs of Ephestia kuehniella Zeller were provided as food, developmental and reproductive fitness of M. pygmaeus reared for over 30 consecutive generations using artificial living and oviposition substrates was similar to that of predators kept on tobacco leaves. Plantless‐reared fifth instars of the predator also had similar predation rates on second instars of the tobacco aphid, Myzus persicae nicotianae Blackman, as their peers maintained on plant materials. In a further experiment, predation on aphid prey by fifth instar M. pygmaeus fed one of two egg yolk‐based artificial diets was compared with that of nymphs fed E. kuehniella eggs. Despite their lower body weights, predators produced on either artificial diet killed similar numbers of prey as their counterparts reared on lepidopteran eggs. Our study indicates that artificial rearing systems may be useful to further rationalize the production of this economically important biological control agent.     

Differential vulnerability of prey to an invading top predator: integrating field surveys and laboratory experiments

Abstract 1. A new top predator, the dragonfly Cordulegaster boltonii, invaded Broadstone Stream (U.K.) in the mid‐1990s. This provided a rare opportunity to assess the impact of a new, large carnivore on a community that has been studied since the 1970s and has one of the most detailed food webs yet published. The vulnerability of the resident species to the invader was assessed by integrating experiments, which examined discrete stages in the predation sequence, with empirical survey data. 2. Although the new predator preyed on nearly every macro‐invertebrate in the food web, vulnerability varied considerably among prey species. Size‐related handling constraints initially set the predator's diet, resulting in strong ontogenetic shifts, with progressively larger prey being added while small prey were retained in the diet, as predators grew. Within the size range of vulnerable prey, encounter rate limited the strength of predation, with mobile, epibenthic species being most at risk. Contrary to most studies of interactions between freshwater predators (usually stoneflies) and prey (usually mayflies), the new predator did not elicit avoidance responses from its prey, probably because it combined a highly cryptic feeding posture with an extremely rapid attack response. 3. The invader exploited its prey heavily in experiments, even at prey densities orders of magnitude above ambient. In the field, electivity reflected prey availability, as determined by mobility and microhabitat use, rather than prey abundance or active predator choice. Consequently, the invader had skewed effects within the prey assemblage, with sedentary, interstitial species being far less vulnerable than more active, epibenthic species, some of which, including a previous top predator, have declined markedly since the invasion. 4. By examining the predation sequence in detail and integrating surveys with experiments, species traits and system characteristics that determine the strength of trophic interactions may be identified, and their potential importance in natural food webs assessed. In so doing, greater insight can be gained into which species (and systems) will be most vulnerable to invading or exotic predators, an imperative in both pure and applied ecology.     

A growth/mortality trade-off in larval salamanders and the coexistence of intraguild predators and prey

Behavioral and morphological traits often influence a key trade-off between resource acquisition and vulnerability to predation, and understanding trait differences between species can provide critical insight into their interactions with other species and their distributions. Such an approach should enhance our understanding of the criteria for coexistence between species that can interact through both competition and predation (i.e. intraguild predators and prey). I conducted a common garden experiment that revealed strong differences between three guild members (larval salamanders Ambystoma laterale, A. maculatum, and A. tigrinum) in behavior, morphology, and growth in the presence and absence of a shared top predator (the larval dragonfly Anax longipes). All three species also reduced their activity and modified their tail fin depth, tail muscle length, and body length in response to non-lethal Anax. Species that act as intraguild predators were more active and could grow faster than their intraguild prey species, but they also suffered higher mortality in laboratory predation trials with Anax. I also used survey data from natural communities to compare the distribution of Ambystoma species between ponds differing in abiotic characteristics and predatory invertebrate assemblages. An intraguild prey species (A. maculatum) was found more reliably, occurred at higher densities, and was more likely to persist late into the larval period in ponds with more diverse invertebrate predator assemblages. Taken together, these results indicate that top predators such as Anax may play an important role in influencing intraguild interactions among Ambystoma and ultimately their local distribution patterns.     

Prey‐mediated changes in the selectivity of the predator Macrolophus pygmaeus (Heteroptera: Miridae)

When foraging in communities with mixed prey, generalist predators may be confronted with prey species that differ in quality, size and mobility and interact with one another. To examine prey selection, predation by Macrolophus pygmaeus (Heteroptera: Miridae) was recorded by providing a diet of either one or two prey species of Myzus persicae (third‐instar nymphs), Aphis gossypii (fourth‐instar nymphs), Trialeurodes vaporariorum (third‐instar nymphs) and Ephestia kuehniella (eggs). In the experiments, prey mobility, prey quality and prey biomass were considered. The biomass consumed by the predator was dependent on the combination of prey species and the quantity of biomass offered. In choice experiments with diets mixed of two prey species at equal densities, the predation to A. gossypii was significantly reduced in the presence of E. kuehniella but the rate of consumption of M. persicae, T. vaporariorum and E.kuehniella was not significantly affected by the coexistence of any other species in the mixed prey diet. When equal amounts of biomass from two prey species were provided in combination, the total consumed biomass was significantly reduced in the mixed prey diets composed of E. kuehniella eggs and aphid nymphs. Thus, under the mixed‐prey situation, prey selection by predators may be affected by interactions among prey species differing in traits such as quality, mobility and size.     

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.     

Prey size and predator density modify impacts by natural enemies towards mosquitoes

1. Interactions between multiple predators can modify prey risk and profoundly alter ecological community dynamics. Further, ontogenic prey size changes are known to mediate prey risk through refuge effects. Understandings of these biotic factors is important for robust quantifications of natural enemy effects on target species, yet their combined influence lacks investigation. 2. Functional responses were used to quantify the predatory impacts of Notonecta glauca (water boatman; Ng) and Gammarus pulex (river shrimp; Gp) towards four different larval instars of Culex pipiens in container-style habitats. Using conspecific pairs of predators, multiple predator effects (MPEs) of both predator species were examined across larval prey sizes, and prey preference tests were applied to examine prey selectivity across predator–prey body size ratios. 3. Both predators were able to feed on C. pipiens across their larval ontogeny; however, Ng consumed significantly more larvae than Gp. Functional responses of Ng were typically Type IIs, whereas Gp trended towards sigmoidal Type IIIs. Predation by pairs of Ng and Gp showed independent MPEs towards first-, third-, and fourth-instar stages (except predation by Gp at higher densities of fourth-instar) stages, whereas, for second-instar stages, Ng showed synergistic MPEs and Gp showed antagonistic MPEs. Both predators preferred late instar mosquitoes (Ng: fourth instar; Gp: third instar). These preferences reflected predator:prey weight–length ratios, showing that relative sizes of predators and prey are important factors in prey selectivity. 4. The results obtained in the present study demonstrate that MPEs, combined with intraspecific prey preferences, may modulate trophic interactions within ecosystems. Therefore, such effects should be increasingly considered to further the understanding of agent efficacies.     

The effect of prey size and prey density on the functional response,survival, growth and development of a predatory pentatomid bug,Podisus maculiventris say

Two experiments on the nymphal predation of Podisus maculiventris were conducted using Spodoptera litura larvae as prey. First experiment: The predator nymphs divided into three groups were reared individually from second instar to adult in a small vessel. Each nymph in the groups 1, 2 and 3 was allowed to attack the serially growing larvae (these were supplied at the rate of one per day) from 3-, 5- and 7-day old after hatching, respectively. The first prey used for the group 1 was so small that it was not only insufficient to satiate the predator but also was difficult to be searched out. But these disadvantages were soon recuperated due to the rapid growth of the prey and all nymphs could survive to adults. The survival rate of third and fourth instar nymphs in the group 3 was severely affected by vigorous counterattack of older prey larvae. Second experiment: The predator nymphs were individually reared either in a small vessel or in a large one at various rates of food supply (the prey larvae of 7-day old were used). The functional response curves obtained for each instar of the predator took a saturation type within a certain range of the prey density. The saturation level specific to each instar was generally higher for the predator reared in the large vessel than in the small one. The functional response of fourth and fifth instar nymphs was accelerated at a high prey density, viz. 16 larvae per vessel. Even at the low rate of food supply, viz. one larva per day per predator, the predator nymphs could survive to adults, but the size of resultant adults were abnormally small.     

The Feeding Behaviour of a Sit and Wait Predator Ranatra dispar (Heteroptera: Nepidae): The Effect of Prey Density and Age Structure on the Number of Prey Eaten

Functional response experiments were performed in the laboratory to examine the effect of prey density (as observed in the field) on feeding behaviour, and to measure handling-times and attack-rates for each instar and adult of Ranatra dispar Montandon (Heteroptera: Nepidae) feeding on five size-classes of its common prey, Anisops deanei Brooks (Heteroptera: Notonectidae). The most generally applicable response was the Type 2, although for both the predator fifth instar and adult female and male feeding on the two smallest prey sizes, the asymptote or plateau was not observed even at the highest prey density given. Generally, the handling-time increased as prey-size increased, and decreased as the predator size increased. The attack-rate surface was far more complex. For the first two predator instars (I and II), the maximum attack-rate occurred on the smallest prey sizes (1 and 2). The maximum attack-rate for predator instar III was almost the same for prey sizes 1 and 2, that of predator instar IV was greater for prey size 2, while in the three largest predator sizes (V, female and male), the maximum attack-rate was found for prey size 3. Predator instar V had the largest attack-rate values over all prey sizes, and both the predator adult female and male had lower attack-rates for various prey sizes than instars V, IV and, to some degree, III. The results support the suggestion that small predator instars will usually compete with large instars for prey, unless they are spatially or temporally separated. Observations in the field indicate that a distinct age-specific spatial distribution exists in R. dispar and the prey, A. deanei, with the smallest individuals being found predominantly in the shallow (littoral zone) water, while the larger individuals are found in the deeper water.     

Feedback between size balance and consumption strongly affects the consequences of hatching phenology in size‐dependent predator–prey interactions

In many size‐dependent predator–prey systems, hatching phenology strongly affects predator–prey interaction outcomes. Early‐hatched predators can easily consume prey when they first interact because they encounter smaller prey. However, this process by itself may be insufficient to explain all predator–prey interaction outcomes over the whole interaction period because the predator–prey size balance changes dynamically throughout their ontogeny. We hypothesized that hatching phenology influences predator–prey interactions via a feedback mechanism between the predator–prey size balance and prey consumption by predators. We experimentally tested this hypothesis in an amphibian predator–prey model system. Frog tadpoles Rana pirica were exposed to a predatory salamander larva Hynobius retardatus that had hatched 5, 12, 19 or 26 days after the frog tadpoles hatched. We investigated how the salamander hatch timing affected the dynamics of prey mortality, size changes of both predator and prey, and their subsequent life history (larval period and size at metamorphosis). The predator–prey size balance favoured earlier hatched salamanders, which just after hatching could successfully consume more frog tadpoles than later hatched salamanders. The early‐hatched salamanders grew rapidly and their accelerated growth enabled them to maintain the predator‐superior size balance; thus, they continued to exert strong predation pressure on the frog tadpoles in the subsequent period. Furthermore, frog tadpoles exposed to the early‐hatched salamanders were larger at metamorphosis and had a longer larval period than other frog tadpoles. These results suggest that feedback between the predator‐superior size balance and prey consumption is a critical mechanism that strongly affects the impacts of early hatching of predators in the short‐term population dynamics and life history of the prey. Because consumption of large nutrient‐rich prey items supports the growth of predators, a similar feedback mechanism may be common and have strong impacts on phenological shifts in size‐dependent trophic relationships.     

Learned Recognition of Intraspecific Predators in Larval Long-Toed Salamanders Ambystoma macrodactylum

The ability of prey to detect predators and respond accordingly is critical to their survival. The use of chemical cues by animals in predator detection has been widely documented. In many cases, predator recognition is facilitated by the release of alarm cues from conspecific victims. Alarm cues elicit anti‐predator behavior in many species, which can reduce their risk of being attacked. It has been previously demonstrated that adult long‐toed salamanders, Ambystoma macrodactylum, exhibit an alarm response to chemical cues from injured conspecifics. However, whether this response exists in the larval stage of this species and whether it is an innate or a learned condition is unknown. In the current study, we examined the alarm response of naïve (i.e. lab‐reared) larval long‐toed salamanders. We conducted a series of behavioral trials during which we quantified the level of activity and spatial avoidance of hungry and satiated focal larvae to water conditioned by an injured conspecific, a cannibal that had recently been fed a conspecific or a non‐cannibal that was recently fed a diet of Tubifex worms. Focal larvae neither reduced their activity nor spatially avoided the area of the stimulus in either treatment when satiated, and exhibited increased activity towards the cannibal stimulus when hungry. We regard this latter behavior as a feeding response. Together these results suggest that an anti‐predator response to injured conspecifics and to cannibalistic conspecifics is absent in naïve larvae. Previous studies have shown that experienced wild captured salamanders do show a response to cannibalistic conspecifics. Therefore, we conducted an additional experiment examining whether larvae can learn to exhibit anti‐predator behavior in response to cues from cannibalized conspecifics. We exposed larvae to visual, chemical and tactile cues of stimulus animals that were actively foraging on conspecifics (experienced) or a diet of Tubifex (naïve treatment). In subsequent behavioral treatments, experienced larvae significantly reduced their activity compared to naive larvae in response to chemical cues of cannibals that had recently consumed conspecifics. We suggest that this behavior is a response to alarm cues released by consumed conspecifics that may have labeled the cannibal. Furthermore, over time, interactions with cannibals may cause potential prey larvae to learn to avoid cannibals regardless of their recent diet.     

Repellent,antifeedant and toxic effects of Ageratum conyzoides (Linnaeus) (Asteraceae) extract against Helicovepra armigera (Hübner) (Lepidoptera: Noctuidae)

Repellent, antifeedant and toxic effect of crude hexane extract of Ageratum conyzoides were investigated against Helicoverpa armigera. In orientation bioassay, the extract exhibited dose-dependent repellency against neonates. Extract significantly increased the mortality and decreased growth of different larval stages when administrated orally in artificial diet. EC₅₀ value was at 0.11% for larval growth inhibition. Toxicity of the extract was manifested by high mortality of first instar larvae after 7 days of feeding on diet containing 0.05–0.4% of extract with LC₅₀ of 0.17%. Under choice bioassay, extract showed strong antifeedant activity against fifth instar larvae with DI₅₀ of 0.21%. In nutritional bioassay, extract significantly reduced RCR, RGR, ECI and ECD of fifth instar larvae with increased AD. When RGR were plotted against RCR, the growth efficiency of larvae fed on treated diet was significantly lower than the control fed larvae suggesting the antifeedant and toxic effect of extract.     

Impact of temperature and prey density on the predacious capacity and behaviour of Stethorus punctillum Weise

Temperature had various effects on the predacious efficacy of immature and mature stages of the coccinellid predator, Stethorus punctillum on the two-spotted spider mite, Tetranychus urticae. In the case of immature stages, food consumption at the lowest tested temperature (15°C) was significantly higher than that at higher temperatures (25 and 35°C). On the contrary, positive correlation between food consumption and temperature was evaluated in the case of adult predator. Regarding predator responses to different prey density, a high positive correlation between food consumption and prey density was evaluated among 4^th instar larvae of the predator, followed by adult predator, while younger instars did not show reasonable increases with increasing prey densities. These results confirm that larval and adult stages of S. punctillum exhibit “Type II” functional response. In conclusion, the 4^th instar larvae and adult predator are the most preferable stages in winter and summer crops to control T. urticae, respectively.     

Multi-predator effects across life-history stages: non-additivity of egg- and larval-stage predation in an African treefrog

The effects of multiple predators on their prey are frequently non‐additive because of interactions among predators. When prey shift habitats through ontogeny, many of their predators cannot interact directly. However, predators that occur in different habitats or feed on different prey stages may still interact through indirect effects mediated by prey traits and density. We conducted an experiment to evaluate the combined effects of arboreal egg‐stage and aquatic larval‐stage predators of the African treefrog, Hyperolius spinigularis. Egg and larval predator effects were non‐additive – more Hyperolius survived both predators than predicted from their independent effects. Egg‐stage predator effects on aquatic larval density and size and age at hatching reduced the effectiveness of larval‐stage predators by 70%. Our results indicate that density‐ and trait‐mediated indirect interactions can act across life‐stages and habitats, resulting in non‐additive multi‐predator effects.