Predation and searching efficiency of a ladybird beetle, Coccinella septempunctata Linnaeus in laboratory environment

The predation and searching efficiency of fourth instar of predatory C. septempunctata at various densities of mustard aphid, Lipaphis erysimi (Kaltenbach) and predator was investigated under laboratory conditions. The feeding rate of predatory stage decreased at increased prey- and predator densities. Highest percent (92.80%) prey consumption was observed at initial prey density and lowest percent (40.86%) prey consumption at highest prey density by the fourth instar, though the total prey consumption increased with increase in either prey- or predator densities. Similarly, the individual prey consumption was also highest at initial predator density and lowest at highest predator density owing to the mutual interference between the predators at higher densities. The area of discovery (searching efficiency) also decreased with increase in prey- and predator densities. Handling time of predator was highest at lower prey densities, which decreased with increased prey densities. The highest percentage of prey consumption at the prey density of 50 revealed that 1:50 predator-prey ratio was the best to reduce the pest population.     

Functional and numerical responses of Propylea dissecta (Col., Coccinellidae)

The functional response of a ladybeetle, Propylea dissecta, to increasing density of aphid, Aphis gossypii, was of the curvilinear shape depicting Holling's type II response with fourth instar larva being the most voracious stage when compared with adult male and female. Prey handling time by different predatory stages decreased from 65.45 to 8.72 min with increase in prey density from 25 to 800. The predator aggregation and high prey density reduces the searching efficiency of the predator. Area of discovery was highest (1.4437) when a single predator was searching at minimum aphid density (25) and lowest (0.0366) when eight predators were searching at a constant aphid density (200). Mutual interference and quest constants were 0.75 and 0.40, respectively. The reproductive numerical response, in terms of eggs laid, increased curvilinearly with prey density and female laid 70.5 ± 5.55 eggs when exposed to highest prey density (400) and 12.3 ± 0.79 eggs at lowest prey density (10). The similar shapes of both functional and reproductive responses indicate that both responses are interlinked and function simultaneously.     

淡色斧瓢虫对烟粉虱的捕食作用

研究了淡色斧瓢虫对烟粉虱的捕食作用.结果表明,淡色斧瓢虫对烟粉虱的功能反应呈Holling Ⅱ型.随着猎物龄期的增加,淡色斧瓢虫成虫的寻找效率(a)逐渐降低,处置时间(T_h)依次延长;随着淡色斧瓢虫幼虫龄期的增加,其对烟粉虱卵的寻找效率逐渐提高,处置时间依次缩短.淡色斧瓢虫成虫对烟粉虱卵的捕食效应随捕食者间干扰作用的增加而下降,捕食作用率(E)与淡色斧瓢虫数量（P）之间呈幂函数关系,即E=0.5205P^-0.6631.温度对淡色斧瓢虫的捕食效应影响显著,寻找效率和处置时间与温度间的函数关系为:a=- 0.0002T³+0.0166T²-0.3492T+3.2329,T_h=4×10^-7T³-3×10^-5T²+0.0006T-0.0009.     

Density-dependent effects of prey defences

In this study, we show that the protective advantage of a defence depends on prey density. For our investigations, we used the predator-prey model system Chaoborus-Daphnia pulex. The prey, D. pulex, forms neckteeth as an inducible defence against chaoborid predators. This morphological response effectively reduces predator attack efficiency, i.e. number of successful attacks divided by total number of attacks. We found that neckteeth-defended prey suffered a distinctly lower predation rate (prey uptake per unit time) at low prey densities. The advantage of this defence decreased with increasing prey density. We expect this pattern to be general when a defence reduces predator success rate, i.e. when a defence reduces encounter rate, probability of detection, probability of attack, or efficiency of attack. In addition, we experimentally simulated the effects of defences which increase predator digestion time by using different sizes of Daphnia with equal vulnerabilities. This type of defence had opposite density-dependent effects: here, the relative advantage of defended prey increased with prey density. We expect this pattern to be general for defences which increase predator handling time, i.e. defences which increase attacking time, eating time, or digestion time. Many defences will have effects on both predator success rate and handling time. For these defences, the predator’s functional response should be decreased over the whole range of prey densities. Received: 15 September 1999 / Accepted: 23 December 1999     

Depletion models can predict shorebird distribution at different spatial scales

Predicting the impact of habitat change on populations requires an understanding of the number of animals that a given area can support. Depletion models enable predictions of the numbers of individuals an area can support from prey density and predator searching efficiency and handling time. Depletion models have been successfully employed to predict patterns of abundance over small spatial scales, but most environmental change occurs over large spatial scales. We test the ability of depletion models to predict abundance at a range of scales with black-tailed godwits, Limosa limosa islandica. From the type II functional response of godwits to their prey, we calculated the handling time and searching efficiency associated with these prey. These were incorporated in a depletion model, together with the density of available prey determined from surveys, in order to predict godwit abundance. Tests of these predictions with Wetland Bird Survey data from the British Trust for Ornithology showed significant correlations between predicted and observed densities at three scales: within mudflats, within estuaries and between estuaries. Depletion models can thus be powerful tools for predicting the population size that can be supported on sites at a range of scales. This greatly enhances our confidence in predictions of the consequences of environmental change.     

Searching behavior and time budgets of the predator Podisus maculiventris

Searching behavior of the predaceous insect Podisus maculiventris (Say) (Heteroptera: Pentatomidae) was investigated in the laboratory to verify assumptions made in a predator search model. Female predators were placed into an arena containing 30 lima bean plants (Phaseolus lunatus L.), each having five numbered leaflets. Prey were third-instar larvae of Mexican bean beetle (Epilachna varivestis Mulsant) at two densities. Predators were observed for 4 h periods as they searched the plant canopy. Results showed that predators searched a greater area and for longer at low prey density than at high prey density. Predators apparently searched plants without using cues, did not search areas of the canopy repeatedly after attacks, and spent approximately 1 h handling prey. Predators spent more time resting than searching, and attack rates were negatively correlated with rest time, but were not correlated with search time. Long resting periods by predators may be a result of energy conservation. The implications for using predators such as P. maculiventris against pests in crops are (i) the predators' searching behavior limits the number of prey attacked, and (ii) the predator may be able to persist at low prey densities better than species with different searching behaviors.     

Behavioral responses to prey density by three acarine predator species with different degrees of polyphagy

Behavioral responses by three acarine predators, Phytoseiulus persimilis, Typhlodromus occidentalis, and Amblyseius andersoni (Acari: Phytoseiidae), to different egg and webbing densities of the spider mite Tetranychus urticae (Acari: Tetranychidae) on rose leaflets were studied in the laboratory. Prey patches were delineated by T. urticae webbing and associated kairomones, which elicit turning back responses in predators near the patch edge. Only the presence of webbing affected predator behavior; increased webbing density did not increase patch time. Patch time increased with increased T. urticae egg density in the oligophagous P. persimilis, but was density independent in the polyphagous species T. occidentalis and A. andersoni. Patch time in all three species was more strongly correlated with the number of prey encounters and attacks than with the actual prey number present in the patch. Patch time was determined by (a) the turning back response near the patch edge; this response decayed through time and eventually led to the abandonment of the patch, and (b) encounters with, and attacks upon, prey eggs; these prolonged patch time by both an increment of time spent in handling or rejecting prey and an increment of time spent searching between two successive prey encounters or attacks. Although searching efficiency was independent of prey density in all three species, the predation rate by P. persimilis decreased with prey density because its searching activity (i.e. proportion of total patch time spent in searching) decreased with prey density. Predation rates by T. occidentalis and A. andersoni decreased with prey density because their searching activity and success ratio both decreased with prey density. The data were tested against models of predator foraging responses to prey density. The effects of the degree of polyphagy on predator foraging behavior were also discussed.     

Foraging behaviour and capture success in perch, pikeperch and pike and the effects of prey density

The effect of school size on capture success in three different piscivores, perch Perca fluviatilis , pikeperch Stizostedion lucioperca and pike Esox lucius , was investigated. Roach Rutilus rutilus were used as prey in a pool experiment where individual predators were presented prey at densities of one, two, four, eight and 16 prey, respectively. Treatments were replicated seven times for each predator species. Perch was at first virtually unable to capture a prey from a school and suffered a significant confusion effect with increasing prey density. The effect, however, was limited in the long run, as the perch was a very effective predator in its hunting strategy where it singled out and repeatedly attacked single prey irrespective of prey density or school size. Pikeperch and pike were able to attack and capture prey at any prey density equally successfully and thus did not suffer from a confusion effect. Neither did these predators receive any apparent advantages from increasing prey density.     

Allee effects,extinctions, and chaotic transients in simple population models

Discrete time single species models with overcompensating density dependence and an Allee effect due to predator satiation and mating limitation are investigated. The models exhibit four behaviors: persistence for all initial population densities, bistability in which a population persists for intermediate initial densities and otherwise goes extinct, extinction for all initial densities, and essential extinction in which "almost every" initial density leads to extinction. For fast-growing populations, these models show populations can persist at high levels of predation even though lower levels of predation lead to essential extinction. Alternatively, increasing the predator's handling time, the population's carrying capacity, or the likelihood of mating success may lead to essential extinction. In each of these cases, the mechanism behind these disappearances are chaotic dynamics driving populations below a critical threshold determined by the Allee effect. These disappearances are proceeded by chaotic transients that are proven to be approximately exponentially distributed in length and highly sensitive to initial population densities.     

Consumption rate, functional response and preference of the predaceous mite Iphiseius degenerans to Tetranychus urticae and Eutetranychus orientalis

The functional response of females of the phytoseiid mite, Iphiseius degenerans (Berlese), to increasing densities of females of its prey, Tetranychus urticae Koch and Eutetranychus orientalis Klein, on bean leaves, were studied under laboratory conditions. Our results indicated that the predator consumed significantly more items of E. orientalis than of T. urticae at all densities treatments. Daily consumption of the predator increased with increasing prey density until a plateau was reached-maximum number of prey consumed was ca. 4 for T. urticae and ca. 12 for E. orientalis. A Type II functional response was determined by a logistic regression model. The highest estimated value a (instantaneous rate of attack) and the lowest value of T ( h ) (handling time) were found for the predator feeding on E. orientalis. Prey selection was evaluated by simultaneously presenting both prey species to the predator in various ratios and at increasing densities. I. degenerans showed a higher predation rate and higher preference for E. orientalis at all the ratios and prey densities tested. This may be due to the smaller size or the inactivity of E. orientalis and the inability of the predator to cope with the webbing of T. urticae. Our results suggest that I. degenerans can be considered a suitable biological control candidate based on its preference for E. orientalis in the Mediterranean region.     

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.     

Effects of predator density and duration of predator occupancy on crustacean abundance and diversity in experimental pools

Keystone predators, by reducing the abundance of competitively superior prey, may have indirect positive effects on weak competitors, possibly increasing their abundance or preventing local competitive exclusion. By analogy to the Intermediate Disturbance Hypothesis, we would expect species diversity to peak at intermediate predator densities. In a replicated artificial pool experiment, we examined the relationships between density of the backswimmer Notonecta maculata (0, 1, 2, and 4 per 30 l pool) and invertebrate taxon diversity over an 11-week period of predator occupancy. Diversity reached high levels at high predator density sooner than at intermediate density. At the end of the experiment, taxon diversity was greatest at densities of 2 and 4 Notonecta per pool. While the overall predator density–diversity curve was in line with the intermediate disturbance hypothesis, the reduction in diversity from intermediate to high predator density was not statistically significant. Density of the preferred prey Daphnia magna decreased with Notonecta density, while densities of the smaller cladocerans Moina brachiata and Ceriodaphnia spp. increased. Suppression of Daphnia at high Notonecta densities may partially explain the increase in Moina and Ceriodaphnia densities. However, most of the relationship between Notonecta and the smaller cladocerans appears to be independent of Daphnia, suggesting complex interactions within the community. Our results suggest that keystone predation plays a strong role in structuring this community. Although diversity did not decrease significantly at the highest predator density as predicted, such a decrease may be more likely for pools with longer durations of predator occupancy or with higher predator densities.     

Consequences of refuge for the functional response of Dermestes ater (Coleoptera: Dermestidae) to Musca domestica (Diptera: Muscidae)

It is well known that a predator has the potential to regulate a prey population only if the predator responds to increases in prey density and inflicts greater mortality rates. Predators may cause such density-dependent mortality depending on the nature of the functional and numerical responses. Yet, few studies have examined the relationship between the addition of refuges and the characteristic of functional response fits. We investigated whether addition of a refuge changed the type of functional response exhibited by Dermestes ater on Musca domestica, comparing the inherent ability of D. ater to kill houseflies in the absence and in the presence of refuge. An additional laboratory experiment was also carried out to assess handling and searching times exhibited by D. ater. Logistic regression analyses revealed a type III functional response for predator–prey interaction without refuge, and results were described by the random predator equation. The mean number of prey killed did not differ between experimental habitats, indicating that the addition of refuge did not inhibit predation. However, predators that interacted with prey without refuge spent less time searching for prey at higher densities, increasing predatory interaction. We concluded that this interaction may be weak, because data from experiments with refuge fitted poorly to models. However, the high variability and the nonsignificance of the data from the experiment with refuge show the importance of refuge for promoting spatial heterogeneity, which may prevent prey extinction.     

Handling time promotes the coevolution of aggregation in predator-prey systems

Predators often have type II functional responses and live in environments where their life history traits as well as those of their prey vary from patch to patch. To understand how spatial heterogeneity and predator handling times influence the coevolution of patch preferences and ecological stability, we perform an ecological and evolutionary analysis of a Nicholson-Bailey type model. We prove that coevolutionarily stable prey and searching predators prefer patches that in isolation support higher prey and searching predator densities, respectively. Using this fact, we determine how environmental variation and predator handling times influence the spatial patterns of patch preferences, population abundances and per-capita predation rates. In particular, long predator handling times are shown to result in the coevolution of predator and prey aggregation. An analytic expression characterizing ecological stability of the coevolved populations is derived. This expression implies that contrary to traditional theoretical expectations, predator handling time can stabilize predator-prey interactions through its coevolutionary influence on patch preferences. These results are shown to have important implications for classical biological control.     

Responses of Episyrphus balteatus DeGeer (Diptera: Syrphidae) in relation to prey density and predator size

Laboratory experiments were conducted to determine the functional and numerical responses of the aphidophagous hover fly Episyrphus balteatus DeGeer (Diptera: Syrphidae) to different densities of 4th instar black bean aphids, Aphis fabae Scopoli (Homoptera: Aphididae), on broad bean, Vicia faba L. (Fabaceae). Two different-sized larvae of predators were tested in different densities of similar-sized prey to determine whether functional response parameters depended on the body sizes of predator and prey. In numerical response experiments, gravid E. balteatus females were exposed individually to different densities of 4th instars of A. fabae on cut sections of the broad bean plant, V. faba L. Logistic regression suggested a type II functional response for both larval sizes of E. balteatus. The searching efficiency (a) of the larger larvae was higher than that of the smaller ones. Prey consumption was higher, and handling time (T_h) was lower for larger larvae than smaller ones. The theoretical maximum number of A. fabae nymphs eaten by the different-sized larvae was 125 and 269 nymphs per day. Larger E. balteatus larvae are more efficient predators for aphid management strategies. The reproductive numerical response, in terms of the number of eggs laid, increased curvilinearly with increasing prey density, but the proportion of eggs laid (egg number/prey density) decreased as the initial density of prey increased.     

Effect of prey density and hunger state on capture of krill, Nyctiphanes australis Sars, by Australian salmon, Arripis trutta (Bloch & Schneider)

When hungry salmon were offered a simulated (unattainable) swarm of krill with high central density, the density of the region first attacked increased with increasing attack readiness (measured as snout contacts). When allowed to capture krill in a range of prey densities, hungry salmon captured krill most successfully (in terms of capture/contacts) in the lowest density and less efficiently with increasing densities. However, salmon still captured a greater number of krill in a given time in high compared to low densities. When salmon in three different hunger states were exposed to a simulated swarm of krill, those with the lowest hunger level most often attacked the lowest prey densities first. Salmon with a high hunger level most often attacked the highest prey density first. Those fish with a moderate hunger level chose an intermediate prey density. This behaviour can be interpreted as a hunger-dependent feeding strategy. The ways in which swarming of the prey might influence capture by the predator are considered.     

Effects of substrate on interactions between juvenile sea scallops (Placopecten magellanicus Gmelin) and predatory sea stars (Asterias vulgaris Verrill) and rock crabs (Cancer irroratus Say)

We investigated the effect of substrate (glass bottom, sand, granule, pebble) on predation of juvenile sea scallops (Placopecten magellanicus) by sea stars (Asterias vulgaris) and rock crabs (Cancer irroratus) at two prey sizes (11-15 mm and 24-28 mm shell height), and two prey densities (10 and 30 scallops per aquarium) in laboratory experiments. Specifically, we quantified predation rate and underlying behaviours (proportion of time a predator spent searching for and handling prey, encounter rate between predators and prey, and various outcomes of encounters). We detected a significant gradual effect of particle size of natural substrates on sea star predation: specifically, predation rate on and encounter rate with small scallops tended to decrease with increasing particle size (being highest for sand, intermediate for granule, and lowest for pebble). Substrate type did not significantly affect predation rates or behaviours of sea stars preying on large scallops or of rock crabs preying on either scallop size classes. Other factors, such as prey size and density, were important in the scallop-sea star and scallop-rock crab systems. For example, predation rate by sea stars and crabs and certain sea star behaviours (e.g. probability of consuming scallops upon capture) were significantly higher with small scallops than with large scallops. As well, in interactions between small scallops and sea stars, predation rate and encounter rate increased with prey density, and the proportion of time sea stars spent searching was higher at low prey density than high prey density. Thus, substrate type may be a minor factor determining predation risk of seeded scallops during enhancement operations; prey size and prey density may play a more important role. However, substrate type still needs to be considered when choosing a site for scallop enhancement, as it may affect other scallop behaviours (such as movement).     

Functional response of Cosmoclopius nigroannulatus (Hem.: Reduviidae) to different densities of Spartocera dentiventris (Hem.: Coreidae) nymphae.

This study evaluated the functional response of the predator Cosmoclopius nigroannulatus on first instar nymphae of Spartocera dentiventris, both species associated with Nicotina tabacum. The experiment was carried out in laboratory conditions: 27 +/- 1 degree C; 80 +/- 5%, RH; 12 h, photophase. Ten newly emerged adults of each sex of C. nigroannulatus were used in each of five densities (5, 15, 25, 35, and 45 individuals) of S. dentiventris nymphae. The predators were observed every 24 h for five days, when the number of dead and/or consumed nymphae was recorded. The results showed a positive correlation between the number of ingested nymphae and the increase in prey density. Females ingested more nymphae than the males. The estimated handling time per prey (Th) was higher in males (3.07 h) than in females (1.93 h), with total handling time (Th x Na) increased with density. Other components of the functional response, such as attack rate (a'), searching time (Ts), and search efficiency (E) showed, in neither males nor females, a negative correlation regarding density. The results indicated a higher predatory efficiency in the females. The components of the functional response fitted significantly the randomic model of the Holling discs equation (Na = N [1 - exp[- a'(T - ThNa)]]), evidencing a functional response of type II.     

低剂量农药作用下拟环纹豹蛛对褐飞虱的功能反应及搜寻行为

在室内研究了低剂量农药作用下拟环纹豹蛛对褐飞虱的功能反应及搜寻行为。结果表明,拟环纹豹蛛对褐飞虱的功能反应均符合Holling Ⅱ模型,且在最适的低剂量农药(V_药∶V_水为2∶5 000甲胺磷,12∶5 000杀虫双和1∶20烯丙菊酯)刺激下,蜘蛛捕食褐飞虱的数量是最多的,蜘蛛用于对1头褐飞虱进行制服和取食等活动所需的时间（处理猎物时间,Th）短,仅分别为未经农药处理的蜘蛛所需时间的0.45、0.32和0.35倍;而施用较高剂量8∶5 000的甲胺磷、28∶5 000的杀虫双和6∶20的烯丙菊酯的蜘蛛,处理猎物时间长得多,依次是经最适低剂量3种农药作用的蜘蛛所用时间的9.70、13.60和13.62倍。拟环纹豹蛛在稻株上的搜寻行为属非随机搜索型,在相同的低剂量农药作用下,蜘蛛的搜寻效应随飞虱密度的增加而下降;但在相同的猎物密度下,蜘蛛的搜寻效应与使用的农药浓度有密切的关系,在上述最适低剂量农药作用下,豹蛛对猎物的寻找效应最佳,其在稻株上的总滞留时间依次是未施药情形的1.56、1.49、1.48倍,总搜寻距离依次是未施药情形的1.36、1.42、1.39倍。据此,进一步证实了低剂量农药增强了稻田蜘蛛的控虫效能和对害虫的相对活力。     

Bifurcation analysis of a predator-prey model with predators using hawk and dove tactics

Most classical prey-predator models do not take into account the behavioural structure of the population. Usually, the predator and the prey populations are assumed to be homogeneous, i.e. all individuals behave in the same way. In this work, we shall take into account different tactics that predators can use for exploiting a common self-reproducing resource, the prey population. Predators fight together in order to keep or to have access to captured prey individuals. Individual predators can use two behavioural tactics when they encounter to dispute a prey, the classical hawk and dove tactics. We assume two different time scales. The fast time scale corresponds to the inter-specific searching and handling for the prey by the predators and the intra-specific fighting between the predators. The slow time scale corresponds to the (logistic) growth of the prey population and mortality of the predator. We take advantage of the two time scales to reduce the dimension of the model and to obtain an aggregated model that describes the dynamics of the total predator and prey densities at the slow time scale. We present the bifurcation analysis of the model and the effects of the different predator tactics on persistence and stability of the prey-predator community are discussed.