Effect of plants on the searching efficiency of a generalist predator: the importance of predator-prey spatial association

In most studies of tritrophic interactions, the effect of plants on predators is confounded with changes in prey and predator behaviors after an encounter event. Here, we estimate how the effect of plants on prey distribution (in the absence of the predator) and on predator foraging behavior (in the absence of prey) may influence predation rate of Orius insidiosus (Say) (Heteroptera: Anthocoridae) in 11 plant by prey species combinations. The within-leaf distributions of O. insidiosus and its prey overlapped most on bean plants. The predator's foraging behavior (e.g., walking speed, turning rate) also differed among plant species. Simulations, using the prey distribution data and predator's foraging patterns on leaf surfaces of each plant species, show that, overall, the searching efficiency of O. insidiosus was higher on leaves of bean and corn than of tomato. However, the predator's searching efficiency was not consistent within plant species. Thus, the combined effect of plants directly on the predator and indirectly through the prey influenced the predator's searching efficiency.     

Feeding behaviour determining differential capture success of evasive prey in underyearling European perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis> L.) and roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis> (L.))

The effect of feeding behaviour on the prey capture efficiency of young-of-the-year European perch and roach was investigated in laboratory experiments using planktonic crustaceans possessing different escape abilities—Daphnia sp. and Cyclops sp. Two sets of experiments were performed. In the first set, the feeding efficiency and behaviour of 270 fish individuals were determined by stomach content analyses and video record evaluations. In the second set of experiments, analysis of attack-effort, which was evaluated as attack-distance and repeated strikes, was undertaken. Except for situations in which Daphnia was offered at high densities, the feeding efficiency of perch was significantly higher compared to roach in all other combinations of prey types and densities. Roach consumed significantly less prey compared to perch when feeding exclusively on the evasive Cyclops and when it was offered in a 1:1 ratio mixture with Daphnia. The mean swimming speed was similar in both fish species, but behavioural differences were evident during prey search and capture. Perch swam through the aquaria in short and fast movements that were interrupted by many stops. Roach exhibited rather continuous swimming that was punctuated by slowdowns instead of stops. The perch attacks were very intensive and repeated strikes occurred, particularly when feeding on evasive Cyclops. On the other hand, roach revealed strong schooling behaviour restricting the fish during inspection of the experimental aquaria. The distinct differences in feeding efficiency between perch and roach were demonstrated to be closely related to differences in their feeding behaviour. Discontinuous searching for prey, vigorous attacks, occurrence of repeated strikes and the absence of schooling increased perch prey capture efficiency, particularly when foraging on evasive copepods.     

The effect of prey type and density on the foraging efficiency of juvenile perch, (Perca fluviatilis)

The foraging efficiency of juvenile perch (Perca fluviatilis), feeding on two types of prey, was studied in laboratory experiments. Waterfleas (Daphnia magna) and phantom midge larvae (Chaoborus flavicans) were offered in a range of densities, either separately or combined. Perch fed more efficiently on each prey type separately than when both were mixed. Foraging efficiency decreased with an increase of mixed prey density with both prey types present in equal numbers, but also when the proportion of Chaoborus increased. This could be caused by the existence of different hunting techniques, each of which is fully efficient in the presence of one prey type only. In the presence of two prey types, the predator constantly has to switch from one hunting technique to another.     

Analysis of the aggregation behavior in the larvae ofHarmonia axyridis Pallas (Coleoptera: Coccinellidae) to prey colony

Summary The searching and handling behaviors ofHarmonia axyridis larvae to the colony ofRhopalosiphum padi were experimentally examined and the processes of their aggregation to the prey colony was analyzed. All the instar larvae searched for the prey at random and they have no preference to the prey colony, but except the 1st instar they tend to aggregate to the plants with prey colonies. The 1st instar larvae tend to stay on the plants they once located. The 2nd to 4th instar larvae often emigrate from the plants without prey colony but seldom emigrate from the plants with prey colonies, and consequently, they aggregate to the plants with prey colonies. The expense of time to eat prey (in the 2nd and 3rd instars) and the change of searching behavior for the prey after feeding (in the 3rd and 4th instars) are responsible for the larval concentration to prey colony as a trapping effect for predators to prey colony.     

The importance of honeydew as food for larvae of Chrysoperla carnea in the presence of aphids

Larvae of the common green lacewing Chrysoperla carnea are predacious and feed on a wide range of small, soft‐bodied arthropods. In addition to their feeding on prey arthropods to cover their nutritional requirements for growth and development, the consumption of non‐prey foods such as honeydew has been reported. It is commonly believed that these food supplements are primarily exploited by the larvae when prey is scarce or of low nutritional quality. Here, we assess whether C. carnea larvae also use honeydew when high‐quality aphid prey are readily available. In a choice experiment, the feeding behaviour of C. carnea larvae was observed in the presence of both aphids and honeydew. The larvae were starved, aphid‐fed, or honeydew‐fed prior to the experiment. The time spent feeding on honeydew compared with feeding on aphids was highest for starved larvae and lowest for honeydew‐fed larvae. Among the three treatments, the aphid‐fed larvae spent the most time resting and the least time searching. In an additional experiment food intake was assessed in terms of weight change when larvae were provided with an ad libitum supply of either aphids or honeydew. Larvae yielded a significant lower relative weight increase on honeydew compared with aphids. The reduced weight increase on honeydew was compensated when larvae were subsequently provided with aphids, but not when honeydew was provided again. This study showed that (i) prior honeydew feeding reduces overall aphid consumption, and (ii) larvae do consume honeydew even after they have been given ad libitum access to aphids. The fact that larvae of C. carnea still use honeydew as a food source in the presence of suitable prey underlines the importance of carbohydrates as foods.     

Ambulatory dispersal behavior ofNeoseiulus fallacis (Acarina: Phytoseiidae) in relation to prey density and temperature

Ambulatory dispersal behavior ofNeoseiulus fallacis (Garman) was studied in the laboratory to evaluate within-plant movement in relation to temperature and prey density. Adult femaleN. fallacis were confined in 2.5-cm-diameter arenas on the abaxial surface of excised corn leaves. Four temperatures (23, 28, 33, and 39° C) and prey densities ranging from 0 to 55 spider mite eggs per cm² were used. The walking paths of these mites were traced, digitized and used to calculate turning angles, walking speeds and turning rates. A computer simulation of walking behavior used this information to model mite ambulatory behavior and predict dispersal rates.Neoseiulus fallacis behavior while on whole corn leaves was quantified to verify the results of the simulation. The results showed thatN. fallacis will follow a leaf or arena edge (edge-walking) at all temperatures and prey densities. In addition, this behavior was used to the exclusion of the other types of behavior such as resting, and random-walk type search when prey egg density was less than 4 eggs per cm². The exclusion of edge-walking behavior from the model caused the model to underestimate substantially the dispersal rates leaves. These data suggest that there are at least two recognizable types of ambulatory search used byN. fallacis—the random-walk type, which is used when prey density is high (searching within prey patches), and the edge-walking behavior, which is used when prey density is low. This behavior allows the mite to travel rapidly from leaf to leaf in search of new prey patches.     

Handling time, prey preference, and functional response for Chrysoperla rufilabris in the laboratory

The predaceous larvae of Chrysoperla rufilabris (Burmeister) exhibited some success-motivated searching, particularly when feeding on Heliothis virescens (F.) eggs, but handling time did not decrease with experience. Handling time for H. virescens larvae was more than twice that for eggs. H. virescens larvae were preferred to cotton aphids (Aphis gossypii (F.)) while aphids were preferred to H. virescens eggs. C. rufilabris larvae exhibited a linear functional response to the three prey types tested, over the prey densities tested.USDA, ARS, Retire. Present address: 200 Highland, College Station, TX 77840, USA     

Feeding by larvae of Hypoatherina tropicalis (Pisces: Atherinidae) and its relation to prey availability in One Tree Lagoon,Great Barrier Reef,Australia

Synopsis Feeding of and food availability for larvae of Hypoatherina tropicalis were investigated in One Tree Lagoon, Great Barrier Reef, Australia, during November 1981 and January 1982. These surface-dwelling larvae and their microzooplankton prey were sampled as near to simultaneously as possible on 12 occasions during the daytime. Larvae of all sizes (5–17 mm SL) fed successfully over the observed range of mean prey densities (12–235 per liter), and the overall feeding incidence was 98.9%. Larger larvae consumed greater numbers and more categories of prey than did smaller larvae. Larvae selected copepods of all sizes, and nauplii, gastropods, bivalves, and foraminiferans that were greater than 75 ¢s mm in width. Tintinnids (mostly 37–74 µm in width) were generally avoided by larvae, but were occasionally important in the diets when they constituted more than 60% of the total available prey, regardless of the density of the selected prey categories. Larvae less than 14 mm SL ingested meroplankton (gastropods, bivalves, foraminiferans, and polychaetes) in direct relation to the densities available, and without regard to the densities of copepods available. However, the largest larvae (14–17 mm SL) ingested meroplankton in inverse relation to the density of copepods available, indicating that larvae consumed more meroplankton when the concentration of copepods was low. Such flexibility and opportunism in feeding behavior may increase the larvae's chances of obtaining adequate nutrition during periods of suboptimal feeding conditions.     

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.     

Feeding by marine fish larvae: developmental and functional responses

Synopsis The relationship between prey consumption rate and prey concentration (functional response), and its change with growth (developmental response) were examined in the laboratory for three species of marine fish larvae: bay anchovy Anchoa mitchilli (Engraulidae), sea bream Archosargus rhomboidalis (Sparidae) and lined sole Achirus lineatus (Soleidae). The major objective was to determine relative predatory abilities of the larvae by fitting feeding rate data to developmental and functional response models. Feeding success, prey capture success, attack rates, handling times and search rates were estimated. Prey consumption rates and attack rates of bay anchovy usually were highest, but at the lowest prey level (50 per liter) first-feeding sea bream larvae had the highest consumption rate. Sea bream could consume prey at near-maximum rates at prey levels lower than those required by the other species. As larvae grew, time searching per attack decreased rapidly for all species, especially at low prey levels. Handling time also decreased, but most rapidly for bay anchovy. Search rates were highest for bay anchovy and lowest for lined sole. Bay anchovy had the best apparent predation ability, but when previous results on larval growth rates, survival rates and growth efficiencies were considered, sea bream larvae were the most efficient predators and the least likely of the three species to be limited by low prey levels.     

The feeding ecology of the pit-making ant lion larva,Myrmeleon bore: Feeding rate and species composition of prey in a habitat

The prey species composition and feeding rate of the pit-making ant lion larva,Myrmeleon bore Tjeder, which inhabits open sandy areas, were examined. Not less than 30 prey species, most of which were ants, were collected during a research period of 1.5 years. First instar larvae most often (81.1%) captured ants. Although 3rd instar larvae captured larger-sized prey than individuals of any other instar, they also captured small prey. The feeding rate of 3rd instar larvae was estimated by using the frequency of observed predation (FOP; (no. of ant lions handling a prey)/(total no. of pits observed)), the prey-handling time and the rhythm of daily foraging activity. FOP ofM. bore larvae was constant on the whole from spring to autumn. It was estimated that each captured 1.25 prey per day on average during this period. This estimate, however, was the feeding rate for days on which there was no rain. Assuming that the larvae cannot capture prey due to pit destruction when there is more than 10 mm of rainfall per day, the figure was reduced to 1.03 prey/day. The estimated feeding rate was evaluated with reference to larval foraging behavior.     

Effects of satiation on feeding and swimming behaviour of planktivores

Hunger affects the feeding and swimming behaviour in fish. After 36 h of food deprivation, the feeding and swimming behaviour of Pseudorasbora parva (Cyprinidae) was studied under different prey densities (0.5, 1, 2, 5, 10 and 25 of Daphnia pulex per liter). The initial feeding rates showed marked variations in relation to prey availability. Under high prey densities, the initial feeding rate of fish was higher and subsequently decreased faster, when compared to those feeding under low prey densities. At higher prey densities, two factors were involved: that of higher prey encounter rates and also the attainment of food satiation at a faster rate. Across all prey densities, the feeding rates of fish reached a plateau after satiation. The swimming speed of fish was found to be negatively related to the prey density and a significant change in swimming speed was noted as being directly related to the level of satiation. It was found that the increasing satiation level greatly influenced the handling time and reactive volume of predator, which finally caused reduced feeding rates.     

Satiation time and predatory behaviour of the dragonfly nymph Mesogomphus lineatus

Summary The dragonfly nymph Mesogomphus lineatus satiated predating 21 larvae of Culex fatigans in 50 minutes; number of attack and predatory efficiency decreased precipitously from 2 attacks/min and 70% during the first 10 minutes of feeding to 0.04 attack/min and 0.01% efficiency respectively during the sixth 10 minutes interval. The nymphs fed after 6, 12, 18, 24, 36 or 42 hours of deprivation consumed 4, 13, 15, 20, 21 or 21 larvae; apparently, the maximum appetite is returned after about 36 hours of deprivation. Satiation time, which was 50 minutes at the density of 15 larvae/aquarium, decreased to 30 minutes in aquaria containing 200 larvae. The nymphs predated increased the number of prey when they were exposed to higher densities of C. fatigans and Anopheles stephansi larvae, and pupa of C. fatigans. They consumed equal weight (but different number) of Culex and Anopheles larvae at all the tested prey densities and selectively selected Culex larva over the pupa or Anopheles larva. Comparative analyses suggest that the dragonfly nymphs deserve serious consideration as larvivorous predators.     

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.     

The effect of invertebrate and vertebrate predators on the foraging movements of Ischnura elegans larvae (Odonata: Zygoptera)

SUMMARY.

• 1 The foraging movements of late instar Ischnura elegans larvae were monitored in laboratory experiments to study the effects of predators on larval feeding behaviour.
• 2 Ischnura larvae are sit-atid-wait, or ambush, foragers, moving occasionally between perches in search of profitable feeding sites. Larval foraging movements, monitored at different densities of Daphnia prey, increased significantly when prey were absent.
• 3 In experiments without prey, larval movement was inhibited by the presence of fish predators, as well as by invertebrate predators (Notonecta glauca), but not by closely related, non-predatory invertebrates (Corixa punctata) or physical disturbance of the water (intermittent air bubbles).
• 4 Further experiments varied Ischnura hunger levels (0–8 days without food) and illumination (light or dark) with and without notonectid predators. Hunger had no consistent effect on penultimate instar behaviour but final instar foraging activity was significantly modified: movements increased after 4 days starvation and decreased again after 8 days. This response was suppressed by the presence of predators. Both larval instars moved significantly less often in the light, even when predators were absent.
• 5 These phenotypically flexible predator-avoidance responses are likely to decrease the risk of predation by both visual and tactile predators. However, predators clearly have an important influence on the feeding niche of Ischnura larvae, and may decrease the overall feeding efficiency, growth rate, and survival of larvae by constraining their movement in search of profitable feeding sites.

     

Prey to predator size ratio influences foraging efficiency of larval Aeshna juncea dragonflies

We investigated foraging behaviour of larval dragonflies Aeshna juncea in order to examine the significance of prey density and body size in predator-prey dynamics. A. juncea were offered separately three size-classes of Daphnia magna at low and high densities. The data were collected with direct observations of the foraging individuals. We found that large A. juncea larvae could better enhance their intake of prey biomass as prey size and prey density increased than their smaller conspecifics. However, increasing feeding efficiency of both larval instars was constrained by declining attack success and search rate with increasing prey size and density. With small D. magna, in contrast to large A. juncea, small A. juncea increased their searching efficiency as prey density increased keeping D. magna mortality rate at a constant level. In a predator-prey relationship this indicates stabilizing potential and feeding thresholds set by both prey density and prey-predator size ratio. Attack success dropped with prey size and density, but did not change in the course of the foraging bout. For both A. juncea sizes prey handling times increased as more medium and large prey were eaten. The slope of the increase became steeper with increasing prey-predator size ratio. These observations indicate that components of the predator-prey relationship vary with prey density, contrary to the basic assumptions of functional response equations. Moreover, the results suggest that the effects of prey density change during the ontogeny of predators and prey.     

The searching behaviour of Anthocoris confusus (Reuter) in relation to prey density and plant surface topography

• 1 The searching behaviour of A.confusus females was investigated in an artificial arena.
• 2 Females showed an increase in the frequency of turning movements following feeding and this concentrated search in a small area. As a result more prey were found in areas where prey distribution was clumped.
• 3 If no prey was encountered within 5–8 min the search track straightened out.
• 4 First and second instar nymphs searching on broad bean plants moved faster on the undersides of leaves which were the sites most likely to support aphid populations. A considerable proportion of available time was wasted in periods of inactivity.
• 5 It was concluded that while plant topography strongly influences search pattern, the underlying trend demonstrated in these experiments was of advantage to predators searching for colonial prey.

     

The effect of rearing the ladybirdHarmonia axyridis onEphestia kuehniella eggs on the response of its larvae to aphid tracks

Larvae ofHarmonia axyridis Pallas (Col., Coccinellidae) exhibited two walking patterns during prey search. Extensive search occurred when searching for prey patches and was characterized by long linear paths and a fast speed. Intesive search, which appeared after the ingestion of a prey in a patch, resulted from a lowering of the linear speed and an increase in the number of stops and angular speed. When larvae reared on the aphidAcyrthosiphum pisum Harris (Hom., Aphidae) crossed an artificial substratum previously contaminated by this prey, they changed their path direction and adopted intensive search. They probably perceived aphid odor tracks and consequently modified their walking pattern. This gustatory capacity probably allowed very mobile larvae to locate prey patches more rapidly and improve encounter with preys in every patch.H. axyridis larvae reared on a substitute prey, the eggs ofEphestia kuehniella Zeller (Lep., Pyralidae), for more than a hundred generations, also changed their path orientation but retained extensive search. The weak response of these larvae to aphid tracks may have resulted from either a decrease in their sensitivity to gustatory aphid stimuli or their difficulty in associating aphid odor with aphid presence. These larvae needed more time and more preliminary encounters than larvae reared on aphids before catching prey.     

龟纹瓢虫幼虫的食物搜索行为

本试验采用Nakamuta(1982)装置研究龟纹瓢虫Propylea japonica(Thunberg)幼虫的食饵搜索行为,结果表明:1.五种刺激均能激发搜索行为由广域型转换为地域集中型;2.摄食时间与GUT呈正相关;3.摄食的最后一个食饵大小决定GUT的长短;4.摄食后0—15秒的搜索速度小、弯曲角度大.     

Group predatory behavior by the assassin bugAgriosphodrus dohrni Signoret (Hemiptera: Reduviidae)

Nymphs of Agriosphodrus dohrniSignoret (Reduviidae) have a strong gregariousness and show group predatory behavior. This study was conducted to clarify adaptive significance of group predation of this species, including laboratory observations and 6-year field surveys. In the laboratory, observations on both solitary and group attacking against armyworms were made at varying prey size classes to compare the capture success rate by solitary predators with that by groups. The efficiency in capturing the prey was significantly higher in group attacking at any prey size class compared. Data obtained from the field surveys indicated the tnedency for searching nymphs to feed in group and to increase the number of predators feeding per prey item with increasing prey size. Average sizes of prey captured were also larger in group feeding throughout the nymphal stage. In particular, it was remarkable that, when prey were “creeping” types, the upper size limit of prey eaten was dramatically increased.