Host odours enhance the responses of adult Rhyzopertha dominica to male-produced aggregation pheromone

The behavioural responses of adult male and female Rhyzopertha dominica (Coleoptera, Bostrichidae) to blends of host volatiles and male-produced aggregation pheromone were observed in a four-arm airflow olfactometer. The odour sources used were five pheromone-releasing males each on a single maize grain (lower maize-volatiles blend), five pheromone-releasing males on 500 g of maize (higher maize-volatiles blend) or the host volatiles emanating from 500g of maize (maize volatiles alone). Multiple-choice tests, in which volatiles from all three odour sources were presented in the exposure chamber at the same time, were used to study odour preferences of the males and females. Both sexes showed strongest attraction to the higher maize volatile blend but there were significant sex differences in response to the odour sources. Males spent significantly more time than females in the zone with only maize volatiles, and females spent significantly more time in the higher maize-volatiles zone. However, when odour sources were offered singly, females gave numerically greater responses than males to all sources although this difference was statistically significant for only the lower maize-volatiles blend. As males are more attracted than females to host odours alone it is suggested that they may be more highly adapted to seek out new hosts while females are more inclined to locate a food source by following the pheromone signals produced by males. However, both sexes responded most strongly to the odour source comprising aggregation pheromone with the higher proportion of maize volatiles.     

Population ecology of Thrips palmi (Thysanoptera:Thripidae) in orchid farms in Thailand

The population dynamics and in-farm distribution of Thrips palmi Karny were evaluated in two orchid farms, one in the Bangkok metropolitan area and the other in Ratchaburi Province, Thailand. Yellow sticky traps were hung above the orchid canopy (three traps/220 m²), and the trapped adults were counted weekly from November 2009 to October 2010 at the first location and from January to December of 2012 at the second location. T. palmi nymphs and adults were both counted weekly on 300 orchid panicles in an area of 2,200 m² at both locations. We found that the T. palmi population dynamics at the first location was represented by a W-shaped curve, with the distribution clumped throughout the year, whereas the curve was J-shaped at the second location, with the distribution mostly clumped. Qualitative surveys of thrips species were conducted on ten different orchid genera in ten locations and on 20 different Dendrobium hybrids in 20 locations. Five random sampling plots (1 × 4 m²) were examined on each farm. The results indicated that only T. palmi was found on every farm. Moreover, quantitative comparison among five orchid types revealed that Dendrobium flowers with darker color are more attractive to T. palmi. However, this phenomenon does not coincide with Oncidium sp. and Mokara sp. because of their unfavorable morphology for thrips behavior.     

Effect of Watermelon Silver Mottle Virus on the Life History and Feeding Preference of Thrips palmi

Thrips-borne tospoviruses cause numerous plant diseases that produce severe economic losses worldwide. In the disease system, thrips not only damage plants through feeding but also transmit causative agents of epidemics. In addition, thrips are infected with tospoviruses in the course of virus transmission. Most studies on the effect of tospoviruses on vector thrips have focused on the Tomato spotted wilt virus–Frankliniella occidentalis system. Thus, we focused on another thrips-borne tospovirus, Watermelon silver mottle virus (WSMoV), to examine the effect of virus infection on its vector, Thrips palmi. In this study, the direct and indirect effects of WSMoV on the life history traits and feeding preference of T. palmi were examined. The survival rate and developmental time of the WSMoV-infected larval thrips did not differ significantly from those of the virus-free thrips. Comparing the developmental time of larval thrips fed on the healthy plants, thrips-damaged plants, and thrips-inoculated plants (the WSMoV-infected plants caused by thrips feeding), feeding on the thrips-damaged plants reduced the developmental time, and the WSMoV infection in host plants partially canceled the effect of thrips damage on the developmental time. In addition, no significant variations between the virus-free and WSMoV-infected adult thrips regarding longevity and fecundity were observed. These results implied that WSMoV did not directly affect the life history traits of T. palmi, but the WSMoV infection indirectly affected the development of T. palmi through the virus-infected plants. Furthermore, feeding preference tests indicated that T. palmi preferred feeding on either the thrips-damaged plants or the thrips-inoculated plants to the healthy plants. The effect of tospoviruses on the life history and feeding preference of vector thrips might vary among host plants, virus species, vector species, and environmental factors.     

In the presence of red light,cucumber and possibly other host plants lose their attractability to the melon thrips <Emphasis Type="Italic">Thrips palmi</Emphasis> (Thysanoptera: Thripidae)

The melon thrips, Thrips palmi Karny (Thysanoptera: Thripidae), is a serious agricultural pest of many crops. Previous studies have shown that red light decreases the number of Thrips palmi in greenhouses. In order to understand how red light affects T. palmi, we examined the behavioral responses to host plants that were irradiated with a red light-emitting diode panel (660 nm) in an environment with natural or fluorescent (normal-white) light. When T. palmi were allowed to move freely around in the experimental arena, we found that fewer individuals were attracted to plants irradiated by red light than to plants under normal light illumination. We then used a sticky trap of green coloration to exclude olfactory and visual stimuli associated with the host plants in order to test binary choice behavior in T. palmi. The number of thrips attracted to the green sticky trap irradiated with red light was approximately half of that without red light irradiation. This is the first study to show that an addition of red light can change the behavior of insects, leading to an avoidance of green targets in an environment of normal illumination.     

Mating Aggregations and Mating Success in the Flower Thrips, Frankliniella schultzei (Thysanoptera: Thripidae), and a Possible Role for Pheromones

Aggregations of Frankliniella schultzei males were observed on the corollas of Hibiscus rosasinensis and Gossypium hirsutum flowers in southeast Queensland. Aggregations were seen only on the upper surfaces of corollas but may have occurred on other flower parts, which were hidden from view. Conspecific females entered aggregations and a small proportion of them mated [18% (n = 163), H. rosasinensis; 30% (n = 181), G. hirsutum]. Most females (87 and 72%, respectively) that did not mate in aggregations walked to other flower parts. Behavior was difficult to observe on these parts, but mating was sometimes observed there. The number of females that landed within aggregations on the upper surfaces of both H. rosasinensis and G. hirsutum corollas was highly correlated with the number of males (r = 0.88, r = 0.93, respectively; P < 0.001). Significantly more mating pairs were observed in high-density aggregations (mean ± SE, 1.10 ± 0.22 and 4.44 ± 0.48, respectively) than in low-density aggregations (0.37 ± 0.11 and 1.67 ± 0.29, respectively) (P < 0.05) on flowers of both species. More F. schultzei females were attracted to sticky traps baited with live conspecific males set among flowering Ipomoea indica (mean ± SE, 8.83 ± 0.32) and G. hirsutum (10.90 ± 0.79) plants than to control traps (0.10 ± 0.05 and 0.70 ± 0.25, respectively) (P < 0.05), presumably in response to male-produced pheromones. Significantly more females were attracted to traps with high male densities than to traps with low densities. We found no statistical evidence that aggregation size influenced mating success (proportion males that mated). Mating success, however, should be evaluated with respect to mating on all flower parts and not just the upper surfaces of corollas. The results of this study constitute the first behavioral evidence for an attractant sex pheromone in thrips.     

An observation on the embryonic development in Thrips palmi (Thysanoptera: Thripidae) eggs obtained by an artificial oviposition setup

Thrips palmi is an important insect pest of vegetables and ornamental crops worldwide. Besides direct damage caused by feeding, it transmits several tospoviruses in a persistent-propagative manner. Eggs of T. palmi are microscopic and embedded within plant tissue by the sharp ovipositor of adult female. In the present study, an artificial oviposition setup has been standardized for T. palmi. Eggs of T. palmi were harvested in sterile water between two thin membranes. The developmental stages of T. palmi embryo were studied starting from oviposition up to hatching by inverted and confocal reflection microscopy. Energids were homogeneously distributed at an early stage of development. The anterior end of the egg curved with a constriction post 38 h. Initiation of tissue organization, mouthparts, appendages, compound eyes were observed at different time points. Appendages were well developed and segmentation was prominent post 70 h. The embryo was completely developed at around 80 h and hatched by 86 h post oviposition at 28 °C temperature. The study first time reports the embryonic development of T. palmi that would be helpful in detailed investigations of thrips developmental biology and evolution.     

Response of a parasitoid fly, Gymnosoma rotundatum (Linnaeus) (Diptera: Tachinidae) to the aggregation pheromone of Plautia stali Scott (Hemiptera: Pentatomidae) and its parasitism of hosts under field conditions

We studied the attraction of a tachinid fly, Gymnosoma rotundatum (Linnaeus) to the male-produced aggregation pheromone of the brown-winged green bug, Plautia stali Scott, its parasitism on the bug, and its seasonal occurrence in the field. The tachinid fly was continuously attracted to the aggregation pheromone from spring to autumn and utilized the bugs as hosts. Our field experiment to clarify the effect of the pheromone on parasitism demonstrated that parasitism occurred only in female bugs baited with synthetic aggregation pheromone and did not occur in females without the pheromone. The parasitoid flies therefore appeared to use the bug’s pheromone as a host-finding kairomone. The pheromone attracted not only female flies but also males. Male flies may increase their chance of encountering pheromone-attracted females by waiting near pheromone sources. The tachinid develops multiple generations in active hosts from spring to autumn and overwinters in dormant hosts. Thus, G. rotundatum seems to be highly adapted to using P. stali as its host, and it is a potentially important biological control agent of P. stali populations in the field.     

Economic thresholds of Thrips palmi (Thysanoptera: Thripidae) for eggplants in a greenhouse

Thrips palmi Karny has recently been implicated in damage to eggplant fruits. This thrips causes cosmetic scars on and deforms fruits, thus lowering their market value. To investigate the relationship between the density of thrips and the resulting damage, experimental plots with initial release densities of 0, 20, 40, 70, and 100 adults thrips per plot were established under greenhouse conditions. Thrips density (for flower sampling = x ₁; for sticky trap = x ₂) was monitored using flower sampling and blue sticky trap counts, and was found to be directly related to the proportion of damaged-fruits (y) and the reduction in fruit yield: significant relationships were found for the flower samples (y = 1.2261x ₁ ? 0.6232, r ² = 0.8582) and for the trap catches (y = 11.667 ln(x ₂) ? 9.5, r ² = 0.8896). The proportion of damaged fruits that could be tolerated from an economic perspective, based on the cost of controlling the thrips population chemically and the market value of the fruit, was 6.67–11.76 %; this translated into economic thresholds of 1.05–1.5 adults and/or larvae of T. palmi per flower or 4.91–10.17 adults T. palmi per four-day sticky card count.     

Male-specific (Z)-9-tricosene stimulates female mating behaviour in the spider Pholcus beijingensis

Chemical signals play an important role in spider sexual communication, yet the chemistry of spider sex pheromones remains poorly understood. Chemical identification of male-produced pheromone-mediating sexual behaviour in spiders has also, to our knowledge, not been reported before. This study aimed to examine whether chemically mediated strategies are used by males of the spider Pholcus beijingensis for increasing the probability of copulation. Based on data from gas chromatography–mass spectrometry analysis, electroantennography assay and a series of behavioural tests, we verified that (Z)-9-tricosene is a male-specific compound in the spider P. beijingensis. This compound acts as an aphrodisiac: it increases the likelihood that a female will mate. Mate-searching males release (Z)-9-tricosene to stimulate sexual behaviour of conspecific females. In the two-choice assay, however, sexually receptive females show no preference to the chambers containing (Z)-9-tricosene. This indicates that the male pheromone of P. beijingensis is not an attractant per se to the conspecific females. This is, to our knowledge, the first identification of a male-produced aphrodisiac pheromone in spiders.     

Sex ratio and female sexual status of the coconut pest, Oryctes monoceros (Coleoptera: Dynastidae), differ in feeding galleries and pheromone-baited traps

Oryctes monoceros is a serious coconut pest, causing up to 40% damage in tropical Africa. Synthetic aggregation pheromone, ethyl 4-methyloctanoate, has been used to lure adults to traps. Traps with pheromone plus decaying palm material captured a high proportion of males. This raises the question whether individuals, which damage palms are receptive to the pheromone. We studied the sex ratio of the insects feeding on coconuts and those attracted to pheromone traps. Sixty two percent of adults from feeding galleries on living coconut palms were females. Pheromone with rotting palm material lured 43% females. To investigate the reasons for this difference, we compared the reproductive system of females lured to the odour traps or feeding in coconut galleries, or present in old rotting stems. Ninety six percent of the females trapped by pheromone had mated, and were sexually mature. In the galleries on living palms, 46% of females were immature, and 24% had not mated. In old rotting stems where eggs are laid and larvae develop, a mixture of 52% mated and 48% virgin females was found. Therefore, the pheromone together with the odour of rotting coconut stems signals a reproduction site to beetles, particularly mature females. In practice, the pheromone-baited traps will help in reducing the dissemination of gravid females, but will not affect directly the numbers of immature ones attacking palms. Our results show that when using pheromones for monitoring or controlling insects, the physiological status of the insects may have unexpected effects on the outcome.     

Modelling mating success of saproxylic beetles in relation to search behaviour, population density and substrate abundance

We compared the efficiency of two mate-finding strategies exploited by representatives of the beetle families Cisidae and Anobiidae (genus Dorcatoma) that live inside fruiting bodies of wood-decaying fungi. In the Cisidae both sexes are attracted to host odour, but no pheromones seem to be present (nonpheromone strategy). In the Dorcatoma species only the females are attracted to host odour, but having found a host they attract males with a sexual pheromone (pheromone strategy). With a simulation model, we compared the efficiency of the two strategies at four densities of trees hosting fungal fruiting bodies and at three relative densities of insects. We found only small differences in efficiency between the two strategies at high relative densities of conspecific individuals, regardless of host tree density. The pheromone strategy was relatively more efficient when the relative density of insects or the density of host trees decreased. Thus, species adopting the nonpheromone strategy are probably more sensitive to habitat fragmentation and more likely to decline and go extinct at low population densities (because of Allee effects) than species using the pheromone strategy. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Disruption of sexual communication in the mirid bug Lygocoris pabulinus by hexyl butanoate

1 The metathoracic scent gland in Lygocoris pabulinus contains mostly hexyl butanoate. As secretions of this gland in Heteroptera may serve as an alarm pheromone, we determined whether hexyl butanoate is released by disturbed bugs, and whether this compound disrupts sexual attraction of L. pabulinus males towards females. 2 Undisturbed males and females, and disturbed males released less than 100 ng/h hexyl butanoate, whereas disturbed females released a highly variable amount, ranging from 25 ng/h to more than 1 μg/h. 3 In the field, traps with virgin females and rubber septa containing 20 mg hexyl butanoate, caught a total of one male in a month. In control traps without hexyl butanoate, 36 males were caught in the same period. 4 In Y‐track olfactometer tests, males were not attracted to virgin females when a dispenser with 20 mg hexyl butanoate was placed in the bottle with females. Males were attracted to females when the dispenser was placed downwind from the females, but upwind from the point of male release. 5 These results suggest that males are not repelled by hexyl butanoate, but that this compound inhibits sex pheromone release in females. Application possibilities for pest management are discussed.     

Detection of the Quarantine Species Thrips palmi by Loop-Mediated Isothermal Amplification

Thrips palmi (from the order Thysanoptera) is a serious insect pest of various crops, including vegetables, fruits and ornamental plants, causing significant economic losses. Its presence constitutes a double threat; not only does T. palmi feed on the plants, it is also a vector for several plant viruses. T. palmi originated in Asia, but has spread to North and Central America, Africa, Oceania and the Caribbean in recent decades. This species has been sporadically noted in Europe and is under quarantine regulation in the European Union. For non-specialists its larval stages are indistinguishable morphologically from another widespread and serious insect pest Frankliniella occidentalis (a non-quarantine species in the European Union) as well as other frequently occurring thrips. In this study, we have developed a loop-mediated isothermal amplification protocol to amplify rDNA regions of T. palmi. The results were consistent whether isolated DNA or crushed insects were used as template, indicating that the DNA isolation step could be omitted. The described method is species-specific and sensitive and provides a rapid diagnostic tool to detect T. palmi in the field.     

Odour-baited traps influence thrips capture in proximal unbaited traps in the field

Two field experiments examined the distance over which an attractant odour of a volatile chemical could influence thrips capture in proximal traps that were without the odour. In each experiment a star‐shaped array of water traps consisted of a centre trap with or without an odour surrounded by odourless traps at 0.5, 1, 2, 5, and 10 m in eight equally spaced radial arms 45° apart. Experiments ran for 47 h (centre trap: ethyl nicotinate) or 7 h (centre trap: ethyl isonicotinate). Each had four replicates. With ethyl nicotinate, more thrips were trapped in the centre‐baited traps than in the unbaited centre traps (63×, 7×, 98× and 200× for total thrips, Thrips tabaci Lindeman ♀, and Thrips obscuratus Crawford ♀and ♂, respectively) (Thysanoptera: Thripidae). More total thrips and T. tabaci♀ were trapped in the centre traps baited with ethyl isonicotinate than in unbaited centre traps (21× for both). For ethyl nicotinate, numbers of T. obscuratus in unbaited traps downwind from the baited centre trap declined by 50% within 0.4 m (♀) and 2 m (♂) and by 95% within 3 m (both ♀ and ♂) based on model predictions. For ethyl isonicotinate, numbers of T. tabaci in unbaited traps downwind from the baited centre trap declined by 50% within 1.3 m and by 95% within 10 m based on model predictions. Wind direction was an important factor in the degree and direction of thrips capture with the highest thrips capture downwind from the centre trap with odour. There was no increase in numbers of T. tabaci in any traps without odour in the ethyl nicotinate‐centred array. Differences in trapping patterns between thrips species and odours indicated that there were thrips species–odour specific interactions. Experiments examining differences between traps with and without a thrips attractant odour need to be designed very carefully to ensure meaningful results especially in enclosed and/or low‐wind indoor situations.     

Optimizing a pheromone lure for the sugar beet root maggot fly,Tetanops myopaeformis

We previously described a putative aggregation pheromone in adults of the sugar beet root maggot, Tetanops myopaeformis (von Röder) (Diptera: Ulidiidae), comprising nine compounds identified from males. Here, we conducted a series of experiments aimed at simplifying the blend of compounds necessary to achieve attraction as well as determining the dose that maximizes captures when formulated into an attractant lure. In all experiments, females showed stronger and more consistent evidence of attraction than males. White sticky traps baited with different blends of pheromone compounds that included the major component, (R)-(−)-2-nonanol, showed significantly higher female captures relative to those baited with blends that excluded the major component. (R)-(−)-2-nonanol alone was at least as effective as any blend that included this compound with other minor pheromone components. Lures using racemic 2-nonanol were as effective as the (R) enantiomer for both females and males, with some evidence of weak attraction to the (S) enantiomer (which is not produced by males) observed as well. Maximum capture rates using racemic 2-nonanol were estimated to occur with doses of ca. 795.5 and 621.6 mg for females and males, respectively. Addition of 2-nonanol lures to standard orange sticky stake traps currently used to monitor flies increased captures of both sexes. The pheromone lure developed here could improve trapping efficiency of current monitoring programs for T. myopaeformis and may also be used to develop other management tools for this important pest of sugar beet.     

Identification of a Male-Produced Pheromone Component of the Citrus Longhorned Beetle,Anoplophora chinensis

The Asian wood-boring beetle Anoplophora chinensis (Forster) (Coleoptera: Cerambycidae) is an important pest of hardwood trees in its native range, and has serious potential to invade other areas of the world through worldwide commerce in woody plants and wood products. This species already has been intercepted in North America, and is the subject of ongoing eradication efforts in several countries in Europe. Attractants such as pheromones would be immediately useful as baits in traps for its detection. Because long-range pheromones are frequently conserved among closely related species of cerambycids, we evaluated two components of the volatile pheromone produced by males of the congener A. glabripennis (Motschulsky), 4-(n-heptyloxy)butan-1-ol and 4-(n-heptyloxy)butanal, as potential pheromones of A. chinensis. Both compounds subsequently were detected in headspace volatiles from male A. chinensis, but not in volatiles from females. Only 4-(n-heptyloxy)butanol elicited responses from beetle antennae in coupled gas chromatography-electroantennogram analyses, and this compound attracted adult A. chinensis of both sexes in field bioassays. These data suggest that 4-(n-heptyloxy)butan-1-ol is an important component of the male-produced attractant pheromone of A. chinensis, which should find immediate use in quarantine monitoring for this pest.     

Predicting the potential geographic distribution of Thrips palmi in Korea,using the CLIMEX model

Thrips palmi Karny, melon thrips was introduced and first recorded in 1993 in Korea. This species has become a serious pest of vegetable and ornamental crops. The CLIMEX simulation was applied to T. palmi to predict its potential geographic distribution in Korea under the Representative Concentration Pathway (RCP) 8.5 climate change scenario. In the CLIMEX simulation, the ecoclimatic index was calculated, and compared in each simulated year and each simulated location. The map comparisons show good agreements between simulated and present distributions of T. palmi, indicating that the CLIMEX model has promising potential for prediction of future distributions of this species in Korea. In the near future, until the year 2020, all the western and eastern parts of Korea show favorable to marginal suitability for T. palmi populations in the fields. After the year 2040, potential distributions shift from no persistence to favorable for establishment and persistence from coastal to interior regions of the Korean peninsula, except for a north‐eastern interior region which is the northernmost part of a high mountainous (Baekdu‐Daegan) area in Korea. Based on the simulation results, the geographical distribution of T. palmi will expand over its current weather restrictions in the near future under a severe climate change scenario. Thus, pest management measures and strategies should be re‐evaluated in Korea, and should include further studies on interspecific competition and ecosystem changes due to climate changes.     

Can Mass Trapping Reduce Thrips Damage and Is It Economically Viable? Management of the Western Flower Thrips in Strawberry

The western flower thrips Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) is a cosmopolitan, polyphagous insect pest that causes bronzing to fruit of strawberry (Fragaria x ananassa). The main aim of this study was to test whether mass trapping could reduce damage and to predict whether this approach would be economically viable. In semi-protected strawberry crops, mass trapping of F. occidentalis using blue sticky roller traps reduced adult thrips numbers per flower by 61% and fruit bronzing by 55%. The addition of the F. occidentalis aggregation pheromone, neryl (S)-2-methylbutanoate, to the traps doubled the trap catch, reduced adult thrips numbers per flower by 73% and fruit bronzing by 68%. The factors affecting trapping efficiency through the season are discussed. Damage that would result in downgrading of fruit to a cheaper price occurred when bronzing affected about 10% of the red fruit surface. Cost-benefit analysis using this threshold showed that mass trapping of thrips using blue sticky roller traps can be cost-effective in high-value crops. The addition of blue sticky roller traps to an integrated pest management programme maintained thrips numbers below the damage threshold and increased grower returns by a conservative estimate of £2.2k per hectare. Further work is required to develop the F. occidentalis aggregation pheromone for mass trapping and to determine the best timing for trap deployment. Mass trapping of thrips is likely to be cost-effective in other countries and other high-value crops affected by F. occidentalis damage, such as cucumber and cut flowers.     

The aggregation pheromone system of a Dendroctonus bark beetle in Guatemala

Analysis of the volatile contents of hindguts from a Dendroctonus bark beetle in Guatemala indicated distinct differences from those of D. frontalis, which it resembles morphologically. The major compound in emergent males, most of which disappears during initial attack of host trees (Pinus maximinoi), was identified as 1-phenylethanol. During initial attack females produce frontalin and trans-verbenol, males produce myrtenol, and both sexes produce 1-heptanol and 2-heptanol. Synthetic frontalin attracted predominantly males to small traps, but the use of larger trapping surfaces increased the proportion of females caught. The heptanols, as well as 1-phenylethanol, apparently had some regulatory effect on the response to frontalin, but myrtenol and trans-verbenol were inactive. The results suggest that frontalin is the key compound responsible for the aggregation of this species, but the response-regulating mechanism differs from that of D. frontalis.     

Time allocation of Orius sauteri in attacking Thrips palmi on an eggplant leaf

Orius sauteri (Poppius) (Heteroptera: Anthocoridae) is a polyphagous predator used as a biological control agent of palm thrips, Thrips palmi (Karny) (Thysanoptera: Thripidae). We studied O. sauteri's searching efficiency, time allocation on a leaf, leaving tendency, and attacking of prey. Approximately 78% of the encountered prey was eaten. Searching for prey was concentrated for 86% of the time on the lower leaf side, where palm thrips are usually found. Patch residence times on empty leaves were different from those on leaves with T. palmi larvae. Walking activity was not affected by the thrips density, and walking took place during 64% of the total search time. The leaving tendency of O. sauteri was affected by the time from patch entry and the presence or absence of palm thrips, but not by the thrips density. If prey were present, the leaving tendency decreased (multiplication factor 0.327), resulting in longer giving‐up times than when no prey was present. The fact that the leaving tendency increases when patch exploitation lasts longer suggests that not much time is wasted on patches where encounters with prey are scarce.