Visual and Chemical Cues Used in Host Location and Acceptance by a Dipteran Parasitoid

Locating potential hosts for egg laying is a critical challenge in the life history of many insects. Female insects in several orders have evolved mechanisms to find hosts by using olfactory and visual signals derived from their hosts. We describe visual and chemical cues used by the dipteran parasitoid Apocephalus paraponerae (Diptera: Phoridae) in the location and acceptance of its host ant Paraponera clavata (Hymenoptera: Formicidae: Ponerinae). Our results show that A. paraponerae uses the visual cue of ant body size when locating hosts at short range and that these flies lay more eggs in ants that retain their surface chemicals than in ants with these chemicals removed. We compare the cues used by A. paraponerae with cues used by tephritid fruit flies in location and acceptance of their hosts, and we suggest further avenues for the study of host location, acceptance, and host discrimination of A. paraponerae and other parasitoids of ants.     

Host Location of a Pupal Parasitoid in a Tritrophic System Compared to a Model Offering Mechanosensory Cues Only

Several species of hymenopteran parasitoids are able to locate concealed pupal hosts by vibrational sounding. However, the specific role of this technique of mechanosensory host location has not yet been investigated in a natural, tritrophic system with multiple cues. Here we compared the host location of the pupal parasitoid Xanthopimpla stemmator in a tritrophic system with corn borer pupae in maize stem to the behavior on a paper model offering mechanosensory cues only. In general, the behavioral pattern and the behavioral states exhibited by host-searching female parasitoid were identical in the model and in the tritrophic system, while quantitative aspects differed. Our results demonstrate that vibrational sounding maintains its significance for host location in an environment with multiple cues, and that additional cues may increase responsiveness of females.     

红火蚁的危害及其防治

一种非常危险的入侵性害虫——红火蚁近年在我国台湾和香港登陆,对中国大陆和亚洲其他地区构成严重威胁。本文介绍了红火蚁的危害以及对红火蚁的防治方法。     

Colony Wide Behavioral Contexts of Stridulation in Imported Fire Ants (Solenopsis invicta Buren)

Red imported fire ants, Solenopsis invicta, possess stridulatory organs and stridulate in a variety of contexts. We used a stethoscope mounted microphone to study stridulation at the colony level in the context of emigration, disturbance, and excavation. In conjunction with preliminary observations of nest and foraging activities, our results suggest stridulation serves multiple functions in S. invicta. Stridulation was not significantly increased in colonies during responses to disturbance, and only marginally during colony emigration. Colonies involved in excavation, however, exhibited a significant increase in stridulatory activity. Four possible explanations for the function of stridulation in this context are discussed in relation to the stridulatory behavior of individuals, solitary wasps, and published literature on formicid stridulation.     

Larviposition, Host Cues, and Planidial Behavior in the Sound-Locating Parasitoid Fly Homotrixa alleni (Diptera: Tachinidae)

Some groups of tachinid flies deposit mobile first-instar larvae (or planidia) on or near their host. Flies within one such group, the tribe Ormiini, parasitize singing species of ensiferan Orthoptera and use sound for long distance host location. However, what induces tachinids to larviposit and whether planidia use any cues actively to locate their host remains poorly known. This paper examines the larviposition and planidial behavior of the ormiine Homotrixa alleni in relation to its bushcricket host, Sciarasaga quadrata. Sound alone was sufficient to elicit larviposition in gravid female H. alleni, where females arriving at an arena placed over a speaker broadcasting host song deposited an equal number of planidia in the presence or absence of a silent S. quadrata. Flies were observed to larviposit by forcibly expelling planidia up to 6 cm in a forward direction from the fly, with less than half of the trials with a host present resulting in physical contact between the host and the fly. In the host's absence, flies walked around the arena significantly more often, remained on the arena for the experimental duration (10 min), and changed orientation frequently. In the host's presence, flies generally maintained a position facing the host, stayed in the quadrant of first approach, and typically flew off the arena within 2 min of arrival. When the oncoming fly approached a forward facing host, more planidia were found in the arena's center (i.e., closer to the host) than in the no-host or rearward-facing host treatment. Planidia experimentally placed on a circular arena averaged 1 cm of movement in 15 min but none of the following cues–host song, host song with song-vibration transmission, a silent host, and a silent host with host movement-vibration transmission–significantly affected the direction or distance planidia traveled. At 20 ± 1°C, over half of the planidia died within 1 h and all died within 2 h of deposition. The significance of these results in relation to reproductive strategies and parasitism is discussed.     

Sex Allocation by a Parasitoid Wasp (Hymenoptera: Ichneumonidae) to Different Host Species: A Question for the Mechanism of Host Size Estimation

Sex allocation by the polyphagous solitary pupal parasitoid wasp Pimpla luctuosa Smith to a small host species, Galleria mellonella (L.), and a large host species, Mamestra brassicae L., was investigated to test whether female wasps responded to hosts of different sizes across different host species. In the experiments, both host species were presented to each test female wasp. Primary and secondary sex ratio experiments revealed that female wasps laid more female eggs in larger pupae of each host species, indicating that female wasps recognized size differences within host species. The wasp sex ratio (male ratio) from M. brassicae, however, was much higher than that expected on the basis of the sex ratio curve from different-sized G. mellonella. Larger hosts of each host species yielded larger wasps, indicating that the host size estimation by female wasps across different host species was incomplete or was not simple. These results suggested that P. luctuosa evaluated host size not only by physical measures such as dimension but also by other unknown measures. A possible explanation for the adaptiveness of different sex ratio responses by Pimpla luctuosa to different host species was discussed.     

Adaptive Host Preference and the Dynamics of Host–Parasitoid Interactions

Models of two independent host populations and a common parasitoid are investigated. The hosts have density-dependent population growth and only interact indirectly by their effects on parasitoid behavior and population dynamics. The parasitoid is assumed to experience a trade-off in its ability to exploit the two hosts. Three alternative types of parasitoid are investigated: (i) fixed generalists whose consumption rates are those that maximize fitness; (ii) “ideal free” parasitoids, which modify their behavior to maximize their rate of finding unparasitized hosts within a generation; and (iii) “evolving” parasitoids, whose capture rates change between generations based on quantitative genetic determination of the relative attack rates on the two hosts. The primary questions addressed are: (1) Do the different types of adaptive processes stabilize or destabilize the population dynamics? (2) Do the adaptive processes tend to equalize or to magnify differences in host densities? The models show that adaptive behavior and evolution frequently destabilize population dynamics and frequently increase the average difference between host densities.     

Post-Release Host-Specificity Testing of Pseudacteon tricuspis, a Phorid Parasitoid of Solenopsis Invicta Fire Ants

Inherent in any biological control program is the risk of nontarget effects. Pseudacteon tricuspisBorgmeier, a parasitoid phorid fly, has been introduced to the United States from South America as a potential biocontrol agent of the red imported fire ant, Solenopsis invictaBuren. We conducted tests of host specificity on introduced populations of P. tricuspis, which are attracted to alarm pheromones released by their hosts during events such as mound disturbances and interspecific interactions. We monitored disturbed mounds of S. invicta and its close congener, S. geminata(F.), during the expansion of P. tricuspis across north Florida and after populations had been established for ~3 years. We also tested host acceptance in established populations of P. tricuspis by offering trays containing S. invicta, S. geminata, and 14 additional ant species representing 12 different non-Solenopsis genera. Although P. tricuspiswas commonly observed to hover over and attempt to oviposit on S. invicta, we never observed any parasitization attempts on any other ant species. As predicted by laboratory tests, released populations of P. tricuspis appear to be highly host specific and pose no obvious threat to nontarget species.     

Host location by the gerrid egg parasitoidTiphodytes gerriphagus (marchal) (Hymenoptera: Scelionidae)

Mechanisms used by a scelionid egg parasitoid to locate its gerrid host eggs were studied using field experiments. Parasitism by Tiphodytes gerriphaguswas higher in exclosures containing adults of Limnoporus dissortisthan in cages without gerrids. Higher densities of adult gerrids were associated with higher rates of parasitism, but dispersion of aquatic vegetation did not apparently affect rates of parasitism. T. gerriphagusmay use visual cues, chemical cues, or a combination of both to locate their hosts. We discuss the possible influence of T. gerriphaguson the reproductive behavior of its hosts.     

Host Age Response in the Parasitoid Wasp Spalangia cameroni (Hymenoptera: Pteromalidae)

Female Spalangia cameroni produced more offspring from younger house fly pupae, both when given a choice of host ages and when not given a choice. Host age did not affect offspring survivorship. Offspring were larger when they had developed on younger hosts and the effect was independent of offspring sex. Having previously parasitized old hosts versus young hosts did not reduce a female's production of offspring in subsequent hosts. Females distinguished between young and old hosts both in the light and in the dark. Females did not distinguish between host ages prior to physical contact with the host but could distinguish by the time they first began exploring a host by tapping it with their antennae; thus, they could distinguish before drilling into a host.     

Host Location by Apterous Aphids After Escape Dropping from the Plant

In response to approaching enemies aphids may drop from their host plant to the ground. The risk of predation on the ground, desiccation and host loss, create the need for efficient host location by the dropping aphids. Most studies have focused on the factors that influence dropping behavior; only a few have addressed host location after dropping from the plant. We assessed post-dropping behavior in apterous Macrosiphoniella artemisiae (Boyer de Fonscolombe) (Aphididae), which feed on Artemisia arborescens L. Vibration of the apical bud induced dropping in 36% of the colony members. Dropping rates were highest in mature aphids (63%). In the experimental arena (without ground predators), nearly all mature aphids that were dropped 13 cm from the plant, found their way back in ca. 40 s. The location process may be based on visual cues, as M. artemisiae is capable of visually discriminating between host and non-host targets and apparently does not react to volatiles emitted from the plant.     

Short Distance Cues Used by the Adult Parasitoid <Emphasis Type="Italic">Microctonus hyperodae</Emphasis> Loan (Hymenoptera: Braconidae,Euphorinae) for Host Selection of a Novel Host <Emphasis Type="Italic">Listronotus oregonensis</Emphasis> (Coleoptera: Curculionidae)

Insect parasitoids use host kairomone to detect their hosts. However, in parasitoid species that attack adult hosts, the mobility of adult insect may mean that the host can move away for kairomone sources. The effect of Listronotus oregonensis LeConte (Coleoptera: Curculionidae) adult sex, feces and movement on host selection behavior by Microctonus’’ hyperodae Loan (Hymenoptera: Braconidae; Euphorinae) females was evaluated in the laboratory. We hypothesized that, in addition of using host kairomones, parasitoids of adult stage should use host movement for host selection. The sex of L. oregonensis did not affect the host selection behavior of M. hyperodae. However, host feces decreased the number of weevil antennations done by M. hyperodae. Microctonus hyperodae stopped less frequently near immobile L. oregonensis than near walking ones and these latter were frequently pursued by M. hyperodae. Host movement was the stimulus that elicited oviposition by M. hyperodae. The adaptive implications of these results are discussed.     

Host specificity ofCostesia yakutatensis [Hym.: Braconidae] on lepidoptera in peppermint and alfalfa

Seven species of Lepidoptera, common to alfalfa and peppermint in western Oregon, were tested as potential hosts forCotesia yakutatensis (Ashmead). Laboratory and field studies demonstrated that onlyAutographa californica (Speyer) andTrichoplusia ni (Hubner) were suitable. Percent parasitism, duration of oviposition, and number of parasitoids per host were highest whenC. yakutatensis was associated withA. californica.     

Host preference and survival in selected lines of a Drosophila parasitoid,Asobara tabida

Host selection behaviour of parasitoids has important fitness consequences, if hosts of different quality are available. Here the host selection behaviour, the ability to distinguish between hosts differing in their suitability, of the Drosophila parasitizing wasp Asobara tabida was studied. Females from five lines selected for higher survival in the encapsulating host species D. melanogaster ( 16 ) were compared with females from control lines. Females from all five selected lines more readily accepted the encapsulating host species for oviposition when offered together with a nonencapsulating host species than females from the control lines. We found no evidence for pleiotropic effects and suggest that host selection behaviour evolved parallel to the ability to escape encapsulation in the hosts. Our results also suggest that given the appropriate selection pressures, host selection behaviour can quickly evolve in parasitoids, enabling them to adapt fast to changing circumstances.     

Role of Caterpillar Silk Thread in Location of Host Pupae by the Parasitoid <Emphasis Type="Italic">Diadromus pulchellus</Emphasis>

Diadromus pulchellus is a solitary ichneumonid parasitoid. Its only known host is the pupa of Acrolepiopsis assectella, a specialist herbivore of Allium species. D. pulchellus females parasitize A. assectella pupae within 48 h after the caterpillars spin their cocoon and begin to pupate. Having observed that the cocoon produced by the leek moth caterpillar stimulates parasitoid egg-laying and that caterpillar leaves a silk thread, we studied the hypothesis that silk thread might be involved in host-finding by the parasitoid. Behavioral tests showed that when D. pulchellus females encounter a host silk thread, they change directions, follow the thread, and quickly locate the host. These findings show that pupal parasitoids can use signals produced by their hosts at the developmental instar preceding the one that they parasitize.     

Influence of Host Deprivation and Egg Expenditure on the Patch and Host-Finding Behavior of the Parasitoid Wasp <Emphasis Type="Italic">Mastrus ridibundus</Emphasis>

The influence of host deprivation and egg expenditure on the host location behavior ofMastrus ridibundus was evaluated under controlled conditions in a walk-in field cage in a greenhouse. The activity of females peaked at a temperature of 25–27°C generating two peaks of activity through the day. Increased host deprivation did not influence patch and host-finding success: females responded to host stimuli even after 9 days of host deprivation. In contrast, the longer M. ridibundus was exposed to excess hosts, the greater was the reduction in patch and host-finding success. Egg depletion expressed as the percentage of realized lifetime fecundity expended, rather than egg load, offered a good match for this decline. These results are discussed in the context of biological control introductions and the choice of pre-release treatment.     

Host Location of Hyssopus pallidus, a Larval Parasitoid of the Codling Moth, Cydia pomonella

The effectiveness of parasitoids as biological control agents depends largely on their host location behavior. In this study we describe the host-searching behavior of Hyssopus pallidus (Askew), a larval parasitoid of the codling moth Cydia pomonella L. We observed parasitoid behavior on mature apples (a potential host patch) and elucidated some of the stimuli which determine host location. Female wasps showed more complex and intensive searching on infested apples than on mechanically damaged or noninfested apples. Wasps were able to enter infested apples through the calyx or the tunnel made by the host larvae and parasitize them. Area-restricted searching was observed on infested apples, in particular on areas contaminated with host frass. In a further bioassay, we confirmed that host frass contains a host location kairomone. The kairomone appears to be produced by the host independently from its diet, even though frass produced by hosts fed on an apple are more attractive than frass produced by hosts fed on a fruit-devoid artificial diet. The capability of H. pallidus to locate its host inside apples makes the foraging strategy of this species potentially useful as a biological control agent.