Host location by parasites can be achieved by either active or passive mechanisms. In spite of their significance, the efficacy of these methods has been little researched. High fecundity in parasites is discussed in terms of the role it plays in dispersal and transmission. Some concepts developed by mainstream behavioural ecologists are outlined and their relevance to parasitology is indicated. ‘Reproductive value’ is recommended as an appropriate measure of the costs and benefits of behavioural acts. Although costs of reproduction have been rarely studied in parasites, they are likely to occur in cosexual insects, nematodes and crustaceans. Experiments using captive hosts and/or in vitro cultivation could help in the construction of realistic optimality models. We suggest that r- and K-selection theory could assist in the study of the evolution of parasite behaviour. We discuss how parasite populations are dispersed and controlled and consider the implications of overdispersion. WTe outline three sources of signals to which parasites may respond and suggest that understanding evolutionary mechanisms and community organisation of parasites and hosts requires evaluation of fundamental behavioural responses to environmental signals. The study of closely related groups of parasites and their hosts may advance our knowledge of the evolution of parasite life cycles and the evolutionary costs and benefits of behavioural acts.

The robber fly Mallophora ruficauda is one of the most important apicultural pests in the Pampas region of Argentina. This species is a parasitoid of scarab beetle larvae. Females lay eggs away from the host, and the larvae perform active search behaviour toward Cyclocephala signaticollis third instar larvae, parasitoid's preferred host. This behaviour is mediated by host-related chemical cues produced in hosts’ fermentation chamber. Also, C. signaticollis larvae are attracted to fermentation chamber extracts. As scarab larvae have microbe-rich fermentation chamber, it has been suggested that microorganisms could be involved in the production of these semiochemicals. The aims of this work were first to ascertain the presence of microorganisms in the fermentation chamber of C. signaticollis larvae and second to determine the role of microorganisms in the orientation response of parasitoid and host larvae. We found that microorganisms-free C. signaticollis larvae showed deterioration in their development and did not produce the attractive semiochemicals. Therefore, we isolated fermentation chamber microorganisms of host larvae by means of different cultures media, and then, assayed different microorganisms’ stimuli by binary choice tests. We were able to isolate microorganisms and determine that M. ruficauda larvae are attracted to semiochemicals from protein degradation in the fermentation chamber. However, C. signaticollis larvae were not attracted to any semiochemicals associated with microorganisms’ activity in the fermentation chamber. Although we were unable to elucidate the exact role of gut microorganisms in host behaviour, we discuss their relevance in parasitoid host-seeking behaviour and host conspecific interaction in M. ruficauda–C. signaticollis system.

Larvae of the robber fly Mallophora ruficauda are ectoparasitoids of white grubs and adults are an important apiculture pest in Argentina. Females oviposit on tall grasses and the second instar larva actively searches and locates hosts. There are nine potential hosts in the distribution area of this parasitoid and Cyclocephala signaticollis (Coleoptera: Scarabaeidae) is the most parasitized in the field. However, M. ruficauda has a certain degree of behavioural flexibility towards different host species, and not being a strict specialist. The conditions under which the parasitoid orientates and accepts different hosts’ species are unknown. We studied the host specificity of M. ruficauda towards three species of Cyclocephala genus and we determined whether this specificity depends on larval age. We also evaluated whether larva orientation towards Cyclocephala species changes with chemical cue concentration. We assessed host specificity measuring the orientation and acceptance behaviours towards kairomones extracts and live individuals of Cyclocephala species using M. ruficauda larvae of low and high life expectancy (i.e., young and aged second instar larvae). We observed that young larvae orientated only towards C. signaticollis chemical stimulus, whereas aged larvae orientated also towards C. modesta, and the same was observed with increasing stimuli's concentration. Both young and aged M. ruficauda larvae orientate towards live C. signaticollis and C. putrida species and rejected C. modesta. Also, we found that larvae accepted all Cyclocephala hosts. In conclusion, our results indicate that specificity in the laboratory, observed through host orientation and host acceptance behaviours, depends not only on the availability of host species, but also on the nature of the host's stimuli combined with parasitoid age.

Wheat bulb fly (WBF) larvae use chemotaxis to orientate towards host-plant root exudates. This study aimed to investigate the role of the primary plant metabolite carbon dioxide (CO2) in host-plant location by WBF. Arena based behavioural experiments were used to identify whether CO2 induced chemotaxis (directional movement in response to a chemical stimulus) or kinesis (non-directional movement in response to a stimulus) from WBF larvae. No chemotactic response was observed when larvae were presented to a point source of CO2. However, elevated levels of CO2 induced kinesis, with both track length and tortuosity (number of twists and turns in the movement path) increasing at elevated CO2 levels of 1000–2000 ppm, demonstrating increased searching behaviour. Soil emission of CO2 was quantified to compare soil levels with those identified as eliciting behavioural effects on the larvae. Samples removed from soil gave a mean CO2 concentration of 557 (±46) ppm, which is lower than the lowest concentration of CO2 found to induce a behavioural response and higher than the lowest CO2 concentration tested, which was found not to alter behaviour. It is proposed that increased CO2 concentrations in the soil act as a behavioural trigger, inducing intensive searching of an area by WBF larvae. This increases the likelihood of finding more host-specific identifiers, such as secondary metabolites when near a potential host-plant.

The life cycle of the parasitic copepod Lernaeocera branchialis involves 2 hosts, typically a pleuronectiform host upon which development of larvae and mating of adults occurs and a subsequent gadoid host, upon which the adult female feeds and reproduces. Both the copepodid and adult female stages must therefore locate and identify a suitable host to continue the life cycle. Several mechanisms are potentially involved in locating a host and ensuring its suitability for infection. These may include mechano-reception to detect host movement and chemo-reception to recognize host-associated chemical cues, or kairomones. The aim of this study was to identify the role of kairomones in host location by adult L. branchialis, by analysing their behaviour in response to fish-derived chemicals. Experiments demonstrated that water conditioned by immersion of whiting, Merlangius merlangus, elicited host-seeking behaviour in L. branchialis, whereas cod- (Gadus morhua) conditioned water did not. Lernaeocera branchialis are considered a genetically homogeneous population infecting a range of gadoids. However, their differential response to whiting- and cod-derived chemicals in this study suggests that either there are genetically determined subspecies of L. branchialis or there is some form of environmental pre-conditioning that allows the parasite to preferentially recognize the host species from which it originated.

The robber fly Mallophora ruficauda is one of the principal apicultural pests in the Pampas region of Argentina. As adults, the flies prey on honey bees and other insects; while, as larvae, they parasitize scarab beetle larvae. Females of M. ruficauda lay eggs away from the host in tall grasses. After being dispersed by the wind, larvae drop to the ground, where they dig in search of their hosts. It is known that second instar larvae of M. ruficauda exhibit active host searching behaviour towards its preferred host, third instar larva of Cyclocephala signaticollis, using host-related chemical cues. Furthermore, previous works show that these chemical cues are produced in the posterior body half of hosts. However, the precise anatomical origin of these cues and whether they mediate any behaviour of C. signaticollis larvae remains yet unknown. In order to determine the precise origin of the chemical cue, we carried out olfactometer assays with different stimuli of extracts of the posterior C. signaticollis body half. Additionally, we tested whether C. signaticollis is attracted to any of the same extracts as in the previous experiments. We found that both second instar of M. ruficauda and third instar of C. signaticollis are attracted to extracts of the fermentation chamber (proctodeum). This is the first report of attraction of conspecific larvae in scarab beetles. We discuss a possible case of system communication exploitation in an immature parasitoid-host system.

Cephalonomia stephanoderis and Prorops nasuta are two bethylid wasps released into several Latin American countries for classical biological control of coffee berry borer, Hypothenemus hampei, the most serious insect pest of coffee worldwide. Recent studies on the host location behaviour of these parasitoids have shown that females of both species are attracted to volatile compounds released by immature stages and dust and frass of H. hampei. In this study, we investigated the role of the contact chemicals present in dust and frass of H. hampei on the behaviour of P. nasuta and C. stephanoderis females. Parasitoids remained longer on patches treated with methanol extracts than on acetone and hexane extracts. Females spent more time on the patch treated with the methanol extract of dust and frass than on the patches treated with the methanol extract of dry coffee and methanol control. The concentration of the methanol extracts from dust and frass influenced the locomotory activity of parasitoids of both species. The time that females spent in the patch tended to increase as the concentration of the methanol extracts increased. A further experiment aimed to identify other behavioural descriptors and gain a deeper understanding of the mechanisms underlying the response of parasitoids to methanol extracts was performed. Females of both species spent more time, covered more distance, turned more (per unit time and per unit distance), and decreased their speed when they contacted patches treated with methanol extracts in comparison to patches treated with methanol control.

Parasites with low host specificity are predicted to search for hosts in ways less specific than highly host-specific species. They may also be better dispersers. Unionicolan mites that parasitize mussels are positively phototactic, with some species becoming negatively phototactic when host substances are present. This response has been interpreted as an aid to host location and occurs in species where adult mites have a relatively permanent association with hosts. Three species of Unionicola (Unionicolidae: Acari) that differ in host specificity co-occur in freshwater mussels (Unionidae) in St Mark's River in north Florida.

Nycteribiids (Diptera: Nycteribiidae) are specific haematophagous ectoparasites of bats, which spend nearly all their adult lives on hosts. However, females have to leave bats to deposit their larva on the walls of the roosts, where they later emerge as adult flies. Nycteribiids had thus to evolve efficient sensorial mechanisms to locate hosts from a distance. We studied the sensory cues involved in this process, experimentally testing the role of specific host odours, and general cues such as carbon dioxide, body heat, and vibrations. As models we used two nycteribiids (Penicillidia conspicua and Penicillidia dufourii) and their primary bat hosts (Miniopterus schreibersii and Myotis myotis, respectively). Carbon dioxide was the most effective cue activating and orientating the responses of nycteribiids, followed by body heat and body odours. They also responded to vibration, but did not orientate to its source. In addition, sensory cues combined (carbon dioxide and body heat) were more effective in orientating nycteribiids than either cue delivered alone. Results suggest that nycteribids have some capacity to distinguish specific hosts from a distance, probably through their specific body odours. However, the strong reliance of nycteribiids on cues combined indicates that they follow these to orientate to nearby multispecies bat clusters, where the chances of finding their primary hosts are high. The combination of sensory cues seems therefore an effective strategy used by nycteribiids to locate bat hosts at a distance.

Fruit flies have evolved mechanisms using olfactory and visual signals to find and recognize suitable host plants. The objective of the present study was to determine how habitat patterns may assist fruit flies in locating host plants and fruit. The tomato fruit fly, Neoceratitis cyanescens (Bezzi), was chosen as an example of a specialized fruit fly, attacking plants of the Solanaceae family. A series of experiments was conducted in an outdoor field cage wherein flies were released and captured on sticky orange and yellow spheres displayed in pairs within or above potted host or non-host plants. Bright orange spheres mimicking host fruit were significantly more attractive than yellow spheres only when placed within the canopy of host plants and not when either within non-host plants or above both types of plants. Additional experiments combining sets of host and non-host plants in the same cage, or spraying leaf extract of host plant (bug weed) on non-host plants showed that volatile cues emitted by the foliage of host plants may influence the visual response of flies in attracting mature females engaged in a searching behaviour for a laying site and in assisting them to find the host fruit. Moreover, the response was specific to mature females with a high oviposition drive because starved mature females, immature females and males showed no significant preference for orange spheres. Olfactory signals emitted by the host foliage could be an indicator of an appropriate habitat, leading flies to engage in searching for a visual image.

Parasitoids employ different types of host-related volatile signals for foraging and host-location. Host-related volatile signals can be plant-based, originate from the herbivore host or produced from an interaction between herbivores and their plant host. In order to investigate potential sex- and species-related differences in the antennal response of parasitoids to different host-related volatiles, we compared the electroantennogram (EAG) responses of both sexes of the specialist parasitoid, Microplitis croceipes (Cresson), and the generalist, Cotesia marginiventris (Cresson), to varying doses of selected plant-based host-related volatiles: two green leaf volatiles (cis-3-hexenol and hexanal) and three inducible compounds (cis-3-hexenyl acetate, linalool, and (E,E)-α-farnesene). Mating had no significant effect on EAG response. Females of both species showed significantly greater EAG responses than conspecific males to green leaf volatiles, which are released immediately after initiation of herbivore feeding damage. In contrast, males showed greater responses than conspecific females to inducible compounds released much later after initial damage. Cotesia marginiventris females and males showed greater EAG responses than counterpart M. croceipes to the tested compounds at various doses, suggesting that the generalist parasitoid shows greater antennal sensitivity than the specialist to the tested host-plant volatiles. These results are discussed in relation to the possible roles of green leaf volatiles and inducible compounds in the ecology of female and male parasitoids.

The morphology, distribution, and hosts of two egg parasitoids, Ufens principalis Owen sp. n. and U. ceratus Owen sp. n. (Hymenoptera: Trichogrammatidae), are described for the first time. These species are compared to U. niger (Ashmead), the only species of Ufens s. str. currently described from the Nearctic, and diagnostic differences are presented. The behavioural biology of U. principalis, and U. ceratus to a smaller extent, is also described for the first time. Ufens principalis exhibited rapid and long-distance response in the form of directional flight toward freshly laid eggs of Homalodisca species, its primary hosts in southern California. Parasitism involved aggregations of female U. principalis on fresh Homalodisca egg masses, which remained attractive to U. principalis for a relatively short time. The level of oviposition by U. principalis females was low during most of the day and peaked before sunset in tandem with a peak in Homalodisca oviposition. Oviposition behaviour of U. principalis is described and the distribution of ovipositor probe durations showed that most probes were generally of very short duration. Mating of both Ufens species occurred on the egg mass, with males showing aggressive behaviour towards each other as they competed for emerging females. Ufens ceratus males displayed greater aggression towards other males than U. principalis males. By contrast, fights among U. principalis males involved more individuals and lasted longer than corresponding fights between U. ceratus males.

Parasitoid fitness depends largely on the capability to locate a host in an ecosystem. A parasitoid of a polyphagous host might not be able to find or to access the host in all its feeding niches. This study evaluated the niche selection of Hyssopus pallidus (Askew), a larval parasitoid of Cydia pomonella (Linnaeus), at the plant level with the goal of assessing its potential for biological control on different fruit crops throughout the plant cycle. Parasitoid behaviour during host location and reproduction rate were investigated on host caterpillars actively feeding on apple, pear, apricot or plum, and on caterpillars diapausing under the bark. Under laboratory conditions, the host searching behaviour of H. pallidus varied depending on the fruit species offered and the infestation of the fruits. Parasitoid females searched longer on apples than on other fruit species, and they searched longer on infested than on uninfested apples. Female wasps were able to locate and parasitize host caterpillars under the tree bark, and their behaviour did not vary with host accessibility. The numbers of caterpillars attacked by H. pallidus depended on the fruit species. The highest numbers of caterpillars were parasitized in apples and apricots. Their accessibility (i.e. position) within the fruit or on the branch did not influence parasitism success. Although hosts were parasitized throughout the season, the best results were achieved with early and late releases. Therefore, the host niche selection behaviour of H. pallidus most likely co-evolved with the host C. pomonella on apples, which renders H. pallidus a valuable biocontrol agent for successful release at different times of the season into apple orchards.

In the first section of this review, the stimuli involved in orienting biting flies to their hosts from a distance are discussed under the following headings: (1) Appetitive search—driven by endogenous rhythms and hunger. (2) Activation—the switchover from search behaviour to active host-location brought about in the insect by host-originating stimuli including, depending on species, carbon dioxide, visual contact, and specific host odours. (3) Attraction—the process of active host-location through which the insect comes into the immediate vicinity of the host. This relies on host-originating stimuli including visual contact and olfactory stimuli such as carbon dioxide, volatiles such as acetone, and specific, usually very potent host-related odours.

The second part of this review discusses the properties of the odour plumes biting flies rely on to locate the host. Possible methods of host-location using air-borne odour plumes are presented and the “positive anemotaxis” model based on moth pheromone work is stressed.

In a recent review of host location by tsetse an attempt was made, using previously published data, to draw some general conclusions regarding the range of detection of warm-blooded hosts by tsetse and mosquitoes. The paper included some misleading and invalid conclusions. This topic and the supporting data are reconsidered. It is suggested that the maximum range of a positive response by mosquitoes or tsetse to stationary cattle baits may be expressed as 0.31 × host weight (kg)0.90 (P < 0.001).

Examples from recent literature are used to discuss the host location behaviour of tsetse in relation to feeding patterns. Tsetse recognise potential hosts by their visual and olfactory characteristics. These, and mechanical stimulation, will activate tsetse and initiate host-oriented responses. Approach to a stationary host is by upwind flight modulated by olfactory stimuli with visual responses only at short range, while the approach to a moving host is largely mediated by visual input. Non-random feeding patterns, even where host and tsetse occupy the same habitat, may be explained by host responses to tsetse attack. There is little evidence of long range discrimination between hosts. Some epidemiological implications of the resulting feeding patterns are discussed, and the host location behaviour of tsetse is compared with that of other insects including mosquitoes.