Gene Flow in the Exotic Colonizing LadybeetleHarmonia axyridisin North America

The introduced parasitoid,Aphytis melinusDeBach (Hymenoptera: Aphelinidae), is used for augmentative biological control of California red scale,Aonidiella aurantii(Maskell) (Homoptera: Diaspididae). Commercially reared wasps are reared on oleander scale,Aspidiotus neriiBouché (Homoptera: Diaspididae). Oleander scale covers lack the chemical,O-caffeoyltyrosine, a kairomone mediating host selection byA. melinus.Wasps reared on oleander scale but individually exposed, or primed, toO-caffeoyltyrosine more readily accepted California red scale for probing in laboratory bioassays and parasitized a greater proportion of available California red scale in the field than wasps reared similarly but not exposed toO-caffeoyltyrosine. Thus, it may be possible to improve host recognition of commercial, insectary-rearedA. melinusby exposing them toO-caffeoyltyrosine prior to release. The goal of this study was to develop an inexpensive but effective means of priming thousands of wasps simultaneously toO-caffeoyltyrosine. The most effective method, but potentially the most expensive, was simply to spray parasitized oleander scale on their host plant with diluteO-caffeoyltyrosine prior to wasp emergence. In additional experiments, using controlled doses of syntheticO-caffeoyltyrosine applied to scale covers, we showed that primed wasps require both a lower minimum dose ofO-caffeoyltyrosine for recognition and also respond to measuredO-caffeoyltyrosine doses more consistently than unprimed wasps. The ability to mass-prime thousands of wasps prior to release is a crucial step toward realizing the concept of behavioral improvement of host selection of parasitoids on a commercial scale.     

Innate and Learned Cues: Scale Cover Selection by Aphytis melinus (Hymenoptera: Aphelinidae)

Aphytis melinus recognizes and accepts covers of its host, California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae), before assessing the size or quality of the scale body beneath. We evaluated the role of a non-volatile kairomone, O-caffeoyltyrosine, and prior experience with hosts on the recognition (antennal drumming) and acceptance (ovipositor probing) of scale covers differing in age and, therefore, size. We tested several hypotheses concerning the role of experience with hosts on host recognition and acceptance. The first predicts that experience with a particular host size leads to increased selection of that host size in the future. The second predicts that the quality of the experience with hosts sets a threshold of quality for future acceptance. We manipulated the quality of wasp experience with hosts by exposing some wasps to high-quality hosts (large scale insects under large covers) and other wasps to low-quality hosts (small scale insects under small covers and small scale insects under large covers). Control (naive) wasps were held without experience with hosts. Wasps were then offered five size classes of covers directly after removal from the scale insects (unmanipulated) or after removing the kairomone from the covers (kairomone-free). For covers with natural levels of kairomone, no effect of experience with hosts on host recognition was observed. When the kairomone was removed, however, any experience with hosts increased recognition compared to naive wasps. Moreover, experienced wasps preferentially recognized covers of the size to which they had experience. Results for host acceptance differed from those for host recognition. Wasps given experience with large hosts accepted more covers with kairomone than wasps in other treatments, but the size preference did not vary among treatments. When the kairomone was removed, however, wasps given experience with large hosts under large covers preferred larger covers. Host recognition and acceptance are controlled by different behavioral mechanisms. Prior experience with hosts does not alter host recognition but does affect the rate of acceptance. The quality of the experience does not affect the acceptance of covers containing natural levels of kairomone but does affect host acceptance when the kairomone is removed. Experience with hosts alters the motivation of wasps to accept covers, and the direction of this effect is determined by the size (quality) of body the wasps were given during the experience.     

Priming Aphytis: behavioral modification of host selection by exposure to a synthetic contact kairomone

The introduced parasitoid, Aphytis melinus DeBach (Hymenoptera: Aphelinidae), utilizes a kairomone, O-caffeoyltyrosine, to recognize California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae). Wasps used in augmentative release programs for California red scale on California citrus are reared on oleander scale, Aspidiotus nerii Bouché (Homoptera: Diaspididae), themselves reared on squash. The goals of this study were: 1) to determine if long-term rearing on oleander scale had caused A. melinus to develop a preference for oleander scale, and 2) to determine if the preference of oleander-reared A. melinus for California red scale might be enhanced by exposing them to synthetic O-caffeoyltyrosine prior to release. Wasps that were removed from their hosts as pupae and allowed to emerge as adults isolated from their hosts retained a strong preference for California red scale regardless of rearing host. This preference was reduced if wasps were allowed to emerge from oleander scale, thus acquiring early adult experience with oleander scale. The preference for California red scale was restored, however, by exposing wasps reared on oleander scale to synthetic O-caffeoyltyrosine prior to bioassay. Exposure of A. melinus reared in commercial insectaries to O-caffeoyltyrosine prior to release may be a means to improve the effectiveness of such wasps in augmentative release programs to control California red scale.     

Environmental variation in physical and chemical cues used by the parasitic wasp,Aphytis melinus, for host recognition

O-caffeoyltyrosine is a host recognition kairomone forAphytis melinus DeBach (Aphelinidae) found in the covers of its host, California red scale,Aonidiella aurantii (Maskell) (Diaspidae). This study tests the hypothesis that the concentration ofO-caffeoyltyrosine and scale cover size are reliable indicators of scale body size, an important component of host quality forA. melinus, over a range of scale rearing conditions. Both scale cover area andO-caffeoyltyrosine concentrations were only qualitatively related to scale body size during the third instar, the scale life stage most suitable forA. melinus. Scale cover area andO-caffeoyltyrosine concentrations were reduced, relative to scale body size, when scale were reared on bark and leaves compared to fruits. Scale cover area andO-caffeoyltyrosine concentration were also relatively reduced when scales were reared in mid-summer compared to spring and fall, and when reared on orange cultivars compared to lemon cultivars in the field. Finally, scale cover area andO-caffeoyltyrosine concentration were reduced when scale were reared at 52% compared to 100% humidity in the laboratory. Scales appear to be chemically conspicuous toA. melinus for a short period of the time in which they are physiologically susceptible. Scales that minimize their cover size and maximize the incorporation rate ofO-caffeoyltyrosine into covers may minimize their conspicuousness toA. melinus. Minimizing scale cover size, but not necessarily incorporation rates, may make scales more vulnerable to predators, however.     

Comparative Development and Performance of Artificially Reared versus Host-RearedDiapetimorpha introita(Cresson) (Hymenoptera: Ichneumonidae) Wasps

Diapetimorpha introita(Cresson) (Hymenoptera: Ichneumonidae), a native ectoparasitoid ofSpodopteraspp. pupae, was reared in the laboratory on an artificial diet devoid of any insect host components. Diet-reared wasps demonstrated a propensity to search for and parasitize natural hosts in a field cage trial. Longevity of the diet-reared wasps was comparable with the longevity of wasps reared on host pupae. Survival rate ofD. introitawas 61.3% when reared on diet and 76.3% when reared on host pupae. Wasps reared on the artificial diet had longer developmental times, reduced fecundity, and reduced adult weights compared to wasps reared on host pupae. These studies suggest that future research efforts should focus on increasing fecundity and weight of diet-reared wasps and decreasing the mortality and developmental time of wasps reared on the artificial diet. The ability to rearD. introitaon an inexpensive, artificial diet significantly enhances the potential of mass rearing this parasitoid for inundative releases againstSpodopteraspp.     

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.     

Host‐size decisions of female parasitoid wasps seeking hidden hosts

1. Choice of host size may play a critical role in parasitoid success, a task that takes on added complications when dealing with concealed hosts, but most such studies of insect behaviour have only taken place in the laboratory. 2. This study investigates the success of a wasp (Alabagrus texanus: Braconidae) in finding host caterpillars Herpetogramma theseusalis (Crambidae) of the most effectively handled size hidden in shelters, in both the field and the laboratory. 3. First, the study tested wasp preference and success in parasitizing large, middle‐sized and small caterpillars (> 5, 3–5, < 3 mm) presented in the open, one at a time, in the laboratory. The wasps attacked (inserted or attempted to insert their ovipositor) a higher proportion of middle‐sized (3–5 mm) caterpillars compared with either small (< 3 mm) or large (> 5 mm) caterpillars. Naïve wasps attacked large caterpillars more often than did experienced wasps. Wasps responded to increasing caterpillar size by increasing the number of legs used to pin their prey rather than by increasing handling time. 4. The frequencies of visits to shelters in the field containing a majority of either large or middle‐sized caterpillars were then compared, followed by a test providing the wasps with similar choices under controlled laboratory conditions. Wasps most frequently visited shelters containing a majority of middle‐sized caterpillars both in the field and under controlled laboratory conditions. 5. The combined results confirmed that the wasps can size‐select their hosts both in the field and in laboratory tests.     

Discrimination of previously searched,host-free sites byMicroplitis croceipes (Hymenoptera: Braconidae)

The ability of free- ranging parasitoids to discriminate between previously visited and unvisited sites containing host kairomone (caterpillar frass) but not hosts was tested. Females of Microplitis croceipes,a host specialist and plant generalist larval parasitoid of Helicoverpa (Heliothis) zea,were allowed to fly freely in a simulated plant patch in a flight chamber. Wasps spent less time searching frass sites previously searched by themselves or by conspecifics than unsearched frass sites. In addition to chemical marking, spatial memory of visual cues was implicated as a mechanism for discriminating against self-visited, host-free sites.     

Odour-mediated foraging by yellowjacket wasps (Hymenoptera: Vespidae): predation on leks of pheromone-calling Mediterranean fruit fly males (Diptera: Tephritidae)

Predation is probably the most important male mortality factor in insect species with courtship displays that render males performing them conspicuous targets of predators. Sexually active Mediterranean fruit fly males, Ceratitis capitata (Wied.), aggregate in leks, where they participate in agonistic encounters and engage in visual, acoustic and pheromone-calling displays to attract receptive females. The objective of this study was to assess: a) whether sexually displaying C. capitata males in leks inside host and non-host foliage are subject to predation by the most prominent predators yellow-jacket wasps, Vespula germanica (F.), and if so, b) whether olfactory, visual or auditive stimuli are used by foraging wasps in locating male C. capitata prey. Studies were carried out in a citrus orchard and surroundings on the island of Chios, Greece. Observations were conducted using perforated containers hung within mulberry, fig or citrus foliage. Living C. capitata flies of different sex and either mature or immature were placed inside. Our results show that the yellowjacket wasps have learned to associate the presence of sexually active medfly males aggregated in leks with their prey's pheromone (kairomone). Foraging wasps, flying through the crowns of host trees, responded to the odour source of C. capitata male pheromone by approaching from downwind. Even inside dense citrus tree foliage, wasps keyed in on aggregations of pheromone-calling males using olfactory stimuli. Stimuli of visual and acoustic male signalling were only used at close range, after having followed the pheromone plume close to its source. Visual cues played a greater role in directing wasp foraging under more open and exposed host foliage conditions. Odour-based foraging of wasps inside host foliage in the mid-morning hours, when medfly male lekking activities peak, shifted gradually to a more visual-based host fruit patrolling in the afternoons to capture ovipositing and feeding medfly females. On ripe fruit, particularly fig, V. germanica visual prey hunting also included the capture of feeding medfly males, other feeding Diptera, as well as medfly larvae extracted from wasp-made perforations in the fruit.     

Host recognition by the specialist endoparasitoidMicroplitis croceipes (Hymenoptera: Braconidae): Role of host- and plant-related volatiles

The specialist parasitoidMicroplitis croceipes Cresson can parasitize only noctuid larvae in the generaHelicoverpa andHeliothis. To be successful in their search for hosts, the ability to distinguish hosts from nonhosts feeding on the same plant is beneficial. In flight tunnel experiments, we found that prior to landing on the odor sourceM. croceipes were able to distinguish volatiles released from frass of host larvae(Helicoverpa zea Boddie) and nonhost larvae (Spodoptera exigua Hübner andSpodoptera frugiperda J. E. Smith) fed on cotton. However, an initial contact experience with frass of cotton-fed host larvae appeared to be critical for this ability. Wasps that had antennated frass of host larvae fed pinto bean diet were equally attracted to frass of host and nonhost larvae fed on pinto bean diet. In short-range walking experiments, wasps located cotton-fed host larvae faster than diet-fed larvae, regardless of their experience. Wasps that had antennated frass of cotton-fed host larvae were less attracted to cotton-fed nonhost larvae, compared to host larvae, and preferred to sting host larvae. Plant-related volatiles in host frass and larvae appear to play a major role in the successful location of host larvae.     

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.     

Recognition of heterospecific parasitism: Competition between Aphidiid (Aphidius ervi) and Aphelinid (Aphelinus asychis) parasitoids of Aphids (Hymenoptera: Aphidiidae; Aphelinidae)

The solitary parasitoids Aphidius erviHaliday (Hymenoptera: Aphidiidae) and Aphelinus asychisWalker (Hymenoptera: Aphelinidae) attacked but generally did not oviposit in pea aphids parasitized by the other species. Wasps selectively oviposited in unparasitized hosts when given a choice. Host discrimination depended on the recognition of internal cues. Females of A. asychiseither could not recognize or ignored A. ervi'sexternal host marking pheromone. Under most conditions, A. ervisurvived in superparasitized hosts, killing competing A. asychislarvae by physical attack and possibly physiological suppression. The outcome of larval competition was not affected by oviposition sequence or age difference between larvae; A. asychissurvived only when it had substantially completed larval development before the host was superparasitized by A. ervi.It is suggested that competition for host resources incurs a cost, for the winner in terms of reduced size or increased development time and for the loser in terms of lost progeny and searching time. Consequently, heterospecific host discrimination can be functional. Internal, and probably general, cues enable wasps to recognize and avoid oviposition in hosts already parasitized by an unrelated species.     

Postemergence Experience Affects Sex Ratio Allocation in a Gregarious Insect Parasitoid

We tested the hypotheses that postemergence experience with plants (“early adult learning”) modifies sex ratio and clutch size allocations of Cotesia congregata (Say) (Hymenoptera: Braconidae), a gregarious larval endoparasitoid of Manduca sexta L. (Lepidoptera: Sphingidae). Emerging wasps were exposed for 2–3 h to (a) one of two host plants (tomato or tobacco) or no plant, and (b) one of two novel plants (arugula or parsley) or no plant. Each female was permitted a single oviposition in a host offered with one of the two plant species 24 h later. Hosts were reared on laboratory diet before and after parasitization. Wasps exposed to either host plant allocated proportionately more females to hosts offered with the plant species experienced at emergence than wasps with the alternate species, but clutch sizes did not differ. Irrespective of plant species, wasps exposed to novel plants allocated proportionately more females to hosts than wasps without plant experience, and larger clutches to hosts offered with parsley than with arugula. Differential responses to host and novel plants suggest inherent recognition of host foodplants by C. congregata. Results demonstrate a direct effect of learning on reproductive potential.     

Influence of aphid size,age and behaviour on host choice by the parasitoid wasp Ephedrus californicus: a test of host-size models

Summary When host quality varies, parasitoid wasps are expected to oviposit selectively in high-quality hosts. We tested the assumption underlying host-size models that, for solitary species of wasps, quality is based on host size. Using Ephedrus californicus, a solitary endoparasitoid of the pea aphid, we evaluated the influence of aphid size (= mass), age and defensive behaviours on host selection. Experienced parasitoid females were given a choice among three classes of 5-day-old apterous nymphs: small aphids that had been starved daily for 4 h (S4) and 6 h (S6) respectively, and large aphids permitted to feed (F) normally. Wasps attacked more, and laid more eggs in, small than large aphids (S6>S4>F). This rank-order for attack did not change when females could choose among aphids of the same size that differed in age; however, wasps oviposited in all attacked aphids with equal probability. Host size did not influence parasitoid attack rates when aphids were anaesthetized so that they could not escape or defend themselves. As predicted by host-size models, wasp size increased with host size (F>S4; S6), but large wasps required longer to complete development than their smaller counterparts (S4E. californicus reflects a trade-off between maximization of fitness gains per egg and the economics of search-time allocation. Because large aphids are more likely to escape parasitization, a wasp must balance her potential gain in fitness by ovipositinng in a high-quality (large) aphid against her potential cost in terms of lost opportunity time if the attack fails.     

Endophytic insect communities of two prairie perennials (Asteraceae: Silphium spp.)

Little is known of the biology of most insects that are endemic to prairie ecosystems of North America, with the exception of large and conspicuous species. In particular, species that are sequestered within plant tissues are commonly overlooked. In this paper, we assess the biodiversity of endophytic insects that inhabit stems of Silphium laciniatum L. and S. terebinthinaceum Jacquin (Asteraceae), endemic plants of tallgrass prairies. Endophytic herbivores, gall wasps Antistrophus rufus Gillette and A. minor Gillette (Hymenoptera: Cynipidae) and stem-boring larvae of the beetle Mordellistena aethiops Smith (Coleoptera: Mordellidae) were attacked by 10 species of natural enemies. We report new host plant associations for herbivores, and new host insect associations for parasitoids. The two plant species differed significantly in their densities of gall wasps and the vertical dispersion of galls within stems. Interactions within and between trophic levels attest to the biodiversity of endophytic insect communities, and the specialized nature of these insects suggests they are highly vulnerable to habitat conservation practices that involve destruction of dead vegetation.     

DNA barcoding of endoparasitoid wasps in the genus Anicetus reveals high levels of host specificity (Hymenoptera: Encyrtidae)

The genus Anicetus includes economically important biocontrol agents that are introduced for control of soft and wax scale insect agricultural pests (Ceroplastes spp.). Understanding of host–parasitoid associations is critical to the successful outcome of their utilization in biological control projects. However, identification of these parasitoids is often difficult because of their small size and generally similar morphological features, and hence, studies on the host–parasitoid associations. Here, nucleotide sequence data were generated from the mitochondrial COI gene and the D2 region of 28S rRNA to assess genetic variation within and between species of Anicetus occurring in China. The results of this study support the use of the COI and the D2 region of 28S rRNA gene as useful markers in separating species of Anicetus, even in cases where morphological differences are subtle. On the other hand, the COI gene is also useful in recognizing species with much variation in morphology. DNA barcoding reveals high levels of host specificity of endoparasitoids wasps in the genus Anicetus. Our results indicate that each Anicetus species is adapted to a limited set of host species, or even are monospecific in their host choice.     

Patch Time Allocation by the Parasitoid Diadegma semiclausum (Hymenoptera: Ichneumonidae). III. Effects of Kairomone Sources and Previous Parasitism

We investigated the effects of kairomone sources and previous parasitism on the patch-leaving behavior of Diadegma semiclausum, a solitary endoparasitoid of larval Plutella xylostella. Individual female wasps were released onto an experimental plant, and were allowed to freely leave for an alternative host plant placed upwind of the experimental plant in a wind tunnel. In one experiment, the experimental plant was either intact, contained host damage alone, or contained both hosts and host damage. In another experiment, the plant was infested with either unparasitized hosts, hosts parasitized previously by the female herself, or parasitized by Cotesia plutellae, another larval endoparasitoid of P. xylostella. We analyzed the influence of kairomone sources, host types, and within-patch foraging experience on the patch-leaving tendency of D. semiclausum by means of the proportional hazards model. Presence of host damage, and unsuccessful host encounters as a result of host defenses decreased the parasitoids' patch-leaving tendency, while successful oviposition, self-superparasitism, and rejection of parasitized hosts increased their patch-leaving tendency. A conceptual model of the parasitoid's patch-leaving behavior is proposed on the basis of the results of current and previous studies.     

Use of contact chemical cues in prey discrimination by Cerceris fumipennis

Cerceris fumipennis Say (Hymenoptera: Crabronidae) uses a wide range of adult buprestid beetles to provision nests. Wasps seldom make ‘mistakes’ and attack beetles in other families. Bioassays showed that the wasps use contact chemical cues in the beetles’ epicuticle to discriminate buprestids from other beetles. Wasps rejected buprestids that had their cuticular hydrocarbons (CHC) removed by being washed in solvents. The washed, rejected buprestids became acceptable to 70% of wasps when they were coated with an extract of buprestids’ own CHC. Washed buprestids coated with extracts of non‐buprestid beetle CHC were not accepted. Analyses of buprestid CHC profiles showed that they are relatively simple, consisting of five classes. Other coleopteran families that may co‐occur in the arboreal hunting habitat of C. fumipennis have a broader range of CHC classes. Experiments adding buprestid hydrocarbons to a non‐buprestid beetle, unwashed Popillia japonica Newman (Coleoptera: Scarabaeidae), suggested that wasps may be deterred by the presence of CHC classes not found in Buprestidae. Adding a synthesized dimethyl‐branched hydrocarbon, a class of CHC found in Chrysomelidae but not Buprestidae, to the cuticle of unwashed buprestid beetles, caused wasps to reject the buprestid beetles. We propose that CHC act as a kairomone for C. fumipennis, triggering attack, whereas classes of CHC not found in Buprestidae disrupt this response, and thus help to determine the prey range of the wasp.     

Host preference and offspring performance are linked in three congeneric hyperparasitoid species

1. The optimisation theory predicts that insect mothers should oviposit on resources on which they attain the highest exclusive fitness. The development of parasitoid wasps is dependent on limited host resources that are often not much larger than the adult parasitoid. 2. In the present study preference and development in three congeneric species of secondary hyperparasitoids attacking cocoons of two congeneric primary parasitoids that differ significantly in size were compared. Gelis agilis (Fabricius) and G. acarorum (L.) are wingless hyperparasitoids that forage in grassy habitats, whereas G. areator (Panzer) is fully winged and forages higher in the canopy of forbs. 3. The three species were reared on cocoons containing pupae of a small gregarious endoparasitoid, Cotesia glomerata (L.), and a larger solitary species, C. rubecula (Marshall), both of which develop in the caterpillars of pierid butterflies. 4. Adult mass was correlated with initial cocoon mass in all three species, whereas development time was unaffected. Wasps were larger when developing in C. rubecula. However, for a given host mass, wasps were larger when developing on the smaller host, C. glomerata. This suggests that there is a physiological limit to hyperparasitoid size that was exceeded when C. rubecula served as host. 5. All three hyperparasitoids strongly preferred to attack cocoons of the larger species, C. rubecula, often avoiding cocoons of C. glomerata entirely. 6. Preference and performance are correlated in the three Gelis species. However, owing to variation in the distribution and thus abundance of their hosts, it is argued that cumulative fitness may be still higher in the smaller host species.     

Volatile cues used by the parasitoid, Aphytis melinus, for host location: California red scale revisited

Aphytis melinus DeBach (Hymenoptera: Aphelinidae) is a biological control agent of diaspidid scale insects. The parasitoid has a narrow host range but its hosts are polyphagous. We determined the source of volatile cues the wasp uses to locate its few host species when those hosts occur on more than one host plant species. We addressed four questions in regard to the use of volatile cues in host location of California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae): (1) Does A. melinus use volatile cues to assist in host location? (2) Are these cues innately recognized or learned? (3) Are cues produced by female California red scale, or from other sources? (4) Are the cues specific to the host or host plant? These questions were tested through the use of a Y-tube olfactometer. Female A. melinus used volatile cues to orient toward both infested and uninfested host plant material. Wasps learned these cues by associating odors from the host plant with host presence. They had no innate preferences for scale insect or host plant volatile stimuli. Contrary to previous studies, we found no evidence of orientation toward the female-produced sex pheromone of California red scale, nor to volatile cues from the attacked host stage. Wasps given experience with scale insects growing on lemon fruit subsequently oriented toward lemon and orange fruit and leaves. The scale species with which the wasp was given experience did not affect this preference. Wasps given experience with California red scale growing on squash did not orient toward infested lemon fruit. The host ranges of the parasitoid and its hosts are used to explain the adaptive value for the evolution of learned rather than fixed responses to cues used in foraging behavior.