Interspecific Host Discrimination Research Articles

Host Patch Use and Potential Competitive Interactions Between Two Egg Parasitoids From the Family Scelionidae, Candidate Biological Control Agents of Bagrada hilaris (Hemiptera: Pentatomidae).

Two egg parasitoids from Pakistan, Gryon sp. nr. gonikopalense Sharma (Hymenoptera: Scelionidae) and Trissolcus hyalinipennis Rajmohana & Narendran (Hymenoptera: Scelionidae), are currently being investigated as potential classical biocontrol agents for Bagrada hilaris Burmeister. The former is the most promising because of its ability to attack B. hilaris eggs in soil, but the latter was recently discovered in California. This study detailed the patch use and oviposition behavior of both species and assessed their relative foraging efficiency. We also investigated possible competitive interactions by assessing 1) the occurrence of intra- and interspecific host discrimination, 2) mutual interference between females (extrinsic competition), 3) the outcome of multiparasitism (intrinsic competition), and 4) the effect of competition on host suppression. Our results showed that T. hyalinipennis females tended to probe repeatedly in the same hosts leading to longer host patch time and lower foraging efficiency. Both species avoided conspecific superparasitism, but not multiparasitism. When the two species foraged simultaneously, G. sp. nr. gonikopalense seemed to be slightly superior in extrinsic competition, while neither species seemed to have an advantage in intrinsic competition. Also, neither species could develop as a facultative hyperparasitoid, but they inflicted non-reproductive mortality on eggs containing 4-d-old larvae of their competitor. Lastly, host mortality inflicted by G. sp. nr. gonikopalense and T. hyalinipennis when acting together appeared to be additive. These results suggest that the presence of T. hyalinipennis in California is unlikely to reduce G. sp. nr. gonikopalense efficiency, but will instead improve the biological control of B. hilaris.

Assessing the competitive interactions between two egg parasitoids of the Eucalyptus snout beetle, Gonipterus platensis, and their implications for biological control

The Eucalyptus snout beetle, Gonipterus platensis (Marelli), is an important pest of Eucalyptus globulus Labill. This insect is partially controlled by the egg parasitoid Anaphes nitens (Girault) in many regions, but the introduction of additional natural enemies can potentially increase pest control. In this study, we evaluate intra- and interspecific competitive interactions between the incumbent A. nitens and the new egg parasitoid Anaphes inexpectatus Huber and Prinsloo. The effects of temperature, order of parasitism, number of parasitoid ovipositions, time interval between ovipositions, and host egg age were analysed. Distinct outcomes of competition were found at different temperatures, with benefit to A. inexpectatus at 20 °C. The first species to parasitise generally prevailed over the second, indicating exploitation competition. However, interference competition was also apparent, namely when A. inexpectatus laid multiple eggs, outcompeting A. nitens, and when the first parasitism occurred six days before. In this case, the second species was able to eliminate the first. Anaphes nitens tended to reject eggs parasitised by A. inexpectatus, whereas A. inexpectatus showed no interspecific host discrimination behaviour towards eggs parasitised by A. nitens. Overall, A. nitens parasitised more hosts and is expected to contribute more to pest control, but it was found to be more susceptible to intraspecific competition. Results suggest that A. inexpectatus and A. nitens should be able to coexist, as asymmetric competition was found to depend on temperature. However, A. inexpectatus establishment in the field in areas where A. nitens is already present may be delayed or even prevented due to interspecific competition. As such, the introduction of A. inexpectatus in a classical biological control programme against G. platensis is advised to be carried out by releasing large numbers of parasitoids in consecutive occasions.

The hyperparasitoid Marietta picta (Hymenoptera: Aphelinidae) mediates competitive interactions between two parasitoids of Paratrioza sinica (Hemiptera: Psyllidae): Tamarixia lyciumi (Hymenoptera: Eulophidae) and Psyllaephagus arenarius (Hymenoptera: Encyrtidae)

Both the ectoparasitoid Tamarixia lyciumi Yang (Hymenoptera: Eulophidae) and the endoparasitoid Psyllaephagus arenarius (Hymenoptera: Encyrtidae) are solitary nymphal parasitoids of Paratrioza sinica (Hemiptera: Psyllidae), a pest of goji berry, Lycium barbarum. Moreover, T. lyciumi is frequently attacked by a hyperparasitoid Marietta picta Andre (Hymenoptera: Aphelinidae) in the field. To analyze competitive interactions between T. lyciumi and P. arenarius mediated by M. picta, we evaluated interspecific host discrimination and larval competition between T. lyciumi and P. arenarius, and the effect of M. picta on their competitive interactions, under laboratory conditions. The results showed that T. lyciumi could discriminate hosts pre-parasitized by P. arenarius, but not vice versa. However, P. arenarius was a superior intrinsic competitor in multiparasitized hosts regardless the order of parasitism. Although multiparasitized hosts produced more P. arenarius and fewer T. lyciumi after exposure to M. picta, the total number of emerging T. lyciumi and P. arenarius did not decline significantly, and sex ratios did not change. Thus, larval superiority would appear to favor the establishment of newly introduced P. arenarius in the Ningxia region, while the inferior competitor T. lyciumi is able to utilize interspecific host discrimination to avoid larval competition. Competitive interactions between T. lyciumi and P. arenarius are mediated by M. picta, and the observed host–parasitoid–hyperparasitoid interactions potentially facilitate their collective coexistence in biological control of P. sinica.

Host discrimination and ovicide by aphid hyperparasitoids Asaphes suspensus (Hymenoptera: Pteromalidae) and Dendrocerus carpenteri (Hymenoptera: Megaspilidae)

Asaphes suspensus (Nees) and Dendrocerus carpenteri (Curtis) are common aphid hyperparasitoids that attack aphidiine primary parasitoids in mummified aphids. In this study, we investigated the self- and interspecific host discrimination and interspecific ovicidal behavior of A. suspensus and D. carpenteri. In choice tests, females of both species laid significantly more eggs on unparasitized hosts than on hosts parasitized by themselves. Interspecific host discrimination was observed in D. carpenteri, which avoided oviposition on hosts previously parasitized by A. suspensus. Asaphes suspensus, however, did not avoid oviposition on hosts parasitized by D. carpenteri. Dissection of mummies after experimental trials of interspecific discrimination revealed that damaged D. carpenteri eggs with a concave puncture site were found only when A. suspensus laid eggs on hosts parasitized by D. carpenteri. Additional observations indicated that most D. carpenteri eggs on parasitized hosts exposed to A. suspensus oviposition behavior were killed (~85 %). The apparent lack of interspecific host discrimination by A. suspensus suggests that the ovicidal ability of this species diminishes its fitness loss by multiparasitism.

How could host discrimination abilities influence the structure of a parasitoid community?

Three related Aphidius parasitoid species share the same host, the grain aphid Sitobion avenae. Among this parasitoid community, Aphidius rhopalosiphi is the most abundant species in the field. Both the interspecific host discrimination of A. rhopalosiphi towards hosts parasitized by the two other species (i.e. A. avenae and A. ervi) and the interspecific host discrimination of the two other species towards hosts parasitized by A. rhopalosiphi were studied here. Results showed that females of A. rhopalosiphi and A. avenae both discriminated between unparasitized hosts and hosts parasitized by the other species. This discrimination occurred only after ovipositor insertion, suggesting the perception of an internal marker of parasitism. Likewise, females of A. rhopalosiphi and A. ervi were able to discriminate between unparasitized hosts and hosts parasitized by the other species. However, in this combination of species, recognition of parasitized hosts occurred before ovipositor insertion, through an antennal perception, suggesting the presence an external cue indicating parasitism. Hence, interspecific host discrimination in the three Aphidius species is based on internal or external cues, which are used either alone or together. Our results showed that the cues used for interspecific host discrimination depend on the specific identity of the interaction. These differences seemed strongly linked to the way the different species respond to defensive behaviours of their aphid hosts. Results are discussed in the context of optimal foraging and possible consequences for community structure.

Efficiency, intrinsic competition and interspecific host discrimination of Copidosoma desantisi and Trichogramma evanescens, two parasitoids of Phthorimaea operculella

Trichogramma evanescens West. (Hymenoptera: Trichogrammatidae) and Copidosoma desantisi Annecke & Mynhardt (Hymenoptera: Encyrtidae) are potential parasitoids of the potato tuber moth (PTM), Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae) in Egypt. Discrimination of a parasitized host from an unparasitized host would prevent wasting of time, eggs and reduce competition with conspecifics or heterospecifics. Therefore, we evaluated interspecific host discrimination, multiparasitism and intrinsic competition between the two wasp species. In a choice test, females of T. evanescens showed high interspecific host discrimination only when they were offered 2-day-old C. desantisi parasitized and unparasitized PTM eggs. In contrast, C. desantisi showed high host discrimination and preferred unparasitized eggs to PTM eggs harboring 2-h- or 2-day-old T. evanescens’ eggs. We also evaluated the effect of different introduction sequences on the efficacy of the two wasps. Dissection data indicated that the two parasitoids had a negative impact on each other. There was a significant reduction in the total number of deposited eggs as well as total number of parasitized hosts by each parasitoid. Regarding the rearing experiment, the total number of T. evanescens-induced black eggs or C. desantisi formed mummies in combined treatments was significantly lower than in single parasitoid treatments (control). Moreover, C. desantisi was inferior and did not develop from any multiparasitized host regardless of oviposition order. It was suggested that combined release of the two wasps would not elevate rate of parasitism over that of single parasitoid treatments and competition between them would reduce their efficacy.

Host discrimination and interspecific competition of Trissolcus nigripedius and Telenomus gifuensis (Hymenoptera: Scelionidae), sympatric parasitoids of Dolycoris baccarum (Heteroptera: Pentatomidae)

Trissolcus nigripedius Nakagawa and Telenomus gifuensis Ashmead (Hymenoptera: Scelionidae) are solitary egg parasitoids of Dolycoris baccarum L. (Heteroptera: Pentatomidae), a polyphagous seed-sucking bug of agricultural crops. Field examinations revealed that the two parasitoids are the most common species, but only a single species emerge from a host egg mass. To explain this observation we tested two hypotheses of interspecific host discrimination and asymmetry in competitive interaction between Tr. nigripedius and Te. gifuensis. Trissolcus nigripedius and Te. gifuensis could discriminate host egg pre-parasitized by either self or conspecific like other scelionid parasitoids that use external mark on host eggs. When provided with host eggs pre-parasitized by each other, both Tr. nigripedius and Te. gifuensis multiparasitized 94% and 100% of the host eggs without interspecific host discrimination, respectively. Interestingly, from the multiparasitized eggs irrespective of oviposition sequence, progeny of Te. gifuensis always survive better than Tr. nigripedius. Telenomus gifuensis is superior in immature competition probably due to shorter egg incubation period after oviposition, hence progeny of Te. gifuensis become first instar faster than that of Tr. nigripedius. However, adult Tr. nigripedius is always superior competitor in possessing and guarding the host eggs even after oviposition against Te. gifuensis. Therefore, the asymmetry in competitive interaction between Tr. nigripedius and Te. gifuensis may explain the emergence of a single species from a host egg mass in the field in spite of no interspecific host discrimination.

Odour‐mediated long‐range avoidance of interspecific competition by a solitary endoparasitoid: a time‐saving foraging strategy

1. In studies on optimal foraging strategies, long-range decisions in the pursuit of resource are rarely considered. This is also the case for sympatric parasitoids, which may be confronted with the decision to accept or reject host larvae that are already parasitized by a competing species. They can be expected to reject already parasitized hosts if it is likely that they will lose the resulting intrinsic competition. However, examples of such interspecific host discrimination are rare. 2. We propose that parasitoids that are not egg-limited should reject inferior hosts only if it saves them time, and that this will be achieved mainly when the parasitoids are able to detect competitors from a distance. We tested this hypothesis using the sympatric parasitoids Cotesia marginiventris (Cresson) and Campoletis sonorensis (Cameron). 3. C. sonorensis was found to be the superior intrinsic competitor but, upon contact with a host larva, both wasps readily accepted hosts that had already been parasitized by the other species. However, in an olfactometer experiment, C. marginiventris females were found to strongly avoid the odour of their superior competitor. 4. These results are in accordance with a time optimization scenario, whereby the inferior competitor accepts competition if it costs only an egg, but avoids competition if it may save time that can be allocated to the search for more profitable hosts. 5. Models on host discrimination strategies in parasitoids had not yet considered discrimination from a distance. Long-range foraging decisions can also be expected for other organisms that have to choose between resources of varying suitability and profitability.

Kairomone stimulates increased probes and host stings in a cleptoparasitoid

Abstract. Eupelmus vuilleti acts as a cleptoparasitoid when encountering a host parasitized by Dinarmus basalis. The encounter of a parasitized host stimulates an increasing number of ovipositor probes directly above the parasitized host, an increasing number of host stings and the destruction of approximately 40% of the D. basalis eggs. The stimulation of ovipositor probes appears to be due to the detection of a stimulus that is different from the stimulus allowing interspecific host discrimination. It appears that a proteinacious substance produced by the D. basalis venom gland and deposited on the edge of the drilled hole induces probing behaviour of E. vuilleti.

Interspecific competition between Xanthopimpla stemmator Thunberg and Dentichasmias busseolae Heinrich (Hymenoptera: Ichneumonidae), pupal parasitoids attacking Chilo partellus (Lepidoptera: Crambidae) in East Africa

Abstract Interspecific competition between Xanthopimpla stemmator and Dentichasmias busseolae was studied using pupae of the invasive crambid stemborer Chilo partellus as the host. While X. stemmator is an old association, D. busseolae formed a relatively new association with C. partellus in East Africa. Two different time intervals between parasitism (0 and 48 h) and two parasitoid sequences [i.e., X. stemmator before D. busseolae (Xs–Db) and D. busseolae before X. stemmator (Db–Xs)] were chosen. In addition, the parasitoids’ performance on pupae in maize stems and ears was assessed. For both X . stemmator and D . busseolae , there was no difference in foraging time between unparasitized pupae and pupae previously parasitized by the other species, indicating that the two species were not capable of interspecific host discrimination. In the Xs–Db sequence, the time interval between parasitism did not have an influence on the percentage of pupae producing either parasitoid species. By contrast, in the Db–Xs sequence, the percentage of pupae producing X. stemmator was almost 8 times higher in the 0-h than the 48-h interval, while for D. busseolae it was the reverse. In the 0-h interval, X. stemmator outcompeted D. busseolae irrespective of whether it parasitized first or second, while in the 48-h interval, the parasitoid parasitizing first won. While D. busseolae successfully searched for and parasitized pupae in both stems and ears, parasitism of pupae in ears by X. stemmator was negligible. It was concluded that the two species could co-exist because they partly exploit different ecological niches.

Intra- and interspecific host discrimination in arrhenotokous and thelytokous Eretmocerus spp.

Bemisia tabaci (Gennadius) is a serious pest of vegetable, ornamental, and agronomic crops throughout the world. To control B. tabaci, Eretmocerus eremicus Rose & Zolnerowich, and Eretmocerus mundus Mercet are considered the most effective parasitoids in dry tropical regions. In parasitoids, choosing the `right` hosts has direct consequences for their reproductive success and efficiency as biocontrol agent. Therefore, being able to discriminate a parasitized host from an unparasitized one would be important to prevent wasting time, eggs, and to reduce the mortality risk for their offspring. We evaluated intra- and interspecific host discrimination and the chance of super-parasitism or multi-parasitism in two populations of E. mundus (sexual and asexual) and E. eremicus. Different combinations and sequences of female introduction were carried out for the various populations and species. Experienced females avoided super-parasitism. However, naive females did lay eggs under hosts that were previously parasitized by conspecific females. E. eremicus females avoided to multi-parasitize hosts parasitized by E. mundus. However, E. mundus females did multi-parasitize the hosts that had been parasitized earlier by E. eremicus. In the case of super-parasitism, the outcome showed that neither of the E. mundus populations was stronger, whereas in the case of multi-parasitism E. mundus appeared stronger than E. eremicus. Since those populations and species are morphologically similar a molecular method had to be developed to identify the outcome of super- or multi-parasitism, which is presented in Appendix A.

Comparison of foraging behavior, interspecific host discrimination, and competition of Encarsia formosa and Amitus fuscipennis

The foraging behavior of Amitus fuscipennis MacGown & Nebeker and Encarsia formosa Gahan was studied on tomato leaflets with 20 Trialeurodes vaporariorum (Westwood) larvae in the first or third stage. Ten of the whitefly larvae were previously parasitized and contained a conspecific or a heterospecific parasitoid egg or larva. The host type (host stage and/or previous parasitization) did not influence the foraging behavior of either parasitoid species. The residence time on these tomato leaflets was about 0.9 h for A. fuscipennis and 1.9 h for E. formosa. Amitus fuscipennis hardly stood still and fed little, while E. formosa showed extensive standing still and feeding. As a result, the time walking while drumming was similar for both parasitoid species. The numbers of host encounters and ovipositions per leaflet were similar for both parasitoid species. However, the residence time of A. fuscipennis was half as long as that of E. formosa so the rate of encounters and ovipositions was higher for A. fuscipennis. Amitus fuscipennis is more efficient in finding and parasitizing hosts under these conditions. The walking activity and host acceptance of the synovigenic E. formosa diminished with the number of ovipositions, but not those of the proovigenic A. fuscipennis. Encarsia formosa is egg limited, while A. fuscipennis is time limited because of its short life span and high egg load. Both parasitoid species discriminated well between unparasitized larvae and self-parasitized larvae, but discriminated poorly those larvae parasitized by a conspecific and did not discriminate larvae parasitized by a heterospecific. Self-superparasitism, conspecific superparasitism, and multiparasitism were observed for both parasitoid species. Superparasitism always resulted in the emergence of one parasitoid and multiparasitism resulted in a higher emergence of one parasitoid of the species that had parasitized first. The data suggest that A. fuscipennis is a good candidate for use in biological control of high-density spots of T. vaporariorum when we consider its high encounter and oviposition rate.

Interspecific host discrimination and within-host competition between Encarsia formosa and E. pergandiella (Hymenoptera: Aphelinidae), two endoparasitoids of whiteflies (Hemiptera: Aleyrodidae)

Interspecific host discrimination and within-host competition between Encarsia formosa Gahan and Encarsia pergandiella (Howard), two endoparasitoids of whiteflies, were studied under laboratory conditions. Interspecific host discrimination was studied at two time intervals (0 h and 72 h after the first species had oviposited). Parasitized and unparasitized Trialeurodes vaporariorum (Westwood) hosts were accepted for oviposition at the same rate by the two parasitoid species. Host type did not affect the handling time of the two parasitoids. The outcome of within-host competition was investigated after females of the two species parasitized the hosts at various time intervals. In four treatments, E. pergandiella was allowed to oviposit 0, 24, 48 and 72 h after E. formosa while in the other two, E. formosa was allowed to oviposit 0 and 72 h after E. pergandiella. In four of these treatments: E. formosa following E. pergandiella at 0 and 72 h, and E. pergandiella following E. formosa at 0 and 24 h, E. pergandiella prevailed. In the host discrimination experiment (72 h interval), 20% of E. pergandiella eggs were killed by E. formosa females. Interspecific ovicide was also observed in the within-host competition experiment, in which 6% of 72-h-old E. pergandiella eggs were killed by E. formosa females.

Self-, Intra-, and Interspecific Host Discrimination in Telenomus busseolae Gahan and T. isis Polaszek (Hymenoptera: Scelionidae), Sympatric Egg Parasitoids of the African Cereal Stem Borer Sesamia calamistis Hampson (Lepidoptera: Noctuidae)

Oviposition behavior, intra- and interspecific host discrimination, and super- and multiparasitism by the scelionids Telenomus busseolae and T. isis were studied using batches of eggs of the noctuid stalk borer Sesamia calamistis as the host. Both Telenomus species were able to discern eggs already parasitized. As a result, self-superparasitism was only 4.0% for T. busseolae and 5.8% for T. isis. Likewise, intraspecific superparasitism was avoided by both species and was significantly higher for T. busseolae than T. isis, and higher when parasitized eggs were offered immediately (0 h) and after 48 h than after 24 h; apparently, the recognition of parasitized eggs at 24 and 48 h was based on the presence of parasitoid larvae rather than a specific marking substance. Multiparasitism was avoided if the female had a choice between unparasitized and parasitized eggs. In a choice experiment, it was 10.2 and 2.5% for T. busseolae and T. isis, respectively. In a nonchoice experiment, multiparasitism did not vary between species and time treatment; it was low, varying between 9.6 and 24.1%. In the 0-h treatment, T. busseolae always outcompeted T. isis, accounting for 63.4 or 91.7% of the offspring, when T. isis or T. busseolae, respectively, was the first species. In the 24-h treatment, the first species emerged more often than the second did but the host egg mortality, i.e., eggs from which neither borer larvae nor parasitoid emerged, was >40%. Whereas T. busseolae is ubiquitous in Africa, T. isis has never been reported from eastern Africa, and it has been proposed for introduction against the prevailing noctuid pest Bussoela fusca. Based on earlier life table and host suitability studies and on the present findings, it is concluded that T. isis would establish in the midaltitudes but not the highlands of eastern Africa.

Intra- and interspecific host discrimination by host-seeking larvae of coleopteran parasitoids

Intraspecific host discrimination is widespread in solitary parasitoids whose adult females forage for and evaluate host suitability, whereas interspecific discrimination is less common. In some parasitoid species, mostly Diptera and Coleoptera, the larva performs the last step of host searching. It has been suggested that host discrimination will rarely occur in such host-seeking larvae because their low mobility results in a low host encounter rate. We determined the extent to which the larvae of Aleochara bilineata Gyllenhal (Coleoptera: Staphylinidae), a solitary parasitoid of aggregated Diptera pupae: (1) discriminated between unparasitized hosts and hosts parasitized by conspecifics; (2) used semiochemical cues to discriminate; (3) were influenced by life expectancy, presence of conspecifics and host availability in their host acceptance decision; and the extent to which (4) A. bilineata and A. bipustulata L., a species exploiting the same hosts and occurring sympatrically, showed interspecific host discrimination. A. bilineata larvae were able to discriminate between unparasitized hosts and hosts parasitized by conspecifics in a choice experiment. Such behavior has never previously been described for a coleopteran parasitoid or for a parasitoid species whose larvae perform host searching. Host discrimination in this species was not based on the presence of visual or tactile cues (e.g., entrance holes) but rather on chemical cues. The life expectancy of A. bilineata larvae was significantly shorter in the presence than in absence of hosts, and older larvae had lower parasitism success than young larvae in a 24-h experiment. However, the host acceptance decision of A. bilineata larvae was not influenced by larval age or the presence of conspecifics when the ratio of hosts per larva was greater than or equal to 1. When hosts were scarce, the degree of superparasitism increased significantly with the number of foraging conspecifics and the age of the larvae. Both species of Aleochara showed intra- and interspecific host discrimination in a choice experiment. In contrast to A. bipustulata, A. bilineata larvae more frequently parasitized hosts parasitized by A. bipustulata than those parasitized by conspecifics. We suggest that host discrimination will be frequent in solitary parasitoids with host-seeking larvae when hosts are aggregated.

Competition between Gyranusoidea tebygi and Anagyrus mangicola, parasitoids of the mango mealybug, Rastrococcus invadens: interspecific host discrimination and larval competition

AbstractThe competition between Gyranusoidea tebygi Noyes and Anagyrus mangicola Noyes (both Hymenoptera: Encyrtidae), exotic parasitoids of the mango mealybug, Rastrococcus invadens Williams (Homoptera: Pseudococcidae) was studied in the laboratory. No significant differences were found in the way each parasitoid species examined, attacked, stung, and oviposited into hosts, unparasitized, or previously parasitized by the other species. This suggests that neither species discriminates against each other. The total number of parasitoids of either species emerging did not significantly differ between competition experiments. When A. mangicola was the first parasitoid to attack a host, it had no significant advantage over G. tebygi. However, when A. mangicola followed G. tebygi by either 4 or 24 h, it clearly won. Overall A. mangicola won the competition in 70.9% of all cases. The level of the competition, either at the egg or larval stage, and factors responsible for the elimination of older larvae by younger ones could not be assessed in these experiments. The coexistence of the two parasitoids as complementary for the biological control of the mango mealybug is discussed.

Ovipositional Behavior and Host Discrimination in Three Scelionid Egg Parasitoids of Stink Bugs

Abstract Ovipositional behavior of the scelionid egg parasitoids Trissolcus basalis (Wollaston), Psix tunetanus (Mineo & Szabo), and two color forms of Trissolcus utahensis (Ashmead) was observed in laboratory experiments that employed a common host, the pentatomid Euschistus conspersus Uhler. Brood sex ratios were female biased and less than 0.20; P. tunetanus deposited a precise sequence of male and female offspring compared to the other species. In host discrimination experiments in which individual females were presented with an egg mass partially parasitized by a conspecific female, T. utahensis showed considerable ovipositional restraint toward parasitized eggs. When presented with an egg mass partially parasitized by a female of another species—either T. basalis or P. tunetanus — T. utahensis displayed little discrimination and oviposited in most of the parasitized eggs that it contacted. When the experiment was reversed— T. basalis and P. tunetanus females presented individually with an egg mass partially parasitized by T. utahensis —both species showed little discrimination and oviposited in most of the parasitized eggs that they contacted. For eggs in which multiparasitism involved the combination of T. utahensis and P. tunetanus, the latter emerged from over 70%, regardless of whether it was the first or second ovipositing female. When super- or multiparasitism involved the combination of the two T. utahensis color forms or the combination of T. utahensis and T. basalis, offspring of the first ovipositing female emerged from over 70% of the eggs. These results are consistent with the view that interspecific host discrimination in parasitoids is less common than conspecific host discrimination and provide some support for the hypothesis that brood sex ratio and competitive ability are inversely related in guilds of quasi-gregarious egg parasitoids. Implications for the selection of parasitoid species for classical and augmentative biological control are discussed.

Interspecific host discrimination and competition inApoanagyrus (Epidinocarsis) lopeziandA.(E.)diversicornis, parasitoids of the cassava mealybugPhenacoccus manihoti

Abstract.Choice experiments on interspecific host discrimination inA.lopeziandA. diversicorniswere carried out on discs of cassava leaf containing four hosts (P.manihoti) that had been parasitized by the other species and four unparasitized hosts.A.lopeziaccepted both host types equally for oviposition, whereasA.diversicornisaccepted fewer hosts that had been parasitized byA.lopezithan unparasitized ones.A.diversicornisis therefore capable of interspecific host discrimination, but such a capability was not demonstrated forA.lopezi.Survival probability in singly parasitized hosts was 0.85 for both parasitoid species. When the time interval between ovipositions was 2 h or less, survival in multiparasitized hosts was 0.68 forA.lopeziand 0.17 forA.diversicornis, irrespective of priority. IncreasingA.lopezipriority to 24±2h did not increaseA.lopezisurvival.A.diversicornissurvival, however, increased to 0.43 whenA.diversicorniswas given 24 ± 2 h priority.A.diversicorniseggs took 19 h longer thanA.lopezieggs to hatch. This could explain the difference in competitive abilities in multiparasitized hosts.The observed difference in host selection behaviour betweenA.lopeziandA.diversicornisis in accordance with the different benefits of multiparasitism:A.lopezigains more thanA.diversicornisbecause of its superior within‐host competitive abilities.Neither species avoided multiparasitism completely. The low survival probability ofA.diversicornisin multiparasitized hosts may partly explain its failure to establish when introduced into Africa as part of a biological control programme ofP.manihoti.

Intra- and interspecific host discrimination in two closely related egg parasitoids.

Intraspecific host discrimination is frequently found in solitary parasitoids, but interspecific host discrimination, where female parasitoids recognize hosts already parasitized by females of other species, is rare. This particular behaviour appears to be adaptive only under specific circumstances. In this paper, we quantified intraspecific host discrimination in Anaphes n. sp. (Hymenoptera: Mymaridae), an endoparasitoid of the eggs of Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) and interspecific host discrimination toward eggs parasitized by Anaphes sordidatus (Girault), a sympatric species competing for the same resource in similar habitats. To examine host discrimination, choice experiments were used where the females had to choose between different categories of eggs (unparasitized, parasitized by Anaphes n. sp. or A. sordidatus). Superparasitism and multiparasitism were avoided in experiments where the female had a choice between unparasitized hosts and hosts parasitized by the same female, by a conspecific or by a female A. sordidatus. When all hosts available were parasitized, conspecific superparasitism occurred more often than self-superparasitism or multiparasitism. These results indicated that females Anaphes n. sp. were capable of self-, conspecific and interspecific discrimination. Self-discrimination followed recognition of an external marking while interspecific discrimination occurred mostly after insertion of the ovipositor. Interspecific discrimination could result from the recent speciation of these species and could be associated with a genotypic discrimination. This behavior appears to be adaptive because of the competition for common hosts between the two parasitoid species.

Competition for hosts between two hyperparasitoids of aphids,Dendrocerus laticeps andDendrocerus carpenteri (Hymenoptera: Megaspilidae): The benefit of interspecific host discrimination

The two hyperparasitoids of aphids, Dendrocerus carpenteri(Curtis) and Dendrocerus laticeps(Hedicke), differ in their interspecific host discrimination behavior: D. carpenterireadily accepts hosts for oviposition previously parasitized by D. laticeps,whereas D. laticepsoften refrains from ovipositing on hosts previously parasitized by D. carpenteri.The two species also differ in their competitive abilities: D. carpenteriis competitively superior to D. laticepswhen the time between ovipositions is 1 h. D. carpenteri's superiority probably arises from (a) the action of the venom produced by the female to paralyze the host, as evidenced by the comparison of emergence patterns from naturally vs artificially parasitized hosts, and (b) its more rapid egg development. Because of its competitive abilities, D. carpenterigains only limited benefit from discriminating interspecifically, at least when encountering a host parasitized by a competitively inferior species, such as D. laticeps.In contrast, D. laticepsgains considerable benefit from recognizing hosts previously parasitized by D. carpenteri,as the probability to produce offspring from such hosts is greatly reduced.