An attelabid weevil (Euops splendida) cultivates fungi

An attelabid weevil,Euops splendida, the female of which makes leaf-roll cradles, is symbiotic with a fungus. She possesses a set of fungus-cultivating structures: spore reservoir, spore incubator, and spore bed, and also comb plate which is used for sowing spores in the nibbled sites on the cradle leaf. Some parts of this paper were presented in a symposium at the 45th Annual Meeting of the Entomological Society of Japan (Sakurai 1985).     

To be suspended or to be cut off? Differences in the performance of two types of leaf-rolls constructed by the attelabid beetle Cycnotrachelus roelofsi

The attelabid beetle Cycnotrachelus roelofsi constructs two types of leaf-rolls (cradles): one is suspended from leaves (suspended type), and the other is severed from the leaves (cut-off type). To evaluate differences in performance between these two cradle types, we monitored densities, survival rates, and mortality factors throughout the active season, from April to June. The proportions of the two cradle types changed over time; the suspended type was dominant early in the season but was gradually replaced by the cut-off type. Irrespective of differences in mortality factors between the two cradle types, the survival rate was always higher in the cut-off type than in the suspended type. Beetle mortality tended to be highest at the egg stage, and the predominant cause of mortality was parasitism by two minute trichogrammatid wasp species, Poropoea morimotoi and P. sp.1. Parasitism by P. morimotoi was significantly higher in cut-off cradles than in suspended cradles, when cut-off cradles were abundant, whereas parasitism by P. sp.1 tended to be higher in suspended cradles when suspended cradles were abundant. These results suggest opposed frequency-dependent attacks by P. morimotoi and P. sp.1. We discuss how these egg parasitoids maintain coexistence of these two cradle types.     

Host suitability and preference studies of Trichogramma cordubensis (Hymenoptera: Trichogrammatidae)

Studies of host suitability and preferences of Trichogramma cordubensis Vargas and Cabello (Hymenoptera: Trichogrammatidae) were performed with eggs of six Lepidoptera (Noctuidae) species: Thysanoplusia orichalcea Fabricius, Peridroma saucia (Hübner), Xestia c-nigrum L., Phlogophora meticulosa (L.), Noctua pronuba (L.), and N. atlantica (Warren). Host suitability was studied by analysing separately the effects of the attacked host species and the influence of the rearing host species on different biological parameters of T. cordubensis. Host preference was analysed by offering eggs of two host species simultaneously to a single female wasp without previous oviposition experience (dual-choice tests). Results show that P. saucia, followed by P. meticulosa were the least suitable hosts for T. cordubensis, since on these species the preimaginal development of the parasitoids was significantly longer and, the number of parasitized eggs as well the number of offspring per female were significantly lower. Contrarily, T. cordubensis parasitized at a higher rate the eggs of the endemic non-target species, N. atlantica. Dual choice tests showed that the option of the first host to be accepted by the wasp was random; however, the mean number of parasitized eggs differed significantly when two host species were offered simultaneously to T. cordubensis, always being the host species with heavier eggs the most parasitized.     

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.     

Competition between the filth fly parasitoids Muscidifurax raptor and M. raptorellus (Hymenoptera: Pteromalidae)

Competition bioassays were conducted with the filth fly pupal parasitoids Muscidurax raptor (Girault & Sanders) and M. raptorellus (Kogan & Legner) (Hymenoptera: Pteromalidae) using house fly Musca domestica L. (Diptera: Muscidae) hosts at different host densities. Muscidifurax raptor had a significant impact on M. raptorellus when hosts were limiting in sequential parasitism tests. Fewer than six M. raptorellus adult progeny emerged from groups of 50 fly pupae that were parasitized by M. raptor at the same time or when M. raptor parasitism preceded M. raptorellus by 48 h, respectively, compared with 42–55 M. raptorellus progeny produced when this species was tested alone. Production of M. raptor was significantly lower when parasitism by this species was preceded by M. raptorellus (25) than when M. raptor was tested alone (43). When the two species parasitized hosts at the same time in different proportions at low host:parasitoid densities (5:1), M. raptorellus produced 13 progeny per parent female when it was the sole species present and fewer than two when M. raptor was present. No negative impact of M. raptorellus on M. raptor was observed. Neither species had a substantial effect on the success of the other at higher host:parasitoid densities.     

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.     

Multiple host use by the egg parasitoid Trissolcus basalis (Wollaston) in a soybean agricultural system: biological control and environmental implications

1 Species of bugs (Hemiptera: Pentatomidae and other families) that are parasitized by the egg parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) in the soybean agricultural system on the Darling Downs in south-eastern Queensland, Australia, are reported. The degree to which eggs of each bug species are used by T. basalis is quantified, which allows assessment of the role of these alternative host species in the biological control of the green vegetable bug Nezara viridula (L.). 2 Egg masses of nine species of pentatomid bugs and one unidentified bug species were collected. Parasitism rates of egg masses of all species averaged 50–70% but were significantly lower for the more important pest species, including the green vegetable bug, than for some of the agriculturally less important species. 3 Trissolcus basalis emerged from nearly all species collected and was the major parasitoid to emerge from most species. A number of native species were parasitized heavily by T. basalis and parasitism of such species may enhance biological control of the green vegetable bug, but may also pose environmental concerns. Parasitism of predatory bugs by T. basalis may have a negative impact on the biological control of other pests, especially lepidopterous pests. 4 Although the impact of T. basalis on native and predatory host species was not directly quantified, these host species remain abundant and therefore do not appear to be affected adversely by such high rates of parasitism.     

Prevalence of parasitism and adult survival time of Aedes albifasciatus (Diptera: Culicidae) parasitized by Strelkovimermis spiculatus (Nematoda: Mermithidae)

We described the carryover of Strelkovimermis spiculatus (Poinar and Camino) (Nematoda: Mermithidae) from mosquito larvae, the primary site of maturation, to adults. We analyzed the survival time of male and female Aedes albifasciatus (Macquart) (Diptera: Culicidae) parasitized by S. spiculatus, the time of emergence of nematodes from adult mosquitoes, and the state of parasitism in the same mosquito cohorts during the immature stages. Mosquito larvae with single and multiple parasitism (up to 11 parasites) were observed. The mortality of mosquito larvae and adults was produced in all cases where at least one mermithid emerged. The mortality of S. spiculatus showed an increasing trend in mosquito larvae with larger numbers of nematodes and was higher in larvae parasitized by eight or more nematodes. Maximum survival of parasitized adult females of Ae. albifasciatus was 38 days, while non‐parasitized adult males and females survived 39 and 41 days, respectively. Strelkovimermis spiculatus mortality was observed in Ae. albifasciatus larvae with single or multiple parasitisms. The spread of mermithid parasitism in adult mosquito populations is discussed.     

The effect of mermithid parasitism on predation of nymphal Baetis bicaudatus (Ephemeroptera) by invertebrates

We investigated how infection by the mermithid nematode Gasteromermis sp. affected predation on its nymphal mayfly host, Baetisbicaudatus, by two invertebrate predators – the stonefly nymphs of Kogotusmodestus and the caddisfly larvae of Rhyacophilahyalinata. Predation trials and behavioral observations were conducted in stream-side, flow-through experimental chambers. When parasitized and unparasitized prey were offered in equal numbers, K. modestus consumed significantly more parasitized than unparasitized nymphs. R. hyalinata consumed equal numbers of both prey types. Behavioral observations of foraging K.␣modestus on parasitized and unparasitized prey suggested that the increased consumption of parasitized nymphs was due to differences in the behavior of infected mayflies in response to the predator. Specifically, parasitized nymphs drifted less often to escape an approaching predator (non-contact encounters) compared to unparasitized nymphs, which increased the number of contact encounters and attacks that occurred between K.␣modestus and parasitized prey. Because all hosts are castrated, these behavioral alterations affect only the fitness of the parasite, which is killed along with its host by invertebrate predation. We present a number of hypotheses to explain why the parasite causes increased predation on its host. These include the large size of the parasite affecting the sensory abilities of the host, the larger energetic costs of escape behavior for parasitized individuals, and natural selection from fish predation against drifting behavior by parasitized individuals. Received: 27 May 1996 / Accepted: 30 September 1996     

The oviposition behavior ofAphidius ervi andMonoctonus paulensis (Hymenoptera: Aphididae) encountering different host species (Homoptera: Aphididae) in sequential patches

Two-day-old mated females ofAphidius ervi Haliday andMonoctonus paulensis (Ashmead) were each provided with two sequential host patches. Patches were comprised of plastic petri dishes containing either 15 pea aphids,Acyrthosiphum pisum (Harris), or 15 alfalfa aphids,Macrosiphum creelii Davis. Both wasp species parasitized more hosts in patches containing pea aphids than in those containing alfalfa aphids, regardless of sequence. Females ofA. ervi also laid more eggs per aphid in patches containing pea aphids than in patches containing alfalfa aphids. When both patches contained alfalfa aphids,M. paulensis females parsitized more aphids in the second patch than in the first. Fewer alfalfa aphids were parasitized in the second patch when the first patch contained pea aphids, and fewer eggs were laid per alfalfa aphid. Parasitoid females of both species exhibited consistently higher rates of oviposition into their preferred host species and adjusted their reproductive allocation to hosts and host patches as a function of their experience in previous patches.     

Effect of Maruca vitrata (Lepidoptera: Crambidae) host plants on life‐history parameters of the parasitoid Apanteles taragamae (Hymenoptera: Braconidae)

Abstract The effect of four host plant species of the herbivore Maruca vitrata Fabricius (Lepidoptera: Crambidae) on development time, longevity, fecundity and sex ratio of the parasitoid Apanteles taragamae Viereck (Hymenoptera: Braconidae) was investigated under laboratory conditions. The larvae were parasitized when in the second instar. Maruca vitrata larvae were fed with flowers of four legumes, that is, Vigna unguiculata (cowpea), Sesbania rostrata, Lonchocarpus sericeus and Pterocarpus santalinoides, or an artificial diet both before and after parasitization. The parasitoid did not develop in hosts feeding on L. sericeus or V. unguiculata at 25°C, or on P. santalinoides at 25°C or 29°C. Apanteles taragamae had the shortest development time on artificial diet at both 25°C and 29°C while the longest development time was recorded on L. sericeus at 29°C. Female wasps took longer to develop compared to males at the two temperatures, regardless of the feeding substrate of their host. The longevity of the wasps at 25°C varied among feeding substrates, but not at 29°C. Survival rate of parasitized larvae depends on the feeding substrate. Moreover, infection of host larvae with Maruca vitrata multi‐nucleopolyhedrovirus (MaviMNPV) killed larger proportions of parasitized larvae at 25°C than at 29°C, which was likely caused by the difference in parasitoid developmental rate. The proportion of female parasitoids was lowest on L. sericeus. The daily fecundity showed a nonlinear trend regardless of the feeding substrate, indicating that A. taragamae is a pro‐ovigenic species. The data support the slow growth–high mortality hypothesis.     

Functional responses of aphid parasitoids,Aphidius colemani (Hymenoptera: Braconidae) and Aphelinus asychis (Hymenoptera: Aphelinidae)

We evaluated the functional responses of two aphid parasitoids: Aphidius colemani on the green peach aphid Myzus persicae (Hemiptera: Aphididae), and Aphelinus asychis on M. persicae and the potato aphid, Macrosiphum euphorbiae (Hemiptera: Aphididae). Parasitoid oviposition occurred at host densities of 5, 10, 20, 30, 50, 80 or 100 aphids for A. colemani and 5, 10, 20, 30 or 50 aphids for A. asychis. More M. persicae were parasitized by A. colemani than by A. asychis at an aphid density of 50. Among the three types of functional response, type III best described the parasitoid response to the host densities both in A. colemani and A. asychis. The estimated handling time was shorter for A. colemani than for A. asychis (0.017 and 0.043 d, respectively). The proportion of aphids that were parasitized exhibited the same characteristic curve among the three host-parasitoid combinations: a wave form that appeared to be a composite of a decelerating (as in type II) response at low host density and an accelerating-and-decelerating (as in type III) response at medium to high host density. We hypothesize that the novel host species (and its host plant), density-dependent superparasitism, and/or density-dependent host-killing may have induced the modified type III response.     

Fitness of Encarsia sophia (Hymenoptera: Aphelinidae) parasitizing Trialeurodes vaporariorum and Bemisia tabaci (Hemiptera: Aleyrodidae)

Abstract Fitness and efficacy of Encarsia sophia (Girault & Dodd) (Hymenoptera: Aphelinidae) as a biological control agent was compared on two species of whitefly (Hemiptera: Aleyrodidae) hosts, the relatively smaller sweetpotato whitefly, Bemisia tabaci (Gennadius) biotype ‘B’, and the larger greenhouse whitefly, Trialeurodes vaporariorum (Westwood). Significant differences were observed on green bean (Phaseolus vulgaris L.) in the laboratory at 27 ± 2°C, 55%± 5% RH, and a photoperiod of 14: 10 h (L: D). Adult parasitoids emerging from T. vaporariorum were larger than those emerging from B. tabaci, and almost all biological parameters of E. sophia parasitizing the larger host species were superior except for the developmental times of the parasitoids that were similar when parasitizing the two host species. Furthermore, parasitoids emerging from T. vaporariorum parasitized more of these hosts than did parasitoids emerging from B. tabaci. We conclude that E. sophia reared from larger hosts had better fitness than from smaller hosts. Those from either host also preferred the larger host for oviposition but were just as effective on smaller hosts. Therefore, larger hosts tended to produce better parasitoids than smaller hosts.     

Reproductive strategies under multiparasitism in natural populations of the parasitoid wasp Nasonia (Hymenoptera)

Parasitoid Nasonia wasps adjust their progeny sex ratio to the presence of conspecifics to optimize their fitness. Another trait under female control is the induction of offspring diapause. We analysed progeny sex ratios and the proportion of diapausing offspring of individual Nasonia females in host patches parasitized by two species, Nasonia vitripennis and Nasonia giraulti, in North American field populations using microsatellite fingerprinting. Both Nasonia species produced similar sex ratios on hosts that were co‐parasitized by their own species as by the other species, indicating that females do not distinguish between con‐ and heterospecific clutches. The sex ratios of the diapause and adult fractions of mixed broods from single females were not correlated. We found further indications that N. vitripennis females take the emergence time of the offspring into account in their sex allocation. The reproductive strategies of Nasonia under multiparasitism are largely adaptive, but also partially constrained by information.     

Ecological and morphological characteristics of endoparasitoids on Elcysma westwoodii (Vollenhoven) (Lepidoptera: Zygaenidae)

Six species of insect endoparasitoids were identified from Elcysma westwoodii, which is the most damaging lepidopteran pest of Prunus yedoensis. From Hymenoptera, two species were identified: a species in Braconidae and Charops striatus in Ichneumonidae. From Diptera, there were four species in Tachinidae: Compsilura concinnata, Exorista sp., Pales sp. and Tachinidae spp. The parasitic ratio was 4.86% (45 of 926 larvae). The hymenopterans were parasitic on 31 individuals of E. westwoodii (68.9%) and the dipterans were parasitic on 14 individuals (31.1%). It was found that parasitoids from the larvae of E. westwoodii were all either endoparasitoids or larval parasitoids. However, Exorista sp. of Tachinidae was found to be either a larval parasitoid or larval-pupal parasitoid. Additionally, all the identified parasitoids were solitary parasitoids, as only one parasite occurred in a larva of E. westwoodii. Because the larva of E. westwoodii eats and molts after it is parasitized, all the parasitoids were identified as koinobionts. There were no big differences in morphological characteristics and life histories between C. striatus and C. concinnata. However, for Exorista sp. and Pales sp., males took 3–5 days longer to emerge from their pupae and had remarkably longer body lengths than females.     

Field establishment of Aphidius colemani Vier. (Hym., Braconidae, Aphidiinae) in the Czech Republic

Abstract: Aphidius colemani Vier. (a Chilean strain) was introduced and became established in the field between 1992 and 2000 in the Czech Republic. The target aphid species, Aphis spiraephaga Muell. on Spiraea ornamentals, and later the Russian wheat aphid, Diuraphis noxia (Kurdj.) on small grains were found parasitized, but an increasing number of other host aphid species in various urban ecosystems, crops and native communities was also detected. The establishment of A. colemani is classified as positive in supplementing the native parasitoid guilds as well as in contributing to the overall biodiversity of hymenopterous parasitoids in the cultivated landscape.     

Biology of Cotesia ( = Apanteles) telengai (Hymenoptera: Braconidae) on its natural host Agrotis segetum (Lepidoptera: Noctuidae)

The host suitability of Agrotis segetum Denis & Schiff., A. ipsilon Hufn., Spodoptera littoralis Boisd, S. exigua Hub., Mythimna loreyi Duponchel and Mamestra oleracea L. for the gregarious braconid Cotesia ( = Apanteles) telengai Tobias was determined under laboratory conditions. The parasitoid only completed its development in larvae of A. segetum. The percentage of successfully parasitized larvae and the mean duration of C. telengai egg‐larval period were inversely related to the age of the host or host instar at parasitization. The mean number of parasitoids which emerged per parasitized larva was positively correlated with the larval age. The sex ratio was consistently high (ca. eight males to one female), independent of the host instar parasitized, as compared to 1:1 as observed frequently in field populations of this wasp. The females of C. telengai were active and produced offspring at temperatures of 15, 20, 25 and 30° C. However, the mean percentage of parasitized larvae increased from 13.1 to 72% and the mean progeny per parasitoid female increased from 14.7 to 129.4 parasitoids, both significant, when the experimental temperature was raised from 15 to 30°C, while their mean development time decreased from 75.5 to 19.2 days. At 25°C, the virgin and mated females continued oviposition until days 16 and 17, with a lifetime total of progeny of 397.6 (SD ±224.7) and 611.1 (SD± 128.8) parasitoids respectively, reaching a maximum of 64.3 and 99.2 on day 2 respectively.     

Interspecific competition between the braconid endoparasitoids Glyptapanteles porthetriae and Glyptapanteles liparidis in Lymantria dispar larvae

The effect of interspecific competition between the solitary endoparasitoid Glyptapanteles porthetriae Muesebeck (Hymenoptera: Braconidae) and the gregarious Glyptapanteles liparidis Bouché (Hymenoptera: Braconidae), was investigated in larvae of Lymantria dispar L. (Lepidoptera: Lymantriidae). Host larvae were parasitized by both wasp species simultaneously in premolt to the 2^nd or the 3^rd host instar or in an additional approach with a 4‐day delay in parasitization by the second wasp species. Host acceptance experiments revealed that both wasp species do not discriminate between unparasitized host larvae and larvae parasitized previously by the same or the other species. In more than 90% female wasps parasitized the larva they encountered first. During the period of endoparasitic development, larvae of the competing parasitoid species never attacked the egg stage of the other species. When host larvae were parasitized simultaneously by both wasp species, the rate of successful development of both species depended on the age of the host larva at the time of its parasitization; G. liparidis emerged successfully from 44% of host larvae parasitized during the premolt to 2^nd instar, G. porthetriae from 28%, and in 20% of the hosts both parasitoid species were able to develop in one gypsy moth larva. However, when host larvae were parasitized simultaneously during premolt to the 3^rd instar, G. liparidis was successful in 90% of the hosts, compared to 8% from which only G. porthetriae emerged. In the experiments with delayed oviposition, generally the species that oviposited first succeeded in completing its larval development. Larvae of the species ovipositing with four days delay were frequently attacked and killed by larvae of the first parasitizing species or suffered reduced growth. As the secondary parasitoid species, G. porthetriae‐larvae were never able to complete their development, whereas G. liparidis developed successfully in at least 12,5% of the multiparasitized host larvae. Thus, multiparasitism of gypsy moth larvae by both Glyptapanteles species corresponds to the contest type; however, G. porthetriae is only able to develop successfully as the primary parasitoid of young host larvae.     

Trichogramma platneri (Hym.: Trichogrammatidae): Host choices between viable and nonviable codling moth,Cydia pomonella,and three-lined leafroller,Pandemis limitata (Lep.: Tortricidae) eggs

In laboratory host-preference studies,Trichogramma platneri Nagarkatti (Hym.: Trichogrammatidae) females, reared on viable codling moth,Cydia pomonella (L.) (Lep.: Tortricidae) eggs, parasitized significantly more viable than nonviable codling moth eggs produced by cobalt 60 irradiated adults. This ovipositional preference was maintained when competition increased per host egg. Similarly, one, three and fiveT. platneri females consistently oviposited more eggs into viable versus nonviable host eggs. A single femaleT. platneri parasitized a similar number of viableC. pomonella and three-lined leafroller,Pandemis limitata (Robinson) (Lep.: Tortricidae) eggs. However, as the number of females per host eggs increased, a significantly larger number of codling moth versus leafroller eggs were parasitized. One, three and fiveTrichogramma females parasitized significantly moreP. limitata versus nonviableC. pomonella eggs. NonviableC. pomonella eggs deflated to less than 25% of their original volume in four and ten days when held under dry and humid conditions respectively. The blackened prepupal stage of the parasitoid generally occurred in the nonviableC. pomonella eggs four days post-parasitization whereas the parasitized viable eggs blackened zero to one day earlier. Incubation under dry or humid conditions did not appear to influence the number of parasitized eggs that were able to blacken and show evidence of parasitism. Summerland Research Centre Contribution No. 938.     

Asolcus bennisi n. sp. (Hymenoptera,Proctotrupoidea) parasite oophage deGraphosoma lineata L. (Het. Pentatomidae)

Summary The Author describes from Morocco a new species ofAsolcus which is closely related to the sibling speciesA. basalis Wollaston andA. nigribasalis Voegelé. The species is namedA. bennisi and parasitizes in nature the eggs ofGraphosoma lineata L. The separation ofA. bennisi from the species of thebasalis group is based on biological tests and on a characteristic pigmentation of the parasitized host eggs reared under optimal laboratory conditions.