Hymenopterous parasitoids of bark beetles [Scolytidae] in Israel: Relationships between host and parasitoid size,and sex ratio

Information on parasitoids of bark beetles infesting conifers and broadleaves is presented to show the relations between host and parasitoid size, and parasitoid sex ratios. Sticky traps were employed to determine the sex ratios of parasitoid species operating on beetle-infested material. A wide range in body length was found for most species. Females ofCerocephala eccoptogasteri Masi,Cheiropachus quadrum Febr.,Eurytoma morio Boheman,Heydenia pretiosa Forster,Metacolus unifasciatus Forster,Rhaphitelus maculatus Walker andRoptrocerus xylophagorum (Ratz.) were usually significantly longer than their males, whereas the males of all fourDendrosoter spp. were significantly longer than the females. Body lengths of the males and females were almost identical inEntedon ergias Walker andEcphylus caudatus Rusch. The sex ratio varied markedly within most species; the mean ratio for 5 of them differed significantly from 1∶1. The sex ratio ofM. unifasciatus andE. morio was significantly related to host size. The sex-ratio of parasitoids trapped on sticky traps varied with time in relation to the phenology of the host beetle. Females were more numerous during the larvae development while males were more abundant between the pupal stage and emergence of the host. Male courtship behaviour ofDendrosoter caenopachoids Rusch.,D. protuberans (Nees),M. unifasciatus andR. xylophagorum is described. The effect of host size, the diversity in sex ratios, and possible interaction between parasitoid size and sex-ratio are discussed. Contribution from the Agricultural Research Organization, Bet Dagan, Israel. N° 1328-E, 1984 series.     

Interactions betweenAulonium ruficorne [Coleoptera: Colydiidae] and other natural enemies of bark beetles [Coleoptera: Scolytidae]

The within tree distribution of some common natural enemies of bark beetles in pine plantations in Israel and some aspects of their feeding habits were studied with special emphasis on the potential impact on the predatorAulonium ruficorne Olivier. A total of 12 predators and 2 parasite species were found associated withA. ruficorne in the natural enemy complex of bark beetles on pines. No secondary parasites were detected. The anthocoridScoloposcelis pulchella (Zetterstedt) and the dipteranMedetera striata Parent were observed feeding on immature stages ofA. ruficorne in the absence of scolytids. The associated Coleoptea:Nemosema elongatum F.,Rhizophagus bipustulatus L.,Corticeus spp.,Plastysoma spp. andPlegaderus discisus Erickson are thought to compete withA. ruficorne on larvae and pupae of bark beetles when the latter are in limited quantities (especially in the lower section of the tree). The parasites, mainlyMetacolus unifasciatuss Forster andDendrosoter caenopachoides Ruschka are assumed to compete withA. ruficorne during the larval period in the smooth bark section of the stem. Competition might occur mainly during spring and fall. Deutonymphs of the miteIpiduropoda sellnicki were detected on the abdomen ofA. ruficorne adults. Larvae of the predator were rarely infected in the field by the bacteriaSerratia sp. while laboratory cultures suffered high rate of mortality caused by this pathogen.      

Arthropods and nematodes co-occurring with the eastern larch beetle,Dendroctonus simplex [Col.: Scolytidae], in Newfoundland

Fifty-four species of insects, 5 of spiders, 5 of mites, 1 harvestman and 1 nematode co-occurred with the eastern larch beetle,Dendroctonus simplex LeConte, on tamarack,Larix laricina (Du Roi) K. Koch, at 2 localities in Newfoundland. The most abundant predators were the flyMedetera gaspensis Bickel and the beetleRhizophagus dimidiatus Mannerheim and the most abundant parasitoids were the HymenopteraSpathius canadensis Ashmead,Rhopalicus tutela (Walker) andRoptrocerus xylophagorum (Ratzburg).D. simplex is recorded as a new host for 9 predator and 3 parasitoid species.      

Effect of parent density on fecundity of two parasitoids [Hym.: Pteromalidae] of the olive beetle,Phloeotribus scarabaeoides [Col.: Scolytidae]

Under laboratory conditions (22±2°C, 60%±5% H.R.; and 16L.: 8D. photoperiod), the rearing of the 2 main parasitoids ofPhloeotribus scarabaeoides (Bernard, 1788) in Southern Spain:Cheiropachus quadrum (Fabricius, 1787) andRaphitelus maculatus Walker, 1834 (Hym.: Pteromalidae) has been carried out. Several densities of population of the parasitoids have been tested in order to determine their reproductive behaviour, and this is significant if the output of a rearing is to be increased. A great competition among the individuals of the same sex has been observed; in the case of the ΦΦ, a potential negative correlation with the increase of density of population, has been noted. The biotic potential belonging to the 2 principal parasitoids species with and without competition, are calculated, and the aspects concerning the parasitoid colonization of the olive tree beetle infestation spots, are discussed.      

Parasites of bark beetles [Col.: Scolytidae] on pine and cypress in Israel

Ten species of hymenopterous parasites were reared from logs of pine and cypress, naturally infested with 5 species of bark beetles,viz., Orthotomicus erosus Woll.,Pityogenes calcaratus Eichh., andTomicus destruens Woll. — on pine;Phloeosinus armatus Reitt. andP. aubei Perris — on cypress, in Israel. The parasites were:Dendrosoter chaenopachoides Rusch,D. hartigii (Ratz.),D. middendorfi (Ratz.),D. protuberans Nees (Braconidae), Eurytoma morio Boheman (Eurytomidae), Cerocephala eccoptogastri Masi,Heydenia pretiosa F?rster,Metacolus unifasciatus F?rster,Rhaphitelus maculatus Walker andRoptrocerus xylophagorum (Ratz.) (Pteromalidae). The most common wereM. unifasciatus on pine and cypress,D. chaenopachoides andR. xylophagorum on pine, andD. protuberans on cypress, occurring in samples collected during most seasons. Less common wasE. morio, which was reared from both conifers.D. middendorfi was found on pine, and only during winter. The other species were found only occasionally.     

Observations on the seasonal abundance ofIcerya purchasi onAcacia baileyana in Adelaide,South Australia

Population estimates of the cottony-cushion scale,Icerya purchasi Maskell were obtained from 2Acacia baileyana F. von Mueller trees in Adelaide, South Australia. Most scales were located within 50 cm of the tip of the twig (sample unit), some adults being found on thick woody branches (5 to 10 cm Ø) and on the tree trunk (>50 cm Ø). Scales were present throughout the year, mostly in very low numbers, and completed 2 annual generations i.e. from April to October and from November to April. This was confirmed by data on the duration of development from a field experiment with 12 sets of monthly cohorts.     

Spruce stands provide natural food sources to adult hymenopteran parasitoids of bark beetles

Adults of synovigenic hymenopteran parasitoids of bark beetles need supplemental feeding to maximise their lifetime realised fecundity, and yet little is known about the natural sources that provide this food. Here we show that, in spite of the reduced diversity of flowering plants expected in even-aged spruce plantations, several plant species are present that might provide nectar and pollen to the hymenopteran parasitoid complex of Ips typographusL. (Coleoptera: Scolytidae), throughout the growing season. Conifer aphids are also abundant in these stands and produce large amounts of honeydew, which might also be consumed by the parasitoids. Laboratory experiments demonstrate that longevity is significantly increased in Coeloides bostrychorum Giraud (Hymenoptera: Braconidae) when the wasps are exposed to flowers of various species commonly found in spruce stands.     

Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

Parasitoids of the gall-forming aphidSmynthurodes betae [Aphidoidea: Fordinae] in Israel

In an ecological investigation of the gall-forming aphid,Smynthurodes betae Westw., we discovered that more than 20% of the fundatrix galls were parasitized byMonoctonia pistaciaecola Stary (Aphidiidae). This is a new host record for this parasitoid. A Pteromalid hyper-parasite was identified asPachyneuron? leucopiscida Mani. Ecological information on the parasitoid is provided, and the low frequency of parasitoids known from gall-forming aphids, compared with freeliving ones, is discussed.     

Food acceptance by a monophagous and an oligophagous insect in relation to seasonal changes in host plant suitability

Food acceptance by larvae of two lepidopteran species feeding on Rosaceae, viz. Yponomeuta evonymellus (monophagous) and Y. padellus (oligophagous), was compared. The influence of seasonal changes in plants as food for both insects was examined, in particular, the effects of nitrogen and sorbitol in leaves. In the laboratory, Y. evonymellus accepts Crataegus monogyna, a host plant of Y. padellus, and Y. padellus accepts Prunus padus, the host plant of Y. evonymellus. P. padus is the most suitable food plant for Y. evonymellus. No difference in food-quality for Y. padellus was found between C. monogyna and P. padus. The performance of both species on P. padus is less influenced by seasonal changes than on Crataegus. The suitability of Crataegus decreases during the season. This is probably caused by the decrease of its nitrogen content, and not by the decrease of sorbitol in the plant. The monophagous, Y. evonymellus, is more sensitive to seasonal changes in its food when fed with a non host plant than the oligophagous Y. padellus. In oviposition experiments both species have a preference for their normal host-plants.     

Association of Geosmithia fungi (Ascomycota: Hypocreales) with pine- and spruce-infesting bark beetles in Poland

Bark beetles (Coleoptera, Scolytinae) are known to be associated with fungi, particularly species of the orders Ophiostomatales and Microascales. However, very little is known about other ectosymbionts of phloeophagous bark beetles on Pinaceae. In this study, we examined the Geosmithia species associated with eight bark beetle species infesting Picea abies and Pinus sylvestris branches in Poland. Fungi were isolated from 1 731 samples collected from 14 study sites. We identified a total of 653 isolates that were sorted into nine taxa based on their phenotypic similarity and phylogeny of their ITS-LSU regions of rDNA, β-tubulin, elongation factor 1α and the second-largest subunit of the RNA polymerase II gene. They represented nine species without formal names. There were large quantitative and qualitative differences in the composition of Geosmithia communities between P. sylvestris and P. abies trees. The proportion of samples infested with Geosmithia species suggests that this association is more widespread among bark beetles infesting branches of P. sylvestris. In addition, these beetles were vectors of different Geosmithia species compared with than the beetles that colonize P. abies. In mixed-conifer forests, the Geosmithia communities were more diverse and richer than in pure spruce or pine stands, where the insects Pityogenes chalcographus and Pityophthorus pityographus with low host-specificity play a distributing role for various Geosmithia species. Among eight bark beetle species examined, only P. bidentatus, P. pityographus, P. chalcographus and Polygraphus poligraphus acted as effective vectors for Geosmithia species. The following hypothesis emerges from these studies: changes in the composition of ectosymbionts of pine- and spruce-infesting bark beetles in Central Europe run along a gradient of thickness of the wood substrata preferred by insects.     

A survey of black scale,Saissetia oleae [Hom.: Coccidae] parasitoids [Hym.: Chalcidoidea] in southern California

A survey of black scaleSaissetia oleae (Olivier) parasitoids present in southern California was conducted between September 1987 and September 1989. From 308 collections of black scale-infested citrus, olive, and oleander twigs from 19 sites in southern California, 1,610 specimens were collected. Nine primary and six secondary parasitoid species were identified. Four primary species were abundant in southern California:Metaphycus bartletti Annecke & Mynhardt,M. helvolus (Compere),Scutellista caerulea (Fonscolombe) (=S. cyanea Motschulsky), andDiversinervus elegans Silvestri. The most common secondary parasitoids wereMarietta mexicana (Howard),Cheiloneurus noxius Compere, andTetrastichus minutus (Howard). In the coastal region of southern California,M. bartletti was the most abundant parasitoid, followed in order byD. elegans, S. Caerulea, andM. helvolus. In the intermediate and interior regions,M. helvolus was most abundant.D. elegans was second most abundant in the intermediate region, but was rare in the interior region.M. bartletti was second in abundance in the interior region and third in the intermediate region.      

Integration of visual and olfactory cues of hosts and non-hosts by three bark beetles (Coleoptera: Scolytidae)

Abstract.  1. There has been a long-standing pre-occupation with how phytophagous insects use olfactory cues to discriminate hosts from non-hosts. Foragers, however, should use whatever cues are accurate and easily assessed, including visual cues.
2. It was hypothesised that three bark beetles, the mountain pine beetle (MPB), Dendroctonus ponderosae Hopkins, the Douglas-fir beetle (DFB), D. pseudotsugae Hopkins, and the western balsam bark beetle (WBBB), Dryocoetes confusus Swaine, integrate visual and olfactory information to avoid non-host angiosperms (e.g. paper birch, trembling aspen), that differ in visual and semiochemical profile from their respective host conifers (lodgepole pine, Douglas-fir, interior fir), and tested this hypothesis in a series of field trapping experiments.
3. All three species avoided attractant-baited, white (non-host simulating) multiple-funnel traps, and preferred attractant-baited black (host-simulating) traps. In experiments combining white, non-host traps with non-host angiosperm volatiles, bark beetles were repelled by these stimuli in an additive or redundant manner, confirming that these species could integrate visual and olfactory information to avoid non-host angiosperms while flying.
4. When antiaggregation pheromones were released from white traps, the DFB and MPB were repelled in an additive-redundant manner, suggesting that beetles can integrate diverse and potentially anomalous stimuli.
5. The MPB demonstrated the most consistent visual preferences, suggesting that it may be more of a 'visual specialist' than the DFB or WBBB, for which visual responses may be more contingent on olfactory inputs.     

Prior encounters modulate subsequent choices in host acceptance behavior by the bark beetle Ips pini

Laboratory bioassays indicate that the bark beetle Ips pini employs flexible, rather than absolute, responses to phytochemicals in its host acceptance behavior. Each beetle's decision to enter substrate was influenced by the types and concentrations of monoterpenes present. However, previous rejection of a simulated host containing a moderate concentration of monoterpenes increased the likelihood that the same concentration would be accepted upon a second or third encounter. This flexibility more than offsets any loss of vigor due to starvation and age that accompanies a process of trial and error. Starvation decreased beetles' total lipid content, but beetles can recover some energetic losses by a small amount of feeding during each trial. In addition to its adaptive value, a flexible host acceptance strategy may yield population level consequences. That is, bark beetles preferentially enter trees having low concentrations of monoterpenes, but may modify their acceptance thresholds when cues associated with stressed trees are not available. This could partially explain how some tree-killing bark beetles colonize a broader physiological range of trees during outbreaks. The adaptive value of relating individual decisions to population density may arise from two ecological relationships: first, as populations rise, the pool of stressed trees is rapidly depleted; secondly, healthy trees are attainable through pheromone – mediated mass attacks when adequate numbers of beetles are present. Flexible host acceptance behaviors may also reduce the advantage of relying exclusively on pre-landing cues to distinguish between susceptible and non-susceptible trees.     

Life history,immature morphology and adult behavior ofTelenomus solitus [Hymenoptera: Scelionidae]

The results of laboratory studies on the life history, morphology of immature stages, effects of food and water on adult longevity, host age preference for oviposition, parthenogenesis, and adult behavior ofTelenomus solitus Johnson are presented.Telenomus solitus completed its development in 10.8 days (egg, 0.3 day; 1^st instar, 2.7 days; 2^nd instar, 3.7 days; and pupa, 4.1 days). The adults lived longest when provided with both honey and water. Without food, they survived less than 24 h. Parasitization of host eggs varied significantly with their age, being highest in 42 h old eggs. Fewer than 10% of eggs older than 60 h were parasitized successfully. The periodicity of oviposition was unimodal; it occurred throughout the 24 h period and peaked in the late photophase. Females ofT. solitus are arrhenotokous. The behavioral sequences during courtship, mating, and oviposition are described.      

Resource utilization and sex allocation in response to host size in two ectoparasitoid wasps on subcortical beetles

Patterns of host resource utilization and sex ratio manipulation in relation to host size were investigated for two solitary ectoparasitoid wasps,Atanycolus initiator andSpathius brevicaudis (Hymenoptera Braconidae). Both species parasitize subcortical beetles on the trunks of Japanese pine trees.A. initiator is on average 8 times larger in body weight and has an ovipositor that is 3.7 times longer than that ofS. brevicaudis. In both parasitoids, the size of emerging wasps was positively correlated with host size, but the host/wasp size regressions were linear for all three major host species inA. initiator, whereas inS. brevicaudis the regression was logarithmic for a relatively large host species. The sex ratios (proportion of males) of both parasitoids emerging from different host species decreased with increasing host size, but the overall sex ratio on each host species was male-biased inA. initiator, while female-biased inS. brevicaudis. How the proportion of host consumed changed in response to host size, differed between the two parasitoids for the same host species. In the field survey, the size and sex ratio of the emerging two parasitoids from a dead tree were closely related to host size. However, the spatial distribution of the two parasitoids depended on the bark thickness of the trunk. The data suggest that differences in the relative evaluation of host size and in ovipositor length may enable the coexistence of the two parasitoid wasps.     

Braconid parasitoids ofTephritidae [Diptera] infesting coffee and other fruits in West-Central Africa

Exploration for parasitoids ofCeratitis capitata (Wiedemann) [Diptera: Tephritidae] was conducted in Cameroon and Togo and parasitoids collected were released in Costa Rica. Collections yielded large numbers ofOpius perproximus Silvestri,Biosteres caudatus Szepligeti, andB. caudatus auct. [Opiinae: Braconidae]. Other Opiines collected wereB. fullawayi (Silvestri),B. desideratus (Bridwell), and an undescribed species ofOpius. Parasitization of tephritids in coffee ranged from 10–56%, with an average 35% parasitism in a research plantation and 17% in a commercial plantation. Parasitism in fruits other than coffee was less than 5%. The most commonly reared tephritids from coffee wereTrirhithrum coffeae Bezzi andC. (Pterandrus) anonae Graham.C. capitata occurred in low frequency in coffee; and adults were rarely observed. Other rearings includedT. albonigrum (Enderlein),T. sp. nr.validum Bezzi,T. nigerrimum auct. nec. Bezzi,C. (Pterandrus) colae Silvestri,C. (Pterandrus) flexuosa (Walker),C. (Ceratalaspis) spp.cosyra (Walker) group,C. (Pardalaspis) sp.punctata (Wiedemann) group,C. (Ceratalaspis) giffardi Bezzi,Dacus (Dacus) bivittatus (Bigot),D. (Didacus) ciliatus Loew, andGymnodacus sp.

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Résumé Une récolte des parasites deCeratitis capitata (Wiedemann) [Diptères: Téphritidés] a été conduite au Cameroun et au Togo, et les specimens, recueillis ont été libérés au Costa Rica. Cette étude a mis, en évidence la présence d'un grand nombre d'Opius perproximus Silvestri, deBiosteres caudatus Szépligeti, et deB. caudatus auct. (Braconidés: Opiinés). Les autres opiinés collectés comprenaientB. fullawayi (Silvestri),B. desideratus (Bridwell), et une espèce non décrite du genreOpius. Le taux de parasitisme des téphritidés collectées sur le café variait de 10 à 56%, avec une moyenne de 35% dans une plantation expérimentale et 17% dans une plantation commerciale. Sur les autres fruits, le taux de parasitisme était inférieur à 5%. Les espèces de téphritidés élevées le plus fréquemment des grains de café étaientTrirhithrum coffeae Bezzi etC. (Pterandrus) anonae Graham.C. capitata n'était présente qu'en faible abondance dans le café et des adultes n'ont que rarement été observés. Les autres espèces rencontrées comprenaientT. albonigrum (Enderlein),T. sp. nr.validum Bezzi,T. nigerrimum auct. nec. Bezzi,C. (Pterandrus) colae Silvestri,C. (Pterandrus) flexuosa (Walker),C. (Ceratalaspis) spp. groupecosyra (Walker),C. (Pardalaspis) sp. groupepunctata (Wiedemann),C. (Ceratalaspis) giffardi Bezzi,Dacus (Dacus) bivittatus (Bigot),D. (Didacus) ciliatus Loew, andGymnodacus sp.

     

The effect of host age ofLymantria dispar larvae [Lep.: Lymantriidae] on the development ofGlyptapanteles liparidis [Hym.: Braconidae]

The endoparasitic development ofG. liparidis was examined in 3 different host stages of gypsy moth larvae. Hatching ofG. liparidis-larvae occurred 3 to 5 days after oviposition in hosts parasitized during their premoulting period, and after 5 to 7 days in those parasitized in the 3^rd midinstar state. The parasites generally moulted to the 2^nd larval instar between the 11^th and 13^th day in the first group, and between the 13^th and 15^th day in the latter, when they had reached a volume of 0.04–0.05 mm³. The positive correlation between host ecdysis and the ecdysis of 1^st stadium larvae to L₂ suggested that host moulting influenced the development of the parasitoid larvae. Emergence from the host larvae occurred at 20°C after 27 days on average, and coincided with the parasites moulting to the 3^rd instar. Five to 7 days after spinning their cocoons near the developmentally arrested host larva, the male, and 1 to 2 days later the female wasps eclosed. Due to the variation in the number of parasites per host, no difference was observed between the hosts parasitized at various stages; however, a tendency for later parasitized hosts to contain more parasite larvae was evident. The nutritional conditions of the moth parental generation influenced both host and parasite development. On the other hand no influence of host age was observed on emergence dates of larvae and wasps.      

Fruit infesting tephritids [Dipt.: Tephritidae] and associated parasitoids in Chiapas,Mexico

A total of 1,302 parasitoids representing 8 species and 4 families were recovered from 9,818 fruit fly host fruits sampled. The most common parasitoid species wasDiachasmimorpha longicaudata (Ashmead). Average percent parasitism ranged between 0.44 and 29.23%. Parasitoid emergence data indicate thatAnastrepha ludens (Loew),A. obliqua (Sein),A. serpentina (Wiedeman),A. striata (Schiner) andToxotrypana curvicauda (Gerstaecker) were subject to parasitism. We provide information on the population fluctuation ofAnastrepha ludens, A. obliqua, A. serpentina, A. distincta (Greene),A. striata, A. fraterculus (Wiedeman),A. chiclayae (Greene),A. montei (Costa Lima),A. leptozona (Hendel) andA. tripunctata (Wulp).Anastrepha ludens andA. obliqua were the most common species, representing 95.3% of all fruit fly species caught in McPhail traps.      

Records on parasitoids of smaller poplar borer,Saperda populnea [Coleoptera,Cerambycidae] along the Danube in Bulgaria

During the period 1986–1988 the parasitoids of smaller poplar borer,Saperda populnea L. (Coleoptera, Cerambycidae) along the Danube in Bulgaria were studied. A total of 898 smaller poplar borer galls were investigated—289 in 1986, 404 in 1987, and 205 in 1988. In the particular years the parasitoids of smaller poplar borer namelyBillaea irrorata Meig.(Diptera, Tachinidae), Dolichomitus populneus Ratz. (Hym.: Ichneumonidae), andIphiaulax impostor Scop. (Hym.: Braconidae) destroyed from 13.1 up to 28.0% of the pest larvae, thus decreasing their frequency together with the other limiting factors by 56.1% up to 81.2%. The tachinid flyB. irrorata decreased the smaller poplar borer larvae frequency by 9.3% up to 18.9%, while the ichneumonoidD. populneus, and the braconoidI. impostor destroyed insignificant percentages of these larvae—0.5% up to 1.7% and 0.5% up to 2.5% resp. The flight period of the pest preceded or almost coincided with the ones ofD. populneus, andI. impostor. The flyB. irrorata always appeared last and its flight period was expanded.