Pine monoterpenes and pine bark beetles: a marriage of convenience for defense and chemical communication

Pine-feeding bark beetles (Coleoptera: Scolytidae) interact chemically with their host pines (Coniferales: Pinaceae) via the behavioral, physiological, and biochemical effects of one class of isoprenoids, the monoterpenes and their derivatives. Pine monoterpenes occur in the oleoresin and function as behaviorally active kairomones for pine bark beetles and their predators, presenting a classic example of tri-trophic chemical communication. The monoterpenes are also essential co-attractants for pine bark beetle aggregation pheromones. Ironically, pine monoterpenes are also toxic physiologically to bark beetles at high vapor concentrations and are considered an important component of the defense of pines. Research over the last 30 years has demonstrated that some bark beetle aggregation pheromones arise through oxygenation of monoterpenes, linking pheromone biosynthesis to the host pines. Over the last 10 years, however, several frequently occurring oxygenated monoterpene pheromone components (e.g., ipsenol, ipsdienol and frontalin) have also been shown to arise through highly regulated de novo pathways in the beetles (reviewed in Seybold and Tittiger, 2003). The most interesting nexus between these insects and their plant hosts involves the late-stage reactions in the monoterpenoid biosynthetic pathway, during which isomeric dimethylallyl diphosphate and isopentenyl diphosphate are ultimately elaborated to stereospecific monoterpenes in the trees and to hydroxylated monoterpenes or bicyclic acetals in the insects. There is signal stereospecificity in both production of and response to the monoterpenoid aggregation pheromones of bark beetles and in response to␣the monoterpenes of the pines. In the California fivespined ips, Ips paraconfusus, we have discovered a number of cytochome P450 genes that have expression patterns indicating that they may be involved in detoxifying monoterpene secondary metabolites and/or biosynthesizing pheromone components. Both processes result in the production of oxygenated monoterpenes, likely with varying degrees of stereospecificity. A behavioral analysis of the stereospecific response of I. paraconfusus to its pheromone is providing new insights into the development of an efficacious bait for the detection of this polyphagous insect in areas outside the western United States. In contrast, a Eurasian species that has arrived in California, the Mediterranean pine engraver, Orthotomicus (Ips) erosus, utilizes both a monoterpenoid (ipsdienol) and a hemiterpenoid (2-methyl-3-buten-2-ol) in its pheromone blend. The stereospecificity of the response of O. erosus to the monoterpenoid appears to be the key factor to the improved potency of the attractant bait for this invasive species.Dedicated to Professor David L. Wood on the occasion of his 75th birthday, January 8, 2006     

Artificial diet and rearing methods for Thanasimus dubius (Coleoptera: Cleridae), a predator of bark beetles (Coleoptera: Scolytidae)

Clerid beetles are common natural enemies of bark beetles, and could potentially be used as biological control agents if they could be reared in sufficient numbers. We developed an artificial diet devoid of insect components for rearing Thanasimus dubius (Fabricius), a clerid that attacks several economically important bark beetles in eastern North America. We reared larvae of this predator using the artificial diet, and then used either natural or factitious prey to feed the adults so produced. Two different methods of presenting the diet were also examined. We then compared the performance of T. dubius reared on the artificial diet with newly-emerged wild individuals collected from the field. Our results suggest that adult predators reared on the diet are near in quality to wild ones, and high R₀ values can be obtained. No difference in prey preference was found between wild and diet-reared individuals after five generations in the laboratory. Sufficient numbers of predators could be generated using these techniques to permit limited field trials of augmentative biological control.     

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.     

Hymenopterous parasitoids of bark beetles [Scolytidae] in Israel: Host relation,host plant,abundance and seasonal history

Twenty six species of parasitoids were recorded from 15 species of bark beetles in Israel. Parasitoids were collected from infested cut branches of broadleaves and trap-trees of pine and cypress. Most of the parasitoids are oligophagous and range fromEcphylus caudatus, a monophagous species exclusive toHypoborus ficus, toHeydenia pretiosa, which develops on 11 species.Scleroderma domesticus, Agrilocida ferrieri andEupelmus nearurozonus are occasional parasitoids of bark beetles.Cephalonomia hypobori, Israelius carthami, S. domesticus, Entedon ergias, Cerocephala eccoptogasteri andRoptocerus xylophagorum are cryptoparasitoids and search for their host in the galleries. Known parasitoids are listed from 15 tree genera belonging to 8 botanical families. A typical range for the common parasitoids is presented with within-tree distribution of pine parasitoids. Relatively monophagous species are usually dominant parasitoids in a specific beetle population, whereas relatively polyphagous parasitoids vary in abundance among different host populations.Calosota spp., 2Eurytoma spp. andDendrosoter hartigii are usually present in small numbers. Information concerning the seasonal history of the parasitoids suggests that the major species may raise about 5 to 7 annual generations. Contribution from the Agricultural Research Organization, Bet Dagan, Israel N^o 1331-E, 1984 series     

Interaction betweenRastrococcus invadens [Hom.: Pseudococcidae] and two natural enemies

Two natural enemies of the mealybugRastrococcus invadens, the parasitoidGyranussoidea tebygi and the pathogenHirsutella cryptosclerotium were tested. Levels of parasitism by the parasitoid were reduced by the pathogen but overall mortality of the mealybug was greater both when agents were acting together.      

Incidence of natural enemies of the Colorado potato beetle,Leptinotarsa decemlineata [Coleoptera: Chrysomelidae] on a native host in Mexico

In central Mexico, the center of origin forLeptinotarsa decemlineata (Say) and its principal host plants,L. decemlineata is associated with a diverse complex of natural enemies. Observations during 2 years in Morelos, Mexico revealed 4 species of asopine pentatomids, 7 species of foliar searching carabids, and 2 coccinellids among the predators ofL. decemlineata. Parasitoids include 3 species of tachinids andEdovum puttleri Grissel, an eulophid egg parasitoid. These natural enemies include several predators that have not previously been reported attackingL. decemlineata. University of Maine Agriculture Experiment Station No. 1553     

Parasitoids and other natural enemies of Maruca vitrata feeding on Sesbania cannabina in Taiwan

Natural enemies of Marucavitrata (Geyer) feeding concealed inside leafwhorls of a green manure legume, Sesbaniacannabina, were monitored during the 1996 and1997 summer seasons in Taiwan. A braconid,Apanteles taragamae, accounted for anaverage of 92% of all parasitoid specimensreared from M. vitrata larvae and pupae.Other parasitoid species included two otherbraconids: Bassus asper and Dolichogenidea sp., three ichneumonids: Trichomma sp., Triclistus sp., and Plectochorus sp., and two unidentifiedtachinids. All parasitoid species attacked M. vitrata larvae and emerged as mature larvaefrom host larvae (braconids) or as adults fromhost pupae (ichneumonids and tachinids). Oneunidentified predatory staphylinid and threeentomopathogenic fungi: Fusarium sp.Paecilomyces sp. and Beauveriabassiana were also observed attacking M.vitrata larvae. Apanteles taragamae is a solitary endoparasitoid. Its parasitismreached as high as 63% of M. vitratalarvae found attacking S. cannabina. Theparasitism was higher during June to August andreduced during September to November when otherparasitoid species were more active. Ninespecies of hyperparasitoids, all belonging toHymenoptera, were recovered, mainly from thebraconids.     

Interactions between an eucoilid [Hymenoptera] and a staphylinid [Coleoptera] parasitoid of the cabbage root fly

The interaction betweenTrybliographa rapae andAleochara bilineata, 2 parasitoids of the cabbage root fly, is discussed. Larvae ofA. bilineata could not differentiate between cabbage root fly pupae containingT. rapae in its endoparasitic state and unparasitized pupae but could recognize pupae containingT. rapae once the latter had reached its ectoparasitic state. Attack byA. bilineata whileT. rapae was still in its endoparasitic state usually resulted in the staphylinid killing the eucoilid. IfT. rapae had reached the ectoparasitic state before the host pupa was attacked byA. bilineata larvae the eucoilid survived attack by the beetle larva. Multiparasitism, however, resulted in increased levels of mortality of both parasitoid populations.      

Notes on natural enemies attackingLepidosaphes species [Homoptera: Diaspididae] associated withCitrus in Costa Rica

Aphytis lepidosaphes Compere (Hymenoptera: Aphelinidae) parasitize two species ofLepidosaphes onCitrus in Costa Rica.Zagloba beaumonti andPentilia discors (Col.: Coccinellidae) also feed on both species.      

Interactions between the predators thanasimus formicarius (Col.: Cleridae) and rhizophagus depressus (Col: Rhizophagidae), and the bark beetle tomicus Piniperda (Col.: Scolytidae)

The occurrences of Thanasimus formicarius (L.) (Cleridae), Rhizophagus depressus (F.) (Rhizophagidae), and Epuraea marseuli Reitter (Nitidulidae) in cut Scots pines, Pinus sylvestris L., attacked by Tomicus piniperda (L.) (Col.: Scolytidae) were recorded in the field, and interactions between the species were studied in caged pine bolts attacked by T. piniperda. T. formicarius eggs and R. depressus adults were abundant in the T. piniperda attacked trees, whereas only a few individuals of E. marseuli were found. T. formicarius and R. depressus, but not E. marseuli, reproduced in the caged bolts. T. piniperda offspring production per unit area of bark was reduced by 41% when reared with R. depressus, by 81% when reared with T. formicarius, and by 89% when all three species were reared together, compared with T. piniperda alone. The interaction between T. formicarius and R. depressus was mutually antagonistic. When both species were present in the same bolt the total number of larvae was reduced by 49% for R. depressus and the number of large larvae (length > 10 mm) was reduced by 35% for T. formicarius compared with their respective production values when each species was present alone with the bark beetle. There was a positive relationship between T. piniperda egg gallery density and the production of R. depressus larvae per m². Larvae of both R. depressus and T. formicarius developed into new adults during the first summer.

---

Résumé Interactions entre les prédateursThanasimus formicarius (Col.: Cleridae) etRhizophagus depressus (Col.: Rhizophagidae) et le scolyteTomicus piniperda (Col.: Scolytidae) La présence deThanasimus formicarius (L.) (Cleridae),Rhizophagus depressus (F.) (Rhizophagidae), etEpuraea marseuli Reitter (Nitidulidae) a été étudiée sur le terrain dans des pins sylvestres (Pinus sylvestiris L) abattus, attaqués partomicus piniperda (L.) (Col.: Scolytidae) et les relations entre les espèces ont été étudiées sur des rondins de pin attaqués parT. piniperda et placés sous cage. Le nombre d’∄ufs deT. formicarius et d’adultes deR. depressus était important dans les arbres attaqués parT. piniperda, alors que seuls quelques individus deE. marseuli ont pu être trouvés.T. formicarius etR. depressus, mais pasE. marseuli, se sont reproduits dans les rondins sous cage. La production de descendants deT. piniperda par unité de surface d’écorce était réduite de 41% quand le scolyte était élevé avecR. depressus, de 81% en élevage avecT. formicarius et de 89% quand les trois espèces sont élevées ensemble, par comparaison avecT. piniperda seul. Les interactions entreT. formicarius etR. depressus étaient mutuellement antagonistes. Quand les deux espèces étaient présentes dans le même rondin, le nombre total de larves était réduit de 49% pourR. depressus et le nombre de grosses larves (longueur > 10 mm) était réduit de 35% pourT. formicarius comparé aux valeurs respectives quand chaque espèce était présente isolément avec le scolyte. Il y a une corrélation positive entre la densité de galeries avec des ∄ufs deT. piniperda et la production de larves deR. depressus par m². Les larves deR. depressus et deT. formicarius se sont transformés en adultes l’été suivant.

     

Note on the natural enemies ofPsylla hyalina [Homoptera: Psyllidae] onAlbizia lebbeck

A survey of the natural enemies associated withPsylla hyalina infestingAlbizia lebbeck has revealed 2 primary parasitoids, 1 hyperparasitoid, 5 coccinellids, 1 lygaeid and 3 fungal pathogens, all being new records forP. hyalina.      

Interactions betweenAcyrthosiphon kondoi [Homoptera: Aphidoidea] andAphelinus asychis [Hymenoptera: Chalcidoidea] and other parasites and hosts

In laboratory trials to investigate the parasite/host spectra of certain aphid pests and hymenopterous parasites, the aphidAcyrthosiphon kondoi Shinji encapsulated the egg of the aphelinid parasiteAphelinus asychis Walker. The resultant brown, sclerotic capsule was formed within 24 h of exposure of the aphid to parasitization and as far as is known prevented the development of the parasite to the larval stage. The capsule remained throughout the life of the aphid, whose longevity and fecundity were apparently not seriously impaired. A small number ofAphelinus escaped encapsulation, especially in aphids already containing capsule(s), and developed into normal, reproductive adults.A. kondoi did not encapsulate, andA. asychis was not encapsulated by any other species. However, thoughA. asychis readily parasitizedAphis citricola van der Goot,A. nerii Boyer de Fonscolombe andToxoptera citricidus (Kirkaldy), most of its progeny ceased development in these aphids before reaching the mummification stage, and died within the dead or dying, non-mummified aphid host.     

Kinetics of larval gregarious behavior in the bark beetleDendroctonus micans (Coleoptera: Scolytidae)

This paper provides a simple mathematical model of the aggregation behavior of the gregarious intracortical-feeding larvae of the bark beetle, Dendroctonus micans. The model's assumptions are that each individual produces pheromones at a constant rate and reacts positively to a pheromone gradient. These hypotheses have been tested by comparing experiments and simulations, both of which showed that (1) homogeneously distributed individuals aggregate rapidly at the center of the experimental or theoretical arena and (2) eccentric, preformed groups succeed in attracting dispersed individuals, provided that the initial size of these groups is sufficiently high. There is good agreement between most of the experimental and theoretical results, providing a link among chemical communication, density of larvae, and random events occurring during the development of a spatial structure.     

Helicosporidium infection of the great European spruce bark beetle, Dendroctonus micans (Coleoptera: Scolytidae)

An infection of the great European spruce bark beetle Dendroctonus micans (from Turkey) by the parasitic green alga Helicosporidium is described. This is the first time that Helicosporidium has been found to infect a bark beetle (Scolytidae) and the first time that a naturally infected beetle has been reported for the Eurasian continent. The typical cysts of Helicosporidium contain three ovoid cells and one helical, filamentous cell. The morphological characteristics are revealed by light and electron microscopy. Distinct electron-dense inclusions with a peculiar ultrastructure may represent pyrenoids. Since D. micans is an important pest, the discovery of a natural pathogen may offer a chance for biological control.     

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.     

Pesticide-induced mortality of natural enemies of the onion maggot,Delia antiqua [Dip.: Anthomyiidae]

Pesticide-induced differential mortality betweenDelia antiqua(MEIGEN)and several other organisms associated with its natural control [D. platura(MEIGEN),Coenosia tigrina (F.),Entomophtora muscae (COHN), andAphaereta pallipes (SAY)] were evaluated under simulated field conditions. Direct and residual differential mortalities were described for 3 herbicides (Chloro-IPC, nitrofen, and CDAA), 3 fungicides (maneb, chlorothalonil, and copper sulfate), and 1 insecticide (malathion). The recommended field application rates of these chemicals produced high positive differential mortality levels betweenD. antiqua and some of the natural mortality agents. Chloro-IPC, a preplant and midseason herbicide, induced 100% mortality ofA. pallipes over the 3-day residual test period. Modifications in behavioral orientation and death patterns ofD. antiqua adults infected withE. muscae were also noted in malathion-treated flies such that subsequent conidial dispersal was highly restricted. Michigan Agricultural Experimental Station Journal Article Number 10133.     

Survival of within-treeIps calligraphus [Col.: Scolytidae]: Effect of insect associates

Mechanical exclusion methods were used on the preferred portion of the hosttree bole to evaluateIps calligraphus brood mortality caused by the insect associate complex with and without the contribution ofMonochamus sawyer foraging. The studies compared brood adultIps calligraphus emergence from beetle infested bolts from which all insect associates were excluded, from which onlyMonochamus sawyers were excluded, and those bolts which were exposed to all insect associates. The least mortality occurred in bolts from which all insect associates were excluded, significantly more mortality occurred in bolts from which only sawyers were excluded, and the greatest mortality occurred in bolts exposed to all insect associates. Average monthly mortality due to insect associates was 89%, with 51% due to sawyer foraging. Insect associates were found to be more effective from May through July, while sawyer foraging produced the greatest within-bolt mortality from August through October.     

Trichogramma semblidis [Chalcidoidea: Trichogrammatidae] as a parasite of the bark beetle eggs [Coleoptera: Scolytidae]

It has been found thatTrichogramma semblidis (Aurv.) parasitizes the eggs ofleperisinus fraxini (Panz.),L. orni (Fuchs) andHylesinus crenatus (Fabr.) in typical variant ofCircaeo-Alnetum Oberdorfer 1953 in the Laski Experimental Forestry near Kepno. Thorough studies were conducted on exposed European ash logs and split wood in 3 methodical variants. The analysis of the galleries ofLeperisinus fraxini has showed that the average egg infestation amounted to 14.1%, 13.0% and 11.1% respectively. The egg infestation of individual gallery systems in the exposed control logs and split wood ranged between 2.7–51.2%, 1.2–44.1% and 1.7–69.2%, respectively, in different variants and the average effectiveness of the parasite amounted to 12.7% of the host egg mortality in 1970. Earlier observations and analyses of the gallery systems made on the ash trunks and branches in 1967 and 1968 showed 80.0% and 98.0% infestation of the eggs ofL. fraxini and 10.0% and 1.5% infestation of the eggs ofL. orni. Separate developmental stages of the parasite, the behaviour of the latter and penetration of the gallery systems were observed too. The most important data on the host species ofT. semblidis have been compared.     

An evaluation of some natural enemies of Spodoptera exigua on sugarbeet in northern California

Eggs and small to medium-sizedlarvae of Spodoptera exigua (Hübner)(Lepidoptera: Noctuidae) are exploited by acomplex of natural enemies in spring-plantedsugarbeet fields in northern California. Fieldstudies revealed that predation on sentinel eggmasses ranged from 20 to 100%egg mass;predation rate was lowest in fields previouslytreated with methomyl and highest innon-treated fields. Predators typicallydestroyed all of the eggs in a given egg mass;percentage predation per egg mass was densityindependent (spatial context). Survival of eggs(to neonate larvae) in cages that excludedpredators ranged from 80 to $>$90%. Theegg-predator guild consisted of adults andnymphs of Orius tristicolor(White)(Anthocoridae), Nabis americoferusCarayon (Nabidae), Lygus hesperus Knight(Miridae), and Geocoris punctipes (Say)(Geocoridae); larvae of Chrysoperlacarnea (Stephens) (Chrysopidae); and adults ofCollops vittatus (Say) (Melyridae). Laboratory evaluation revealed that largelarvae of C. carnea and the adults of theother species (except for O. tristicolor)could consume 100 eggs of S. exigua in a48 h period. The parasite guild associatedwith small and medium-sized larvae consisted ofthree species: Hyposoter exiguae(Viereck) and Pristomerus spinator (F.)(Ichneumonidae), both larval endoparasites; andChelonus insularis Cresson (Braconidae),an egg-larval endoparasite. Parasitization in field samples ranged from 30 to 65%. Smalland medium-sized larvae were also infected witha nuclear polyhedrosis virus (NPV); rates ofNPV infection ranged from 0 to 35% in fieldsamples. These results are consistent withanecdotal evidence that natural enemies,primarily generalist predators, are largelyresponsible for maintaining populations ofS. exigua at relatively low levels innontreated sugarbeet fields.     

Habitat preferences of aphidophagous coccinellids [Coleoptera]

From May to July 1982 and 1983, coccinellids were sampled in habitats that differed in aphid density, insolation, and host plant type. The number of adults ofCoccinella 7-punctata (CS),C. 5-punctata (CQ),Propylea 14-punctata (PQ),Adonia variegata (AV),Adalia bipunctata (AB),A. 10-punctata (AD), andCalvia 14-guttata (CA) were recorded. AB, AV and CS preferred high, while CQ tolerated low aphid density. CQ, CS, AB and AV preferred sunny, while AD and CA tolerated shaded conditions. CS and AV preferred herbaceous plants, AD and CA large plants, particularly trees, AB showed little specialization, CQ preferred trees early in the season and sparse herbaceous plants later. PQ appeared to be a generalist with a wide tolerance of all these factors, but with a tendency to move from trees to herbaceous plants as the season progressed. Habitats with a high abundance of a species were assumed to be those with on optimum combination of the preferred conditions. There were slight differences in the breadth of habitat preferences, and a little habitat overlap between species. The tendency to aggregate was greater in AB than in other species. Annual variations in species abundance influenced the number of habitats occupied, but not the abundance in the favoured places.