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.

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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.

     

Prolonged development time of the bark beetle predator Thanasimus formicarius (Col.: Cleridae) in relation to its prey species Tomicus piniperda (L.) and Ips typographus (L.) (Col.: Scolytidae)

1 The generation time of the bark beetle predator Thanasimus formicarius (L.) (Col.: Cleridae) was found to be predominantly two years both in the field and in rearing experiments conducted with two of its main prey species, the pine shoot beetle Tomicus piniperda (L.) and the spruce bark beetle Ips typographus (L.) (Col.: Scolytidae). 2 Emergence of T. formicarius adults in the first summer was only observed in one of the two rearing experiments, and these individuals represented only 6% of that generation. 3 All individuals not emerging as adults in the first summer remained as larvae in their pupal chambers until the second summer. Pupae were found starting around mid-June, and adults (in pupal chambers) were found from late July through to the end of August. 4 Newly emerged adults had to feed in order to survive hibernation. 5 The existence of T. formicarius races, specialized on certain bark beetle species and with phenologies matching their hosts, could not be demonstrated. After hibernation there was no difference in feeding activity, timing of egg-laying or proportion of egg-laying females between the T. formicarius adults reared as larvae on T. piniperda (flight period in April) and those reared as larvae on I. typographus (main flight period generally starting in late May or early June) when exposed to a temperature and day-length typical of the early spring conditions prevailing during the flight period of T. piniperda. 6 T. formicarius was parasitized by Enclisis vindex (Tschek) (Hym.: Ichneumonidae) in the pupal chamber. 7 The importance of these findings for the population dynamics of bark beetles is discussed.     

The influence of natural enemies on brood production inIps typographus (Col. scolytidae) with special reference to egg-laying and predation byThanasimus formicarius (Col.: Cleridae)

The predatorThanasimus formicarius (L.) (Coleoptera, Cleridae) and its preyIps typographus (L.) (Coleoptera, Scolytidae) were studied in the laboratory and the field. In the laboratory, 11T. formicarius laid 71–132 eggs (mean=162) during 66–123 days. During this time they ate 66–132I. typographus adults per pair (male +female). The number of eggs laid per female was not correlated with life span or the number ofIps eaten. In the field, predation byT. formicarius larvae and other natural enemies onI. typographus brood was studied in the last year of an outbreak. Caged and uncaged spruce bolts attacked byI. typographus were used, and pairs ofT. formicarius were released in the cages. The treatments were: uncaged bolts, caged bolts withoutT. formicarius, caged bolts with 4T. formicarius pairs, and caged bolts with 8T. formicarius pairs. The productivity ofI. typographus was highest in the caged bolts withoutT. formicarius (mean=4.5 offspring/female) and lowest in the uncaged bolts (mean=0.9 offspring/female). The density ofI. typographus galleries was similar in the different treatments. Hence, the variation in productivity between treatments could not have been due to differences in the levels of intraspecific competition. There was no difference in bark beetle productivity or density ofT. formicarius larvae between bolts with 4 pairs ofT. formicarius and bolts with 8 pairs (mean=2.5 offspring/female). This indicates that some kind of interference occurred betweenT. formicarius individuals (e.g. cannibalism) and that a maximum level of predation was reached. Predation by larvae ofMedetera spp. (Diptera, Dolichopodidae),Thanasimus spp. and other beetles, and parasitism by wasps (Hymenoptera, Pteromalidae) probably caused the low productivity in the uncaged bolts.      

Natural enemies ofTomicus Piniperda andIps acuminatus (Col,scolytidae) onPinus sylvestris near orléans,france: Temporal occurrence and relative abundance,and notes on eight predatory species

Ips acuminatus (Gyllenhall) andTomicus piniperda (L.) were the main scolytid beetles in Scots pine,Pinus sylvestris L., outbreak areas near Orléans, France during 1978-1979.Ips acuminatus attacked thin-bark logs whileT. piniperda attacked thick-bark logs. More than 150 species of insects were associated with these scolytids. Forty-five species (31 predators and 14 parasitoids) were confirmed as natural enemies of bark beetles. Three predators were abundant:Thanasimus formicarius L. (Col.: Cleridae),Rhizophagus depressus (F.) (Col.: Rhizophagidae) andMedetera spp. (Dipt.: Dolichopodidae). The first two were especially frequent in galleries ofT. piniperda, whileMedetera spp. were more abundant in galleries ofI. acuminatus. The main parasitoids wereRhopalicus tutela (Walker) andR. brevicornis Thomson (Hym.: Pteromalidae),Coeloides abdominalis Zetterstedt andC. melanostigma Strand,Dendrosoter middendorfi Ratzeburg andD. hartigii Ratzeburg, andSpathius rubidus Rossi (Hym.: Braconidae). These parasitoids are polyphagous. However,C. abdominalis was obtained fromT. piniperda but not fromI. acuminatus, while 5.rubidus andD. hartigii were obtained fromI. acuminatus but not fromT. piniperda. Temporal and relative abundance of natural enemies ofT. piniperda andI. acuminatus were studied. The predatorsR. depressus andT. formicarius had complementary limiting effects on bark beetles as adults and larvae ofR. depressus fed mainly on scolytid eggs, whileT. formicarius adults attacked scolytid adults and the clerid larvae preyed upon scolytid larvae. The predatory pressure applied byT. formicarius andR. depressus toT. piniperda early during the spring was later complemented by a series of parasitoids and other predators. This sequential timing of the natural enemies ofT. piniperda suggests that the interactions among the guild are more complementary than adverse. Differential timing of the natural enemies in logs infested byI. acuminatus and some evidence of competition between the predatorsMedetera spp. and parasitoids suggested a less effective control of this scolytid. Number and duration of larval instars were determined, and measurements of predatory activity were made forThanasimus formicarius, Rhizophagus depressus,Hypophloeus fraxini Kugelann (Col.: Tenebrionidae),Platysoma frontale Paykull (Col.: Histeridae), andXylocoris cursitans (Fallen) (Het.: Anthocoridae). Predatory activity was evaluated for last instar larvae and adults ofScoloposcelis obscurella (Zetterstedt) (Het.: Anthocoridae), and for last instar larvae ofMedetera sp. andLonchaea collini Hackman (Dipt.: Lonchaeidae)     

Semi‐field experiments investigating facilitation: arrival order decides the interrelationship between two saproxylic beetle species

1. Acanthocinus aedilis (Linnaeus) and Rhagium inquisitor (Linnaeus) both colonise the cambial layer in newly dead Scots pine Pinus sylvestris L. and thus are potential competitors. 2. Species interactions and in particular priority effects were investigated in an experiment with a replacement series design. Four pairs of adult beetles were released on each of 91 caged pine bolts (0.35 m long, 13 replicates). To test for priority effects, a 2‐week interval was used to separate the species. 3. The interrelationship in simultaneous coexistence was positive for A. aedilis and negative for R. inquisitor (+,?). Acanthocinus aedilis produced 161% more offspring per female in coexistence than alone. Rhagium inquisitor had lower offspring quality in coexistence where its larvae weighed 39% less than in one‐species bolts. 4. The interrelationship depended on arrival order. When A. aedilis had priority the interaction was again advantageous to A. aedilis (+,?) but when R. inquisitor had priority no species interaction occurred (0,0). Both species therefore fared better when having priority. 5. Facilitation in cerambycids is novel and the facilitative effect on A. aedilis could be oviposition incitement or resource enhancement by R. inquisitor.     

Impacts of Tomicus minor on distribution and reproduction of Tomicus piniperda (Col., Scolytidae) on the trunk of the living Pinus yunnanensis trees

This study investigated the impacts of Tomicus minor on Tomicus piniperda when the two Tomicus species coexist in the trunks of living Yunnan pine (Pinus yunnanensis L.) trees growing in the Kunming region, in south-western China. Tomicus piniperda mostly locates in the mid- and upper trunks of Yunnan pine tree; whereas T. minor mainly attacks the mid- and lower trunks. In the mid-trunk area from 1.0 to 5.0 m above ground, there are overlapping attack zones for the two Tomicus species, which accounts for an average of 80% of the entire zone occupied by T. piniperda and an average of 70% of the zone occupied by T. minor. In correspondence with their attack distributions, the average attack densities of the two species varied with height along the trunk, with 165.3 egg galleries per m² at a height of 6 m for T. piniperda, and 138.2 egg galleries per m² at the 1 m height for T. minor. It is suggested that T. minor adjusts its attack pattern with respect to T. piniperda, and thereby minimizes interspecific competition. No remarkable difference of average T. piniperda egg gallery length was found between the zone in which only T. piniperda occurred and the zone in which T. piniperda and T. minor coexisted; this is suggested to be due to low host quality in the upper trunk region where only T. piniperda was present. The number of T. piniperda larval galleries was highest when only T. piniperda was present, and decreased as T. piniperda and T. minor coexisted, particularly in the case when the density of T. piniperda was less than that of T. minor. Average larval density was 1649 larval galleries per m² where only T. piniperda occurred. However, when T. piniperda coexisted with T. minor, T. piniperda larval density averaged 1010 per m² when T. piniperda density was higher than T. minor, and averaged 442 per m² when T. piniperda density was less than T. minor, which led to the conclusion that T. minor makes a negative impact on T. piniperda reproduction when the two Tomicus species jointly colonize the same trunk of Yunnan pine tree.     

Oviposition behavior and reproductive success of the cerambycid Acanthocinus aedilis in the presence and absence of the bark beetle Tomicus piniperda

The influence of Tomicus piniperda (L.)(Col.:Scolytidae) attacks on Acanthocinus aedilis (L.)(Col.:Cerambycidae) (1) oviposition behavior, (2) breeding material preference and (3) reproductive success was investigated in caged pine bolts with and without egg galleries of T. piniperda. In addition, A. aedilis oviposition behavior was studied in the field. In the presence of T. piniperda attacks most A. aedilis eggs were laid through the entrance holes of the bark beetle egg galleries. In the absence of bark beetle attacks most A. aedilis eggs were laid at places where the outer bark had been damaged and the phloem was exposed. In cages, A. aedilis preferred to oviposit in bolts with bark beetle attacks. Thus, A. aedilis oviposited in all 17 of bark beetle attacked bolts but in only four of nine bolts without T. piniperda attacks. Neither the number of A. aedilis offspring nor the body size of emerging adults differed significantly between bolts attacked by bark beetles and unattacked bolts. Nor was there any significant relationship between the density of bark beetle egg galleries and the production of A. aedilis offspring.     

Parasitism byAphidius ervi (Hym.: Aphidiidae): Preference for pea aphid and blue alfalfa aphid (Hom.: Aphididae) and competition withA. Smithi

The pea aphidAcyrthosiphon pisum (Harris) and the blue alfalfa aphidA. kondoi Shinji are pests of alfalfaMedicago sativa L. worldwide. The parasitoidAphidium ervi Haliday attacks both species andA. smithi is host specific to pea aphid Experiments were conducted to determine the preference ofA. ervi forA. pisum andA. kondoi at three densities of hosts using the percentages of parasitism and superparasitism, and the distribution of eggs among hosts as criteria. Also examined was the outcome of competition between the two parasitoids when exposed to the same hosts. A. ervi attacks moreA. pisum when provided alone (no choice test) and when provided together in equal numbers withA. kondoi (choice test). There was no significant difference in the number of progeny produced by either parasitoid when a female of each species was introduced simultaneously into the same test unit containingA. pisum hosts. The effects of the pathogenPandora neoaphidis on this competition is reviewed.     

Host preferences ofTrichogrammatoidea bactrae fumata (Hym.: Trichogrammatidae) an egg parasitoid of leafrollers (Lep.: Tortricidae)

Trichogrammatoidea bactrae fumata Nagaraja is a naturally occurring egg parasitoid of some leafrollers in New Zealand kiwifruit orchards. The parasitoid showed a distinct preference for younger host-eggs ofEpiphyas postvittana, in which it achieved a higher rate of successful development. Such a preference is not as apparent withCtenopseustis obliquana. Host-species preference experiments found that the parasitoid rarely attackedPlanotortrix octo, and successful wasp emergence from parasitised eggs of this host was low.Epiphyas postvittana was the preferred host whenT. bactrae fumata was reared on this species. When reared onC. obliquana, the parasitoid showed no preference forE. postvittana orC. obliquana.     

Development of an improved attractive lure for the pine shoot beetle, Tomicus piniperda (Coleoptera: Scolytidae)

Abstract 1 The pine shoot beetle, Tomicus piniperda (L.) (Coleoptera: Scolytidae), is an exotic pest of pine, Pinus spp., and was first discovered in North America in 1992. 2 Although primary attraction to host volatiles has been clearly demonstrated for T. piniperda, the existence and role of secondary attraction to insect‐produced pheromones have been widely debated. 3 Currently, commercial lures for T. piniperda include only the host volatiles α‐pinene in North America and α‐pinene, terpinolene and (+)‐3‐carene in Europe. Several potential pheromone candidates have been identified for T. piniperda. 4 We tested various combinations of host volatiles and pheromone candidates in Michigan, U.S.A., and Ontario, Canada, to determine an optimal blend. 5 Attraction of T. piniperda was significantly increased when trans‐verbenol (95% pure, 3.2%cis‐verbenol content) was added with or without myrtenol to α‐pinene or to blends of α‐pinene and other kairomones and pheromone candidates. 6 Our results, together with other research demonstrating that trans‐verbenol is produced by T. piniperda, support the designation of trans‐verbenol as a pheromone for T. piniperda. A simple operational lure consisting of α‐pinene and trans‐verbenol is recommended for optimal attraction of T. piniperda.     

纵坑切梢小蠹对云南松蛀害研究

在昆明地区,纵坑切梢小蠹Tomicus piniperda L.表现出枝梢聚集、树干蛀害等重要的行为学特征,形成三种基本蛀害模式。横坑切梢小蠹、蓝色伴生真菌参与了纵坑切梢小蠹危害过程,并在其中发挥积极作用。上述因素的综合影响,加强了纵坑切梢小蠹对云南松Pinus yunnanensis寄主树木的危害能力。     

Attack dynamics of the pine shoot beetle, Tomicus piniperda (Col.; Scolytinae) in Scots pine stands defoliated by Bupalus piniaria (Lep.; Geometridae)

Abstract 1 After a 1‐year, extensive pine looper (Bupalus piniaria) outbreak, plots were laid out to study tree susceptibility to attack, and performance of Tomicus piniperda in pine trees suffering from varying levels of defoliation. 2 Tomicus piniperda was the dominating stem‐attacking species among the primary stem colonizers, and 82% of all trees that died had been colonized by T. piniperda. 3 Beetle attacks primarily struck severely defoliated trees, i.e. trees that suffered from 90% to 100% defoliation. 4 Beetle attacks peaked in the second year after cessation of the outbreak, and suppressed trees were both more frequently attacked and more susceptible to beetle attack than intermediate and dominant trees. 5 Trees surviving beetle attacks carried more foliage than trees that did not survive the attacks. 6 A single year of severe defoliation is enough to render pine trees susceptible to secondary pests, such as T. piniperda.     

Differences in photoperiod‐induced diapause plasticity among different populations of the bark beetle Ips typographus and its predator Thanasimus formicarius

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Observations on the reproductive biology of two parasites ofHylesinus varius andPhloeotribus scarabaeoides (Col: Scolytidae): Cheiropachus quadrum (Hym: Pteromalidae) andDendrosoter protuberans (Hym: Braconidae)

Observations on the biology ofCheiropachus quadrum (Hym: Pteromalidae) andDendrosoter protuberans (Hym: Braconidae), were conducted. Both species are the main parasites of the olive bark beetlesHylesinus varius andPhloeotribus scarabaeoides (Col: Scolytidae) in the South of Spain. Results have shown that an increase in body size of the host does not imply an increase in parasite efficiency. In fact, host size inversely affects parasite efficiency forC. quadrum. Bearing in mind this fact, the abundance of the host and the ease of its rearing in the lab, it is therefore advisable to useP. scarabaeoides as the host for mass rearing of the parasites studied here. On the other hand, the presence of white light is a negative factor for parasite longevity and fecundity. The pupae and all larval instars are parasitised.C. quadrum does not have a preference for any particular stage or larval instar of the host whilst there is a preference for the third and fifth larval instar byD. protuberans. With respect to the sex ratio of parasites, an increase in the number of males increases the fecundity of the females. The results obtained in this study can be considered essential in the development of a biological control system for olive bark beetle pests based on an increase in the population ofC. quadrum andD. protuberans.     

A new species of Bracon (Hymenoptera: Braconidae) parasitic on alien sweetpotato weevils in the south-west islands of Japan

A new species of braconid, Bracon yasudai Maeto et Uesato sp. nov., is described here from the south‐west islands of Japan. It is a solitary idiobiont ectoparasitoid of the larvae of the West Indian sweetpotato weevil, Euscepes postfasciatus (Curculionidae), and the sweetpotato weevil, Cylas formicarius (Brentidae), both feeding on Ipomoea batatas (L.) (Convolvulaceae). The percentage parasitism of the braconid on E. postfasciatus in the vines of I. batatas was 19.9–40.7% in the field. Bracon cylasovorus (Rohwer) reared from C. formicarius in the Philippines is also described.     

Efficiency of four heteroptera as predators ofAphis gossypii andMacrosiphum euphorbiae (Hom.: Aphididae)

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LIFE CYCLE OF THANASIMUS FORMZCARZUS (COLEOPTERA: CLERIDAE) IN SOUTHERN NORWAY*

Abstract Observations on the life cycle and bionomics of Thanasimus formicarius (L.) were made under natural conditions and in the laboratory. The studies provide information on mass flight, development periods of eggs, larvae, pupae and prepupae, and adult maturation stage based on monthly field investigations. The results of the research suggest that the overall life cycle of T. formicarius covers approximately two years and four months in southern Norway. During the first year of the life cycle, the clerid larvae overwinter in the pupal chambers at the bases of attacked spruce trees; adults emerge and mature sexually in the second year; and the mature adults overwinter inside the bark crevices of trunk and oviposit in the spring of the third year. The studies also expose that the clerid occurence coincides well with the bark beetle attacks occurring in spring and autumn in northern Norway.     

Interactions between Spathius agrili (Hymenoptera: Braconidae) and Tetrastichus planipennisi (Hymenoptera: Eulophidae), larval parasitoids of Agrilus planipennis (Coleoptera: Buprestidae)

Three hymenopteran parasitoids native to China are being released in the United States as biological control agents for the emerald ash borer (EAB), Agrilus planipennis Fairmaire, an Asian buprestid species responsible for mortality of ash trees (Fraxinus spp.) in North America. Two of these hymenopterans, Spathius agrili Yang (Braconidae), a larval ectoparasitoid, and Tetrastichus planipennisi Yang (Eulophidae), a larval endoparasitoid, prefer late-instar EAB larvae. This overlapping host preference raises concerns that interspecific competition following field releases may compromise establishment of one or both species. In a series of laboratory and field experiments, we found S. agrili and T. planipennisi exhibited similar parasitism rates when presented alone with EAB larvae for 12–14 days. However, S. agrili was more efficient at locating and parasitizing hosts within the first 27 h, possibly explaining why S. agrili excluded T. planipennisi in the laboratory trials and nearly excluded T. planipennisi in field trials when the two species were presented together with EAB larvae. We found that S. agrili parasitized larvae previously parasitized by T. planipennisi but not the reverse. However, S. agrili offspring failed to complete development on hosts that were previously parasitized by T. planipennisi. We recommend releasing these species separately in time or space to avoid the antagonistic interactions observed in this study.     

Morphological and genetic identification of the three pine pests of the genus Tomicus (Coleoptera, Scolytidae) in Europe

Abstract

• 1 Morphological characters were elaborated and part of the mitochondrial COI gene was sequenced in order to facilitate the determination of the three European pine bark beetles Tomicus piniperda, T. destruens and T. minor. The sequence information also provided the first information on the phylogenetic and phylogeographical relationships of these species.
• 2 Three hair rows were found on the antennal club of T. destruens between the second and third suture. Tomicus piniperda had only one row. Three different hair types were detected on the elytra – two hair types were found on T. piniperda, whereas the third hair type was only detected on the elytra of T. destruens.
• 3 The COI region (445 bp) revealed high sequence divergence among T. destruens, T. piniperda and T. minor. The three species proved to be monophyletic species with 16.98–19.23% sequence divergence. A phylogenetic approach placed T. minor and T. destruens as sister taxa, which contradicts the morphological findings.
• 4 European populations of T. piniperda shared two haplotypes, indicating a homogenous distribution of the genotypes. In the American populations only one of these European haplotypes was found. The Greek, Italian and Spanish T. destruens populations revealed three population‐specific haplotypes, indicating restricted gene flow.
• 5 Species‐specific primers were designed to allow a rapid and definitive determination of the two sibling Tomicus species by PCR.