Effect of nutrition of parasites with nectar of melliferous plants on parasitism of the elm bark beetles (Col., Scolytidae)

In the forest of Kljestevica, the elm is attacked by four species of elm bark beetles: Scolytus multistriatus (Marsh.), Scolytus scolytus (Fab.), Scolytus pygmaeus (Fab.) and Scolytus ensifer (Rich.) (Col., Scolytidae). Scolytus multistriatus and S. pygmaeus are the most numerous species. Elm bark beetle is parasitized by four species of parasites: Dendrosoter protuberans (Nees), Ecphylus silesiacus (Ratz.), Coeloides scolyticida (Wesm.) (Hym., Braconidae) and Entedon leucogramma (Ratz.) (Hym., Eulophidae). Ecphylus silesiacus and D. protuberans are the most numerous species. The nectar of mustard ( Sinapis alba L., Brassicaceae) flowers, sweet basil ( Stachys recta L., Lamiaceae) flowers and of wild carrot ( Daucus carota L., Umbelliferae), has an important effect on the number of parasites and on the parasitism of the elm bark beetles. Wild carrot is the most attractive plant to the parasites for additional nutrition, especially for D. protuberans . Mustard and sweet basil flowers are suitable for additional nutrition of E. silesiacus imagos. On meadow flowers growing spontaneously in forest clearings, the lowest number of parasite imagos was collected by an entomological catcher, and the lowest percentage of parasitized larvae and eggs of the elm bark beetle was recorded in elm control catch trunks that were not near the cultivated melliferous plants.     

Biological control of Dutch elm disease: Larvicidal activity of Trichoderma harzianum, T. polysporum and Scytalidium lignicola in Scolytus scolytus and S. multistriatus reared in artificial culture

A method is described for the culture of larvae of Scolytus multistriatus and Scolytus scolytus on an artificial medium following exposure to cultures of microorganisms. In control cultures, a natural mortality rate of 21.2% was found for S. multistriatus and 17.6% for S. scolytus. The effects of Trichoderma harzianum, T. polysporum and Scytalidium lignicola on fifth instar larvae of S. scolytus and S. multistriatus reared on the artificial medium were studied. The fungi were larvicidal and larval mortality was increased to more than 80% by inoculation of the larvae with the fungi. Another fungus, Phomopsis oblonga, had little effect on larvae of S. scolytus. The results are discussed in relation to mechanism of pathogenicity of the fungi and their potential use in the control of Dutch elm disease. It is proposed that with modifications, the method is applicable to other bark beetle pests.     

Phoretic mites and nematode associates of Scolytus multistriatus and Scolytus pygmaeus (Coleoptera: Scolytidae) in Austria

Abstract 1 The species assemblages and abundance of phoretic mites and nematodes associated with the elm bark beetles, Scolytus multistriatus and Scolytus pygmaeus, were studied in Austria. 2 A total of 3922 individual mites were recorded from 144 adults of S. multistriatus and 178 adults of S. pygmaeus. The species spectrum was identical and the relative abundance of mites was very similar for both species of scolytids. Nine mite species, Pyemotes scolyti, Pseudotarsonemoides eccoptogasteri, Trichouropoda bipilis, Tarsonemus crassus, Proctolaelaps eccoptogasteris, Proctolaelaps scolyti, Chelacheles michalskii, nr. Eueremaeus sp. and Elattoma sp. were detected. Two of the nine species, nr. Eueremaeus sp. and Elattoma sp., are documented here as new associates of Scolytus spp. 3 Pyemotes scolyti was the most frequent mite species, and Ps. eccoptogasteri and T. bipilis were relatively common, whereas the other mites occurred occasionally or were rare. 4 The trophic roles of most of the mites associated with S. multistriatus and S. pygmaeus are poorly known, but they may include fungivores, parasitoids of bark beetle broods, predators of bark beetle broods and/or mites and/or nematodes. 5 Besides phoretic mites, two nematode associates were seen on the investigated insects. A species of Cryptaphelenchus occurred under the elytra of both scolytid species, whereas the adults of a Neoparasitylenchus sp. were present inside abdomens of S. multistriatus, but absent from S. pygmaeus.     

Fusarium solani, a weak pathogen of the larval stages of the large elm bark beetle Scolytus scolytus (Coleoptera: Scolytidae)

Fusarium solani, a common colonist of beetle-infested elm bark, has been isolated from dead larvae and adults of the large elm bark beetle, Scolytus scolytus. Two infectivity tests indicated that this fungus is probably a weak pathogen of the larval stages of the beetle. Total mortality was not achieved in either test, but the mortality associated with fungal inoculation was occasionally significantly greater than the mortality in the uninoculated controls in both experiments (P<0.05, experiment A; P<0.025, experiment B). Disease symptoms suggest that the larvae attacked by F. solani may have died from bacterial infection.     

The internal microbial flora of the elm bark beetle, Scolytus scolytus, at all stages of its development

Adult Scolytus scolytus emerged from the pupal stage with sterile guts although microorganisms were present within the pupal body. Gut sterility was maintained in a large proportion of newly formed adults while they were boring out of an elm log. The adult gut microbial flora is mainly acquired during subsequent feeding. Differences were found between the microbial gut flora of both larvae and adult S. scolytus, and of both male and female beetles.     

Fauna of phloemo-xylophagous insects,their parasitoids and predators on <Emphasis Type="Italic">Ulmus minor</Emphasis> in Serbia

The fauna of phloemo-xylophagous insects, their parasitoids and predators was studied on Ulmus minor in Serbia. Sixty insect species were recorded, of which 22 species were phloemo-xylophagous insects, 33 species were their parasitoids and 5 species were their predators. Among phloemo-xylophagous insects, the most common and most abundant species were Scolytus pygmaeus (59.93%), Magdalis armigera (11.79%), S. multistriatus (9.24%), Exocentrus punctipennis (8.68%), S. ensifer (5.48%) and S. kirschii (1.28%). The most frequent and most abundant parasitoid was the species Ecphylus silesiacus (65.95%). Also, the parasitoids Dendrosoter protuberans (7.46%), Rhaphitelus maculatus (6.25%), Cheiropachus quadrum (5.49%), Acrocormus semifasciatus (3.68%), Entedon ergias (1.65%), Spathius rubidus (1.53%), Eubazus augustinus (1.46%) and Eurytoma morio (1.37%), were of major importance.     

The Banded Elm Bark Beetle, Scolytus schevyrewi Semenov (Coleoptera, Curculionidae, Scolytinae) in North America: a taxonomic review and modifications to the Wood (1982) key to the species of Scolytus Geoffroy in North and Central America

In 2003, an Asian bark beetle, Scolytus schevyrewi Semenov (Coleoptera: Curculionidae: Scolytinae), the banded elm bark beetle, was detected for the first time in North America. This paper modifies the Wood (1982) key to the species of Scolytus Geoffroy to enable identification of Scolytus schevyrewi in North and Central America. Variation of diagnostic characters in Scolytus schevyrewi is discussed.     

Comparative laboratory studies on three fungal pathogens of the elm bark beetle Scolytus scolytus: Pathogenicity of Beauveria bassiana,metarhizium anisopliae, and Paecilomyces farinosus to larvae and adults of S. scolytus

The entomogenous fungi Beauveria bassiana (nine isolates), Metarhizium anisopliae (seven isolates), and Paecilomyces farinosus (four isolates) were tested as pathogens of larvae of the elm bark beetle, Scolytus scolytus. Single isolates of B. bassiana and M. anisopliae were also tested against adult beetles. Of the 21 isolates tested as conidial suspensions against larvae, all proved pathogenic. The three most and least virulent isolates were, respectively, isolates of B. bassiana and M. anisopliae. The other isolates fell between these two extremes, with the four P. farinosus isolates all moderately virulent. Spore retention on larvae following inoculation was estimated by washing conidia off the larvae. From the results it was possible to relate larval mortality to the approximate spore dose causing infection at different spore concentrations. Thus, application of spores of the three pathogens at a concentration of 10³ spores/ml resulted in limited mortality. At this concentration, an average of only a single spore was recovered from the inoculated larva. Adult bark beetles also proved susceptible to infection by isolates of B. bassiana and M. anisopliae. They were exposed to discs of elm bark dipped in a conidial suspension. It was estimated that a dose of less than 100 spores could cause infection of beetles following feeding on the elm bark discs.     

The DD-136 strain of Neoaplectana sp. as a potential biological control agent for the European elm bark beetle,Scolytus scolytus

When logs infected with Scolytus scolytus were sprayed with the DD-136 strain of Neoaplectana sp. in solutions of either distilled water, glycerin, or P.B.I. wetting agent, the nematodes were efficiently dispersed and entered the logs through the insect entrance holes in the bark.Logs treated with Neoaplectana sp. were found to contain a significantly higher proportion of dead, nematode-infected insects. No difference was found between the various methods of nematode application. The results indicate that Neoaplectana sp. may have a possible role in the control of S. scolytus in elm logs.     

脐腹小蠹形态特征和生物学特性

脐腹小蠹Scolytus schevyrewi Seme.是榆树的一种重要蛀干害虫。本研究在宁夏盐池县通过室内观察和林间调查相结合,研究了脐腹小蠹的形态特征及其生活史、成虫寿命、习性、交尾行为等生物学特性。结果显示:脐腹小蠹虫在宁夏盐池县一年发生2代,以老熟幼虫或蛹越冬,老熟幼虫约占98.2%,蛹约占1.8%。越冬幼虫于5月上旬开始化蛹,5月中旬为化蛹盛期,5月下旬开始羽化,6月上旬为羽化盛期。第2代幼虫于6月底、7月初开始化蛹,7月下旬达羽化盛期,8月上旬羽化结束;脐腹小蠹雌成虫平均寿命为(27.14±3.74)d,雄成虫的平均寿命为(16.86±4.13)d,差异显著(P<0.05);此外,脐腹小蠹无滞育现象;脐腹小蠹的4种天敌,分别是榆小蠹长茧蜂Elachistocontrum sp.、虱形螨Pedieuloids ventricosus Newport、拟截斑郭公虫Thansimus sp.和蠼螋Labidura sp.。     

Laboratory studies on the fungus Verticillium lecanii, a larval pathogen of the large elm bark beetle (Scolytus scolytus)

From 1972 to 1974, estimates of the natural larval mortality (> second instar) of elm bark beetles caused by pathogenic organisms were always below 7'5 % of the beetle population. The pathogenic fungus Verticillium lecanii was frequently isolated from field-collected dead larvae, and in the laboratory all larvae were killed in 5 days when exposed to spore concentrations of 4·5 × 10⁶ spores/ml. V. lecanii begins to lose its pathogenicity after prolonged culture on artificial media. The time taken for V. lecanii to kill Scolytus scolytus larvae when exposed to a logarithmic series of spore dilutions from 9·1 × 10⁷/ml to 9·1 × 10³/ml increased with decreasing amounts of inoculum. Even at spore concentrations as low as 9·1 × 10³/ml the mortality of treated larvae was greater than that of untreated individuals. At 100% r.h. all treated larvae were killed over a temperature range of 5–30 °C; those maintained at 25 °C were killed most rapidly and those kept at 5 °C the slowest.     

Some effects of freezing temperatures on overwintering larvae of the large elm bark beetle (Scolytus scolytus)

Overwintering final instar larvae of Scolytus scolytus were tested for coldhardiness by exposure to a range of sub-zero temperatures (–7°Cto –31 °C) in a frost-gradient apparatus for 7 days. The Lt₅₀ for larvae removed from the bark (– 20.5 °C) was significantly different (P < 0.01) from the Lt50 (–18.3 °C) for larvae insulated by the bark (thickness of 7 mm ± 2 mm). Larvae with food in their digestive tract were more susceptible to freezing than the overwintering final instars which had voided their stomach contents. Duration and intensity of cold did not cause any adverse long-term effects. Most of the larvae which survived the sub-zero treatments were able to pupate or reach the adult stage. The mean supercooling point of the overwintering larvae (–30.85 °C) confirmed their cold-hardiness.     

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.     

Dendrosoter protuberans (Hymenoptera: Braconidae), An Important Elm Bark Beetle Parasitoid

The most favourable period for Dendrosoter protuberans development occurred when wasps were introduced 16 days after the beginning of Scolytus multistriatus development (the greatest number of emerged wasps per a single parasitoid female with a very positive sexual index and high percentage of parasitised elm bark beetle larvae). No significant differences were observed in parasitoid efficiency between the 11^th and the 21^st day. The most unfavourable period for elm bark beetle parasitisation was six days after the beginning of S. multistriatus development (a low reproduction of D. protuberans, a negative sexual index and a low number of destroyed S. multistriatus larvae). The reproduction of both the elm bark beetle and D. protuberans was greater in the first, than in the second generation. The developmental period of D. protuberans in the first generation was approximately two days longer than in the second generation. Furthermore, the development of females was 1-2 days longer than the development of males. D. protuberans developed most successfully on S. multistriatus (the most abundant parasitoid reproduction with the longest developmental period and a very positive sexual index that resulted in a high percentage of parasitised larvae) and then on S. ensifer, S. pygmaeus and Pteleobius kraatzi. On the other hand, S. scolytus larvae were the poorest host of all elm species - the reproduction of D. protuberans in this species was the scarcest with the shortest period of development and a negative sexual index, hence larval parasitisation was the lowest.     

Additions to the check list of bark beetles (Coleoptera,Curculionidae: Scolytinae) from Leningrad Province,Russia

The field work of the authors during the last 16 years in both the western and the eastern parts of Leningrad Province, Russia, has resulted in the addition of 7 species to the regional list of bark beetles, namely: Hylurgus ligniperda F., Crypturgus subcribrosus Egg., Pityogenes irkutensis Egg., Scolytus mali Bechst., S. scolytus F., S. pygmaeus F., and Xyleborinus attenuatus Blandf. Of these species, P. irkutensis occurs only in the eastern districts of Leningrad Province, C. subcribrosus, over the whole territory, and the remaining species, only in the western districts. We suggest excluding Polygraphus proximus Blandf. from the regional faunal list. As a result, the updated list of the bark beetle species of Leningrad Province comprises 75 taxa. New data on findings of rare species in the region are considered.     

Host Acceptance and Larval Competition in the Banded and European Elm Bark Beetles, <Emphasis Type="Italic">Scolytus schevyrewi</Emphasis> and <Emphasis Type="Italic">S. multistriatus</Emphasis> (Coleoptera: Scolytidae): Potential Mechanisms for Competitive Displacement between Invasive Species

A recent survey revealed that the newly invasive banded elm bark beetle, Scolytus schevyrewi, was much more abundant than the long-established European elm bark beetle, S. multistriatus, in areas of Colorado and Wyoming, USA. This study was initiated to determine whether competitive displacement of S. multistriatus by S. schevyrewi might be mediated by a sequence of behavioral interactions between the species on and below the bark surface. At the first step in the sequence, host acceptance behavior (i.e., time to tunnel into the host) was monitored among female S. schevyrewi and S. multistriatus. There was no substantial difference in host acceptance behavior when females were offered small cut logs (bolts) of Siberian elm, Ulmus pumila, infested with con- or heterospecific females. At the second step, mating and oviposition usually follow after a female has excavated a gallery. Mean oviposition per parental female in 1 wk was not significantly different among treatments: S. schevyrewi at medium density of 2.86 parental females per dm², at high density of 5.71 per dm²; S. multistriatus at medium density, at high density; and mixed species (i.e., S. schevyrewi and S. multistriatus combined, with each at medium density). At the third step, progeny production and progeny size were monitored among the same density treatments, but the number of parental beetles and size of bolts were doubled. Differences in progeny production would reflect larval competition, since the number of eggs initially laid was not different. Both S. schevyrewi and S. multistriatus were sensitive to intraspecific competition because size of progeny was smaller in high than medium density treatments, but total progeny production was similar at both densities. Scolytus schevyrewi was a stronger interspecific competitor than S. multistriatus. Smaller S. multistriatus progeny, and four-fold more S. schevyrewi progeny were produced when both species developed on the same host. The contributions of these proximal behavioral events toward the mechanism for competitive displacement are described in the context of the host colonization behavior of Scolytus spp. The competitive advantage of S. schevyrewi larvae established through this study, and the more rapid response of S. schevyrewi to elm hosts established through other investigations, may be the key mechanistic components that facilitate the displacement of S. multistriatus by S. schevyrewi.     

Phloemophagous and xylophagous insects, their parasitoids, predators and inquilines in the branches of the most important oak species in Serbia

Altogether 26 species of phloemophagous and xylophagous insects, 47 species of parasitoids, 14 species of predators and 7 species of inquilines were identified on the branches of Quercus cerris, Q. frainetto, Q. petraea and Q. robur, diameter 3–15 cm, at 24 sites in Serbia over the period 1992–1996. The greatest number of the identified species were taken from Q. petraea branches (66), followed by Q. cerris (49), Q. frainetto (48) and Q. robur (43). Among the identified phloemophagous and xylophagous insects, the most frequent and the most abundant species were Scolytus intricatus, Agrilus angustulus and Xylotrechus antilope. The most frequent and the most abundant parasitoid was Ecphylus silesiacus. In some samples, the species Entedon ergias, Cheiropachus quadrum and Dendrosoter protuberans were also among the more abundant parasitoids. The most significant predator was the species Tilloidea unifasciata, and the most significant inquilines were the species Poecilothrips albopictus and Asynapta pectoralis.     

Feeding by Scolytus bark beetles to test for differently susceptible elm varieties

Dutch elm disease (DED), caused by the fungi Ophiostoma ulmi and O. novo‐ulmi, has reduced elm populations severely in Europe and North America. Breeding programmes are in action to find less susceptible elm varieties suitable for re‐establishing elm stands. Bark beetles, mainly Scolytus spp., are the only known natural vectors of DED. During twig feeding, beetles transfer Ophiostoma spores to healthy elms. Thus, less palatable elms should run a lower risk of DED infections. In feeding preference bioassays, we offered twigs from elms exhibiting different degree of susceptibility to O. novo‐ulmi, together with non‐host trees to Scolytus beetles. Scolytus multistriatus preferred wych elm, Ulmus glabra, to 100% in two‐choice tests, whereas S. laevis did not discriminate between a tolerant and a susceptible variety of field elm, U. minor. We suggest that the feeding assay is useful as a low‐tech method in breeding programmes for evaluating the suitability of promising elm genotypes to vector insects.     

Significance of parasitoids in the reduction of oak bark beetle Scolytus intricatus Ratzeburg (Col., Scolytidae) in Serbia

Abstract:  As the vector of vascular fungi of the genus Ophiostoma the oak bark beetle, Scolytus intricatus , is one of the significant links in the chain of agents of oak forest decline in Serbia. It is known that the adults of this bark beetle, which develop under the bark of the trees infected by fungi of the genus Ophiostoma , transport the spores of these fungi. During the maturation feeding, the spores are transmitted to healthy tree crowns, where they germinate and cause infection. In period 1992–96 at 27 localities in Serbia, the significance of parasitoids in the reduction of this bark beetle was examined, as the intensity of spore transmission depends upon the number of oak bark beetle adults in nature. In the research of parasitoids of S. intricatus 20 species were identified in five families of Hymenoptera: Braconidae (six species), Eurytomidae (one), Pteromalidae (10), Eupelmidae (one) and Eulophidae (two). Among these parasitoids the greatest influence on the abundance of oak bark beetle was the species Ecphylus silesiacus . It was identified in 90.91% of study samples, its domination was 39.15% and the percentage of oak bark beetle parasitism was 5.66%. In addition, the species Entendon ergias , Dendrosoter protuberans and Cheiropachus quadrum were significant in reduction of S. intricatus . The average percentage of parasitism of oak bark beetle by E. ergias was 2.74%, by D. protuberans it was 2.63% and by Ch. quadrum 1.63%. The significance of other parasitoids in the reduction of oak bark beetles was low. In the study period the total parasitism of oak bark beetle varied between a minimum of 8.64% in 1992 and a maximum of 19.64% in 1996. The average for the whole study period was 14.49%.     

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.