Effects of sunlight exposure and log size on pine engraver (Coleoptera: Curculionidae) reproduction in ponderosa pine slash in Northern Arizona, U.S.A.

1 Abiotic conditions within logs can affect pine engraver Ips pini (Say) reproductive success, and slash management techniques have been developed that exploit these relationships to reduce brood production. In the present study, we investigated the affect of sunlight exposure on phloem temperature and moisture in logs of two diameters and the subsequent effects on pine engraver reproduction.
2 Logs, 30 cm in length, with diameters of 10 and 15 cm, were cut, left in the field for natural colonization by I pini , and then placed in an open meadow and under shade cloth, providing 27% and 66% shade, until offspring beetles had left the logs. Phloem temperature and moisture were recorded over the duration of the experiment and, at the end of the field experiment, logs were dissected and galleries were measured to gauge beetle reproductive success.
3 As sunlight exposure increased, phloem temperatures increased and potentially lethal temperatures were often reached in the high-sunlight exposure but seldom in the low-sunlight. Smaller diameter logs had drier phloem than larger diameter logs. All logs dried with time but sunlight level did not affect desiccation rates. Ips pini preferred attacking larger logs and the bottom side of logs. Sunlight exposure had a significant effect on net reproductive success in smaller diameter logs, with very little net reproductive success in high-sunlight exposed logs, and the highest reproductive success was found in small diameter logs in the low-sunlight treatments.
4 Management implications of these results are discussed.     

Attacks on living spruce trees by the bark beetle Ips typographus (Col. Scolytidae) following a storm-felling: a comparison between stands with and without removal of wind-felled trees

Abstract 1 To maintain biodiversity in forests more wind‐felled trees must be left where they fall. However, there is concern among forest owners that this may result in higher tree mortality caused by the spruce bark beetle, Ips typographus (L.) (Col.: Scolytidae). 2 In the 5 years following a major storm disturbance the number of standing spruces killed by I. typographus was determined in a total of 53 stands. In five of the stands all wind‐thrown trees were left (unmanaged stands) and in 48 of the stands, which were situated at distances of 1.4–10.0 km from each focal unmanaged stand, the wind‐felled trees were removed directly after the storm (managed stands). In the winter preceding the fifth summer new storm‐fellings occurred in the study area. 3 In the 4‐year period between the first and second storm‐fellings, 50–322 standing trees were killed by I. typographus per unmanaged stand. There was a direct linear relationship between the number of storm‐felled spruces colonized by I. typographus and the number of trees subsequently killed in the unmanaged stands. 4 Tree mortality caused by I. typographus in the unmanaged stands was almost nil in the first year, peaked in the second or third year, and decreased markedly to a low level in the fourth and fifth years. 5 In the 4‐year period between the first and second storm‐fellings twice as many trees were killed per ha in the unmanaged stands than in the managed stands: the average difference being 6.2 killed trees per ha, equivalent to 19% of the number of spruce trees felled by the first storm in the unmanaged stands. 6 Much higher numbers of trees were killed per ha in the stand edges than in the interiors of both the unmanaged and the managed stands.     

Recapture of Ips typographus L. (Col., Scolytidae) with attractants of low release rates: localized dispersion and environmental influences

1 The dispersal of Ips typographus L. (Col., Scolytidae) was studied using a mark–release–recapture approach in a grid of traps equipped with pheromone lures of release rates of about 8.4 mg/day of 2‐methyl‐3‐buten‐2‐ol (MB) and 0.29 mg/day of (S)‐cis‐verbenol (cV) in experiment 1, and 1.2 mg/day of MB and 0.04 mg/day of cV in experiment 2. 2 We investigated whether beetle dispersal reflected the simple diffusion pattern observed in previous I. typographus experiments, for which attractant release rates generally approached 50 mg/day of MB and 1 mg/day of cV. We also examined how environmental parameters (wind) and human activities (felling) could influence the beetles' flight. 3 The recapture percentage was higher in experiment 1 than in experiment 2: respectively, 7.0% (with 64 traps) and 2.3% (with 100 traps) of the beetles that took off were caught in the traps. 4 With the higher release rate (experiment 1), trap catches decreased with increased distance, whereas with the lower release rate (experiment 2), trap catches rose between 50 and 100 m then decreased with increasing distance. 5 Flight was little orientated by prevailing wind directions, a feature probably explained by the low wind speeds (0–1.2 m/s) observed throughout the study. 6 High trap catches of unmarked beetles close to areas undergoing thinning activities suggest that the presence of freshly cut spruce and larch material could have an influence on dispersal, attracting the beetles into the felling area. Spatial analyses show that capture patterns were autocorrelated up to distances of about 250 m.     

Competition between larvae of Ips typographus

A series of laboratory experiments were conducted to provide parameters for a simulation model for Ips typographus L. (Coleoptera, Scolytidae). This model illustrates larval competition and predicts brood survival in bolts. Brood survival depended upon larvae attaining a weight of at least 1.6 mg dry weight. The pattern of larval foraging in a bolt is hypothesized to be regulated by mutual avoidance among larvae. An analytical solution is presented for the construction of the pattern of larval paths originating from a single maternal gallery.

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Zusammenfassung Die Konkurrenz zwischen Larven des Borkenkäfers Ips typographus L. in Rundhölzern von Picea abies Karsten wurde im Laboratorium studiert. Um die Annahme zu testen, dass die Mortalität bei Borkenkäferlarven hauptsächlich von Futterkonkurrenz stammt, wurden die Parameter dieses Vorgangs geschätzt, die Zahl der Überlebenden vorausgesagt und verglichen mit der tatsächlichen Zahl Überlebender in jedem Rundholz. Die erforderlichen Voraussagen wurden mit dem Simulationsmodell von De Jong & Saarenmaa (1985) gemacht. In diesem Modell wird angenommen, dass die Larven Futter aufnehmen, in dem sie sich in der zweidimensionalen Rindenschicht bewegen und dass sie sterben oder sich vorzeitig in untergewichtige Käfer verwandeln, wenn sie in ein Gebiet gelangen, dessen Futter erschöpft ist.Es wurde geschlossen, dass der wichtigste Aspekt der Larvenbewegung im Hinblick aufs Überleben das gegenseitige Ausweichen ist. Die Muster der Larvengänge, wie sie aus einem Muttergang aufgrund gegenseitigen Ausweichens entstehen, wurden berechnet; sie stimmten sehr gut überein mit den beobachteten Mustern.Das Wachstum und das Überleben wurden in separaten Versuchen untersucht. Es wurde gezeigt, dass vorzeitiger Entwicklungsabschluss als Folge von Nahrungsknappheit eintritt. Das Überleben hängt davon ab, ob die Larve genug Futter zum Verpuppen erhalten hat. Dass die Larve eine bestimmte Grösse erreicht (Wachstum), war proportional zur Bewegung über eine bestimmte Distanz in der Rindenschicht. Unter den Versuchsbedingungen nahm das potentielle Trockengewicht der Puppen um 0,08 mg zu mit jedem mm Larvengang. Die minimale zur Verpuppung erforderliche Futtermenge war 20 mm Ganglänge, was proportional zu 1,6 mg Puppentrockengewicht ist.Aus diesen Resultaten wird geschlossen, dass Futterkonkurrenz die hauptsächliche Mortalitätsursache unter Larven von I. typographus in Rundhölzern im Laboratorium ist. Zudem wurde das Simulationsmodell erfolgreich überprüft; deshalb scheint es, dass Futterkonkurrenz entsprechend der vorigen Hypothese erfolgt. Die Annahme scheint vertretbar, dass dieser letzte Schluss auch für I. typographus unter natürlicheren Bedingungen zutrifft.

     

Performance of the tree‐killing bark beetles Ips typographus and Pityogenes chalcographus in non‐indigenous lodgepole pine and their historical host Norway spruce

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Interactions within and between species at different densities of the bark beetleIps typographus and its predatorThanasimus formicarius

Interactions between the bark beetleIps typographus and one of its predators,Thanasimus formicarius, were investigated in caged spruce logs containing both species in eight different density combinations. Productively (offspring per female) ofIps was adversely affected by highIps gallery density as well as highThanasimus density.Thanasimus productivity was enhanced by highIps gallery density but negatively affected by highThanasimus density.Ips productivity andThanasimus developmental rate differed between tree individuals, probably owing to tree-related differences in phloem thickness. Relative predator-causedIps mortality was ca 20% higher at high gallery density (ca 300 egg galeries per m²) than at low gallery density (ca 100 egg galleries per m²), indicating that mortality was density dependent. This difference was due to the fact thatThanasimus larval density was positively related toIps gallery density. Mortality increased by ca 0.4% with each additionalThanasimus larva per m², independently of gallery density. Relative population levels ofIps andThanasimus were monitored with pheromone traps in two regions differing in theirIps typographus outbreak history. Absolute catch as well as theThanasimus/Ips catch ratio were ten times greater in the outbreak region than in the non-outbreak region. Coupled with the results in the caging experiment, this indicates thatT. formicarius responds numerically to changes inI. typographus numbers per unit bark area as well as toIps population changes at the regional level. The findings suggest that predation under bark may be a significant factor in supressingI. typographus outbreaks.     

Can Norway spruce trees be ‘vaccinated’ against attack by Ips typographus?

1 A field experiment was carried out to test the hypothesis that treatment of Norway spruce trees with the Ips typographus-transmitted blue-stain fungus Ceratocystis polonica enhances tree resistance to later mass attack by this bark beetle. 2 Twenty-five mature trees were pretreated by inoculating a non-lethal dose of the fungus into the bark, while 18 trees served as untreated controls. Three and a half weeks after treatment a bark beetle attack was initiated by attaching dispensers with I. typographus pheromone to the tree trunks. 3 A significantly larger proportion (67%) of the control trees than of the pretreated trees (36%) were killed by the beetle attack. The result is discussed in relation to recent results regarding defence mechanisms in Norway spruce trees.     

Effect of varying monoterpene concentrations on the response of Ips pini (Coleoptera: Scolytidae) to its aggregation pheromone: implications for pest management and ecology of bark beetles

Abstract 1 Host plant terpenes can influence attraction of conifer bark beetles to their aggregation pheromones: both synergistic and inhibitory compounds have been reported. However, we know little about how varying concentrations of individual monoterpenes affect responses. 2 We tested a gradient of ratios of α‐pinene, the predominant monoterpene in host pines in the Great Lakes region of North America, to Ips pini's pheromone, racemic ipsdienol plus lanierone. 3 Ips pini demonstrated a parabolic response, in which low concentrations of α‐pinene had no effect on attraction to its pheromone, intermediate concentrations were synergistic and high concentrations were inhibitory. These results suggest optimal release rates for population monitoring and suppression programmes. 4 Inhibition of bark beetle attraction to pheromones may be an important component of conifer defences. At terpene to pheromone ratios emulating emissions from trees actively responding to a first attack, arrival of flying beetles was low. This may constitute an additional defensive role of terpenes, which are also toxic to bark beetles at high concentrations. 5 Reduced attraction to a low ratio of α‐pinene to pheromone, as occurs when colonization densities become high and the tree's resin is largely depleted, might reflect a mechanism for preventing excessive crowding. 6 Thanasimus dubius, the predominant predator of I. pini, was also attracted to ipsdienol plus lanierone, but its response differed from that of its prey. Attraction increased across all concentrations of α‐pinene. This indicates that separate lures are needed to sample both predators and bark beetles effectively. It also provides an opportunity for maximizing pest removal while reducing adverse effects on beneficial species. This disparity further illustrates the complexity confronting natural enemies that track chemical signals to locate herbivores.     

Characterization of the esterase isozymes of Ips typographus (coleoptera,scolytidae)

Four esterase isozymes hydrolyzing α-naphthyl acetate (α-NA) were detected screening whole body homogenates of larvae and adults of Ips typographus by electrophoresis. Two of the four isozymes (isozymes 3 and 4) were not detected by α-NA staining in the pupal stage, but topical application of juvenile hormone III (JH III) on the pupa induced these isozymes. The JH esterase (JHE) activity on the gel was associated with the proteins of isozyme 2. The compounds OTFP, PTFP, and DFP inhibited this catalytic activity of isozyme 2 on the gel at low concentrations, whereas the proteins of isozyme 3 and 4 were affected only at higher concentrations. A quantitative developmental study was performed to characterize which of the esterases hydrolyzed JH III, using a putative surrogate substrate for JH (HEXTAT) and α-NA. The I₅₀ of several esterase inhibitors and the JH metabolites were also defined. All findings supported the results that a protein associated with isozyme 2 is catabolizing JH and that isozymes 3 and 4 are the main contributors to the general esterase activity on α-NA. The JHE from Tenebrio molitor was purified by affinity chromatography. Although the recovery was low, an analytical isoelectric focusing gel showed that the JHE activity of the purified enzyme. T. molitor cochromatographed at the same pl as the JHE activity of I. typographus. Arch. Insect Biochem. Physiol. 34:203–221, 1997. © 1997 Wiley-Liss, Inc.     

Mutualism is not restricted to tree-killing bark beetles and fungi: the ecological stoichiometry of secondary bark beetles,fungi, and a scavenger

1. All bark beetles are in symbiosis with fungi. Although obligate mutualisms with fungi are common with tree-killing bark beetles (primaries), fungi associated with non-tree-killing bark beetles (secondaries) are usually dismissed as commensals. 2. Using an ecological stoichiometric approach, we show secondaries are also involved in nutrition-based mutualisms, some of which appear obligate, and that differences in symbiont provisioning efficiency have a potent effect on beetle carbon (C): nitrogen (N): phosphorus (P) ratios. 3. Some secondary beetles have high P contents and require efficient P provisioning via fungi, while others have low P contents that may allow them to exploit less efficient fungi or a broader range of species with variable efficiencies. A co-occurring scavenger that feeds on nutrient-poor bark beetle frass (excrement/boring residues) exhibited the lowest phosphorus content yet recorded for an invertebrate. 4. Our results generally support the growth-rate hypothesis that posits differences in C:P and N:P ratios in consumers are due to differential allocation of P to P-rich RNA to support growth. However, while the beetle species that accumulated the most biomass was considerably enriched in P and that with the least biomass was P-poor, one beetle species that was P-rich was also small possibly due to limitation by an element other than P. 5. Our results indicate that fungi are important to a broader range of bark beetles than previously recognised. Additional research is needed to describe how these various symbioses influence forest ecosystems via differential effects of fungi on host beetle fitness.