Density-mediated responses of bark beetles to host allelochemicals: a link between individual behaviour and population dynamics

Abstract  1. Bark beetles (Coleoptera: Scolytidae) accept or reject host conifers based partly on concentrations of phloem monoterpenes. They colonise trees in aggregations, in response to pheromones that attract flying beetles to trees undergoing colonisation. A series of entry and gallery construction assays was conducted to determine whether responses by individual beetles to monoterpenes are altered by pheromones and/or the presence of other beetles.
2. Entry into the amended media by Ips pini and the length of time until entry were not influenced by the presence of aggregation pheromones.
3. Entry into amended media was influenced by the presence of other beetles on the surface of, or constructing galleries in, the substrate. The effects of alpha-pinene and limonene on host entry behaviour were mediated by the density of beetles on the surface of the assay arena, and by the density of beetles constructing galleries within the medium.
4. The percentage of beetles entering medium amended with higher concentrations of monoterpenes increased with increased density of beetles on the surface of the assay arena, until a threshold density of three or four beetles per assay arena, after which entrance rate declined.
5. The presence of other beetles constructing galleries elicited more rapid entry by the test beetles.
6. Gallery lengths were generally higher in the presence of aggregation pheromones.
7. Gallery lengths increased with increased density of beetles within the assay arena.
8. These results suggest a link between the density of bark beetles and responses of individuals. This linkage may partially explain behavioural changes observed during population eruptions.     

Response of the pine engraver, Ips pini (Say) (Coleoptera: Scolytidae), to conophthorin and other angiosperm bark volatiles in the avoidance of non-hosts

1 Seventeen non‐host angiosperm bark volatiles, seven of which are antennally active to Ips pini (Say), the pine engraver (PE), were tested for their ability to disrupt the response of the PE to pheromone‐baited traps. 2 Four green leaf volatiles (GLVs) were tested [1‐hexanol (Z)‐3‐hexen‐1‐ol, hexanal, and (E)‐2‐hexenal]. None had any disruptive effect singly, as a group or in all possible blends based on functional groups, despite the fact that the two aldehydes were antennally active. These compounds may have, in some instances, actually masked the disruptive effect of other compounds. The PE thus differs in its response from other Scolytidae, including other Ips spp. 3 Eight non‐host volatiles that were antennally active to other bark beetles, but not to PEs, had no disruptive effect, validating the use of coupled gas chromatographic‐electroantennographic detection analyses to detect compounds with potential behavioural activity. 4 The bicyclic spiroacetal conophthorin, (E)‐7‐methyl‐1,6‐dioxaspiro[4.5]decane, was disruptive when tested alone. When blends of two aldehydes [salicylaldehyde and nonanal] plus an alcohol and a phenol [benzyl alcohol and guaiacol] were combined with conophthorin, an enhanced disruptive effect was revealed. No single compound, other than conophthorin, disrupted the pheromone‐positive response and no blend that did not contain conophthorin was consistently disruptive to both sexes. Conophthorin seems to be a critical component in the non‐host angiosperm message for I. pini during its host selection phase. 5 Combination of the repellent synomones, verbenone and ipsenol, with the five disruptive non‐host volatiles may provide a potent treatment to protect trees, logs or stands from attack by the PE.     

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.     

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.