Interspecific competition between the principal larval parasitoids of the pine sawfly,Neodiprion sertifer (Geoff.) (Hym.: Diprionidae)

Summary Interspecific competition between the 3 principal larval parasitoids of the pine sawfly, Neodiprion sertifer, is of common occurrence when total larval parasitism, and hence multiple attack rates, are high. At the intrinsic level, the ectoparasitoid Exenterus abruptorius is superior to the 2 endoparasitoids, Lophyroplectus luteator and Lamachus eques, respectively, whereas L. luteator is superior to L. eques only. During mass outbreaks of the host competition between Exenterus and Lophyroplectus is most intense, whilst Lamachus fails to build up in the presence of its 2 competitors (Fig. 1). However, due to its superior host finding ability, Lamachus dominates in areas of low host densities where the 2 other species are less efficient (Table 1). L. luteator frequently sustains heavy losses in direct competition with E. abruptorius, but these can be largely compensated in the next generation because its fertility is about 5-times higher than that of Exenterus.     

Mitonuclear discordance is caused by rampant mitochondrial introgression in Neodiprion (Hymenoptera: Diprionidae) sawflies

We investigate the pervasiveness of hybridization and mitochondrial introgression in Neodiprion Rohwer (Hymenoptera; Diprionidae), a Holarctic genus of conifer-feeding sawflies. A phylogenetic analysis of the lecontei species group revealed extensive discordance between a contiguous mitochondrial region spanning three genes (COI, tRNA-leucine, and COII) and three nuclear loci (EF1alpha, CAD, and an anonymous nuclear locus). Bayesian tests of monophyly and Shimodaira-Hasegawa (SH) tests of topological congruence were consistent with mitochondrial introgression; however, these patterns could also be explained by lineage sorting (i.e., deep coalescence). Therefore, to explicitly test the mitochondrial introgression hypothesis, we used a novel application of coalescent-based isolation with migration (IM) models to measure interspecific gene flow at each locus. In support of our hypothesis, mitochondrial gene flow was consistently higher than nuclear gene flow across 120 pairwise species comparisons (P < 1 x 10(-12)). We combine phylogenetic and coalescent evidence to identify likely cases of recent and ancient introgression in Neodiprion, and based on these observations, we hypothesize that shared hosts and/or pheromones facilitate hybridization, whereas disparate abundances between hybridizing species promote mitochondrial introgression. Our results carry implications for phylogenetic analysis, and we advocate the separation of high and low gene flow regions to inform analyses of hybridization and speciational history, respectively.     

From where are insects recruited? A new model to interpret catches of attractive traps

Abstract 1 Two new concepts describing the origin of insects caught in an attractive trap are presented. 2 Male European pine sawflies Neodiprion sertifer Geoffroy (Hymenoptera: Diprionidae) were marked and released from 50, 100, 200, 400 and 800 m in the four cardinal directions around a centrally placed pheromone trap. 3 Based on linear regression of transformed data, we calculated the seasonal sampling range (r_s) as 1040 m. 4 We estimated the previously defined ‘effective sampling area’ (α) at 4.9 ha, assuming that the insects are evenly distributed around the trap and that they are attracted from a circular area around it. This is the area from which all insects originate if the trap is 100% effective within the area but captures nothing outside of it. The effective sampling area reveals nothing about the origin of the insects caught. We defined the Cumulative Proportional Catch (CPC) that gives the proportion of the trap catch that originates from an area within a distance r from the trap. At r = r_s CPC = 1, and in our study 50% of the captured insects originated up to 450 m from the trap. Thus, for the trap used in this study, a relatively large proportion of the catch originates some distance from the trap. 5 We also defined the Catch Concentration (CC), which is the ratio of the radius of the effective sampling area (r_α) to r_s. For our data, CC = 0.12, which is intermediate to high compared to the few other studies that we have extracted information from. If r_α is considerably lower than r_s, then only a small proportion of the insects caught originate from close proximity to the trap. When r_α is close to r_s, the catch adequately mirrors the population within most of its sampling range. 6 By using these two new concepts, we will better understand why monitoring traps mirror the local population in some cases but not in others. This will help in designing more reliable monitoring programmes.     

Mating duration and Frequency in a Pine Sawfly

Newly emerged females of the European pine sawfly, Neodiprion sertifer Geoffroy (Hymenoptera: Diprionidae), were placed on pine twigs in the field and their behavior was recorded. Twenty to 25% of the females attracted wild males and mated on the day of release. One-third to one-half of these females remained and oviposited, while the rest disappeared from their twigs after mating. The mating frequency peaked at noon, and on average the mating lasted for 19 ± 13 (SD) min. Eleven to 30% of the females that remained on their twigs after mating remated, occasionally up to five times. Most released females disappeared from their twigs on the first day. Disappearances included both predation and dispersal. Only a few dispersals >5 m were recorded, because the sawflies were difficult to follow during flight. By color marking, dispersal up to 20 m was recorded.     

Different photosynthesis-nitrogen relations in deciduous hardwood and evergreen coniferous tree species

Rapidly induced responses can alter host plant suitability for insect growth and survival. The effects of defoliation on the suitability of potted 5-year-old red pine, Pinus resinosa Ait., for the sawfly Neodiprion sertifer (Hymenoptera: Diprionidae), were measured in two experiments. In the first, overall larval growth rate increased on seedlings within 8 days of low (<15%) defoliation. Suitability varied among larval age groups: defoliation increased the performance of older larvae, but not that of young larvae. In the second experiment, larval survival and weight varied non-linearly with defoliation intensity 8 weeks after treatment. Similar responses were observed following artificial and natural defoliation, and on early- and late-season seedlings. These results suggest that some evergreen conifers can respond rapidly to defoliation injury, and that herbivores may simultaneously acclimate to the induced response as they develop. The relative importance of induced response rates to plant-insect interactions is discussed.     

Host stress influences oviposition preference and performance of a pine sawfly

Abstract.

• 1 The effect of ponderosa pine (Pinus ponderosa Doug]. ex Laws) water stress, measured by xylem water potential, on oviposition preference and egg development of Neodiprion fulviceps was determined by two laboratory experiments in 1987 and 1988.
• 2 In the first experiment, adult females were allowed to choose between foliage from water-stressed, watered, and untreated control trees.
• 3 Significant differences were found between treatments in xylem water potential in 1987 but not in 1988.
• 4 Females chose water-stressed trees and a positive correlation was found between female preference and xylem water potential in 1987 but not in 1988.
• 5 In the second experiment, females were placed in globe cages with a single branch from each treatment and allowed to oviposit.
• 6 Significant differences were found among treatments in egg survival in 1987 but not in 1988.
• 7 These results indicate that sawflies can distinguish between stressed and non-stressed trees and that this behaviour is adaptive in that egg survival was greatest on the preferred treatment.

     

Loblolly pine grown under elevated CO2 affects early instar pine sawfly performance

Seedlings of loblolly pine Pinus taeda (L.), were grown in open-topped field chambers under three CO₂ regimes: ambient, 150 l l^–1 CO₂ above ambient, and 300 l l^–1 CO₂ above ambient. A fourth, non-chambered ambient treatment was included to assess chamber effects. Needles were used in 96 h feeding trials to determine the performance of young, second instar larvae of loblolly pine's principal leaf herbivore, red-headed pine sawfly, Neodiprion lecontei (Fitch). The relative consumption rate of larvae significantly increased on plants grown under elevated CO₂, and needles grown in the highest CO₂ regime were consumed 21% more rapidly than needles grown in ambient CO₂. Both the significant decline in leaf nitrogen content and the substantial increase in leaf starch content contributed to a significant increase in the starch:nitrogen ratio in plants grown in elevated CO₂. Insect consumption rate was negatively related to leaf nitrogen content and positively related to the starch:nitrogen ratio. Of the four volatile leaf monoterpenes measured, only -pinene exhibited a significant CO₂ effect and declined in plants grown in elevated CO₂. Although consumption changed, the relative growth rates of larvae were not different among CO₂ treatments. Despite lower nitrogen consumption rates by larvae feeding on the plants grown in elevated CO₂, nitrogen accumulation rates were the same for all treatments due to a significant increase in nitrogen utilization efficiency. The ability of this insect to respond at an early, potentially susceptible larval stage to poorer food quality and declining levels of a leaf monoterpene suggest that changes in needle quality within pines in future elevated-CO₂ atmospheres may not especially affect young insects and that tree-feeding sawflies may respond in a manner similar to herb-feeding lepidopterans.     

Ecosystem alteration modifies the relative strengths of bottom-up and top-down forces in a herbivore population

1. Ecosystem alterations can affect the abundance, distribution and diversity of plants and animals, and thus potentially change the relative strength of bottom-up (the plant resource) and top-down (natural enemies) trophic forces acting on herbivore populations. 2. The hypothesis that alterations of the forest ecosystem associated with precommercial thinning have contributed to the increased severity of outbreaks of Neodiprion abietis (Harris), a sawfly defoliator, through the reduction of trophic forces acting on N. abietis larvae, was tested using exclusion techniques. 3. The relative contributions to N. abietis larval mortality of bottom-up and top-down forces both increased with increasing levels of defoliation and were both reduced by thinning. The reduction of bottom-up and top-down forces caused a 58% mean increase in N. abietis larval survival in thinned compared with untreated stands, which is less than would be expected by the sum of the effects of thinning on each source of mortality. Evidence indicates that the partly compensatory, partly additive nature of the mortality associated with trophic forces in the system under study is responsible for this discrepancy. 4. To our knowledge, this is the first study to show the impact of ecosystem alterations on the balance between bottom-up and top-down forces acting on an eruptive herbivore population along a gradient of host-plant defoliation, and how this can lead to increased outbreak severity. It is stressed that accurate estimates of the relative contributions of bottom-up and top-down forces to mortality cannot be obtained if the additive or compensatory nature of the mortality associated with these trophic forces is overlooked.     

Genome-based phylogeny of dsDNA viruses by a novel alignment-free method

Sequence alignment is not directly applicable to whole genome phylogeny since several events such as rearrangements make full length alignments impossible. Here, a novel alignment-free method derived from the standpoint of information theory is proposed and used to construct the whole-genome phylogeny for a population of viruses from 13 viral families comprising 218 dsDNA viruses. The method is based on information correlation (IC) and partial information correlation (PIC). We observe that (i) the IC-PIC tree segregates the population into clades, the membership of each is remarkably consistent with biologist's systematics only with little exceptions; (ii) the IC-PIC tree reveals potential evolutionary relationships among some viral families; and (iii) the IC-PIC tree predicts the taxonomic positions of certain “unclassified” viruses. Our approach provides a new way for recovering the phylogeny of viruses, and has practical applications in developing alignment-free methods for sequence classification.     

Effects of associational resistance and host density on woodland insect herbivores

1. Specialist herbivores often become less abundant per unit of host tissue as host density increases (resource dilution). They usually become less abundant when non-host species are mixed with their host plants (associational resistance). Most studies of these trends have involved herbaceous host plants and have not examined both trends for the same herbivores. 2. Three hypotheses were tested for the response of insect specialists to host plant density: resource concentration, plant apparency and resource dilution. Two hypotheses were tested for the response of herbivores to non-host plants: associational resistance and plant apparency. 3. From 1992 to 2007, I examined the responses of three monophagous insect herbivores to the densities of their host, Pinus edulis, and of two non-hosts, Pinus ponderosa and Juniperus spp. 4. Herbivore loads increased with host density, though the correlations were weak and often variable between generations. These results were consistent with the resource concentration and plant apparency hypotheses, but not with resource dilution. 5. Herbivore loads decreased as non-host density increased, consistent with the associational resistance hypothesis. This and other studies have shown that associational resistance is important in many types of plant communities. 6. The absence of resource dilution on woodland trees contrasted with studies of herbaceous host plants. Further comparisons of woody and herbaceous host plants are needed to elucidate the reasons for this difference.