Influence of inter-tree variation in time of budburst of white spruce on herbivory and the behaviour and survivorship of Zeiraphera canadensis

Abstract.

• 1 Field studies were carried out during 4 years to determine the effects of inter-tree variability in the time of budburst of white spruce, Picea glauca Moench (Voss.), on the distribution, behaviour and survivorship of, and herbivory by, the spruce bud moth, Zeiraphera canadensis Mutt. & Free.
• 2 Differences between trees in herbivory, bud moth density and survivorship were inversely related to the mean date of budburst. The most and least susceptible half-sib‘families’evaluated were the first and last to flush, respectively. Differences in herbivory for trees in two families with similar phenologies were partially due to oviposition preference.
• 3 Oviposition and larval survivorship were highest on trees in the most susceptible families. Although egg hatch was more closely synchronized with budburst in warm than in cool springs, between tree differences in the time of budburst were consistent over years.

     

Genotype and environment interact to influence acceptability and suitability of white spruce for a specialist herbivore, Zeiraphera canadensis

Abstract.

• 1 The independent and interacting effects of plant genotype and site (i.e. environment) on the acceptability of white spruce, Picea gluaca (Moench) Voss, to the spruce bud moth, Zeiraphera canadensis Mut. & Free. and on plant suitability for egg development, were studied at four sites in New Brunswick, Canada.
• 2 A greater proportion of shoots on trees in two half-sib families, previously designated as highly susceptible, were partially eaten by spruce bud moths than shoots on trees in two half-sib families with low susceptibility.
• 3 At the site with the highest bud moth population, oviposition was highest on trees in susceptible families and on branches damaged by bud moth larvae. Oviposition was not higher on trees in susceptible families at the other three sites, resulting in a strong tree genotype × site interaction for oviposition.
• 4 Although there was a significant tree genotype × site interaction for egg predation, egg survival was higher on trees in susceptible families at all sites, due to lower levels of egg parasitism and predation.
• 5 Egg densities were positively but weakly correlated to shoot length and diameter. There were no consistent relationships between shoot length, shoot diameter, needle length or needle density and per cent egg survival, parasitism or predation.
• 6 Because egg survival was higher on trees in susceptible families at all sites but egg densities were only higher on trees in susceptible families at one site, host plant acceptability and suitability were positively related at only one site. We speculate that Z.canadensis does not distinguish between hosts of different suitability until trees are heavily damaged.

     

Preformed and neoformed extension of shoots and sylleptic branching in relation to shoot length in Tsuga canadensis

Summary Shoot systems developed over 3 successive years were investigated on 55 understorey Tsuga canadensis (L.) Carr. trees. Paired comparisons of preformed-leaf content of terminal buds and numbers of leaves produced on new shoots showed that neoformed leaves were produced in large numbers. Parent-shoot character was not useful in predicting numbers of preformed leaves, was better related to total leaves produced, but left the majority of the variation unexplained. This reflected the capacity of any terminal bud to produce a shoot with more or less neoformation, depending on conditions for growth. All shoots over 6 cm long produced sylleptic shoots that bore from two to many leaves and were arranged in a mesitonic pattern along the parent. Some of the longer sylleptic shoots produced lateral buds or second-order sylleptic shoots. Monopodial second-year extensions of sylleptic-shoot axes followed an acrotonic pattern, as did proleptic shoots from the few lateral buds borne on the parent shoots. Such lateral buds were more frequent on shorter parent shoots: they typically occurred near the proximal and distal ends. Duration of shoot extension was positively correlated with shoot length: terminal buds became evident as shoot extension neared cessation.     

Herbivory modifies conifer phenology: induced amelioration by a specialist folivore

Herbivory by Zeiraphera canadensis Mut. & Free. (Lepidoptera: Tortricidae), an early season folivore of white spruce [ Picea glauca(Moench) Voss], has been associated with a shift in the timing of bud burst by its host during the subsequent year. We tested the hypothesis that a herbivory-induced shift in the phenology of bud development improves the window for colonisation of white spruce buds by Z. canadensis. Feeding on cortical tissue of elongating shoots caused the destruction of apical buds and an interruption of apical dominance in the year following herbivory. White spruce compensated for damage with the activation of dormant buds; mainly at proximal positions along shoots. As a result, half of all active buds on previously damaged branches were located immediately adjacent egg sites (i.e. previous year's bud scales), whereas <10% of active buds on intact shoots were situated there. More than 40% of newly emerged larvae colonised the basal buds of damaged shoots versus just 10% for intact shoots. Previous herbivory also influenced the initiation of bud burst. All buds flushed 2 days earlier on damaged shoots and date of bud burst was inversely correlated to bud density, indicating that short damaged shoots with large numbers of buds were stronger sinks for nutrients required for bud development. Egg hatch was best synchronized with early bursting buds on damaged branches. As a consequence, 89% of first-instar larvae successfully colonised buds on damaged branches while only 55% were successful on undamaged branches. Improved survival of larvae in the year following herbivory was a direct result of the evolved response by white spruce to the interruption of apical dominance. The pattern of herbivory by Z. canadensis may have evolved as a strategy to enhance the quality of white spruce for their offspring.     

Variation in the ability of larvae of phytophagous insects to develop on evolutionarily unfamiliar plants: a study with gypsy moth Lymantria dispar and Eucalyptus

• 1 By examining variation in the abilities of polyphagous insects to develop on host plants with secondary metabolites that they have never encountered previously, we may be able to gain some insights into the nature of evolution of biochemical mechanisms to process plant secondary metabolites by phytophagous insects.
• 2 The present study aimed to examine variation in the ability of gypsy moth larvae Lymantria dispar (Lymantriidae) to complete development on different species of the plant genus Eucalyptus (Myrtaceae). Leaves of at least some Eucalyptus species contain formylated phloroglucinol derivatives. These are secondary metabolites that are evolutionarily unfamiliar to the gypsy moth.
• 3 Larvae of gypsy moth showed extremely variable responses in larval performance between Eucalyptus species, between individual trees within host plant species, between moth populations, and between individuals within moth populations.
• 4 Larval survivorship was in the range 0–94%, depending on the host. Failure of at least some larvae to complete development on some Eucalyptus species indicates that gypsy moth larvae have a limited ability to process secondary metabolites in eucalypt leaves.
• 5 At least some individuals, however, appear to already possess biochemical mechanisms that process the secondary metabolites in leaves of Eucalyptus species, and therefore the abilities of larvae to complete development on phylogenetically and chemically unfamiliar hosts are already present before the gypsy moth encounters these potential hosts.

     

Predicted changes in the synchrony of larval emergence and budburst under climatic warming

Summary The impact of climatic warming on the synchrony of insect and plant phenologies was modelled in the case of winter moth (Operophtera brumata) and Sitka spruce (Picea sitchensis) in the Scottish uplands. The emergence of winter moth larvae was predicted with a thermal time requirement model and the budburst of Sitka spruce was predicted from a previously published model (Cannell and Smith 1983) based on winter chilling and thermal time. The date of emergence of winter moth larvae was predicted to occur earlier under climatic warming but the date of budburst of Sitka spruce was not greatly changed, resulting in decreased synchrony between larval emergence and budburst. The general question of how a change of climate might affect phenological synchrony and insect abundance is discussed.     

Outbreaks of the winter moth on Sitka Spruce in Scotland are not influenced by nutrient deficiencies of trees,tree budburst,or pupal predation

Summary Since the early 1980s, the winter moth, Operophtera brumata L. (Lepidoptera: Geometridae) has emerged as a serious pest of Sitka Spruce, Picea sitchensis Bong. plantations in southern Scotland. Outbreaks are characterised by susceptible sites within plantations which can occur immediately adjacent to resistant sites. We investigated the level of some nutrients in the trees, the date of budburst of the trees, and the numbers of some potential predators of winter moth pupae. None could satisfactorily explain outbreak patterns. Although foliage analysis demonstrated that many trees were marginal or deficient in phosphorus, nitrogen and potassium, these deficiencies were not related to the susceptibility of a site. Within sites, the numbers and weights of O. brumata were positively related to phosphorus content and negatively related to calcium content of foliage. Other evidence suggests, however, that these correlations may not represent direct effects of phosphorus and calcium on larval growth and survival. Date of budburst, which commonly determines susceptibility of deciduous hosts to O. brumata, was unrelated to density, and pupal predators were more, not less, abundant in susceptible sites. Although it is difficult to distinguish between factors that initiate outbreaks and those that maintain them, these data suggest that nutrient deficiencies of trees, budburst date, and the distribution of pupal predators of the winter moth cannot explain patterns of outbreak of the winter moth on spruce.     

Genetic differentiation as a result of adaptation to the phenologies of individual host trees in the galling aphid Kaltenbachiella japonica

Abstract.

• 1 The adaptation of the non-migratory galling aphid Kaltenbachiella japonica (Matsumura) to the budburst phenologies of individual host trees (Ulmus davidiana var. japonica Nakai) was investigated. There was a large variation in budburst time between individual host trees. We tested a hypothesis that aphid populations on respective host trees are genetically differentiated through adaptation to the budburst phenologies of host trees.
• 2 There was significant correlation between the budburst time of eight host trees in two successive years.
• 3 Tree-associated populations differed significantly in egg-hatching time, and the mean hatching time was significantly correlated with the mean budburst time of respective host trees.
• 4 Mating was made between male and female sexuales of the same clone (i.e. self-mating) to obtain selfed eggs. Hatching patterns of selfed eggs showed that there was a substantial amount of genetic variance in hatching time between clones (galls). Nested ANOVA demonstrated that the between-tree component contributed more to the phenotypic variance than the within-tree or within-gall components.
• 5 Reciprocal crosses between males and females of different clones furthermore demonstrated significant differences in hatching time between cross combinations, but no significant differences between the two reciprocals within combinations. This result suggests that there are no maternal effects for the timing of egg hatch and that the differences between selfed lines are attributable to genetic variance.
• 6 The fine-scale adaptation hypothesis was supported by egg-hatching experiments, which further suggest that budburst phenology varying between individual trees is acting as intense selective pressure on the egg-hatching time of associated insects.

     

Within-crown distribution of male and female cones on clonal Sitka spruce trees

The within-crown distribution of cones (strobili) was mapped on 48 mature grafts of Sitka spruce [Picea schensis (Bong.) Carr.]. The total number of cones per tree was increased by a mainstem injection of 20 mg GA_4/7 and 8 mm wide bark ring. The cones and buds were classified as either lateral or terminal on each individual branch. The distribution of lateral female, terminal female, lateral male and terminal male cones, showed a general progression from the upper distal to lower proximal regions of the tree crown in all 12 clones. Evidence for preferential allocation of cones to particular branch types is presented. The region of the tree with the greates number of cones varied with treatment and reflected differences in the sex ratio of the individual trees. There is evidence for a relationship between branch length, position within the crown and the type of cone produced.     

Microgeographic variation in recruitment under adult trees: arrival of new genotypes or perpetuation of the existing ones?

• Investigating spatial variation in the relative importance of sexual reproduction and clonal propagation is critical to obtain more accurate estimates of future effective population sizes and genetic diversity, as well as to identify ecological correlates of clonality.
• We combined a stratified sampling scheme with microsatellite genetic analyses to estimate variation in the proportion of sexual versus clonal recruits among saplings in five populations of the tree Pyrus bourgaeana. Using a likelihood framework, we identified clones among the genotypes analysed and examined variation among populations regarding the proportion of saplings coming from clonal propagation. We also examined the relationship between the relative abundance of clonal shoots across the studied populations and their herbivory levels.
• Our results revealed that one third of the saplings examined (N = 225 saplings) had a probability above 0.9 of being clones of nearby (<10 m) trees, with the ratio between clonal propagation and sexual recruitment varying up to eight‐fold among populations. A small portion of these putative clonal shoots reached sexual maturity. Relative abundance of clonal shoots did not significantly relate to the herbivory by ungulates.
• Our results call into question optimistic expectations of previous studies reporting sufficient levels of recruitment under parental trees without animal seed dispersal services. Nevertheless, given that some of these clonal shoots reach sexual maturity, clonal propagation can ultimately facilitate the long‐term persistence of populations during adverse periods (e.g. environmental stress, impoverished pollinator communities, seed dispersal limitation).

     

Variation in budburst phenology of Douglas-fir related to western spruce budworm (Lepidoptera: Tortricidae) fitness

Variation in budburst phenology among individual trees of interior Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) may influence their susceptibility to western spruce budworm (Choristoneura occidentalis Freeman) defoliation. We tested the hypothesis that phenological asynchrony between Douglas-fir and the western spruce budworm is a mechanism of resistance using clones derived from parent trees that showed resistance versus susceptibility to C. occidentalis defoliation in the field. Susceptible clones had earlier budburst phenology compared with resistant clones when they were grown in a common greenhouse environment, demonstrating a genetic basis for parallel phenological differences exhibited by the parent trees. We tested the importance of phenological asynchrony as a factor influencing fitness of C. occidentalis using two different greenhouse bioassay experiments. One experiment compared western spruce budworm performance on equivalent phenological stages of susceptible and resistant clones by matching larval feeding to the columnar (fourth) bud development stage of each clone. Larvae reared on resistant clones had greater realized fitness (i.e., number of F1 offspring produced) than those reared on susceptible clones when the influence of variation in budburst phenology was minimized. In the other experiment, western spruce budworm larvae were placed on all trees on the same date when approximately 50% of all terminal buds in the population were in the yellow (second) budburst stage. Larvae reared on susceptible clones had greater realized fitness than those reared on resistant clones when the influence of phenological asynchrony was expressed. Our results suggest that resistant phenotypes of Douglas-fir have negative effects on survival and reproduction of C. occidentalis under the natural conditions that insects and trees experience in the field. Genetic variation among trees in budburst phenology has an important influence on interactions between the western spruce budworm and Douglas-fir.     

Effects of grafting on host plant resistance in ash (Fraxinus spp.) to emerald ash borer (Agrilus planipennis Fairmaire)

相似文献   

Intra-tree variation in foliage quality drives the adaptive sex-biased foraging behaviors of a specialist herbivore

Some herbivorous insects enhance their fitness using foraging strategies that allow them to find and colonize the best of available resources within heterogeneous plants. The yellowheaded spruce sawfly, Pikonema alaskensis (Roh.) (Hymenoptera: Tenthredinidae), is a common defoliator that oviposits and feeds on the developing foliage of young open-grown black spruce [Picea mariana (Mills) B.S.P.]. While female and male eggs are both laid throughout the crown, most eggs laid in the upper crown are female, and more female than male late-instar larvae disperse acropetally, from the lower and mid-crown to the upper crown, to complete juvenile development. Here we present results from 4 years of manipulative sleeve-cage experiments that were carried out to evaluate the hypothesis that sex-biased oviposition-site selection and acropetal dispersal by P. alaskensis are adaptive responses to intra-tree variation in foliage quality. Survival and proportion of survivors that were female were either the same (2 years) or significantly higher (2 years) for groups placed in the lower or mid-crown during early instars, and then transferred acropetally during late instars to complete development in the upper crown, compared with those forced to feed exclusively in the lower crown. This suggests that females benefited most from acropetal dispersal. Sex ratios of survivors that had been forced to develop exclusively in the upper crown were usually more female biased than those of survivors that developed exclusively in the lower crown, suggesting higher survival for female than for male larvae emerging from eggs laid in the upper crown. Sex-biased egg allocation and larval dispersal appear to increase the survival of P. alaskensis by accounting for differential effects on male versus female larvae of phenology-independent temporal and spatial variations in the quality of foliage within the heterogeneous crown of black spruce.     

Patterns of on-shoot positioning of seed cones in relation to shoot length and position in the crowns of young Picea mariana

Summary Patterns of distribution of seed cones along the length and around the circumference of shoots were determined for young black spruce [Picea mariana (Mill.) B.S.P.] trees aged 9–17 years. Seed cone production averaged 5, 30, 62, 95, and 112 per tree of ages 9, 11, 13, 15, and 17 years, respectively. Cone distribution was restricted to distal and medial 1-year-old branches at age 9 and spread to distal 3-year-old branches by age 17. Sequentially, shoots of 0–25, 26–100, 101–250, 251–325, and 326–375 mm bore more cones in terminal, distal, medial, proximal, and basal positions along shoots, respectively. The uppermost and longest distal 1-year-old shoots bore most of their cones proximally and medially. On the shortest and innermost proximal order 2 shoots, cones were often terminal. The percentages of cones borne on upper, lower, and each of the two side surfaces of shoots were 31, 22, and 21, respectively, and 5% of the cones were terminal. Cones on lower surfaces were most common on the longest shoots near the apex of trees. Cones on upper surfaces became prevalent downward and outward in the crown. On-shoot positioning therefore varied in a predictable manner with shoot length, and thus with pattern of shoot-length distribution in the crown.     

The phenological window for western spruce budworm: seasonal decline in resource quality

• 1 Western spruce budworm Choristoneura occidentalis Free. (Lepidoptera: Tortricidae) emerge in the spring before budburst and then face a rapidly deteriorating host quality each season.
• 2 Measures of fitness, survival and fecundity, were made on cohorts of final‐instar spruce budworms deployed on host trees at several times during the season in four field locations in coastal and interior British Columbia, Canada.
• 3 Survival and fecundity were strongly correlated throughout the season and varied as much as four‐fold from maxima at mid‐season to minima at the end of the season.
• 4 Fitness values overall were greatest in the coastal compared with interior locations. Among interior locations, fitness was greatest at the highest elevation and least at the lowest elevation. Both cohort and sample‐based estimates of survival of wild, final‐instar budworms were relatively high in these outbreak populations.
• 5 The influence of the phenological window and degree of synchrony with the host plant on herbivore abundance often depends on other processes affecting population rates of change.

     

Effects of herbivory and genotype on growth and survivorship of sand-dune willow (Salix cordata)

Abstract.

• 1 The response of different clones of sand-dune willow, Salix cordata, to herbivory by a specialist herbivore, Altica subplicata, was studied in three glasshouse experiments. Plants were caged and exposed to three herbivory treatments: no beetles, low number of beetles, and high number of beetles.
• 2 Plants consistently had significantly higher growth rates in the absence of herbivory than under conditions of low or high herbivory (1.5–6 times higher).
• 3 Herbivore treatment influenced mortality from drought stress; more plants from the low and high herbivory treatments (40% and 80%) died from drought stress than did control plants (0%).
• 4 Clone genotype significantly influenced growth rates and the susceptibility of plants to drought stress. However, clones showed similar growth responses to herbivory, suggesting a lack of genetic variation in tolerance or resistance to herbivory.

     

The Effect of Explant Type on the Establishment of Sitka Spruce [Picea sitchensis (Bong.) Carr.] in Culture

Five different types of shoot tip explant from three-year-oldand 13-year-old Sitka spruce trees were established on a rangeof media and shoot growth was recorded. While Schenk and Hildebrandtmedium gave the greatest shoot elongation, Webb and Street mediumproduced the healthiest-looking shoots. Poorest growth was recordedfor the explants cultured without the crown or any vasculartissues from the previous year's growth. [¹⁴C]sucrose tracerstudies indicated that poor uptake was not responsible for thereduced growth and it was suggested that damage due to inadequatewater uptake may be responsible. Results demonstrated that peelingthe protective scales from resting buds and trimming the shoottissues below the crown to a wedge shape and excluding the barkwas a reliable method of explant preparation for establishingmature and juvenile Sitka spruce in culture. Picea sitchensis, Sitka spruce, mature, juvenile, crown, [¹⁴C]sucrose, micropropagation     

Plant module size influences the intra‐tree distribution and abundance of a shoot‐boring sawfly in young balsam fir

Field surveys were carried out to assess the effects of intra‐tree variation in developing shoot length within and among crown levels on the density and abundance of the balsam shoot‐boring sawfly, Pleroneura brunneicornis Rohwer (Hymenoptera: Xyelidae), in young balsam fir, Abies balsamea (L.) Mill. (Pinaceae). Overall, cardinal direction had no influence on shoot‐borer density or abundance; however, the highest percentage and abundance of bored shoots occurred on intermediate‐sized shoots within the crown (i.e., in the mid‐crown and on the distal‐lateral and medial‐lateral shoots). Comparatively, few shoot borers occurred in the upper or lower crown levels, or on the relatively large terminal shoots within branches. This distribution appears indicative of the higher suitability of intermediate‐sized shoots within hosts for either egg lay or larval performance. Results of this study are most consistent with predictions of the ‘optimal module size’ hypothesis, which posits that herbivore responses to plant module size should reflect the balance of tradeoffs between utilizing relatively large, nutritious shoots vs. small, more easily exploited shoots.     

Foraging on buds and bark of mulberry trees by Japanese monkeys and their range utilization

The Japanese monkey (Macaca fuscata) winter range utilization and the effects of foraging on mulberry trees (Morus bombycis) were studied in the Shimokita Peninsula during four winter seasons. The monkeys ate mainly winter dormancy buds when they visited the mulberry tree clumps for the first time within the winter, but they ate mainly bark when they visited for the second or third times. In the areas utilized by the monkeys over the recent three years, the mulberry trees compensated for the decrease in their number of shoots by producing longer shoots with more buds against the monkey foraging. In the areas used every year for more than four years, however, the mulberry trees were unable to compensate for the foraging pressure. Thus, although the monkeys had apparently operated prudent herbivory within three years, they did not do so on a longer time-scale. They shifted their utilization ranges after having over-exploited the mulberry trees.     

Intraspecific variation in food plant favourability to phytophagous insects: psyllids on Eucalyptus blakelyi M.

Abstract.

• 1 A combination of variables reflecting leaf quality provided the best predictive model explaining differences in population trend of Glycaspis spp. among individuals of the food plant Eucalyptus blakelyi, measured under field conditions on an egg to egg basis. The equation incorporated the product of: (1) proportion of foliage in each age class; (2) probability of foliage escape from herbivory; (3) preference for the foliage class measured as the proportion of eggs oviposited on it by the female; these variables being summed over the young and mature age classes.
• 2 In addition, significant and consistent differences in insect survivorship among trees were demonstrated by transfer experiments. These experiments established that young foliage was preferable to mature foliage for first instar survival. Differences in insect survivorship also showed a consistent pattern among the study trees. The most probable explanation for this consistent pattern was the influence of the trees on the insects, particularly physiological or biochemical differences among trees. It was not, however, possible to identify the cause of the differences.