Effects of previous defoliation on pine looper larval performance

1 During outbreaks of the pine looper, Bupalus piniarius, its host, Pinus sylvestris, is severely defoliated. The larvae of this geometrid normally feed on mature needles. However, because trees are totally defoliated during outbreaks, the next generation is forced to feed on current-year needles. 2 Bupalus piniarius larvae feeding on previously defoliated trees may show lower performance either because of induced resistance or because larvae have to feed on needles not normally fed upon (current instead of mature). 3 These hypotheses were tested in an experiment where larvae were reared on (i) shoots naturally defoliated the previous year, and thus, bearing only current-year needles, (ii) non-defoliated shoots where larvae had access only to current-year needles, and (iii) control shoots with access to both current and mature needles. 4 There was no support for the induction hypothesis. Survival was lower on naturally defoliated shoots than on control shoots (81.3 vs. 90.9%), but survival was lower also on non-defoliated shoots where larvae had access only to current-year needles (78.8%). Data on larval feeding distribution showed a strong preference for mature needles. 5 Needle nitrogen concentration of current-year needles was 38% higher on defoliated trees than on non-defoliated trees. 6 It is concluded that defoliation affected larval performance primarily through the removal of the preferred type of needles and not because of an induced resistance. Effects of increased concentrations of allelochemicals in defoliated shoots, if present, were probably cancelled out by increased nitrogen concentrations.     

Interaction between individuals during the colonisation of Douglas fir needles by sistentes of Adelges cooleyi

Summary A laboratory study of Adelges cooleyi sistentes emerging from diapause shows that the sistentes redistribute themselves over the needles in the form of aggregations. At low densities needle bases are preferred to the tips, this preference becoming less marked as sistentes density increases. Within each needle aggregation a certain amount of spacing out occurs which minimises the number of sistentes settling in close proximity. Over a given period of time adult sistentes produce more eggs at the base of the needles than at the tips. The settling behaviour ensures that at low densities the most favourable parts of the needle at the base are utilised first while the spacing out factor provides sufficient space for the development of the egg masses and minimises the possibility of over-exploitation of one part of the needle in relation to the remainder.     

Factors determining the distribution of the green spruce aphid, Elatobium abietinum, on young and mature needles of spruce

1. The green spruce aphid Elatobium abietinum , was less active and more likely to feed when on mature rather than young needles of Sitka spruce.
2. Compared with survival on mature needles, survival of aphids on young needles was significantly reduced, especially on newly flushed needles, but became less so as the needles matured. The survival of aphids kept close to, but not in contact with, young needles was similarly reduced.
3. On young needles stripped of epicuticular wax, aphids were less active and fed and survived for longer than aphids on untreated young needles. In contrast, the supply of additional amino acids to shoots bearing young needles did not significantly improve the survival of aphids on these needles.
4. In a choice test, aphids were able to discriminate between epicuticular wax from young and mature needles. The reciprocal transfer of wax between young and mature needles reversed the aphids' marked preference for mature needles.
5. These results indicate that the aphid is deterred from settling and feeding on young needles by a volatile substance present in the epicuticular wax.     

Performance of pine looper Bupalus piniarius larvae under population build-up conditions

In the first year of an outbreak, Bupalus piniarius larvae, encounter intense crowding. In the later stages of larval development, they are forced to feed on the non-preferred current-year needles of Scots pine or even on alternative hosts. It was hypothesized that larval feeding on a non-preferred resource (current-year needles) will negatively affect B. piniarius performance. It was also hypothesized that larval mutual interference (crowding without competition for food) will have negative additive effects. These hypotheses were tested in laboratory and field experiments. Fourth instar larvae were reared singly and under crowded conditions in cohorts of ten. Larvae in both situations were reared on control branches (containing both mature and current-year shoots) and branches containing only current-year shoots. Crowded larvae were reared also on Norway spruce, an alternative host. Crowding and feeding on a non-preferred resource had opposite effects on B. piniarius larval performance. Crowding in the late larval instars enhanced larval performance while absence of the preferred resource had negative effects. Larval growth rate was higher and development time was shorter for larvae exposed to crowded conditions than for solitary larvae. There was, however, no difference between the groups in final pupal weights or survival. Survival was 25% lower for larvae feeding on non-preferred current-year needles and pupal weights 9% lower, compared with results for larvae feeding on mature needles. Larvae feeding on Norway spruce suffered greatly extended development time, 82% lower survival, and resulted in 60% lighter pupae compared with conspecifics on Scots pine. It was concluded that not only quantity but also quality of the available food resource is critical for B. piniarius development.     

Morph determination in Melittobia, a eulophid wasp

Abstract. 1. The Caribbean and Central American eulophid wasp, Melittobia sp. 1 ( hawaiiensis complex), (Hymenoptera: Chalcidoidea), whose larvae are exophagous on the prepupae of solitary wasps and bees, has three adult female and two adult male morphs.
2. The males, which constituted only 5% of the progeny, had morphs with, and without, ocelli.
3. In the females, brachypterous, negatively phototactic 'crawlers' developed mainly at low densities (<8.9 eggs cm^-2), while above this density an increasing number and percentage of macropterous, positively phototactic 'fliers' were produced. Intermediate 'jumpers' were positively phototactic but could not fly.
4. In a typical progeny twenty to thirty 'crawlers', twenty to forty 'jumpers' and up to 400 'fliers' emerged in an overlapping sequence.
5. Experiments demonstrated that the proportions of the three morphs emerging were unrelated to the sequence in which the eggs were laid, to the species of the host and to the quality and absolute amount of food given, but related only to the density experienced by the young larvae.
6. We relate this trimorphism of dispersive female morphs to the three levels of ecological space (food item, patch and distribution of patches) postulated by Hassell & Southwood (1978) and suggest the general ecological conditions necessary for it to evolve.     

Endophytic fungi decrease available resources for the aphid Rhopalosiphum padi and impair their ability to induce defences against predators

Abstract.  1. The production of winged morphs is a well known mechanism of induced defence in aphids to escape from natural enemies, and is also a reaction to poor resource quality.
2. Host plants of aphids often associate with endophytic fungi that have been shown to reduce the fitness of some species of aphids.
3. It was hypothesised that endophyte infection of host plants that represent a low quality plant resource should increase the aphid's induced response to a predator because both low plant quality and predator presence represent a stronger cue for wing production than predator presence alone.
4. In a laboratory experiment, bird cherry-oat aphids Rhopalosiphum padi L. were exposed to the factors predator threat and endophyte infection and the effects of these factors on the proportion of winged morphs produced by the aphid colonies was analysed.
5. The presence of endophytic fungi strongly decreased aphid colony sizes. When a predator threat was present, all colonies on endophyte-free grasses produced winged morphs whereas only a few colonies were able to produce winged morphs on endophyte-infected grasses. However, these few colonies produced larger proportions of winged morphs than colonies on endophyte-free grasses. Without a predator threat, no colonies on endophyte-infected grasses produced any winged morphs.
6. These results show that aphids in stressed conditions and with reduced fitness will only invest in inducible defences when predators are present but are unable to produce winged morphs in response to endophyte presence.     

Adaptive loss of color polymorphism and character displacements in sympatric <Emphasis Type="Italic">Mnais</Emphasis> damselflies

A geographical survey of two Mnais damselfly species in the Kinki area of Japan showed evidence for character displacements when the two species were found in sympatry. Mnais costalis, a species that has polymorphic male mating types of orange-winged territorial and clear-winged non-territorial morphs (hereafter abbreviated to orange and clear morphs respectively) in allopatry often shifted to having monomorphic orange morphs in sympatry. The mean body size of orange morphs was consistently larger than that of clear morph in allopatry. The mean body size of the sympatric orange morphs was even larger than that of allopatric orange morphs. By contrast, Mnais pruinosa, a species that also has two morphs of large orange and small clear morphs in allopatry, shifted to having monomorphic clear morphs in sympatry. The mean body size of the sympatric clear morphs was smaller than that of allopatric clear morphs. Divergence was also detected in the preference for habitat insolation conditions between sympatric Mnais damselflies. Both species in allopatric regions prefer half-light forest habitats, while in sympatric regions they showed diversified habitat preference: M. costalis preferred sunny habitats while M. pruinosa preferred shady habitats. Multiple character displacements in signal traits and habitat preference emerged in heterogeneous forest light environments are likely to have synergistic effects on the reproductive isolation of the two species.     

Nitrogen distribution in young Norway spruce (Picea abies) trees as affected by pedospheric nitrogen supply

In numerous locations in Europe spruce trees are exposed to high loads of nitrogen. The present study was performed to characterize the distribution of nitrogen compounds under these conditions. For this purpose Norway spruce ( Picea abies [L.] Karst.) trees were cultivated under close-to-natural conditions of a forest understory in soil from an apparently nitrogen-limited field site in the Black Forest either with, or without supplementation of nitrogen as ammonium nitrate. After 11 and 20 months, growth, total nitrogen contents of the biomass, and total soluble non-proteinogenic nitrogen compounds (TSNN, i.e. nitrate, ammonium, soluble proteinogenic and non-proteinogenic amino compounds) in needles, xylem sap and phloem exudate were analysed. After 20 months of growth, N-fertilization had slightly enhanced the biomass of current-, but not of 1-year-old shoots. At both harvests, total N-content of 1-year-old needles was increased by N-fertilization, whereas current-year needles were not significantly affected. By contrast, TSNN was elevated by N-fertilization in both current-year and 1-year-old needles. The increase in TSNN was mainly attributed to an accumulation of arginine. Xylem sap analysis showed that the increase in TSNN of the needles was a consequence of enhanced nitrogen assimilation of the roots rather than the shoot. Since also TSNN in phloem exudates was enhanced, it appears that N-fertilization elevates the cycling pool of amino compounds in young Norway spruce trees. However, this pool seems to be subject to metabolic interconversion, since mainly glutamine and aspartate are transported in the xylem from the roots to the shoot, but arginine accumulated in the needles and the phloem.     

“Natural” and NH3-induced variation in epicuticular needle wax morphology of Pseudotsuga menziesii (Mirb.) Franco

Summary Changes in epicuticular wax morphology of Pseudotsuga menziesii needles were studied with scanning electron microscopy throughout the growing season in current-year and older needles in 20 trees from two sites, Kootwijk and Garderen (The Netherlands). Fusion of crystalline wax rods leading to a reticulate structure and ultimately to degradation of the regular three-dimensional porous structure started several weeks after bud break and reached a high level at the end of the first growing season. The increase in amorphous (solid) wax showed a similar, but slightly slower development. In 1- and 2-year-old needles the degradation of crystalline wax and increase in amorphous wax had progressed only slightly further. The rates of change in crystalline wax morphology were very similar for sun-exposed and shaded needles, for the two sites, and for the two tree vitality classes included in the samples. The development of epiphyllous fungi also appeared to be largely needle age dependent. The effects of NH₃ on needle wax morphology were studied in young trees used in fumigation experiments. Short-term fumigation (approx. 5 weeks) did not affect wax morphology in current-year needles, but 1-year-old needles which had been exposed to different concentrations from bud break onwards showed a severe degradation of the crystalline wax, regardless of the NH₃ concentration used. In 2-yearold needles the effect of NH₃ could not be traced and was overshadowed by the natural ageing process. Ambient O₃, SO₂ and NO_x levels did not effect epicuticular wax morphology. It is suggested that the variation recorded for the two forest stands does not show effects of local pollution levels.     

Compensatory photosynthesis as a response to partial debudding in ezo spruce, <Emphasis Type="Italic">Picea jezoensis</Emphasis> seedlings

The effects of partial debudding on photosynthesis, stomatal conductance and nitrogen contents of 1-year-old needles and newly grown needles were studied in Picea jezoensis (Sieb. et Zucc.). Seventy-five percent of the buds of P.jezoensis seedlings were removed soon after bud break. Gas exchange was measured three times for 1-year-old needles and twice for current-year needles. The photosynthetic rates of 1-year-old needles were greater in debudded seedlings than in control seedlings, and the difference increased as the growing season progressed. This greater photosynthetic rate in debudded seedlings was accompanied by greater stomatal conductance. However, neither the photosynthetic rates nor the stomatal conductance of current-year needles differed between debudded and control seedlings after the needles had fully expanded. Debudding also had no effect on mass-based nitrogen contents in either the 1-year-old or the current-year needles. Area-based nitrogen in the 1-year-old needles did not differ between debudded and control seedlings, but was greater in debudded seedlings than in control seedlings in current-year needles. These results suggest that the enhanced photosynthetic rate is more likely a result of an increased root/leaf ratio that reduces the stomatal limitation of photosynthetic rate than a result of altered sink-source relationships or increased leaf nitrogen content.     

Adaptive significance of facultative paedomorphosis in Triturus alpestris (Amphibia, Caudata): resource partitioning in an alpine lake

1. Facultative paedomorphosis is a polymorphism that has important evolutionary implications in promoting morphological differentiation and variation in habitat use. It occurs in several urodele species throughout the world. Several hypotheses based on life-history theory have been proposed to explain the wide range of habitats in which facultative paedomorphosis occurs, suggesting multifactorial causes.
2. In harsh habitats, such as mountain lakes, paedomorphosis might promote niche partitioning between morphs. This hypothesis was tested by comparing micro-habitat use and diet of two coexisting morphs in an alpine lake.
3. Paedomorphs occupied all microhabitats in the lake while metamorphs occurred only along the shoreline or at the water surface. Paedomorphic newts were frequent in deep water, where they foraged mainly on plankton. Plankton was poorly exploited by metamorphic newts, which fed mainly on terrestrial insects. Difference between morphs in microhabitat use, and consequently in the diet, were consistent in both sexes and in juveniles.
4. In adults, the mass and energy value of stomach contents did not differ between morphs, suggesting a similar food availability in the habitats occupied.
5. This study indicates habitat partitioning between morphs involving dietary differences. Specific benefits and costs of each ontogenetic pathway may allow their coexistence in this deep and fishless lake. Paedomorphosis benefits individual newts by making new food resources available and presumably reducing competition at the shore of the lake. However, the proximate causes of such an ontogenetic switch remain unclear.     

不同密度下沙地樟子松碳、氮、磷化学计量及养分重吸收特征

以樟子松纯林为对象,研究了6种密度(490、750、1110、1550、1930、2560株.hm-2)下不同器官(当年生叶、一年生叶、当年生枝、一年生枝和细根)的C、N、P化学计量特征及叶片N、P重吸收效率.结果表明:随着林分密度增加,当年生和一年生叶C含量及当年生和一年生枝P含量呈降低趋势(1550株.hm-2除外...     

Effect of defoliation on tracheid production and the level of indole-3-acetic acid in Abies balsamea shoots

To simulate feeding by the spruce budworm ( Choristoneura fumiferana Clem.), the apical current-year shoots on 1-year-old branches in the uppermost whorl of 6-year-old balsam fir [ Abies balsamea (L.) Mill.] trees were either removed completely by debudding before the start of the growing season or defoliated 0, 50, 90 or 100% shortly after budbreak. Debudded branches were treated at the apical end with 0, 0.1 or 1.0 mg of indole-3-acetic acid (IAA) (g lanolin)⁻¹. Ninety % of the 1-year-old needles were also removed from some of the experimental branches. After ca 4 weeks of growth, the radial width of new xylem and the level of IAA were determined in the 1-year-old internode. The IAA content was measured by radioimmunoassay.
The removal or defoliation of current-year shoots inhibited tracheid production and decreased the IAA level. Exogenous IAA stimulated tracheid production and increased the IAA level in debudded branches. Current-year shoot defoliation also inhibited current-year shoot elongation. The inhibitory effect of current-year needle removal on all parameters generally increased with increasing intensity of defoliation. The removal of 1-year-old needles did not affect the IAA level or current-year shoot elongation, nor did it influence tracheid production in branches with current-year shoots. However, removal of 1-year-old needles inhibited tracheid production in debudded branches supplied with exogenous IAA. The results indicate that (1) IAA is involved in the control of tracheid production in the 1-year-old internode, (2) IAA is supplied primarily by current-year shoots, and (3) defoliation by the spruce budworm inhibits tracheid production partly by decreasing the supply of IAA.     

An association between two species of pine aphid, Schizolachnus pineti and Eulachnus agilis

Abstract. 1. Two aphid species feed exclusively on pine needles, Schizolachnus pineti (Fabr.) occupying only the outer (curved) surface with Eulachnus agilis (Kalt.) showing a preference for the inner (flat) surface.
2. Despite E.agilis favouring and S.pineti avoiding senescing needles, both species show a significant tendency to share shoots and needles in the field.
3. The same association is also apparent under laboratory conditions, although E.agilis is randomly distributed between needles in the absence of S.pineti.
4. E.agilis was found to benefit through increased survival and faster growth rates, by feeding on the same shoots and needles as S.pineti.
5. It is suggested that E.agilis may be able to detect and exploit an improvement in the nutritive quality of needles brought about by the feeding activity of S.pineti colonies.     

Carbohydrate Accumulation in Siberian Spruce Needles of Various Ages

The content of mono- and oligosaccharides, hemicelluloses, and water-soluble polysaccharides was studied in Siberian spruce (Picea obovata Ledeb.) needles of various ages. Accumulation of carbohydrates during the growth period and the ratio of their individual fractions were determined by the proportions between the growth rate and assimilate export in developing needles. Throughout the growth period, the content of monosaccharides in needles of all ages was much higher than that of oligosaccharides. Unlike the period of active growth, the dormancy period was characterized by a wider diversity of oligosaccharides and their rather high content. Hemicelluloses that accumulated during the previous growth season and water-soluble polysaccharides of 2- and 3-year-old needles were involved into metabolism at the time of active growth of current-year needles. Analysis of carbohydrate accumulation in 1- to 9-year-old needles showed that the monosaccharide concentration was higher in needles of the first two years. The content of hemicelluloses A and B and water-soluble polysaccharides was higher in older needles. During the growth period, the assimilates of 2- and 3-year-old needles were partly used for the formation of young needles. The increase in relative content of polysaccharides during needle senescence reflects the involvement of photosynthetic products into the secondary pool of carbon metabolites.     

Choice of host plant as a factor in reproductive isolation of the aphid genus Cryptomyzus (Homoptera, Aphididae)

Abstract. 1. Host plant preference experiments were conducted with closely related taxa of the aphid genus Cryptomyzus. Males, and presexual morphs (sexuparae and gynoparae), were used to determine the impact of host plant choice on reproductive isolation. In the case of host-alternating species these morphs are migratory and so will select the host plant.
2. Host plant preference of two closely related taxa of C. alboapicalis (Theobald) was found to promote their reproductive isolation. The preference of sexuparae of these monoecious taxa was more pronounced than that of the males.
3. Host plant preference and subsequent production of oviparae showed that C. galeopsidis (Kaltenbach) consists of two host races restricted to Ribes rubrum L. and R. nigrum L., respectively. The existence of clones, intermediate in their preference and reproductive performance on these plants, suggests that hybridization occurs.     

Grazing by a native and an exotic crayfish on aquatic macrophytes

1. We compared grazing by native noble crayfish ( Astacus astacus ) and the exotic signal crayfish ( Pacifastacus leniusculus ) on seedling or well-established macrophytes.
2. In a pool experiment, seedlings of emergent Scirpus lacustris and floating-leaved Potamogeton natans were heavily grazed by adult signal crayfish, whereas established plants of the same species sustained only minor damage.
3. In a preference experiment two submerged macrophytes ( Chara vulgaris and Elodea canadensis ), and both seedlings and established plants of S. lacustris and P. natans , were presented pairwise to signal and noble crayfish. There was no significant difference in preference by the two crayfish species. Chara vulgaris was preferred to all other plants presented, established plants of S. lacustris and P. natans were never preferred, and seedlings of P. natans were preferred to established P. natans .
4. An aquarium experiment was conducted in which the consumption of Chara by signal and noble crayfish was measured in relation to water temperature. Signal crayfish consumed significantly more Chara than noble crayfish, especially at higher temperatures.
5. Our results indicate that the signal crayfish is the more voracious grazer especially at higher temperatures. There may be negative effects on vegetation (emergents and floating-leaved, as well as submerged species) when the signal crayfish is introduced. Chara species are particularly susceptible, since they are preferred by crayfish and the genus includes a large number of rare species. Stocking of crayfish therefore could lead to the decline or removal of submerged species in order of crayfish feeding preferences and could prevent the spread or cause a decline of emergent and floating-leaved vegetation.     

Grazing by a native and an exotic crayfish on aquatic macrophytes

1. We compared grazing by native noble crayfish ( Astacus astacus ) and the exotic signal crayfish ( Pacifastacus leniusculus ) on seedling or well-established macrophytes.
2. In a pool experiment, seedlings of emergent Scirpus lacustris and floating-leaved Potamogeton natans were heavily grazed by adult signal crayfish, whereas established plants of the same species sustained only minor damage.
3. In a preference experiment two submerged macrophytes ( Chara vulgaris and Elodea canadensis ), and both seedlings and established plants of S. lacustris and P. natans , were presented pairwise to signal and noble crayfish. There was no significant difference in preference by the two crayfish species. Chara vulgaris was preferred to all other plants presented, established plants of S. lacustris and P. natans were never preferred, and seedlings of P. natans were preferred to established P. natans .
4. An aquarium experiment was conducted in which the consumption of Chara by signal and noble crayfish was measured in relation to water temperature. Signal crayfish consumed significantly more Chara than noble crayfish, especially at higher temperatures.
5. Our results indicate that the signal crayfish is the more voracious grazer especially at higher temperatures. There may be negative effects on vegetation (emergents and floating-leaved, as well as submerged species) when the signal crayfish is introduced. Chara species are particularly susceptible, since they are preferred by crayfish and the genus includes a large number of rare species. Stocking of crayfish therefore could lead to the decline or removal of submerged species in order of crayfish feeding preferences and could prevent the spread or cause a decline of emergent and floating-leaved vegetation.     

Host plant variation in plant-mediated indirect effects: moth boring-induced susceptibility of willows to a specialist leaf beetle

Abstract.  1. We examined the plant-mediated indirect effects of the stem-boring moth Endoclita excrescens (Lepidoptera: Hepialidae) on the leaf beetle Plagiodera versicolora (Coleoptera: Chrysomelidae) in three willow species, Salix gilgiana , S. eriocarpa , and S. serissaefolia.
2. When the stem-boring moth larvae damaged stems in the previous year, willows were stimulated to produce vigorously growing lateral shoots on these stems. These new lateral shoots were significantly longer and the upper leaves had significantly higher nitrogen and water content than current-year shoots on unbored stems, although the carbon content and leaf dry mass were not different between lateral and current-year shoots.
3. In the field, leaf beetle larvae and adults had significantly greater densities on lateral shoots of bored stems than on current-year shoots of unbored stems. A laboratory experiment showed that female beetles had significantly greater mass and fecundity when fed on leaves of newly-emerged lateral shoots. Thus, the stem-boring moth had a positive effect on the temporally and spatially separated leaf beetle by increasing resource availability by inducing compensatory regrowth.
4. The strength of the indirect effects on the density and performance of the leaf beetle differed among willow species, because there was interspecific variation in host quality and herbivore-induced changes in plant traits. In particular, we suggest that the differences in magnitude of the changes among willow species in shoot length and leaf nitrogen content greatly affected the strength of the plant-regrowth mediated indirect effect, coupled with host-plant preference of the leaf beetle.     

Magnesium deficiency treatment causes reductions in photosynthesis of well-nourished Norway spruce

In order to investigate effects of magnesium deficiency on Norway spruce [Picea abies (L.) Karst.] photosynthesis, 100 well-nourished 5-year-old spruce trees were grown in sand culture, individually supplied with circulating nutrient solutions. Mineral nutrients were added to the nutrient solutions in optimal quantities and optimal relations to nitrogen. Magnesium was supplied at 0.203, 0.041 and 0.005 mM in order to simulate optimal nutrition, moderate deficiency and severe deficiency. Parameters of photosynthetic gas exchange, chlorophyll, magnesium and starch concentrations were determined in current-year and 1-year-old needles during one growing season. By mid May — 6 months after onset of the Mg deficiency treatments in late autumn — CO₂-assimilation rates of 1-year-old needles were significantly decreased independent of the severity of the deficiency treatment, whereas the chlorophyll concentrations did not differ from the controls. The occurrence of yellowing symptoms during July did not further influence the Mg deficiency effect on photosynthesis. In contrast to 1-year-old needles, significant reductions of photosynthesis and chlorophyll in current-year needles were only caused by severely deficient Mg supply. Mg deficiency affected carboxylation efficiency but not light use efficiency. From the accumulation of starch in the needles, up to 30-fold of the controls, the conclusion has been drawn that reactions of CO₂-fixation were affected by reduced carbohydrate export. The light-dependent pigment reduction, leading to the typical tipyellowing of needles, clearly reflects a secondary effect of Mg deficiency.