Within-tree and among-tree variation in growth of Epirrita autumnata on mountain birch leaves

Abstract.

• 1 We studied among-tree and within-tree variation in the growth of larvae of a geometrid, Epirrita autumnata, on mountain birch (Betula pubescens ssp. tortuosa) leaves at different hierarchical levels: among trees (genets), among ramets within trees, among branches within ramets, among shoots within branches and among leaves within shoots. We used only short shoot leaves, which burst simultaneously in spring.
• 2 Trees accounted for most of the variation in larval growth rate, but there was substantial variation also among ramets within trees, among branches within ramets, and among short shoots within branches. Variation among leaves within short shoots was negligible. When the probabilities from different experiments were combined, the differences were statistically significant at the tree, ramet and branch levels, and approached significance at the shoot level.
• 3 In different experiments, larval growth was from 9% to 54% lower on the worst tree than on the best tree of the experiment. On average, larval growth rate was in different experiments from 11% to 32% lower on the worst ramet than on the best ramet within tree, from 8% to 18% lower on the worst than on the best branch within ramet, and from 12% to 30% lower on the worst than on the best shoot within branch.
• 4 The amount of among-and within-tree variation shown by our results may have ecological and evolutionary implications: among-tree variation should select for discrimination by ovipositing females and dispersing larvae, within-tree variation should select also for optimal foraging behaviour of larvae.

     

Systematic within-tree variation in mountain birch leaf quality for a geometrid, Epirrita autumnata

Abstract.

• 1 We studied within-tree variation in leaf quality of the mountain birch, Betula pubescens ssp. tortuosa, for larvae of the autumnal moth, Epirrita autumnata.
• 2 The purpose of the study was to determine the possible occurrence of systematic differences in larval growth on short shoot leaves (i.e. leaves of the same age): among leaves facing in different compass directions, between leaves of lower and upper branches, among leaves on different positions within a branch and among leaves of different sizes within a short shoot. We also measured larval growth between short shoot and long shoot leaves (i.e. between leaves of different age).
• 3 The larvae grew best on leaves on the north side of trees and most poorly on south side leaves, the east and west sides being intermediate. Leaves from the upper branches supported larval growth better than leaves from the lower ones. The larvae grew better on the smallest leaf of each short shoot and were able to utilize it more efficiently than the two larger leaves. Short shoot leaves from the basal and middle parts of the upper branches of the trees were of better quality for the larvae than short shoot leaves from the tip part of the branches. The larval growth rate did not differ between short shoot and long shoot leaves. In general, within-tree variation in the larval growth rate was lower than variation among different trees.
• 4 Damage to leaves can decrease leaf quality for herbivores in the same year (rapidly inducible responses) or the following year(s) (delayed inducible responses). Our results show that systematic within-tree variation in larval growth can be as great as the effects of rapidly inducible responses and that variation among individual trees can be as great as the mean effects of delayed inducible responses.

     

Stand age-structure influence in a low population peak of Epirrita autumnata in a mountain birch forest

Population densities of Epirrita autumnata caterpillars were recorded from 1984 to 1990 m four mountain birch stands of two age categories, young and old Caterpillar populations peaked m 1986–87, and peak densities were higher in the old than in the young stands Recorded peak densities were low compared with synchronous populations with outbreak densities in neighbouring areas Differences in caterpillar densities between young and old stands could not be explained by differences in survival of early instar caterpillars or in parasitoid-caused larval mortality Instead, tree-age related differences in amounts of suitable oviposition sites and possible changes in food quality due to ageing and lower vigour of mature trees are suggested to explain why E autumnata populations tend to be higher in old stands     

Growth performance of Epirrita autumnata (Lepidoptera: Geometridae) on mountain birch: trees,broods, and tree x brood interactions

Summary The outcome of herbivore-host plant interactions is partly a function of variation within the two populations. We partitioned variance in herbivore growth performance into components attributable to differences between trees, differences between (full-sib) insect broods, and tree x brood interactions. Growth performance of Epirrita larvae feeding on a small (0.25 ha) population of mountain birch was greatly influenced by differences between individual trees. Up to 49% of the variation in insect growth rate was due to tree effects; 5th instar growth rates ranged from 0.38 to 0.56 mg·mg^-1·day^-1 across a sample of 8 trees. About 25% of the variation in pupal weights and larval periods was due to tree effects; on low quality trees larvae required a longer time to attain lower pupal weights. Differences between trees were also evident in physical and chemical characteristics of the leaves. Insect broods differed in the duration of the larval period (15% of the variance) which led to differences in the pupal weight attained (13% of the variance). However, brood-specific differences in growth rate were modest (6% in the 4th instar) or nonexistent (5th instar). There was no evidence for tree x brood interactions, which refutes the possibility of fine scale adaptation to particular tree phenotypes. Hypotheses to explain the existence of this variability, and to predict its evolutionary and ecological consequences, are advanced.     

Long-term dynamics of leaf miners, Eriocrania spp., on mountain birch: alternate year fluctuations and interaction with Epirrita autumnata

Abstract.

• 1 Abundance of leaf-mining larvae of the outbreak species Eriocrania spp. was monitored in northern Sweden in 1955–67 and 1984–92 in a mountain birch forest rejuvenated by an Epirrita autumnata outbreak in 1954–55.
• 2 Eriocrania mine density fluctuated in a regular biennial pattern, probably due to a semivoltine life cycle. The alternate year fluctuations had shifted phase between the two study periods.
• 3 The density fluctuated at a significantly higher mean level and with a lower amplitude in the c. 10-year-old forest in the 1960s than in the 30-year-old forest in the 1980s. However, no difference was found in mine density between c. 30- and >60-year-old stands in the 1980s.
• 4 Significant correlations between the density of mines in high-density years and date of budburst, and between rate of change between high-density years and the time between snow-melt and budburst indicate effects of weather. No correlation was found between yearly average mine density and date of budburst, precipitation or temperatures in May or June.
• 5 No correlation was found between the mean densities of Eriocrania mines and E.autumnata caterpillars in the same and the two preceding years. However, in the first (1955) and highest of three studied E.autumnata peaks a negative effect on population density of Eriocrania was indicated.
• 6 A significant, negative correlation between the number of Eriocrania mines and E.autumnata caterpillars on single branches was found in three out of eighteen years. The separation of caterpillars and mines at branch level indicates an effect of avoidance at intermediate population densities.

     

Foliar oxidases as mediators of the rapidly induced resistance of mountain birch against <Emphasis Type="Italic">Epirrita autumnata</Emphasis>

Induced resistance of the mountain birch against its main defoliator Epirrita autumnata is a well-characterized phenomenon. The causal mechanism for this induced deterioration, however, has not been unequivocally explained, and no individual compound or group of traditional defensive compounds has been shown to explain the phenomenon. Phenolic compounds are the main secondary metabolites in mountain birch leaves, and the biological activity of phenolics usually depends on their oxidation. In this study, we found that the activity of polyphenoloxidases (PPOs), enzymes that oxidize o-diphenols to o-diquinones, was induced in trees with introduced larvae, and bioassays showed that both growth and consumption rates of larvae were reduced in damaged trees. PPO activity was negatively associated with both larval growth and consumption rates in trees with bagged larvae, but not in control trees. Our results suggest that the oxidation of phenolics by PPOs may be a causal explanation for the rapidly induced resistance of mountain birch against E. autumnata. This finding also helps to explain why correlations between insect performance and phenolics (without measuring indices explaining their oxidation) may not produce consistent results.     

Climate change and outbreaks of the geometrids Operophtera brumata and Epirrita autumnata in subarctic birch forest: evidence of a recent outbreak range expansion

1. Range expansions mediated by recent climate warming have been documented for many insect species, including some important forest pests. However, whether climate change also influences the eruptive dynamics of forest pest insects, and hence the ecological and economical consequences of outbreaks, is largely unresolved. 2. Using historical outbreak records covering more than a century, we document recent outbreak range expansions of two species of cyclic geometrid moth, Operophtera brumata Bkh. (winter moth) and Epirrita autumnata L. (autumnal moth), in subarctic birch forest of northern Fennoscandia. The two species differ with respect to cold tolerance, and show strikingly different patterns in their recent outbreak range expansion. 3. We show that, during the past 15-20 years, the less cold-tolerant species O. brumata has experienced a pronounced north-eastern expansion into areas previously dominated by E. autumnata outbreaks. Epirrita autumnata, on the other hand, has expanded the region in which it exhibits regular outbreaks into the coldest, most continental areas. Our findings support the suggestion that recent climate warming in the region is the most parsimonious explanation for the observed patterns. 4. The presence of O. brumata outbreaks in regions previously affected solely by E. autumnata outbreaks is likely to increase the effective duration of local outbreaks, and hence have profound implications for the subarctic birch forest ecosystem.     

No regulatory role for adult predation in cyclic population dynamics of the autumnal moth,Epirrita autumnata

1. Multiannual population cycles of geometrid moths are thought to be driven by trophic‐level interactions involving a delayed density‐dependent component. Predation on adult moths has been a little‐studied mechanism of this phenomenon. 2. Using 29 daytime and 61 night‐time predation trials in the field, we exposed living autumnal moth (Epirrita autumnata Borkhausen, Lepidoptera: Geometridae) females to their natural predators during each autumn throughout the 10‐year population cycle. 3. In our northern study location (70°N), insectivorous passerines had already migrated, and harvestman Mitopus morio Fabricius (Opiliones: Phalangiidae) was found to be the main predator of the adult moths. The predation mortality occurred mainly at night and was positively correlated with the minimum temperature measured during the predation trial. 4. Despite high annual variability in the degree of adult predation, both direct and delayed density‐dependent effects were weak and indicate that predation on adult moths in the autumn does not have any regulatory role in cyclic population dynamics of the autumnal moth in northern Fennoscandia.     

Within-plant variation in induced defence in developing leaves of cotton plants

According to optimal defence theory (ODT) plants should invest in stronger defence in the most valuable parts, such as reproductive or young tissue. Cotton plants are known to express high resistance to herbivore feeding in the developing leaves at the top of the plant after herbivore-damage. Cotton plants also have developing leaves on side shoots from the nodes all along the plant. This allowed us to investigate within-plant distribution of defence between younger tissues at different locations on the plant. We found that all developing leaves showed increased resistance to feeding by larvae of the generalist moth Spodoptera littoralis after herbivore damage to leaves of the plant. An increase in the concentration of secondary compounds, terpenoid aldehydes, was found in developing leaves both at the top of the plant and on the side shoots. However, the resistance was stronger in the top leaves than in the side shoot leaves, showing that there is intra-plant variation in the induced response between different leaves of the same age. After the initial damage, larval feeding mainly occurred on the older, fully developed true leaves. Furthermore, the herbivore-induced resistance in the developing leaves reduced upward movement of feeding larvae on the plant and reduced the feeding on the upper parts of the plant over a period of at least 6 days. The plant thus benefits from defending all developing leaves by directing feeding to older, less valuable leaves and lower parts of the plant. The observed distribution of defence within cotton plants supports ODT.     

Consequences of herbivory in the mountain birch (Betula pubescens ssp tortuosa): importance of the functional organization of the tree

Summary Three types of experiments indicate that the functional organization of the mountain birch may influence the ways in which the tree responds to simulated or natural herbivory. The first experiment showed that herbivory to both short and long shoot leaves affects plant development but, because growth largely proceeds by resources of the previous year, is manifested only in the year following the damage. The second experiment showed that even partial damage to a single long shoot leaf caused the axillary bud of that leaf to produce a shorter shoot the next year. Therefore, the value of a leaf depends also on the organ which it is subtending. In the third experiment we manipulated the apical dominance of shoots in ramets and caused improvement to leaf quality in extant shoots. Ramets within a tree responded individually, probably mediated by disturbance of the hormonal control because removal of apical buds elicited the response although removal of the same number of basal buds did not. Induced amelioration is a different response to induced resistance. The two responses are triggered by different cues and may occur in the same plant. By altering hormonal balance of shoots it is potentially possible for herbivores to induce amelioration of food quality. The ways in which herbivory is simulated may explain variability of results obtained when herbivory-induced responses in plants have been studied.     

Individual and combined effects of disturbance and N addition on understorey vegetation in a subarctic mountain birch forest

Questions: What are the effects of repeated disturbance and N‐fertilization on plant community structure in a mountain birch forest? What is the role of enhanced nutrient availability in recovery of understorey vegetation after repeated disturbance? How are responses of soil micro‐organisms to disturbance and N‐fertilization reflected in nutrient allocation patterns and recovery of understorey vegetation after disturbance? Location: Subarctic mountain birch forest, Finland. Methods: We conducted a fully factorial experiment with annual treatments of disturbance (two levels) and N‐fertilization (four levels) during 1998–2002. We monitored treatment effects on above‐ground plant biomass, plant community structure and plant and soil nutrient concentrations. Results: Both disturbance and N‐fertilization increased the relative biomass of graminoids. The increase of relative biomass of graminoids in the disturbance treatment was over twice that of the highest N‐fertilization level, and N‐fertilization further increased their relative biomass after disturbance. As repeated disturbance broke the dominance of evergreen dwarf shrubs, it resulted in a situation where deciduous species, graminoids and herbs dominated the plant community. Although relative biomass of deciduous dwarf shrubs declined with N‐fertilization, it did not cause a shift in plant community structure, as evergreen dwarf shrubs remained dominant. Both disturbance and N‐fertilization increased the N concentration in vascular plants, whereas microbial biomass N and C were not affected by the treatments. Concentrations of NH₄⁺, dissolved organic N (DON) and dissolved organic C (DOC) increased in the soil after N‐fertilization, whereas concentrations of NH₄⁺ and DON decreased after disturbance. Conclusions: Disturbances caused by e.g. humans or herbivores contribute more to changes in the understorey vegetation structure than increased levels of N in subarctic vegetation. Fertilization accelerated the recovery potential after repeated disturbance in graminoids. Microbial activities did not limit plant growth.     

Genetic variability in a population of arbuscular mycorrhizal fungi causes variation in plant growth

Different species of arbuscular mycorrhizal fungi (AMF) alter plant growth and affect plant coexistence and diversity. Effects of within-AMF species or within-population variation on plant growth have received less attention. High genetic variation exists within AMF populations. However, it is unknown whether genetic variation contributes to differences in plant growth. In our study, a population of AMF was cultivated under identical conditions for several generations prior to the experiments thus avoiding environmental maternal effects. We show that genetically different Glomus intraradices isolates from one AMF population significantly alter plant growth in an axenic system and in greenhouse experiments. Isolates increased or reduced plant growth meaning that plants potentially receive benefits or are subject to costs by forming associations with different individuals in the AMF population. This shows that genetic variability in AMF populations could affect host-plant fitness and should be considered in future research to understand these important soil organisms.     

Fluctuating asymmetry as an indicator of elevation stress and distribution limits in mountain birch (<Emphasis Type="Italic">Betula pubescens</Emphasis>)

Fluctuating asymmetry (FA) has been suggested as a useful indicator of elevation stress and, hence, distribution limits in plants. However, no plant studies have been carried out to test (i) whether FA shows a gradual increase towards the alpine distribution limit and (ii) whether FA responds to elevation stress independent of other stressors which is necessary for FA to be a useful indicator in this context. To test these two hypotheses, this 2-year field study investigated the dose–response relationship between elevation stress and FA in mountain birch (Betula pubescens) under contrasting levels of insect attack in northern Norway. The results showed that FA increased linearly from sea level towards the tree line in both years independent of insect attack, which had no observable effect on FA, i.e. insect attack did not appear to disturb the FA-elevation relation. Thus, in mountain birch, FA appeared to be a reliable indicator of elevation stress. Further investigation is now needed in order to develop this hypothesis.     

Variation in leaf trichomes and nutrients of Wigandia urens (Hydrophyllaceae) and its implications for herbivory

Leaf trichome variation was studied in a population of Wigandia urens (Hydrophyllaceae) in relation to water availability, diversity of herbivorous insects, and grazing rates. Plants of W. urens have glandular and urticant (stinging hairs) trichomes, and it is possible to distinguish two types of leaves: smooth leaves with only glandular trichomes and bristly leaves with both types of trichomes. Density of urticant trichomes was negatively correlated with weekly mean precipitation. Fourteen species of insects in five different orders feed on leaves of W. urens throughout the year. Grazing rates varied according to type and age of leaves. Grazing rates were lower in smooth than bristly young leaves but the rates were similar for mature leaves. Secondary metabolites, screened using qualitative methods, showed similar classes of compounds in both types of leaves. Bristly leaves had significantly higher concentrations of nitrogen, phosphorous and water and these nutritional differences were well correlated with the differences in grazing rates between bristly and smooth leaves.     

Postembryonal growth and its variability of the three marine fishes with special reference to the mechanism of growth variation in fishes

1. To study the relation of growth variation to the behaviour pattern, newly hatched Hemiramphus sajori, Chrysophrys major and Zebrias zebra were reared in 30 litter aquaria during one month, and body length was measured four times.
2. In Hemiramphus sajori coefficient of variation of body length (CV) first increased but showed a tendency to decrease with development of schooling behaviour.
3. In Chrysophrys major CV markedly increased and mode of the distribution curve of body length inclined towards the left side, associated with appearance of aggression and cannibalism.
4. In Zebrias zebra, when behaviour pattern changed from planktonic to demersal some of the small individuals nearly ceased growing. At that time CV of body length remarkably increased and the mode of the distribution curve of body length shifted towards the right side.
5. In all the species a positive correlation was seem between specific growth rate and initial body length soon after intensive feeding began. An negative correlation was observed in most of individuals of Hemiramphus sajori after the appearance of schooling behaviour. When a conspicuous increase in CV and skewness in distribution curve took place, a separate type of correlation was seen in large and small group in the population.
6. Based on above mentioned and other materials a general mechanism of growth variation in fishes was discussed.

     

Consequences of climate variability for the performance of bison in tallgrass prairie

Climate variability is a major structuring factor in grassland ecosystems, yet there is great uncertainty in how changes in precipitation affect grazing herbivores. We determined how interannual variation in the timing and magnitude of precipitation affected the weight gain of free-roaming bison in their first and second year. Bison weights were analyzed for 14 years for Konza Prairie, Kansas, and 12 years for Tallgrass Prairie Preserve, Oklahoma. Greater late-summer precipitation increased bison weight gain. For every 100 mm precipitation, weight gain increased 6.4–15.3 kg depending on age classes and site. In contrast, greater midsummer precipitation decreased weight gain. For every additional 100 mm precipitation, weight decreased 9.7–17.3 kg depending on age class and site. The decreased weight gain of bison with greater midsummer precipitation was associated with increased grass stem production during the period for each of three dominant grasses at Konza Prairie. Although greater stem production increases the quantity of aboveground biomass, it should decrease the overall nutritional quality of biomass to grazers, which would reduce weight gain. With offsetting effects of mid- and late-summer precipitation on weight gain, these results show that predicting the effects of climate change on grazers must incorporate both the timing and magnitude of changes in precipitation and their effects on both the quantity and quality of biomass.     

Physiological bases of genetically determined variation in growth of marine invertebrate larvae: A study of growth heterosis in the bivalve Crassostrea gigas

Many species of marine animals have larval stages whose rates of growth in the plankton are regulated by complex combinations of biological and environmental factors. In this study, we focus on the physiological bases that underlie endogenous variation in growth potential of larvae. Our approach was based on experimental crosses of gravid adults from pedigreed families of the Pacific oyster, Crassostrea gigas. This produced large numbers of larvae with different growth rates when reared under similar environmental conditions of food and temperature. A total of 35 larval families were reared to test hypotheses regarding the physiological bases of growth variation. Growth rate of these larval families varied over a five-fold range, from 3.4 (± 0.5, S.E.) to 17.6 (± 0.6) μm day^− 1. The suite of integrated measurements applied to study growth variation included size, biochemical compositions, rates of particulate and dissolved nutrient acquisition, absorption efficiencies, respiration rates and enzyme activities. We show that a complex set of physiological processes regulated differences in genetically determined growth rates of larvae. One-half of the energy required for faster growth came from an enhanced, size-specific feeding ability. Differences in absorption rates were not significant for slow- and fast-growing larvae, nor were differences in size-specific respiration rates. Metabolic processes accounted for the additional 50% of the energy “savings” required to explain enhanced growth rates. We propose that different protein depositional efficiencies could account for this energy saving. Quantitative analyses of the endogenous physiological factors that cause variation in growth rate will allow for a more sophisticated understanding of growth, survival and recruitment potential of larvae.     

Individual, nightly, and seasonal variation in calling behavior of the gray tree frog, Hyla versicolor: implications for energy expenditure

We investigated individual, nightly, and seasonal variationin calling behavior of a population of gray tree frogs (Hylaversicolor) from Connecticut, USA. Repeated recordings of individualmales on seven nights revealed significant differences amongmales in calling rate on all but one night and differences innumber of pulses per call and number of pulses produced perhour (pulse effort) on four nights. Most males reduced callingactivity late at night (after 2230 h), but some maintained arelatively steady rate of call production before dropping outof the chorus. Data collected for 26 individuals recorded onthree or more nights throughout the breeding season revealedsignificant differences among males in calling rate, numberof pulses per call, and pulse effort, but repeatabilities forall three variables were low (0.17, 0.35, and 0.12, respectively).The highest repeatability was for number of pulses per call,a variable strongly influenced by proximity to calling neighbors,probably because males often interacted with neighbors at similardistances on several successive nights. Males tended to reducethe number of pulses per call as the season progressed and thedistance between neighbors decreased, but they showed no clearseasonal change in calling rate or pulse effort. There was asubstantial seasonal decline in the number of hours of chorusactivity, resulting in a median decrease of 43% in nightly energyexpenditure by calling males.     

Sources of variation affecting cashmere grown in the Pamir mountain districts of Tajikistan and implications for industry development

We aimed to quantify the sources of variation contributing to the main quality attributes of cashmere produced from goats in the Pamir mountain districts of Murghab, Shugnon and Vanj in Tajikistan. In early spring 2010, mid-side samples were taken from 194 adult females, 43 adult males and 20 castrates belonging to 58 farmers and pastoralists in 14 villages. For 57 goats, samples were also taken from the shoulder and hip sites. Mean fibre diameter (MFD), fibre curvature (FC) and cashmere staple length (SL) data were examined using a general linear model to determine the relationships between fleece attributes and other possible effects. For females, the mean (s.d.) for MFD, FC and SL were: 16.5 (1.70) μm; 46 (12.1)°/mm; 53 (22.9) mm. MFD was affected by district, SL and age of goat. SL was affected by district, MFD, gender, age of goat and village. FC was affected by district, MFD, shade of cashmere, age of goat and farmer. Cashmere from Vanj district was finer and shorter than cashmere from Murghab and Shugnon. Cashmere grown on the mid-side and hip sites was finer and had higher FC than cashmere grown on the shoulder. Cashmere grown on the hip was shorter than cashmere grown on the mid-side and shoulder. About 50% of the cashmere sampled was < 16.4 μm and potentially suitable for knitwear. Of this fine cashmere, 53% was 34 mm or longer. A further 37% of the cashmere was 16.4-18.5 μm, and suitable for weaving as 97% was longer than 36 mm. Almost 12% of samples were > 18.5 μm and may only be suitable for weaving or, if cashgora, will have little commercial value. Most of the cashmere was coloured. There are cashmere goats in the Murghab, Shugnon and Vanj districts of Tajikistan which produce the finest qualities of cashmere, comparable to premium grades of Chinese cashmere. There is substantial scope to increase the commercial value of cashmere produced by goats in Tajikistan, in particular increasing SL for fine cashmere, reducing MFD for the longest cashmere and ensuring cashmere has acceptable FC and white colour.     

Nitrogen in cell walls of sclerophyllous leaves accounts for little of the variation in photosynthetic nitrogen-use efficiency

Photosynthetic rate per unit nitrogen generally declines as leaf mass per unit area (LMA) increases. To determine how much of this decline was associated with allocating a greater proportion of leaf nitrogen into cell wall material, we compared two groups of plants. The first group consisted of two species from each of eight genera, all of which were perennial evergreens growing in the Australian National Botanic Gardens (ANBG). The second group consisted of seven Eucalyptus species growing in a greenhouse. The percentage of leaf biomass in cell walls was independent of variation in LMA within any genus, but varied from 25 to 65% between genera. The nitrogen concentration of cell wall material was 0.4 times leaf nitrogen concentration for all species apart from Eucalyptus , which was 0.6 times leaf nitrogen concentration. Between 10 and 30% of leaf nitrogen was recovered in the cell wall fraction, but this was independent of LMA. No trade-off was observed between nitrogen associated with cell walls and the nitrogen allocated to ribulose 1·5-bisphosphate carboxylase/oxygenase (Rubisco). Variation in photosynthetic rate per unit nitrogen could not be explained by variation in cell wall nitrogen.