A tree in the eyes of a moth – temporal variation in oak leaf quality and leaf-miner performance

When examined at any moment in time, different parts of an individual oak tree exhibit almost as large differences in quality as different trees. But how consistent are such patterns in time? In this paper, we use intraclass correlations to assess the temporal consistency of host plant quality at several spatial scales. As measures of quality, we use both individual chemical attributes (phenolic contents) and the overall performance (larval survival) of the host‐specific leaf‐miner Tischeria ekebladella. Concentrations of 24 phenolic compounds were monitored on seven trees throughout a season. Variation in mine initiation and larval survival rates was assessed for individuals transplanted to another set of trees early versus later in the season, while year‐to‐year variation in larval survival was studied through stratified surveys of wild individuals during three years. At all time scales considered, measures of host quality were moderately consistent: a tree favourable in quality at one point in time often remained so, but there was abundant variation around this relationship (ρ=0.4–0.6). One hierarchical level deviated from this general pattern: on individual branches, larval survival rates varied randomly among years (ρ=0). Our study suggests that the quality of trees, and in particular of smaller units within trees, may be difficult to predict both in space and in time. To account for this, insects might benefit from adopting a bet‐hedging strategy when selecting resources.     

Seasonal variation in leaf chemistry of the coast live oak Quercus agrifolia and implications for the California oak moth Phryganidia californica

Summary The perennial foliage of the California coast live oak (Quercus agrifolia Nee) permits herbivores to feed on this oak species throughout the year. Patterns of herbivory for a two-year period on Q. agrifolia were observed in relation to seasonal and age-related changes in the nutritional and defensive characteristics of leaves. Nitrogen and phosphorus contents were higher in new leaves compared to mature foliage. Structural compounds (e.g., cellulose) in leaves rapidly increased with age. Concentrations of tatal phenolics (Folin-Denis) and astringency were higher in new foliage, and concentrations of condensed tannins gradually increased as the leaves matured. Peaks of herbivore damage were observed in June and in September–October, and were caused by outbreaks of the California oak moth (Phryganidia californica). P. californica, a bivoltine oak specialist, exhibited feeding preferences in June for old leaves over emerging leaves, and showed no preferences for leaf classes in September. These results suggest that P. californica is adapted to survive on nutritionally poor foliage and to circumvent quantitative defenses such as condensed tannins.     

Spatial and temporal variation in RGR and leaf quality of a clonal riparian plant: Arundo donax

Arundo donax L. is a tall perennial reed classified as an emergent aquatic plant. In California, it has invaded riparian zones, where it acts as a transformer species. Because plant growth and leaf quality influence the effectiveness of management techniques, we sought to determine if these characters varied temporally and spatially in a northern California population of A. donax. Tissue C and N content and C:N ratio varied during the growing season. Leaf N was higher in spring and in plants that were closer to a stream. It was significantly negatively related to the clump's distance from the stream but not related to its elevation relative to the stream. Plants near the stream produced taller stems with more leaves per stem than those more distant from the stream. RGR differed across time and space. It was highest in spring prior to the appearance of flowers on a few stems that were >1 year old within the clumps. Decline in RGR as the growing season progressed coincided with the appearance of branches and flowers on stems <1 year old on a few plants within the studied population. RGR was significantly related to the N content and C:N ratio of leaves on mature stems (>1 year old). This implies that the decrease in stem growth reflected changes in nutrient availability within the entire A. donax clump and not just in the growing stems (<1 year old). These findings have implications for timing of management techniques.     

Selective oviposition by a leaf miner in response to temporal variation in abscission

Summary Responses to humidity of net photosynthesis and leaf conductance of single attached leaves were examined in populations of herbs from wet soil sites in Beltsville, Maryland and Davis, California, USA. Plants were grown in controlled environments under three conditions which differed in the magnitude of the day-night temperature difference and in daytime air saturation deficit. No population differences in response were found in Abutilon theophrasti. In Amaranthus hybridus stomatal conductance and net photosynthesis were more reduced by increasing leaf to air water vapor pressure difference (VPD) in the population from Beltsville, but only for the growth condition with a constant 25°C temperature. In Chenopodium album, stomatal conductance was more sensitive to VPD in the population from Davis, but only for the growth condition with 28/22°C day/night temperatures. Population differences in the sensitivity to VPD of leaf conductance were associated with differences in leaf area to root weight ratio. The relative reduction of net photosynthesis as VPD increased was greater than, equal to, or less than the relative decrease in substomatal carbon dioxide partial pressure. The pattern depended on species, and on growth condition. From these results one can not conclude that environmental humidity has been a strong selective force in determining sensitivity to humidity of stomatal conductance.     

Intraspecific variation in spring leaf phenology and duration of leaf expansion in relation to leaf habit and leaf size of temperate tree species

Plant Ecology - Spring leaf phenology has been intensively studied in temperate deciduous broad-leaved tree species, but the phenology of evergreen broad-leaved tree species has seldom been focused...     

Fitness consequences of host use in the field: temporal variation in performance and a life history tradeoff in the moth <Emphasis Type="Italic">Rothschildia lebeau</Emphasis> (Saturniidae)

That fitness varies as a function of using different hosts is a basic premise of theory addressing the ecology and evolution of oviposition behavior and host selection. Few data exist demonstrating: (1) the effects of different hosts on fitness in the field, and (2) how these effects vary spatially or temporally. Cohorts of caterpillars were followed from hatching to adulthood to test the hypotheses that: (1) hosts have significant effects on herbivore performance in nature, and (2) host "quality" for performance varies predictably (i.e., the rank order is consistent) across herbivore generations. In total, the fates of >2,000 caterpillars were followed on 238 individual host trees. Host species had significant effects on most, but not all, measured components of caterpillar performance in the field. Variation among generations was mainly quantitative rather than qualitative, with few changes in the rank order of hosts in their effects on performance. There was also a strong seasonal effect on performance such that caterpillar growth and survival were higher in the early wet season compared to the late wet season. Using estimates derived from these data, correlations among larval growth rate, larval survival, total development time, and final adult size were examined at the level of host plant species. Across generations, larval survival was consistently poor, development time was long, but final adult size attained was large on the host Spondias mombin. The converse was true for the host Exostema mexicanum. Relative performance on the host Casearia nitida was variable between the other two hosts. Overall, the data suggest that host use involves a predictable tradeoff between larval survival and final adult size, but argue that which is the "better" host from the female perspective will depend on the fitness consequences of producing a few, relatively large offspring versus producing more, relatively small offspring.     

Spatial and temporal dynamics of Siberian silk moth large-scale outbreak in dark-needle coniferous tree stands in Altai

The spatial and temporal dynamics of fir stands damage caused by a large-scale outbreak of the Siberian silk moth (Dendrolimus sibiricus Tschetv.) in the Altai Mountains has been studied using remote sensing and GIS methods. It is ascertained that forest damages are dissimilar relative to the surface features. The pest outbreak initially occurred on the southwest slopes with mean values of 10° and elevation of 400 m asl. The damages further extended both upward and downward, involving slopes of high steepness and the eastern exposure. The total area of the dead stands comprised 6000 ha, 45% of which were lost due to secondary pest (xylophagous insects) attacks. It is indicated that the use of remote sensing made it possible to determine the beginning of a pest outbreak with one-decade precision. The Siberian silk moth large-scale outbreak occurred against an increase in air temperatures, a decrease in precipitation, and a reduction in late frosts. The tree plants weakened by the Siberian silk moth and water stress were affected by xylophagous insect attacks. The observed and predicted warming and climate aridity increase will facilitate Siberian silk moth outbreaks both within its range and northward of the current margins of the range.     

The effects of group size, leaf size, and density on the performance of a leaf-mining moth

1. The effects of two factors, leaf size and group size, on the performance of the Tupelo leafminer, Antispila nysaefoliella (Lepidoptera: Heliozelidae), were examined by fitting growth models to mine expansion data using nonlinear mixed-effects models. 2. The rate of mine expansion served as a proxy for larval performance because of its correlation with both feeding activity and growth rate and is also the means by which a larva achieves its final mine size (or total consumption). 3. Leaf size was used as a measure of resource availability, and was expected to reduce the impact of resource competition and enhance larval performance. 4. In contrast to the unidirectional effects expected for leaf size (i.e. more resources should enhance performance), the direction for the effects of group size was expected to depend on the mechanism(s) driving the effect. For example, if there is resource competition among larvae in a group, then this could increase the feeding rates of some larvae or reduce the total consumption of others. However, if leaf mining induces host plant chemical defences, then larger groups might elicit a greater defensive response by the host plant (at the leaf), and hence, be characterized by reduced feeding and growth rates. 5. To investigate these interactions, two growth models, the Gompertz model and a modified version of the von Bertalanffy growth equation, were fitted to time series of the sizes of individual leaf mines using nonlinear mixed-effects models. Linear and nonlinear associations of each factor (group size or leaf size) with model parameters were then evaluated using a hierarchical testing procedure by determining: (i) whether inclusion of the factor produced a better-fit model, and (ii) if it did, the form of that relationship (i.e. linear or nonlinear). 6. Three patterns were detected with these analyses. (i) Leaf size had a significant positive, linear relationship with mine expansion rate. (ii) Group size had a significant quadratic relationship with mine expansion rate. (iii) The effects of leaf and group size on the maximum mine size were opposite to those found with growth rate.     

High variation in foliage and leaf litter chemistry among 45 tree species of a neotropical rainforest community

Distinct ecosystem level carbon : nitrogen : phosphorus (C : N : P) stoichiometries in forest foliage have been suggested to reflect ecosystem-scale selection for physiological strategies in plant nutrient use. Here, this hypothesis was explored in a nutrient-poor lowland rainforest in French Guiana. Variation in C, N and P concentrations was evaluated in leaf litter and foliage from neighbour trees of 45 different species, and the litter concentrations of major C fractions were also measured. Litter C ranged from 45.3 to 52.4%, litter N varied threefold (0.68-2.01%), and litter P varied seven-fold (0.009-0.062%) among species. Compared with foliage, mean litter N and P concentrations decreased by 30% and 65%, respectively. Accordingly, the range in mass-based N : P shifted from 14 to 55 in foliage to 26 to 105 in litter. Resorption proficiencies indicated maximum P withdrawal in most species, but with a substantial increase in variation in litter P compared with foliage. These data suggest that constrained ecosystem-level C : N : P ratios do not preclude the evolution of highly diversified strategies of nutrient use and conservation among tropical rainforest tree species. The resulting large variation in litter quality will influence stoichiometric constraints within the decomposer food web, with potentially far-ranging consequences on nutrient dynamics and plant-soil feedbacks.     

Warmer springs disrupt the synchrony of oak and winter moth phenology

Spring temperatures have increased over the past 25 years, to which a wide variety of organisms have responded. The outstanding question is whether these responses match the temperature-induced shift of the selection pressures acting on these organisms. Organisms have evolved response mechanisms that are only adaptive given the existing relationship between the cues organisms use and the selection pressures acting on them. Global warming may disrupt ecosystem interactions because it alters these relationships and micro-evolution may be slow in tracking these changes. In particular, such shifts have serious consequences for ecosystem functioning for the tight multitrophic interactions involved in the timing of reproduction and growth. We determined the response of winter moth (Operophtera brumata) egg hatching and oak (Quercus robur) bud burst to temperature, a system with strong selection on synchronization. We show that there has been poor synchrony in recent warm springs, which is due to an increase in spring temperatures without a decrease in the incidence of freezing spells in winter. This is a clear warning that such changes in temperature patterns may affect ecosystem interactions more strongly than changes in mean temperature.     

Seeing the trees for the leaves – oaks as mosaics for a host-specific moth

From the perspective of a specialist herbivore, how homogenous are individual tree crowns as patches of habitat? We partitioned variation in physical and chemical host leaf traits and in the abundance and performance of a specialist oak leaf miner, Tischeria ekebladella, into variation at different hierarchical levels. For the phenolic contents of the leaves, we examined variation among oak stands, among trees within stands and among branches within trees. For leaf size and water content, we assessed variation among trees within a single stand, among shoots within trees, and among leaves within shoots. For moth abundance and performance, we examined variation across all levels: among oak stands, among trees within stands, among branches within trees, among shoots within branches and among leaves and insect individuals. For measures of phenolic contents, we found little variation among stands but substantial variation among individual trees. Yet, a tree particularly rich in a given compound was often comparatively poor in another. At a finer spatial scale, the phenolic composition of individual parts of a single tree was quite consistent, whereas leaf weight and water content varied widely within individual tree crowns. Moth abundances varied more among shoots within branches than at any other spatial level, whereas moth survival showed equal levels of variation within individual shoots as among separate oak stands. Likewise, for four other measures of larval performance (assessed at the level of trees and lower), we found more variation within than between trees. In conclusion, the large variation observed in the performance of a specialist moth and in the physical traits of the leaves among different parts of single tree crowns refutes the image of an oak tree as an ‘island’ of internally homogeneous quality. Hence, we may expect little evolutionary adaptation of T. ekebladella at the scale of individual trees. The moths may instead evolve to behaviourally select their resource at a very fine scale. Large variation within trees also calls for extensive replication within trees in ecological sampling designs and/or the sampling of maximally similar leaves.     

The effect of fodder plant genotype on the variation of larval fitness traits in genotype classes of green oak leafroller moth

The effect of genetic variation of oak (Quercus pubescens L. and Q. petraea L.) on the genotype fitness components in green oak leafroller moth larvae (Tortrix viridana L.) at esterase (Est-4) and protease (Pts-4) loci was studied. The samples of larvae were collected from nine oak trees, whose genetic variation was assayed by RAPD-PCR using primer OPA14. The contributions of the factors of oak species/genotype and green oak leafroller moth genotype and their interaction to the variation of important size-related traits of the larvae were evaluated by two-way ANOVA. It was shown that the same larval genotype can display maximum fitness on the trees of one species or genotype and minimum, on the trees of other species or genotype. The interactions between the oak genotype and green oak leafroller moth genotype factors lead to the relationships that appear in statistically significant associations between genotype classes of green oak leafroller moth and oak. These results are discussed from the standpoint of a recently developed new field, community or ecosystem genetics.     

Environmental controls and the influence of tree species on temporal variation in soil respiration in subtropical China

Background and aims

The knowledge of individual tree species impacts on soil respiration based on rigorous experimental designs is limited, but is crucial to help guide selection of species for reforestation and carbon (C) management purposes.

Methods

We assessed monthly soil respiration and its components, litterfall input, fine root production and mortality under 19-year-old native coniferous Cunninghamia lanceolata and broadleaved Mytilaria laosensis plantations in sub-tropical China.

Results

Total soil respiration from October 2011 to March 2013 was significantly lower under the C. lanceolata than the M. laosensis plantation. The difference in respiration rates derived from fine roots and the litter layer explained much of the variation of total soil respiration between the two tree species. We used an exponential equation and base temperature (10 °C) to normalize soil respiration rate and its components (R₁₀) and determined the correlation between R₁₀ and soil moisture. Although soil moisture had a positive relationship with R₁₀ derived from roots or litter under both C. lanceolata and M. laosensis forests, these positive correlations were masked by negative relationships between soil moisture and R₁₀ derived from root-free soil, which resulted in a neutral correlation between total R₁₀ and soil moisture under C. lanceolata forests. Monthly litterfall input was associated with variation in concurrent total soil respiration rate under the M. laosensis plantation and respiration rate lagging 3 months behind under the C. lanceolata plantation, which may suggest that litterfall input from M. laosensis can more rapidly produce C substrates for microbial respiration than litterfall from C. lanceolata.

Conclusions

This study highlighted that tree species-induced variation in the quality and quantity of fine roots and litterfall can impact not only the soil respiration rate but also the seasonal variation model of forest soil respiration.     

Temporal variation in leaf nitrogen partitioning of a broad-leaved evergreen tree, <Emphasis Type="Italic">Quercus myrsinaefolia</Emphasis>

We examined temporal changes in the amount of nitrogenous compounds in leaves from the outer and inner parts of the crown of Quercus myrsinaefolia growing in a seasonal climate. Throughout the leaf life span, metabolic protein and Rubisco content closely correlated with total nitrogen content, while structural protein content was relatively stable after full leaf expansion. Chlorophyll content was affected by shading as well as total nitrogen content in outer leaves that were overtopped by new shoots in the second year. Outer leaves showed a large seasonal variation in photosynthetic nitrogen-use efficiency (PNUE; the light-saturated photosynthetic rate per unit leaf nitrogen content) during the first year of their life, with PNUE decreasing from the peak in summer towards winter. Outer and inner leaves both showed age-related decline in PNUE in the second year. There were no such drastic changes in leaf nitrogen partitioning that could explain seasonal and yearly variations in PNUE. Nitrogen resorption occurred in overwintering leaves in spring. Metabolic protein explained the majority of nitrogen being resorbed, whereas structural protein, which was low in degradability, contributed little to nitrogen resorption.     

The effect of fodder plant genotype on the variation of larval fitness traits in genotype classes of green oak leafroller moth

The effect of genetic variation of oak (Quercus pubescens L. and Q. petraea L.) on the genotype fitness components in green oak leafroller moth larvae (Tortrix viridana L.) at esterase (Est-4) and protease (Pts-4) loci was studied. The samples of larvae were collected from nine oak trees, whose genetic variation was assayed by RAPD-PCR using primer OPA14. The contributions of the factors of Yoak species/genotype and green oak leafroller moth genotype and their interaction to the variation of important size-related traits of the larvae were evaluated by two-way ANOVA. It was shown that the same larval genotype can display maximum fitness on the trees of one species or genotype and minimal, on the trees of other species or genotype. The interactions between the oak genotype and green oak leafroller moth genotype factors lead to the relationships that appear in statistically significant associations between genotype classes of green oak leafroller moth and oak. These results are discussed from the standpoint of a recently developed new field, community or ecosystem genetics.     

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.

     

Influence of genetic variation of oaks as forage substrate on the fitness components of green oak leaf roller

The article is focused on the study of influence of genetic variation in DNA (RAPD-PCR, primer OPA 14) of oaks (Quercus pubescens L. and Q. petraea L.) on the variation of some fitness-linked traits in different genotype classes of the Tortrix viridana adults. The results show that the leafroller individuals from the same genotype class may possess the maximum fitness (fecundity and body size components) on the trees of one genotype class and the minimum fitness on the trees of another genotype class. The interaction of the "oak genotype"--"leafroller genotype" factors results in rise of the ties which are expressed in statistically reliable associations between the leafroller and oak genotype classes. The results are discussed from the point of view of the newly developed field of community or ecosystem genetics.     

Differences between understorey and canopy in herbivore community composition and leaf quality for two oak species in Missouri

1. From July 1994 to September 1995, at six censuses, the herbivore community associated with understorey (< 2.5 m height) and canopy (15–20 m) leaves of Quercus alba and Q. velutina was sampled in south-eastern Missouri, U.S.A. 2. Across all censuses, herbivore densities were not significantly different between canopy and understorey for Q. alba and Q. velutina, except in August 1994 when herbivore densities were 60% higher in the canopy on Q. alba. Little significant spatial variation in herbivore densities or community composition was found during the study years. 3. The herbivore community was diverse, consisting of 138 species of leaf-chewing insects. Species richness was significantly greater (by 5–20%) in the understorey than in the canopy for both tree species, and the relative abundance of the main families, different feeding guilds, and most common species differed significantly between understorey and canopy. 4. To determine the extent to which leaf quality explained the observed patterns, percentage nitrogen and protein binding capacity were measured in canopy and understorey leaves of Q. alba and Q. velutina. Per cent nitrogen was higher in canopy leaves for Q. velutina while protein binding capacity was higher in canopy leaves for Q. alba. 5. These results suggest that the herbivore community associated with these two species of Quercus comprises species that appear to respond individually to environmental and biological conditions encountered in the understorey and the canopy.