The structure of the littoral: effects of waterlily density and perch predation on sediment and plant‐associated macroinvertebrate communities

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Density-related foraging behaviour in Closterocerus tricinctus, a parasitoid of the leaf-mining moth, Cameraria hamadryadella

• 1 We examined the foraging behaviour of the parasitoid wasp, Closterocerus tricinctus (Ashmead) (Chalcidoidea: Eulophidae), as it visited larvae of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae), in an outbreak population.
• 2 We tracked females of C.tricinctus, recording the time spent searching for mines and handling host larvae. The density of leaf-mines (host larvae) and their condition were recorded for each leaf visited. A subset of leaves visited by C.tricinctus was enclosed in fine mesh bags so that foraging success could be determined by rearing or dissection. The average density of mines and the average leaf-area mined was estimated for a random sample of leaves from each tree.
• 3 The selection of leaves upon which to forage appears to be density-dependent. C.tricinctus visits leaves with leaf-mine densities twice the average, and when switching leaves lands directly on leaf-mines 5 times more often than expected assuming random landings.
• 4 The total time spent foraging on a leaf, the average time spent handling hosts, and the total search time within leaves tend to decline on leaves with many hosts, but the observed declines are not statistically significant.
• 5 The proportion of leaf-mines visited within a leaf is strongly inversely density-dependent. 30% of visits to leaf-mines are re-visits and 29% of handling time is spent re-handling previously visited hosts. Furthermore, only 21% of visits to mines lead to successful parasitism. We suggest that self-interference and the avoidance behaviour of the host may reduce the number of visits of leaf-mines by C.tricinctus within a leaf.
• 6 The effect of the strongly inversely density-dependent foraging investment within leaves is to offset the observed density-dependent pattern of leaf visitation making the overall spatial pattern of visitation by C.tricinctus to mines of C.hamadryadella inversely density-dependent.
• 7 We suggest that the uncertainty of C.tricinctus surviving on multiply mined leaves because of density-dependent host mortality due to intraspecific competition in high-density host populations, the rarity of high-density host populations, and the rarity of multiply-mined leaves in low-density host populations combine to select against an aggregative response within leaves by C.tricinctus.

     

Impacts of defoliation by tussock moths (Orgyia vetusta) on the growth and reproduction of bush lupine (Lupinus arboreus)

Abstract.

• 1 Outbreaks of tussock moths (Orygia verusta Bdv.) on bush lupines (Lupinus arboreus Sims) may be very intense, although spatially localized, and may last >10 years. To understand better how such outbreaks may persist, we defoliated lupine bushes using variable realistic numbers of tussock moth larvae, and measured the immediate and delayed impacts of defoliation on the growth and seed production of lupine bushes.
• 2 Immediate effects of defoliation included the production of new leaves that were significantly smaller in diameter and biomass, and lower in water content, than the new leaves of undamaged bushes. Also, seed output was reduced by up to 80% on heavily damaged bushes.
• 3 In the longer term, surviving bushes recovered remarkably well, despite having received the full range of possible (survivable) insect densities. After 1 or 2 years’rapid growth, surviving bushes were not affected in terms of height, basal stem diameter or volume by their previous defoliation. However, there was a significant tendency for juvenile bushes to produce more seeds the more heavily they had been attacked the previous year.
• 4 The ability of the host plant to recover, in terms of its biomass, helps to explain the sustained nature of the insect outbreak.

     

Perfect is best: low leaf fluctuating asymmetry reduces herbivory by leaf miners

Fluctuating asymmetry (FA) represents small, random variation from symmetry and can be used as an indicator of plant susceptibility to herbivory. We investigated the effects of FA of two oak species, Quercus laevis and Q. geminata, and the responses of three herbivore guilds: leaf miners, gallers, and chewers. To examine differences in FA and herbivory between individuals, 40 leaves from each tree were collected, and FA indices were calculated. To examine differences in FA and herbivory within-individuals, we sampled pairs of mined and unmined leaves for asymmetry measurements. Differences in growth of leaf miners between leaf types were determined by tracing 50 mines of each species on symmetric leaves and asymmetric leaves. Asymmetric leaves contained significantly lower concentrations of tannins and higher concentrations of nitrogen than symmetric leaves for both plant species. Both frequency of asymmetric leaves on plants and levels of asymmetry positively influenced the abundance of Brachys, Stilbosis and other leaf miners, but no significant relationship between asymmetry and herbivory was observed for Acrocercops. Brachys and Stilbosis mines were smaller on asymmetric leaves, but differences in mine survivorship between symmetric and asymmetric leaves were observed only for Stilbosis mines. This study indicated that leaf miners might use leaf FA as a cue to plant quality, although differential survivorship among leaf types was not observed for all species studied. Reasons for the different results between guilds are discussed.     

The influence of host plant genotype on variation in population densities of the gorse thrips,Sericothrips staphylinus (Thysanoptera: Thripidae), and its consideration in relation to release strategies

Large variations in population densities of Sericothrips staphylinus were observed between gorse bushes at a field site in an earlier Tasmanian study. A replicated potted plant experiment was conducted to assess whether genetic variation between gorse bushes could affect densities of S. staphylinus. There were significant differences in the number of S. staphylinus recovered from the individual bushes sampled within sites and individual bushes across all sites. This provides evidence that genetic variation between gorse bushes is at least partly responsible for the observed variability in field populations. The result validates the current release strategy used for the field establishment of S. staphylinus in south-eastern Australia. This strategy involves making multiple small releases (ca. 250 adults per bush per site) instead of a small number of large releases (ca. 1000 per bush per site), thereby reducing variation in numbers due to genetic affects. Genetic variation between individual plants of the same species and associated effects on herbivore densities should be considered when devising release strategies for biological control agents.     

Natural selection drives leaf divergence in experimental populations of Senecio lautus under natural conditions

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The effects of host plant phenology on the demography and population dynamics of the leaf-mining moth, Cameraria hamadryadella (Lepidoptera: Gracillariidae)

Abstract.

• 1 We examined the effects of variation in the timing of spring leaf production and autumn leaf fall on the survival, mortality and abundance of Cameraria hamadryadella on Quercus alba and Q.macrocarpa.
• 2 We monitored and manipulated the timing of foliation on field and potted Q.alba trees and observed the abundance of C.hamadryadella on those trees. We also monitored and manipulated the timing of leaf fall on Q.alba and Q.macrocarpa trees in the field and observed its effects on survival, mortality and abundance of C.hamadryadella.
• 3 Variation in the timing of spring leaf production has no effect on C. hamudryadella abundance. However, a warm winter and spring in 1991 led to accelerated development and the imposition of a facultative third generation in one out of ten years of observation.
• 4 In 1989, leaves fell relatively early and leaf fall in the autumn accounted for more than 50% of the mortality of C.hamudryadella. in 1990 and 1991 leaves fell relatively late and leaf fall induced mortality was substantially reduced and overwinter survival was markedly increased.
• 5 The abundance of C.hamadryadella remained constant in the spring and summer of 1990 following the previous autumn's relatively early leaf fall, but increased by 10-fold in the spring of 1991 following the relatively late leaf fall of autumn 1990. The abundance of C.hamadryadella also increased 4-fold between the summer of 1991 and the spring of 1992 after another autumn of relatively late leaf fall. We attribute these increases in abundance in part to reduced mortality because of later leaf fall.
• 6 Variation in the timing of autumn leaf fall may be responsible for initiating outbreaks of C.hamadryadella.

     

DEVELOPMENTAL STABILITY IN LEAVES OF CLARKIA TEMBLORIENSIS (ONAGRACEAE) AS RELATED TO POPULATION OUTCROSSING RATES AND HETEROZYGOSITY

Four natural populations of Clarkia tembloriensis, whose levels of heterozygosity and rates of outcrossing were previously found to be correlated, are examined for developmental instability in their leaves. From the northern end of the species range, we compare a predominantly selfing population (t? = 0.26) with a more outcrossed population (t? = 0.84), which is genetically similar. From the southern end of the range, we compare a highly selfing population (t? = 0.03) with a more outcrossed population (t? = 0.58). We measured developmental stability in the populations using two measures of within-plant variation in leaf length as well as calculations of fluctuating asymmetry (FA) for several leaf traits. Growth-chamber experiments show that selfing populations are significantly more variable in leaf length than more outcrossed populations. Developmental instability can contribute to this difference in population-level variance. Plants from more homozygous populations tend to have greater within-plant variance over developmentally comparable nodes than plants from more heterozygous populations, but the difference is not significant. At the upper nodes of the plant, mature leaf length declines steadily with plant age, allowing for a regression of leaf length on node. On average, the plants from more homozygous populations showed higher variance about the regression (MSE) and lower R² values, suggesting that the decline in leaf length with plant age is less stable in plants from selfing populations than in plants from outcrossing populations. Fluctuating asymmetry (FA) was calculated for four traits within single leaves at up to five nodes per plant. At the early nodes of the plant where leaf arrangement is opposite, FA was also calculated for the same traits between opposite leaves at a node. Fluctuating asymmetry is significantly greater in the southern selfing population than in the neighboring outcrossed population. Northern populations do not differ in FA. Fluctuating asymmetry can vary significantly between nodes. The FA values of different leaf traits were not correlated. We show that developmental stability can be measured in plants using FA and within-plant variance. Our data suggest that large differences in breeding system are associated with differences in stability, with more inbred populations being the least stable.     

Effect of leaf litter and density on fitness and population performance of the hole mosquito Aedes triseriatus

Abstract.

• 1 We tested the effects of food, in the form of leaf litter, and density on population growth and fitness correlates of the tree hole mosquito Aedes triseriatus. Our field experiment used a 2 x 2 randomized block design, with three holes as blocks. In cages within three holes, we manipulated densities of Aedes triseriatus larvae, and presence of leaf litter. Our laboratory experiment used a 2 x 3 factorial design in which we also manipulated densities of larvae and availability of leaf litter within similar cages (inside, outside, or absent). For both experiments we determined effects on survival, days to and mass at eclosion of both sexes, and a composite index of population performance λ′, that estimates finite rate of increase.
• 2 In the field experiment, we found significant and large effects of leaf litter and of density on the performance of A. triseriatus. The effect of density did not depend on leaf litter availability. We documented significant variation among tree holes for most correlates of fitness. Often, the effects of treatments varied significantly depending on the tree hole tested.
• 3 In the laboratory experiment we found significant effects of density and litter, and that direct browsing on leaf litter is necessary for the effect of litter on performance of A. triseriatus. In the laboratory, the negative effect of increased density was dependent on leaf litter availability.
• 4 Our results demonstrate that leaf litter has important effects on population performance of tree hole mosquitoes in natural tree holes. Direct browsing on the surface of leaf litter is the most likely mechanism by which litter enhances population growth. The impact of litter on populations varies among tree holes.

     

Growing larger with domestication: a matter of physiology,morphology or allocation?

• Domestication might affect plant size. We investigated whether herbaceous crops are larger than their wild progenitors, and the traits that influence size variation.
• We grew six crop plants and their wild progenitors under common garden conditions. We measured the aboveground biomass gain by individual plants during the vegetative stage. We then tested whether photosynthesis rate, biomass allocation to leaves, leaf size and specific leaf area (SLA) accounted for variations in whole‐plant photosynthesis, and ultimately in aboveground biomass.
• Despite variations among crops, domestication generally increased the aboveground biomass (average effect +1.38, Cohen's d effect size). Domesticated plants invested less in leaves and more in stems than their wild progenitors. Photosynthesis rates remained similar after domestication. Variations in whole‐plant C gains could not be explained by changes in leaf photosynthesis. Leaves were larger after domestication, which provided the main contribution to increases in leaf area per plant and plant‐level C gain, and ultimately to larger aboveground biomass.
• In general, cultivated plants have become larger since domestication. In our six crops, this occurred despite lower investment in leaves, comparable leaf‐level photosynthesis and similar biomass costs of leaf area (i.e. SLA) than their wild progenitors. Increased leaf size was the main driver of increases in aboveground size. Thus, we suggest that large seeds, which are also typical of crops, might produce individuals with larger organs (i.e. leaves) via cascading effects throughout ontogeny. Larger leaves would then scale into larger whole plants, which might partly explain the increases in size that accompanied domestication.

     

Seed germination and seedling allogamy in Rosmarinus officinalis: the costs of inbreeding

• Self‐pollination by geitonogamy is likely in self‐compatible plants that simultaneously expose a large number of flowers to pollinators. However, progeny of these plants is often highly allogamous. Although mechanisms to increase cross‐pollination have been identified and studied, their relative importance has rarely been addressed simultaneously in plant populations.
• We used Rosmarinus officinalis to explore factors that influence the probability of self‐fertilisation due to geitonogamy or that purge its consequences, focusing on their effects on seed germination and allogamy rate. We experimentally tested the effect of geitonogamy on the proportion of filled seeds and how it influences germination rate. During two field seasons, we studied how life history and flowering traits of individuals influence seed germination and allogamy rates of their progeny in wild populations at the extremes of the altitudinal range. The traits considered were plant size, population density, duration of the flowering season, number of open flowers, flowering synchrony among individuals within populations and proportion of male‐sterile flowers.
• We found that most seeds obtained experimentally from self‐pollination were apparently healthy but empty, and that the proportion of filled seeds drove the differences in germination rate between self‐ and cross‐pollination experiments. Plants from wild populations consistently had low germination rate and high rate of allogamy, as determined with microsatellites. Germination rate related positively to the length of the flowering season, flowering synchrony and the ratio of male‐sterile flowers, whereas the rate of allogamous seedlings was positively related only to the ratio of male‐sterile flowers.
• Rosemary plants purge most of the inbreeding caused by its pollination system by aborting the seeds. This study showed that the rates of seed germination and allogamy of the seedlings depend on a complex combination of factors that vary in space and time. Male sterility of flowers, length of the flowering season and flowering synchrony of individuals within populations all favour high rates of cross‐pollination, therefore increasing germination and allogamy rates. Flowering traits appear to be highly plastic and respond to local and seasonal conditions.

     

Defensive role of leaf trichomes in resistance to herbivorous insects in Datura stramonium

This study assessed the role of leaf trichome density as a component of resistance to herbivores, in six populations of Datura stramonium. Phenotypic selection on plant resistance was estimated for each population. A common garden experiment was carried out to determine if population differences in leaf trichome density are genetically based. Among population differences in leaf trichome density, relative resistance and fitness were found. Leaf trichome density was strongly positively correlated to resistance across populations. In 5 out of 6 populations, trichome density was related to resistance, and positive directional selection on resistance to herbivores was detected in three populations. Differences among populations in mean leaf trichome density in the common garden suggest genetic differentiation for this character in Datura stramonium. The results are considered in the light of the adaptive role of leaf trichomes as a component of defence to herbivores, and variable selection among populations.     

The effects of temperature on leaf growth of sugar beet varieties

Leaf growth of nine varieties of sugar beet (Beta vulgaris L.) was studied at constant temperatures of 7, 11, 15 and 20·C, using generalised logistic curves fitted to the data to estimate the parameters of growth. The rate of leaf appearance increased linearly with temperature and was the same in all varieties. There were differences between varieties in the weighted mean rates of expansion of leaf area per plant (ā), the temperature coefficient of ā and the leaf area duration (D); these differences were caused more by differences in rates of expansion and final sizes of individual leaves than by differences in rates of leaf production. The growth of the first six leaves produced by each plant was examined in detail. The greater size of successive leaves of plants and genotypic differences between comparable leaves were more attributable to differences in the rate than differences in the duration of leaf expansion. Increasing temperatures increased leaf size because they accelerated the rate of expansion more than they shortened the duration of the expansion phase. It is inferred that all effects arose through differences in the initial sizes of leaves before they unrolled from the shoot apex. Dry matter production was proportional to D but was partitioned more to the storage root at the colder temperatures. This may have been related to the differential effects of temperature on cell division and expansion and the relative contribution of these two processes to the final sizes of the leaves and storage root.     

Leaf age and larval performance of the leaf beetle Paropsis atomaria

ABSTRACT.

• 1 Larval performance of the leaf beetle Paropsis atomaria Oliver was determined for larvae raised on both new and mature leaves of Eucalyptus blakelyi Maiden. Larvae were transferred to mature leaves at different ages; control larvae stayed on new leaves through all instars.
• 2 Only larvae reared on new leaves through the third instar survived to pupate on mature leaves; developmental time was prolonged by 20% and pupal weight was reduced by 50% in these larvae compared with larvae reared entirely on new leaves. Almost all larvae died when transferred to mature leaves as first, second or third instars.
• 3 Low survival and slow development on mature leaves was mainly due to failure by larvae to feed. Larvae palpated leaves and could discriminate among leaf ages immediately, without biting into the leaf tissue.
• 4 New leaves had higher concentrations of oil and tannins than old leaves, while there were no significant differences in nitrogen concentrations in the two types of leaves. Mature leaves were more than 3 times tougher than new leaves, in terms of g mm^?2 of penetrometer force.
• 5 In drought years E. blakelyi may not produce sufficient new leaves to supply specialist herbivores with their preferred food resource. We infer that drought years reduce P. atomaria larval performance significantly, and influence the population dynamics of the insect.

     

High levels of fecundity in small and isolated populations of a self‐compatible Aloe pollinated by opportunistic birds and bees

• Plant species that are effective colonisers of transient habitats are expected to have a capacity for uniparental reproduction and show flexibility in pollination systems. Such traits may enable populations to be established from a small number of founding individuals without these populations succumbing to reductions in fecundity arising from pollinator limitation.
• We tested these predictions for Aloe thraskii (Xanthorrhoeaceae), a succulent treelet that colonises shifting coastal dunes and has both bird and bee pollinators. We performed hand‐pollination experiments, and selectively excluded bird visitors to determine differences in pollinator effectiveness. We measured pollinator visitation rates and fecundity in populations varying in their size, density and isolation distance.
• Controlled hand‐pollinations revealed that unlike most other Aloe species, A. thraskii is self‐compatible and thus capable of uniparental reproduction. The species does however depend on pollinators and is visited by various bird species as well as by bees. Fruit and seed set are not affected by selective exclusion of birds, thus indicating that bees are effective pollinators. Bird visitation rates increased with increasing plant height and population size, while bee visitation rates increased with increasing population size and density. We found that seed set per flower was lower in large populations than in small populations.
• These results suggest that establishment of populations of A. thraskii from a small number of individuals is unlikely to be limited by the fecundity of individual plants.

     

Plant‐mediated effects of host plant density on a specialist herbivore of Solanum carolinense

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Effects of trichomes on the behavior and distribution of Platyprepia virginalis caterpillars

Trichomes are an important physical resistance mechanism of plants, as they reduce insect herbivore movement, feeding, and digestion. However, we know little about how trichomes influence herbivore distributions and populations. We conducted laboratory and field experiments to evaluate the preferences of Platyprepia virginalis (Boisduval) (Lepidoptera: Arctiidae) caterpillars to natural and manipulated densities of trichomes on their primary food, Lupinus arboreus Sims (Fabaceae). We then conducted field surveys to determine whether variation in trichome density among lupine bushes affected caterpillar spatial distribution on the landscape. Platyprepia virginalis caterpillars preferred lupine leaves with fewer trichomes in choice and no‐choice experiments. In the field, caterpillar feeding damage was found more often on leaves with fewer trichomes. These preferences scaled up to the level of bushes in the landscape such that more caterpillars were found on bushes with lower trichome densities than on bushes with higher trichome densities. This is one of few studies to show the potential for trichome density to influence herbivore population size and distribution in a natural system at a landscape level. The results are consistent with trichomes functioning as a resistance mechanism with consequences for herbivore choice, performance, and distribution.     

Spatial patterns and infestation processes in the horse chestnut leafminer Cameraria ohridella: a tale of two cities

Spatial patterns of the horse chestnut leafminer Cameraria ohridella Deschka & Dimic (Lepidoptera: Gracillariidae) population density was analysed in the cities of Bern and Brussels in order to explore its spatial population dynamics. The surveys were carried out in Bern in 1998 and 2000 and in Brussels in 2001 to assess population density in relation to local characteristics. In Brussels, population density was also measured using pheromone traps distributed over the city and collected twice per moth generation. A quantitative relationship was found between local population density (measured by pheromone traps and survey observations) and the amount of leaves left on the ground the previous fall. Several other factors were related to observed infestation levels: the occurrence of the pathogen Guignardia aesculi was inversely related to infestation by C. ohridella in Bern in 1998 and 2000, the number of horse chestnut trees within 800 m distance was positively related to infestation level in Bern in 2000, and the proportion of green areas within 100 m and the number of other horse chestnut trees within 2000 m were positively related to infestation levels in the 2001 Brussels survey. The pattern of infestation levels as a function of distance to potential population reservoirs suggested that C. ohridella re‐invades areas where overwintering leaves have been cleaned from refuge areas such as parks or urban forests. Our results indicate that the removal of leaves is a feasible first aid control measure to reduce moth population densities. However, leaf removal may not reduce moth densities when done improperly. In places where proper leaf removal is not feasible, other control measures are needed.     

Variability in grape phylloxera preference and performance on canyon grape (Vitis arizonica)

We tested the deme-formation hypothesis experimentally with four populations of leaf-galling grape phylloxera, Daktulosphaira vitifoliae, and its host, canyon grape (Vitis arizonica). An experiment designed to examine preference and performance showed that phylloxera populations did not significantly prefer their original host relative to other hosts in the percent available leaves galled. There were significant herbivore population effects (P<0.01), host effects (P<0.001), and population x host interaction effects (P<0.001). Herbivore populations had different colonizing abilities (performance, as measured in the mean number of galls per leaf) on an individual host (P<0.001), but there was no host effect. Host genotype significantly affected phylloxera performance, measured as survivorship (P<0.01), but a phylloxera population did not necessarily have higher survivorship on its original host. Differences in fecundity, an-other measurement of performance, were due to intrinsic differences among herbivore populations (P<0.05), and not related to host genotype. There was no correlation between distance from a phylloxera population in the field and a host's susceptibility to attack. There was a significant positive relationship between levels of infestation on a clone in the field and its susceptibility to colonization experimentally (P<0.05), suggesting inherent differences in host resistance and susceptibility. These results did not support the deme-formation hypothesis. In a second experiment, host clone x water treatment interactions affected phylloxera survivorship (P<0.05) and fecundity (P<0.05). We conclude that host genotype x environment interactions may prevent sessile, parthenogenetic herbivores from locally adapting to individual host genotypes.     

Localized outbreak of a willow leaf beetle: plant vigor or natural enemies?

The possible roles of plant quality (vigor) and natural enemies in the development of a localized out-break of the leaf beetle Galerucella lineola (Coleoptera: Chrysomelidae) in a stand of Salix cinerea were investigated. Caged and uncaged larvae on six bushes in the outbreak area were compared with caged and uncaged larvae on six bushes in an adjacent nonoutbreak area in terms of performance. In 1997, when the studies were performed, the natural density of the insect (beetles plus eggs) was six times higher in the outbreak area compared with the nonoutbreak area. Even though the vigor (measured as shoot length) of bushes in the outbreak was 72% higher than that of bushes in the nonoutbreak area, we found no difference between areas in the performance (survival, developmental time, pupal weight) of caged larvae or in the willingness of caged females to lay eggs. Among larvae exposed to natural enemies, the disappearance rate was significantly higher in the nonoutbreak area. The density of generalist predators was significantly higher in the nonoutbreak than in the outbreak area. We conclude that differences in plant quality, despite the observed difference in plant vigor, could not explain the observed difference in beetle density between areas. Lower predation pressure in the outbreak area could, however, not be excluded as a possible reason for the higher density of leaf beetles in this area. Received: October 18, 1999 / Accepted: February 4, 2000