Differences In Herbivory Research Articles

Not so simple after all: searching for ecological advantages of compound leaves

Leaves come in many sizes and shapes, and the relationships between leaf traits and the environments they occur in are better understood every day. However we still know very little about the ecological consequences of plants having either compound or simple leaves. We attempted to address this knowledge gap by comparing chemical and physical characteristics (leaf area, length:width ratio, water content, leaf mass per area, ‘toughness’ and C:N ratio), as well as rates of herbivory between compound and simple leaves across 34 species in adjacent rainforest, open woodland and wet sclerophyll (tall open forest) vegetation in northeastern Australia. We found C:N ratio to be lower in simple leaves, but this was the only leaf trait that differed significantly between leaf types and did not stand up under phylogenetic analysis. Overall, we found no differences in herbivory between simple and compound leaves. While it remains unclear what the advantages of having one leaf type over another might be, the differences do not seem to lie in construction, or in vulnerability to herbivores, at least in the Australian Wet Tropics.

Total phenolics and individual phenolic acids vary with light environment inLindera benzoin

The understory shrub Lindera benzoin L. experiences lower rates of herbivory in sun environments than in shade environments. The production of secondary metabolites (e.g., phenolic compounds with known plant defense properties) is one likely contributor to these observed differences in herbivory. This work determined the total phenolic content as well as the concentrations of several individual phenolic acids in L. benzoin leaves found in sun and shade habitats. Total phenolic concentrations were determined to be higher in leaves from sun plants than in those from shade plants (47.5 ± 2.4 vs. 28.6 ± 1.3 gallic acid equivalents, respectively). High-performance liquid chromatography with diode array detection was used to separate and quantify several individual phenolic acids, and specific compounds were identified based on their retention times and ultraviolet spectra. The concentrations of vanillic, chlorogenic, p-coumaric, and ferulic acids were shown to be statistically higher in leaves from sun plants than in those from shade plants (P < 0.05), whereas 2,5-dihydroxybenzoic acid and caffeic acid were not significantly different in L. benzoin leaves from sun versus shade habitats.

Differential herbivory on disk and ray flowers of gynomonoecious asters and goldenrods (Asteraceae)

The selective advantage of gynomonoecy, the sexual system wherein plants produce a mixture of female and bisexual flowers, is poorly understood. One hypothesis for the evolution of this system is that the absence of androecia from female flowers reduces herbivore damage to the gynoecia of these flowers. Here, we examined patterns of herbivore damage in 53 collections representing 25 species of asters and goldenrods from Massachusetts, USA. In these taxa flowers are crowded into compact capitula, with bisexual flowers occupying the centre and female flowers situated on the periphery. Damage to gynoecia of bisexual flowers was significantly greater than damage to gynoecia of female flowers overall, and in about half of the individual populations. We also compared damage to central and peripheral flowers in the heads of 16 collections of other Asteraceae that produce only bisexual flowers to see whether the location of flowers rather than their sex might determine the patterns of herbivory. In only one of these 16 collections did we find a significant difference in herbivory between flower positions. We conclude that herbivore damage is influenced by flower type in asters and goldenrods, a pattern consistent with a role for herbivory in the evolution and maintenance of gynomonoecy. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 544–552.

Explaining differential herbivory in sun and shade: the case of Aristotelia chilensis saplings

Differential herbivory in contrasting environments is commonly explained by differences in plant traits. When several plant traits are considered, separate correlation analyses between herbivory and candidate traits are typically conducted. This makes it difficult to discern which trait best explain the herbivory patterns, or to avoid spurious inferences due to correlated characters. Aristotelia chilensis saplings sustain greater herbivory in shaded environments than in open habitats. We measured alkaloids, phenolics, trichomes, leaf thickness and water content in the same plants sampled for herbivory. We conducted a multiple regression analysis to estimate the relationship between herbivory and each plant trait accounting for the effect of correlated traits, thus identifying which trait(s) better explain(s) the differential herbivory on A. chilensis. We also estimated insect abundance in both light environments. Palatability bioassays tested whether leaf consumption by the main herbivore on A. chilensis was consistent with field herbivory patterns. Overall insect abundance was similar in open and shaded environments. While saplings in open environments had thicker leaves, lower leaf water content, and higher concentration of alkaloids and phenolics, no difference in trichome density was detected. The multiple regression analysis showed that leaf thickness was the only trait significantly associated with herbivory. Thicker leaves received less damage by herbivores. Sawfly larvae consumed more leaf tissue when fed on shade leaves. This result is consistent with field herbivory and, together with results of insect abundance, renders unlikely that the differential herbivory in A. chilensis was due to greater herbivory pressure in open habitats.

Correlated evolution of defensive and nutritional traits in native and non-native plants

We performed a comparative analysis of defensive and nutritional plant traits responsible for differential herbivory in a series of experimental feeding trials with generalist herbivores. We measured three defensive traits (leaf strength, leaf mass per unit area and endophytic fungal infection) and two nutritional traits (foliar nitrogen and water) for 26 native and eight non-native plant species from coastal California shrublands. Our feeding trials involved three species of generalist herbivore (beet armyworm, cabbage looper and the garden snail) in two types of laboratory feeding trial (single plant species and preference tests). All traits were significantly related to the amount of leaf area consumed, with foliar nitrogen followed by leaf strength explaining most of the variation in herbivore damage. Defensive and nutritional traits were tightly correlated with one another. These correlations were still apparent after incorporation of the phylogenetic relationships of species using independent contrasts, suggesting that there has been repeated selection for certain trait combinations. Non-native species had lower defensive traits and greater nutrient content and therefore experienced greater herbivory damage than natives. Poorly defended, nutrient-rich species (like most of the non-natives in our study) may be better suited for rapid growth and nutrient acquisition, thus reducing the cost of replenishing leaf material lost to herbivores.

Variability in leaf traits, insect herbivory and herbivore performance within and among individuals of four broad-leaved tree species

Abstract Individual plants may vary in their suitability as hosts for insect herbivores. The adaptive deme formation hypothesis predicts that this variability will lead to the fine-scale adaptation of herbivorous insects to host individuals. We studied individual and temporal variation in the quality of leaves of the tree species ash, lime, common oak, and sycamore in the field as food for herbivores. We determined herbivore attack and leaf consumption and performance of the generalist caterpillars of Spodoptera littoralis in the laboratory. We further assessed the concentrations of carbon, nitrogen and water in the leaves. All measures of leaf tissue quality varied among and within individuals for all tree species. The level of herbivory differed among the tree individuals in lime, oak and sycamore, but not in ash. Within host individuals, differences in herbivory between the upper and lower crown layer varied in direction and magnitude depending on tree species. In feeding experiments, herbivore performance also varied among and within tree individuals. However, variation in palatability was not consistently related to the leaf traits measured or to herbivory levels in the field. The ranking of individuals with respect to the quality of leaf tissue for herbivorous insects varied between years in lime and oak. Thus, trees of both species might present moving targets for herbivores which prevents fine-scale adaptations. In contrast, among individuals of ash and sycamore the pattern of insect performance remained constant over 2 years. These species may be more suitable hosts for the formation of adapted demes in herbivores.

Increased per capita herbivory in the shade: Necessity, feedback, or luxury consumption?

Leaf chemistry and physiology vary with light environment and are often thought to directly affect herbivory patterns. Biotic (e.g., parasitoids and predators) and abiotic (e.g., temperature, relative humidity) factors known to influence herbivory also co-vary with light environment. Irrespective of mechanism, light-based differences in herbivore damage must be the result of variable herbivore abundance, per capita effects, or both. We examined the effect of light environment on leaf defence and leaf nutritional quality in Lindera benzoin (Lauraceae) and relate this to the abundance and impact of its lepidopteran herbivore Epimecis hortaria (Lepidoptera: Geometridae). In this system we consistently observe greater natural field herbivory in shade habitats relative to high light habitats, despite similar herbivore abundances; differences in herbivory are therefore most likely attributable to different per capita impacts of herbivores across environments. Potential herbivore behaviours responsible for the observed field pattern include increased foraging per day and longer developmental periods in shade habitats. A more complete understanding of observed herbivory patterns requires incorporating variation in herbivore behaviour as influenced by abiotic or biotic factors that co-vary with the different light environments.

DIRECT AND INDIRECT EFFECTS OF CO2, NITROGEN, AND COMMUNITY DIVERSITY ON PLANT–ENEMY INTERACTIONS

Resource abundance and plant diversity are two predominant factors hypothesized to influence the amount of damage plants receive from natural enemies. Many impacts of these environmental variables on plant damage are likely indirect and result because both resource availability and diversity can influence plant traits associated with attractiveness to herbivores or susceptibility to pathogens. We used a long-term, manipulative field experiment to investigate how carbon dioxide (CO2) enrichment, nitrogen (N) fertilization, and plant community diversity affect plant traits and the amount of herbivore and pathogen damage experienced by the common prairie legume Lespedeza capitata. We detected little evidence that CO2 or N affected plant traits; however, plants growing in high-diversity treatments (polycultures) were taller, were less pubescent, and produced thinner leaves (higher specific leaf area). Interestingly, we also detected little evidence that CO2 or N affect damage. Plants growing in polycultures compared to monocultures, however, experienced a fivefold increase in damage from generalist herbivores, 64% less damage from specialist herbivores, and 91% less damage from pathogens. Moreover, within diversity treatments, damage by generalist herbivores was negatively correlated with pubescence and often was positively correlated with plant height, while damage by specialist herbivores typically was positively correlated with pubescence and negatively associated with height. These patterns are consistent with changes in plant traits driving differences in herbivory between diversity treatments. In contrast, changes in measured plant traits did not explain the difference in disease incidence between monocultures and polycultures. In summary, our data provide little evidence that CO2 or N supply alter damage from natural enemies. By contrast, plants grown in monocultures experienced greater specialist herbivore and pathogen damage but less generalist herbivore damage than plants grown in diverse communities. Part of this diversity effect was mediated by changes in plant traits, many of which likely are plastic responses to diversity treatments, but some of which may be the result of evolutionary changes in response to these long-term experimental manipulations.

Comparing Effects of Nutrients on Algal Biomass in Streams in Two Regions with Different Disturbance Regimes and with Applications for Developing Nutrient Criteria

Responses of stream algal biomass to nutrient enrichment were studied in two regions where differences in hydrologic variability cause great differences in herbivory. Around northwestern Kentucky (KY) hydrologic variability constrains invertebrate biomass and their effects on algae, but hydrologic stability in Michigan (MI) streams permits accrual of high herbivore densities and herbivory of benthic algae. Multiple indicators of algal biomass and nutrient availability were measured in 104 streams with repeated sampling at each site over a 2−month period. Many measures of algal biomass and nutrient availability were positively correlated in both regions, however the amount of variation explained varied with measures of biomass and nutrient concentration and with region. Indicators of diatom biomass were higher in KY than MI, but were not related to nutrient concentrations in either region. Chl a and % area of substratum covered by Cladophora were positively correlated to nutrient concentrations in both regions. Cladophora responded significantly more to nutrients in MI than KY. Total phosphorus (TP) and total nitrogen (TN) explained similar amounts of variation in algal biomass, and not significantly more variation in biomass than dissolved nutrient concentrations. Low N:P ratios in the benthic algae indicated N as well as P may be limiting their accrual. Most observed responses in benthic algal biomass occurred in nutrient concentrations between 10 and 30 μg TP l−1 and between 400 and 1000 μg TN l−1.

Spatially variable propagule pressure and herbivory influence invasion of chaparral shrubland by an exotic grass

Although numerous studies have identified mechanisms that either resist or facilitate biological invasions, few studies have explicitly tested how resisting and facilitating mechanisms interact to drive invasion success. In California, USA, undisturbed Mediterranean-type shrublands have resisted invasion by the perennial tussock grass Cortaderia jubata. In some cases, however, this resistance has been spectacularly breached even in the absence of large-scale disturbance. I tested the hypothesis that these invasions are facilitated by local reductions in the strength of biotic resistance. I evaluated invasive success using C. jubata seed and seedling additions at different microhabitats: the edge of a chaparral stand, under shrub canopy at different distances from the stand edge, and in canopy gaps within the stand. When left exposed to mammalian herbivores, seedling survivorship decreased sharply from nearly 40% on the stand edge to zero just 10 m into the stand. When transplants were protected from herbivory, however, distance from the edge had no significant influence on transplant survivorship. Seedling emergence was also greater on the edge and in canopy gaps than under the canopy, but these differences were not caused by differences in herbivory. The flux of invasive propagules reaching the soil surface was immense and greater along the edge and within gaps than under the stand canopy. Mirroring these patterns, naturally occurring seedling abundance declined dramatically with distance from the stand edge, and seedlings were far more common within stand gaps than would be expected given gap frequency within the stand. Despite strong biotic resistance to invasion within the stand, the cover of C. jubata has increased 20% over the last 9 years. These results suggest that the relative amount of susceptible edge habitat and the supply of invasive propagules can facilitate invasion even in the face of strong local biotic resistance.

Factors limiting the intertidal distribution of the mangrove species Xylocarpus granatum.

The tree species Xylocarpus granatum is commonly described as occurring in the upper intertidal zone of mangrove forests, but mature trees are occasionally found at lower elevations. In the Utwe River basin, on the Pacific island of Kosrae, we investigated the relative importance of several biotic and abiotic factors that may control the intertidal distribution of X. granatum. Factors we evaluated included differential seed predation across the lower, mid, and upper intertidal zones and seedling responses to salinity, tidal flooding, and shade. Seed predation was 22.4% over the first 34 days and varied little among zones or in gaps versus under the forest canopy. By day 161, there were still no differences in seed mortality, but a significant difference was found in seedling establishment, with much greater establishment in the upper intertidal plots. X. granatum seedlings in a greenhouse experiment exhibited greater growth in freshwater than seedlings in 23 ppt salinity, which is typical of salinity levels found in the mid intertidal zone in our field study sites in Micronesia, where mature X. granatum trees are generally absent. Seedlings grown in 23 ppt salinity, however, exhibited few visible signs of stress associated with patterns in growth. Seedlings grown in a simulated tidal flooding treatment (with 23 ppt salinity) also showed few signs of stress. Growth declined dramatically under 80% shade cloths, but there were few interactions of shading with either 23 ppt salinity or simulated tidal flooding. Differential seed predation is not likely to be the primary factor responsible for the intertidal distribution of X. granatum on Kosrae. However, seedling tolerance of flooding or salinity may be more important, especially relative to a potential contribution to secondary stress mortality. Other factors may ultimately prove to be more critical, such as physiological effects of salinity on seed germination, effects of tides on seed dispersal and rooting, or differential herbivory on seedlings.

Effects of four mono- and sesquiterpenes on the consumption of alfalfa pellets by sheep.

Effects of individual terpenes on alfalfa pellet intake by lambs were examined in four experiments. Forty-five lambs (nine lambs/treatment) were individually fed alfalfa pellets sprayed with either camphene, myrcene, caryophyllene oxide, or beta-pinene at one of five concentrations (one terpene per experiment). Treatments (0, 0.5, 1, 2, and 10x) were multiples of the concentration (x) of a specific terpene in tarbush. Terpenes were applied to alfalfa pellets (0.64 kg x lamb(-1) x d(-1), DM basis), and consumption was measured during a 20-min interval for 5 d. Lambs were maintained and fed alfalfa pellets in one group (except during 20-min tests) at a mean total daily intake of 4.7% of BW (DM basis). Camphene and caryophyllene oxide tended to decrease intake (linear contrasts were P = .0651 and P = .0504, respectively), whereas myrcene and beta-pinene exerted no effect on the consumption of alfalfa pellets by lambs. Camphene and caryophyllene oxide may be involved in the differential herbivory of individual tarbush plants by livestock.

Flowering, growth and defence in the two sexes: consequences of herbivore exclusion for Salix polaris

Summary For a long time, dioecious plants have been a model system for understanding the interactions between plants and herbivores. Differences in growth rate and, consequently, investment in defence between sexes may lead to skewed sex ratios due to differential herbivory. In this study we evaluated the applicability of this idea to polar willow (Salix polaris), which in the study site, Svalbard, displays a female‐biased sex ratio. Excluding reindeer for 3 years increased the abundance of male flowers in one of two vegetation types investigated. Growth rates differed only slightly between the sexes, with females investing more in inflorescences. The concentration of chemical defence compounds (phenolics and condensed tannins) did not differ between the sexes. On the basis of these findings, the idea that growth rate‐dependent herbivory caused the unbalanced sex ratio in S. polaris has to be rejected. Possibly an interaction of niche differentiation between male and female willows, in combination with reindeer grazing, produced the observed female‐biased sex ratio, but the mechanism remains unclear.

Do floral herbivores respond to variation in flower characteristics in Gelsemium sempervirens (Loganiaceae), a distylous vine?

We tested the hypothesis that traits traditionally thought to function in pollination can also influence attack by floral herbivores. Because distylous species produce two different flower types, we used Gelsemium sempervirens, a distylous vine of southeastern USA, to examine the frequency and pattern of floral herbivory in relation to floral characteristics. Flowers of the short-styled morph had larger corollas but showed no difference in the volume or concentration of nectar produced. Over the 2 yr of this study, 20-30% of plants suffered floral herbivory. The pattern of damage was morph-specific. Long-styled flowers were more likely to have damaged pistils, while stamens were more often damaged in short-styled flowers. In this distylous species, exserted flower organs were more likely to be eaten. Such differential herbivory based on the position of floral organs within a flower may result in reduced fitness via either male (short-styled morph) or female function (long-styled morph).

Herbivory in a fragmented tropical forest: Patterns from islands at Lago Gatún, Panama

By imposing density-dependent mortality upon their hosts, specialistinsect herbivores are thought to contribute to the maintenance of tree diversityin tropical forests. Forest fragmentation may alter patterns of herbivory,however, which may have important implications for tree species diversity inforest remnants. To explore effects of fragmentation on patterns of herbivory,we assessed folivory by Lepidopteran larvae on saplings of four focal treespecies on eight artificial, forested islands at Lago Gatun, Panama. Weexplored the importance of island area, distance to larger land, exposure to dryseason winds, tree species, and season in determining proportions of new leavesdamaged by caterpillars, and proportions of leaf area lost to caterpillars,during two dry and wet seasons. We found that both measures of herbivoryincreased markedly with island area, that island isolation had no apparenteffect on herbivory, and that interactions between season and exposure, andbetween tree species and season, were important determinants of herbivory rates.In addition, we observed species-specific differences in herbivory among hostplants under various conditions imposed by fragmentation. We conclude thatpatterns of herbivory by Lepidopteran larvae are sensitive to fragmentation inthis tropical forest. Differential herbivory among the four tree speciesconsidered here may have important implications for tree species dynamics on theislands of Lago Gatun.

Vigour of a dioecious shrub and attack by a galling herbivore

Summary1. The pattern of attack by the leaf‐galling insect Neopelma baccharidis (Homoptera: Psyllidae) was studied in three populations of the dioecious shrub Baccharis dracunculifolia (Asteraceae) in south‐eastern Brazil. The plant vigour hypothesis, which predicts higher rates of attack and increased herbivore performance on the longest plant shoots, was tested. This work also provides further information for the study of differential herbivory in dioecious plants.2. In total, 9200 shoots were collected randomly from 46 male and 47 female plants belonging to the three populations. Shoot length, number of leaves per shoot, rate of galling, and survival of psyllids did not differ between male and female plants. Another population on the Campus of the Federal University of Minas Gerais was used only to determine the pattern of shoot growth.3. The hypothesis of sex‐mediated herbivory was not corroborated in this study.4. The frequency of galling increased with increasing shoot length, as predicted by the plant vigour hypothesis. Nevertheless, the number of oviposition sites (leaf buds) increased with shoot length.5. The performance of the galling herbivore was not related to shoot length in the plant populations studied.6. In conclusion, Neopelma baccharidis did not select shoots based on length only.

Leaf Phenology and Leaf Damage of Saplings in the Luquillo Experimental Forest, Puerto Rico1

ABSTRACTChanges in light or water availability can result in synchronous leaf production, concentrating food availability for herbivores of young leaves to only a few months. To determine the importance of food availability on herbivory, leaf phenology and leaf damage were studied in the Luquillo Experimental Forest (LEF) of Puerto Rico. We studied 20 individuals of eight species for two years. Every month, new leaves were marked; the following month, leaf area and area of damage were measured. Over two years, comparison of leaf production and percent herbivory were performed for each species, and for all species taken together. More than 30 percent of the annual leaf production occurred in May and June. Leaf production was associated with an increase in PFD (photon flux density) and was not related to the patterns of rainfall. Although leaf production was synchronous, there were no differences in herbivory between the peak and non‐peak periods of leaf production. Possible explanations for the constant levels of herbivory throughout the year are the presence of a generalist herbivore community, the ability of herbivores to track changes in food availability, or high densities of herbivore predators that control herbivore populations.

Effect of individual terpenes on consumption of alfalfa pellets by sheep.

We examined effects of individual terpenes on alfalfa pellet intake of lambs in five experiments. Forty-five lambs (nine lambs/treatment) were individually fed alfalfa pellets sprayed with either p-cymene, alpha-humulene, 1,8-cineole, 3-carene, or sabinene at one of five concentrations (one terpene per experiment). Treatments (0, .5, 1, 2, and 10x) were multiples of the concentration (x) of a specific terpene in tarbush that was related to differential herbivory by livestock in previous studies. Terpenes were applied to alfalfa pellets (.64 kg x lamb(-1) x d(-1), DM basis), and consumption was measured during a 20-min interval for 5 d. Lambs were adapted to handling and pen feeding for 10 d and were maintained and fed alfalfa pellets in one group (except during 20-min tests) at a mean total daily intake of 4.7% of BW (DM basis). None of the five compounds decreased alfalfa pellet consumption during the 20-min interval. These five mono- and sesquiterpenes do not seem to be responsible for differential herbivory of individual tarbush plants by livestock.

Glucosinolates and differential herbivory in wild populations of Brassica oleracea

Glucosinolates are known to elicit responses from Brassica herbivores in laboratory studies. To study their importance in interactions with herbivores in the field, glucosinolate profiles and levels of herbivory were ascertained for wild cabbage plants growing in four neighboring populations in the UK. Glucosinolate profiles differed between plant populations, but not between different habitats within populations. Within habitats, there was no link between individual plant glucosinolate profiles and herbivory by Pieris spp., slugs and snails, flea beetles or aphids. Plants attacked by the micromoth, Selania leplastriana, contained higher levels of 2–hydroxy-3–butenylglucosinolate and 3–indolylmethylglucosinolate than plants within the same population that were not attacked. It is concluded that the differences in glucosinolate profiles between the plant populations are unlikely to be due to differential selection pressures from herbivores feeding on the mature plants over the two years studies.

Sex ratio variation in dioecious plant species: A comparative ecological study of six species of Lindera (Lauraceae)

Sex ratio variation was investigated in natural populations of six dioecious shrub species of Lindera in Japan. Interspecific differences in sex ratio were examined in relation to patterns of population structure, floral dimorphism, fruit production and intersexual differences in herbivory. Sex ratios tended towards equality or bias in favour of males, except for populations of L. glauca, in which no male plants were found. Sex switching in individual plants was not observed. Although male flowers were generally larger in size than flowers of conspecific females, costs of flowering did not appear to influence sex ratio. However, a relation was found between high allocation to fruit production and increasing male fraction of populations. In addition, girths of flowering plants tended to be larger in females, which may indicate delay in age of the first reproduction compared to males. No intersexual difference was found in degree of leaf damage due to herbivory. Results of this study are discussed in the light of other studies on sex ratio variation in dioecious plant species.