Effects of forest edges, fruit display size, and fruit colour on bird seed dispersal in a New Zealand mistletoe,

This study examined how forest edges, fruit display size, and fruit colour influenced rates of seed dispersal in an endemic, bird-dispersed, New Zealand mistletoe species, Alepis flavida. To examine rates of seed dispersal, fruit removal rates were compared between plants growing on forest edges and in forest interior, and also between two morphs of plants with different coloured fruits. Two aspects of fruit display size were examined: plant size and the neighbourhood of conspecific plants. There was no overall difference in fruit removal rates on forest edges and in forest interior, but birds removed fruits from red-fruited plants at a faster rate than from orange-fruited plants. Proximity of plant neighbours interacted with edges to influence fruit removal rates. The smaller the distance to nearest neighbours, the greater the fruit removal rates for orange-fruited plants in both habitats, but this relationship was significant for red-fruited plants only in the interior. Plant size affected fruit removal rates for orange-fruited plants, but not for red-fruited plants, and these differences were consistent in both habitats. Thus, fruit colour had the strongest effects on rates of fruit removal in this system, but forest edges also affected fruit removal rates, via altering the effects of neighbouring plants. Although birds prefer red fruits, there appears to be little selection pressure against orange-fruited plants because fruit removal rates are very high for both morphs.     

Natural and artificial floral damage induces resistance in Nemophila menziesii (Hydrophyllaceae) flowers

Resistance to leaf herbivory is well-documented in plants. In contrast, resistance to herbivory in flowers has received very little attention, even though reproductive tissues are often essential for plant reproduction. Plants may protect reproductive tissues with a range of defenses from constitutive to induced, although ecological costs associated with constitutive defense or resistance are expected to be higher than costs associated with induced responses. Induced responses in flowers may be effective against floral herbivores while minimizing the negative impacts of resistance on pollinators. This study examines induced responses in Nemophila menziesii (Hydrophyllaceae), a plant that frequently receives high levels of floral herbivory. I report that natural caterpillar herbivory increased levels of resistance against caterpillars later in the season. Similarly, artificial clipping to flowers consistently reduced natural damage to flowers vs unclipped controls over two years. Neither whole-plant nor individual seed set was affected by the reduction of floral damage. Induced resistance in reproductive tissues may benefit plants that are exposed to both floral herbivory and pollinator activity and can be an important link between plant antagonists and plant mutualists.     

Heterogeneity in vertebrate and invertebrate herbivory and its consequences for New Zealand mistletoes

Vertebrate herbivores generally have greater effects than invertebrates on plants. However, few studies have investigated the effects of both invertebrate and vertebrate herbivores on a single plant species. In New Zealand, nationwide declines in mistletoe populations have often been attributed to possum herbivory, but never to insect herbivory. The main goal of the present study was to document levels of vertebrate and invertebrate herbivory on endemic New Zealand mistletoe plants to suggest whether herbivory is leading to mistletoe decline. In the present study, the annual amount of leaf loss from herbivory by the brushtail possum (Trichosurus vulpecula), insect herbivory and leaf abscission were measured in two populations each of three mistletoe species (Alepis flavida, Peraxilla colensoi, and Peraxilla tetrapetala, Loranthaceae). In two populations of each species from February 1997 to February 1998, abscission accounted for the most leaf loss (range 10–84% of total mean leaf area, mean 33%), whereas insects and possums usually removed small and similar amounts (less than 3%). Possum browse caused large amounts of abscission in only one population (A. flavida at Eglinton). Observed possum browse was more heterogeneous than insect browse among branches within a plant (possum coefficient of variation = 2.63, insect CV = 1.98, P < 0.001), among plants in a population (possum CV = 2.15, insect CV = 0.69, P < 0.001), and between populations (possum CV = 1.36, insect CV = 1.09). Moreover, insects damaged 100% of the study plants but never removed more than 16% of leaf area on a single plant, whereas possums only browsed 32% of the study plants but severely defoliated some plants. Thus, while the mean amount of biomass removed across a population may have important consequences for mistletoe survival, the effect of possums on mistletoe populations may also depend on the heterogeneity of browse among individuals in the population.     

The Seedling Community of Tropical Rain Forest Edges and Its Interaction with Herbivores and Pathogens1

Edge effects alter biotic interactions and forest regeneration. We investigated whether edge creation affected the seedling community and its interactions with herbivores and leaf‐fungal pathogens. In forest edges and interior sites in Chiapas, Mexico, we counted all woody seedlings and species (10–100 cm tall) present in 1‐m² plots, measured their size (height and leaf number), and examined them for the occurrence of herbivory and pathogen damage. We investigated relationships between levels of damage and size, species richness and density. Species composition and biotic damage varied greatly among sites and habitats (edge vs interior). Late‐successional species dominated the community, but richness was lower in interior sites and species similarity was greater among edges than among interiors. Nearly all species (95%) present at edges and interiors showed herbivory damage, whereas 76 percent of the species in edge plots and 68 percent in interior plots showed pathogen damage. Although leaf area damaged by herbivores was similar between habitats (average 9.2%), pathogen damage was three times greater in edge plots (1.85%) than in interior plots (0.57%). Size was positively related to biotic damage at both habitats. Relationships between herbivory and pathogen damage and between pathogen damage and leaf number were significant only for edges. Biotic damage was not related to density or species richness. Overall, plant community structure was similar between habitats, but biotic damage was enhanced at edges. Thus, disease spread at edges may arise as a threat to tropical rain forest vegetation.     

Effects of two types of herbivores on the population dynamics of a perennial herb

The joint effects of multiple herbivores on their shared host plant have received increasing interest recently. The influence of herbivores on population dynamics of their host plants, especially the relative roles of different types of damage, is, however, still poorly understood. Here, we present a modelling approach, including both deterministic and stochastic matrix modelling, to be used in estimating fitness effects of multiple herbivores on perennial plants. We examined the effects and relative roles of two specialist herbivores, a pre-dispersal seed predator, Euphranta connexa, and a leaf-feeding moth, Abrostola asclepiadis, on the population dynamics and long-term fitness of their shared host plant, a long-lived perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). We collected demographic data during 3 years and combined these data with the effects of natural levels of herbivory measured from the same individuals. We found that both seed predation and leaf herbivory reduced population growth of V. hirundinaria, but only very high damage levels changed the growth trend of the vigorously growing study populations from positive to negative. Demographic modelling indicated that seed predation had a greater impact on plant population growth than leaf herbivory. The effect of leaf herbivory was weaker and diminished with increasing level of seed predation. Evaluation of individual fitness components, however, suggested that leaf herbivory contributed more strongly to host plant fitness than seed predation. Our results emphasize that understanding the effects of a particular herbivore on plant population dynamics requires also knowledge on other herbivores present in the system, because the effect of a particular type of herbivory on plant population dynamics is likely to vary according to the intensity of other types of herbivory. Furthermore, evaluating herbivore impact from using individual fitness components does not necessarily reflect the long-term effects on total plant fitness.     

Influence of leaf herbivory,root herbivory,and pollination on plant performance in Cucurbita moschata

Abstract 1. Plants experience herbivory on many different tissues that can affect reproduction directly by damaging tissues and decreasing resource availability, or indirectly via interactions with other species such as pollinators. 2. This study investigated the combined effects of leaf herbivory, root herbivory, and pollination on subsequent damage, pollinator preference, and plant performance in a field experiment using butternut squash (Cucurbita moschata). Leaf and root herbivory were manipulated using adult and larval striped cucumber beetles (Acalymma vittatum F.), a cucurbit specialist. 3. Leaf herbivory reduced subsequent pistillate floral damage and powdery mildew (Sphaerotheca fuliginea) infection. In spite of these induced defences, the overall effect of leaf herbivory on plant reproduction was negative. Leaf herbivory reduced staminate flower production, fruit number, and seed weight. In contrast, root herbivory had a minimal impact on plant reproduction. 4. Neither leaf nor root herbivory altered pollinator visitation or floral traits, suggesting that reductions in plant performance from herbivory were as a result of direct rather than indirect effects. In addition, no measured aspect of reproduction was pollen limited. 5. Our study reveals that although leaf herbivory by the striped cucumber beetle can protect against subsequent damage, this protection was not enough to prevent the negative impacts on plant performance.     

Effects of above- and belowground herbivory on growth,pollination, and reproduction in cucumber

Plants experience unique challenges due to simultaneous life in two spheres, above- and belowground. Interactions with other organisms on one side of the soil surface may have impacts that extend across this boundary. Although our understanding of plant–herbivore interactions is derived largely from studies of leaf herbivory, belowground root herbivores may affect plant fitness directly or by altering interactions with other organisms, such as pollinators. In this study, we investigated the effects of leaf herbivory, root herbivory, and pollination on plant growth, subsequent leaf herbivory, flower production, pollinator attraction, and reproduction in cucumber (Cucumis sativus). We manipulated leaf and root herbivory with striped cucumber beetle (Acalymma vittatum) adults and larvae, respectively, and manipulated pollination with supplemental pollen. Both enhanced leaf and root herbivory reduced plant growth, and leaf herbivory reduced subsequent leaf damage. Plants with enhanced root herbivory produced 35% fewer female flowers, while leaf herbivory had no effect on flower production. While leaf herbivory reduced the time that honey bees spent probing flowers by 29%, probing times on root-damaged plants were over twice as long as those on control plants. Root herbivory increased pollen limitation for seed production in spite of increased honey bee preference for plants with root damage. Leaf damage and hand-pollination treatments had no effect on fruit production, but plants with enhanced root damage produced 38% fewer fruits that were 25% lighter than those on control plants. Despite the positive effect of belowground damage on honey bee visitation, root herbivory had a stronger negative effect on plant reproduction than leaf herbivory. These results demonstrate that the often-overlooked effects of belowground herbivores may have profound effects on plant performance.     

Interactive effects of soil fertility and herbivory on Brassica nigra

Soil nutrient availability may affect both the amount of damage that plants receive from herbivores and the ability of plants to recover from herbivory, but these two factors are rarely considered together. In the experiment reported here, I examined how soil fertility influenced both the degree of defoliation and compensation for herbivory for Brassica nigra plants damaged by Pieris rapae caterpillars. Realistic levels of defoliation were obtained by placing caterpillars on potted host plants early in the life cycle and allowing them to feed until just before pupation on the designated plant. Percent defoliation was more than twice as great at low soil fertility compared to high (48.2% and 21.0%, respectively), even though plants grown at high soil fertility lost a greater absolute amount of leaf area (38.2 cm² and 22.1 cm², respectively). At both low and high soil fertility, total seed number and mean mass per seed of damaged plants were equivalent to those of undamaged plants. Thus soil fertility did not influence plant compensation in terms of maternal fitness. However, the pathways used to achieve compensation in seed production were different at low and high soil fertility. At low soil fertility, relative leaf growth rates (area added per inital area per day) of damaged plants were drastically reduced over the second week of caterpillar feeding. Damaged plants recovered the leaf area lost to herbivory in the two weeks following insect removal by increasing leaf relative growth rates above the levels seen for undamaged plants, but the replacement of leaf tissue lost to herbivory came at the expense of stem biomass. At high soil fertility, relative leaf growth rates of damaged plants were similar to those of undamaged plants both over the second week of caterpillar feeding and following caterpillar removal, and stem biomass was not affected by herbivory. These results suggest that higher levels of soil nutrients increased the ability of plants to stay ahead of their herbivores as they were being eaten. Because damaged plants at high soil fertility were able to maintain leaf growth rates to a greater extent than damaged plants at low soil fertility, they did not fall as far behind undamaged plants over the period of insect feeding and did not have as much catching up to do after feeding ended to compensate for herbivory.     

Leaf herbivory and drought stress affect floral attractive and defensive traits in Nicotiana quadrivalvis

Adaptive phenotypic plasticity allows sessile organisms such as plants to match trait expression to the particular environment they experience. Plasticity may be limited, however, by resources in the environment, by responses to prior environmental cues, or by previous interactions with other species, such as competition or herbivory. Thus, understanding the expression of plastic traits and their effects on plant performance requires evaluating trait expression in complex environments, rather than across levels of a single variable. In this study, we tested the independent and combined effects of two components of a plant’s environment, herbivory and water availability, on the expression of attractive and defensive traits in Nicotiana quadrivalvis in the greenhouse. Damage and drought did not affect leaf nicotine concentrations but had additive and non-additive effects on floral attractive and defensive traits. Plants in the high water treatment produced larger flowers with more nectar than in the low water treatment. Leaf damage induced greater nectar volumes in the high water treatment only, suggesting that low water limited plastic responses to herbivore damage. Leaf damage also tended to induce higher nicotine concentrations in nectar, consistent with other studies showing that leaf damage can induce floral defenses. Our results suggest that there are separate and synergistic effects of leaf herbivory and drought on floral trait expression, and thus plasticity in response to complex environments may influence plant fitness via effects on floral visitation and defense.     

Leaf damage induces twining in a climbing plant

Successful climbing by vines not only prevents shading by neighbouring vegetation, but also may place the vines beyond ground herbivores. Here we tested the hypothesis that herbivory might enhance climbing in a vine species, and that such induced climbing should be greater in the shade. We assessed field herbivory in climbing and prostrate ramets of the twining vine Convolvulus arvensis. We evaluated plant climbing after mechanical damage in a glasshouse under both sun and shade conditions, and determined whether control and damaged plants differed in growth rate or photosynthetic capacity. Plants experienced greater herbivory when growing prostrate than when climbing onto companion plants, in both an open habitat and a shaded understorey. Experimental plants increased their twining rate on a stake after suffering leaf damage, in both high- and low-light conditions, and this induced climbing was not coupled to an increase in growth rate. Increased photosynthesis was associated with enhanced twining rate only in the shade. Herbivory may be an ecological factor promoting the evolution of a climbing habit in plants.     

Synergistic interactions between volicitin,jasmonic acid and ethylene mediate insect-induced volatile emission in Zea mays

Plants display differential responses following mechanical damage and insect herbivory. Both caterpillar attack and the application of caterpillar oral secretions (OS) to wounded leaves stimulates volatile emission above mechanical damage alone. Volicitin ( N- 17-hydroxylinolenoyl- l -glutamine), present in beet armyworm (BAW, Spodoptera exigua ) OS, is a powerful elicitor of volatiles in excised maize seedlings ( Zea mays cv. Delprim). We consider some of the mechanistic differences between wounding and insect herbivory in maize by examining the activity of volicitin, changes in jasmonic acid (JA) levels, and volatile emission from both intact plant and excised leaf bioassays. Compared to mechanical damage alone, volicitin stimulated increases in both JA levels and sesquiterpene volatiles when applied to intact plants. In a bioassay comparison, excised leaves were more sensitive and produced far greater volatile responses than intact plants following applications of both volicitin and JA. In the excised leaf bioassay, volicitin applications (10–500 pmol) to wounded leaves resulted in dose dependent JA increases and a direct positive relationship between JA and sesquiterpene volatile emission. Interestingly, volicitin-induced JA levels did not differ between intact and excised bioassays, suggesting a possible interaction of JA with other regulatory signals in excised plants. In addition to JA, insect herbivory is known to stimulate the production of ethylene. Significant increases in ethylene were induced only by BAW herbivory and not by either wounding or volicitin treatments. Using intact plant bioassays, ethylene (at 1 µl l⁻¹ or less) greatly promoted volatile emission induced by volicitin and JA but not mechanical damage alone. For intact plants, wounding, elicitor-induced JA and insect-induced ethylene appear to be important interacting components in the stimulation of insect-induced volatile emission.     

Effects of Patch Size and Floral Herbivory on Seed Set in Trichogoniopsis adenantha (Asteraceae) in Southeastern Brazil1

The effects of patch size and floral herbivory on reproductive success of Trichogoniopsis adenantha (Asteraceae) were studied in a semideciduous forest in southeastern Brazil. Plants in large patches and capitula not attacked by endophagous insects had higher seed set. The consequences of living in large or small patches for plant reproductive success are discussed.     

Floral trait expression and plant fitness in response to below- and aboveground plant–animal interactions

Although plant–animal interactions like pollination and herbivory are obviously interdependent, ecological investigations focus mainly on one kind of interaction ignoring the possible significance of the others. Plants with flowers offer an extraordinary possibility to study such mutualistic and antagonistic interactions since it is possible to measure changes in floral traits and fitness components in response to different organisms or combinations of them. In a three factorial common garden experiment we investigated single and combined effects of root herbivores, leaf herbivores and decomposers on flowering traits and plant fitness of Sinapis arvensis. Leaf herbivory negatively affected flowering traits indicating that it could significantly affect plant attractiveness to pollinators. Decomposers increased total plant biomass and seed mass indicating that plants use the nutrients liberated by decomposers to increase seed production. We suggest that S. arvensis faced no strong selection pressure from pollen limitation, for two reasons. First, reduced nutrient availability through leaf herbivory affected primarily floral traits that could be important for pollinator attraction. Second, improved nutrient supply through decomposer activity was invested in seed production and not in floral traits. This study indicates the importance of considering multiple plant–animal interactions simultaneously to understand selection pressures underlying plant traits and fitness.     

Floral herbivore effect on the sex expression of an andromonoecious plant,Isomeris arborea (Capparaceae)

Flower-feeding insects may influence the reproductive behavior of their host plant. In plants with labile sex expression, the ratio of maternal to paternal investment may change in response to damage, an effect that goes beyond the direct reduction of plant gametes. We examined the effects of floral herbivory by the beetle Meligethes rufimanus (Nitidulidae) on the ratio of hermaphroditic flowers to male flowers in an andromonoecious shrub, Isomeris arborea (Capparaceae) in southern California. Plants exposed to herbivory had a greater rate of flower bud abortion than those protected from herbivory. Exposed plants produced a greater proportion of hermaphroditic flowers to male flowers, although damaged inflorescences still produced fewer fruit. An additional manipulative experiment showed that the removal of pistils on inflorescences led to an increase in the proportion of hermaphroditic flowers. This suggests that the presence of fruit may lead to pistil suppression in developing flowers. Adaptive responses to herbivory which favor andromonoecy thus include the continued production of hermaphroditic flowers when floral damage is high (and hence low fruit set), and a switch to male flower production when floral damage is low (and fruit production increases). The consequences of an altered six ratio induced by insect herbivores may lead to indirect effects on both the male and female reproductive success of this plant.     

Phenotypic plasticity in seedling defense strategies: compensatory growth and chemical induction

Phenotypic plasticity in growth (leading to compensation) and secondary chemical production (leading to induction) in response to herbivory are key defense strategies in adult plants, but their role in seedling defense remains unclear. A pair of greenhouse studies was conducted to investigate compensation and induction in seedlings and juvenile plants, using Plantago lanceolata (Plantaginaceae) and the specialist buckeye caterpillar Junonia coenia (Nymphalidae) as a model system. Plants received 50% defoliation at two and four weeks of age, and groups of plants were harvested one week after herbivory and six to eight weeks after herbivory to investigate the duration of the responses. Plants damaged at two weeks showed no chemical induction and fully compensated for the lost leaf tissue by ten weeks of age. Plants damaged at four weeks showed a significant reduction in iridoid glycosides one week after herbivory and achieved full shoot compensation by ten weeks of age at the expense of root biomass. These results indicate that P. lanceolata seedlings use compensation, but not chemical induction, as a defense strategy. This research highlights the importance of considering ontogeny in studies of plant–herbivore interactions and suggests that seedling defense may differ markedly from adult plant defense.     

Bird predation enhances tree seedling resistance to insect herbivores in contrasting forest habitats

According to the associational resistance hypothesis, neighbouring plants are expected to influence both the insect herbivore communities and their natural enemies. However, this has rarely been tested for the effects of canopy trees on herbivory of seedlings. One possible mechanism responsible for associational resistance is the indirect impact of natural enemies on insect herbivory, such as insectivorous birds. But it remains unclear to what extent such trophic cascades are influenced by the composition of plant associations (i.e. identity of ‘associated’ plants). Here, we compared the effect of bird exclusion on insect leaf damage for seedlings of three broadleaved tree species in three different forest habitats. Exclusion of insectivorous birds affected insect herbivory in a species-specific manner: leaf damage increased on Betula pendula seedlings whereas bird exclusion had no effect for two oaks (Quercus robur and Q. ilex). Forest habitat influenced both the extent of insect herbivory and the effect of bird exclusion. Broadleaved seedlings had lower overall leaf damage within pine plantations than within broadleaved stands, consistent with the resource concentration hypothesis. The indirect effect of bird exclusion on leaf damage was only significant in pine plantations, but not in exotic and native broadleaved woodlands. Our results support the enemies hypothesis, which predicts that the effects of insectivorous birds on insect herbivory on seedlings are greater beneath non-congeneric canopy trees. Although bird species richness and abundance were greater in broadleaved woodlands, birds were unable to regulate insect herbivory on seedlings in forests of more closely related tree species.     

Florivory: the intersection of pollination and herbivory

Plants interact with many visitors who consume a variety of plant tissues. While the consequences of herbivory to leaves and shoots are well known, the implications of florivory, the consumption of flowers prior to seed coat formation, have received less attention. Herbivory and florivory can yield different plant, population and community outcomes; thus, it is critical to distinguish between these two types of consumption. Here, we consider the ecological and evolutionary consequences of florivory. A growing number of studies recognize that florivory is common in natural systems and in some cases surpasses leaf herbivory in magnitude and impact. Florivores can affect male and female plant fitness via direct trophic effects and through altered pathways of species interactions. In particular, florivory can affect pollination and have consequences for plant mating and floral sexual system evolution. Plants are not defenceless against florivore damage. Concepts of resistance and tolerance can be applied to plant–florivore interactions. Moreover, extant theories of plant chemical defence, including optimal defence theory, growth rate hypothesis and growth differentiation–balance hypothesis, can be used to make testable predictions about when and how plants should defend flowers against florivores. The majority of the predictions remain untested, but they provide a theoretical foundation on which to base future experiments. The approaches to studying florivory that we outline may yield novel insights into floral and defence traits not illuminated by studies of pollination or herbivory alone.     

Above‐ and belowground herbivory jointly impact defense and seed dispersal traits in Taraxacum officinale

Plants are able to cope with herbivores by inducing defensive traits or growth responses that allow them to reduce or avoid the impact of herbivores. Since above‐ and belowground herbivores differ substantially in life‐history traits, for example feeding types, and their spatial distribution, it is likely that they induce different responses in plants. Moreover, strong interactive effects on defense and plant growth are expected when above‐ and belowground herbivores are jointly present. The strengths and directions of these responses have been scarcely addressed in the literature. Using Taraxacum officinale, the root‐feeding nematode Meloidogyne hapla and the locust Schistocerca gregaria as a model species, we examined to what degree above‐ and belowground herbivory affect (1) plant growth responses, (2) the induction of plant defensive traits, that is, leaf trichomes, and (3) changes in dispersal‐related seed traits and seed germination. We compared the performance of plants originating from different populations to address whether plant responses are conserved across putative different genotypes. Overall, aboveground herbivory resulted in increased plant biomass. Root herbivory had no effect on plant growth. Plants exposed to the two herbivores showed fewer leaf trichomes than plants challenged only by one herbivore and consequently experienced greater aboveground herbivory. In addition, herbivory had effects that reached beyond the individual plant by modifying seed morphology, producing seeds with longer pappus, and germination success.     

The effect of shading on photosynthesis,growth, and regrowth following defoliation for Bromus tectorum

Summary The effect of full sunlight, 60%, or 90% attenuated light on photosynthetic rate, growth, leaf morphology, dry weight allocation patterns, phenology, and tolerance to clipping was examined in the glasshouse for steppe populations of the introduced grass, Bromus tectorum. The net photosynthetic response to light for plants grown in shade was comparable to responses for plants grown in full sunlight. Plants grown in full sunlight produced more biomass, tillers and leaves, and allocated a larger proportion of their total production to roots than plants grown in shade. The accumulation of root and shoot biomass over the first two months of seedling growth was primarily responsible for the larger size at harvest of plants grown in full sunlight. Plants grown under 60% and 90% shade flowered an average of 2 and 6 weeks later, respectively, than plants grown in full sunlight. Regrowth after clipping was greater for plants grown in full sunlight compared to those grown in shade. Even a one-time clipping delayed flowering and seed maturation; the older the individual when leaf area was removed, the greater the delay in its phenology. Repeated removal of leaf area was more frequently fatal for plants in shade than in full sunlight. For plants originally grown in full sunlight, regrowth in the dark was greater than for shaded plants and was more closely correlated to non-flowering tiller number than to plant size. This correlation suggests that etiolated regrowth is more likely regulated by the number of functional meristems than by differences in the size of carbohydrate pools. Thus, shading reduces the rate of growth, number of tillers, and ability to replace leaf area lost to herbivory for B. tectorum. These responses, in turn, intensify the effect of competition and defoliation for this grass in forests. B. tectorum is largely restricted to forest gaps at least in part because of its inability to acclimate photosynthetically, the influence of shade on resource allocation, and the role of herbivory in exacerbating these effects.     

Effects of below- and above-ground herbivores on plant growth,flower visitation and seed set

Separate and combined effects of root and leaf herbivores on plant growth, flower visitation and seed set were tested in a factorial experiment using potted mustard, Sinapis arvensis, at an old fallow field. A 50% leaf removal by cabbageworms (Pieris rapae) when the seedlings had their first four leaves reduced plant height and shoot mass, and delayed the onset of flowering. Root herbivory by two wireworms (Agriotes sp.) over the whole experiment changed flower visitation; the number of flower visitors per plant was higher in plants with root herbivores than in plants without root herbivores. Combined leaf and root herbivory affected flowering period, number of fruits per plant and number of seeds per fruit. Plants attacked by leaf and root herbivores had a shorter flowering period and produced fewer fruits per plant than plants with root herbivores only. Although the experimental plants faced major herbivore-induced growth changes, plant reproduction (seed set and weight per plant) was similar in all treatments, documenting their ability to effectively compensate for leaf and root herbivory.