Interactions of plant stress and herbivory: intraspecific variation in the susceptibility of a palatable versus an unpalatable seaweed to sea urchin grazing

Summary Interactions among environmental stresses, plant defensive characteristics, and plant nutrient status may significantly affect an alga's susceptibility to herbivores. Following desiccation, the palatable seaweed Gracilaria tikvahiae was less susceptible to grazing by the sea urchin Arbacia punctulata while the unpalatable alga Padina gymnospora became more susceptible. Increased grazing on desiccated Padina appeared to result from a loss of chemical defenses following desiccation. Palatable plants treated with organic extracts from desiccated Padina plants were consumed at more than twice the rate of plants treated with extracts from undesiccated plants. Increased susceptibility of Padina did not correlate with changes in protein content of the alga; reduced grazing on desiccated Gracilaria was associated with a decrease in protein content. When Padina was grazed by Arbacia or mechanically damaged to mimic urchin grazing, its susceptibility to Arbacia decreased within 1 to 5 days. These results demonstrate that history of physical or biological stress may affect a plant's susceptibility to herbivory. We hypothesize that urchins cue primarily on attractiveness features (e.g. nutrient content) of highpreference algae and deterrent features (e.g. chemical defenses) of low-preference algae. Stresses may therefore increase, decrease, or not affect a plant's susceptibility to herbivory depending upon the primary feeding cues used by the herbivore, the defensive mechanisms used by the plant, and the way these are altered by various environmental stresses.     

Plant genotypic diversity reduces the rate of consumer resource utilization

While plant species diversity can reduce herbivore densities and herbivory, little is known regarding how plant genotypic diversity alters resource utilization by herbivores. Here, we show that an invasive folivore—the Japanese beetle (Popillia japonica)—increases 28 per cent in abundance, but consumes 24 per cent less foliage in genotypic polycultures compared with monocultures of the common evening primrose (Oenothera biennis). We found strong complementarity for reduced herbivore damage among plant genotypes growing in polycultures and a weak dominance effect of particularly resistant genotypes. Sequential feeding by P. japonica on different genotypes from polycultures resulted in reduced consumption compared with feeding on different plants of the same genotype from monocultures. Thus, diet mixing among plant genotypes reduced herbivore consumption efficiency. Despite positive complementarity driving an increase in fruit production in polycultures, we observed a trade-off between complementarity for increased plant productivity and resistance to herbivory, suggesting costs in the complementary use of resources by plant genotypes may manifest across trophic levels. These results elucidate mechanisms for how plant genotypic diversity simultaneously alters resource utilization by both producers and consumers, and show that population genotypic diversity can increase the resistance of a native plant to an invasive herbivore.     

Effects of plant diversity and time of colonization on an herbivore-plant interaction

Summary Experimental field plantings showed that plant diversity strongly affected the population dynamics of a specialist herbivore, the striped cucumber beetle, Acalymma vittata (Fab.) (Coleoptera: Chrysomelidae). Population densities over time were characterized by two peaks in numbers (from colonization and reproduction, respectively) and were consistently higher in cucumber monocultures (Cucumis sativus L.) than in polycultures of cucumbers, corn (Zea mays L.), and broccoli (Brassica oleracea L.). Greater abundances in monocultures appear to result from two factors: (1) per individual reproductive rates were greater in monocultures than in polycultures, and (2) mark-recapture studies confirmed that beetles stay in monocultures for a longer period of time than in polycultures. Differences in predation did not appear to contribute to the overall differences in herbivore abundances.The primary impact of A. vittata on its host plant, C. sativus, is the dissemination of bacterial wilt disease, Erwinia tracheiphila (E.F.Sm.). Greater numbers of beetles led to greater plant mortality in monocultures. It is suggested that factors other than numbers of beetles (e.g., shading, allelopathy, microclimate) are more important in influencing plant reproduction, since cucumber plants in monocultures had greater yields than did plants grown in polycultures. However, time of beetle colonization strongly affected plant parameters, indicating that the length of time during which herbivores are interacting with plants is of critical importance to plant survivorship, and thus reproduction.     

Parasitism in diversified agroecosystems: Phenology ofTrichogramma minutum [Hymenoptera: Trichogrammatidae]

Parasitism ofOstrinia nubilalis egg masses byTrichogramma minutum was observed in maize-bare ground monocultures and polycultures of maize/bean/squash and maize/clover. Parasitism rates were 1.9 times higher in monocultures than in polycultures; seasonal phenology of parasitism, however, was similar in both. Parasitism first occurred during the late whorl/early tassel stage of maize; peak parasitism occurred during the mid to late tassel stage, and then dropped off rapidly. We speculate that direct predation of egg masses, and ammensal preemptive competition for egg masses byColeomegilla maculata may have contributed to the decline in parasitism rates in both monocultures and polycultures during the latter part of the season.      

The effects of chrysomelid beetle grazing and plant competition on the growth of Rumex obtusifolius

Summary The interaction between grazing by Gastrophysa viridula and interspecific plant competition was investigated for Rumex obtusifolius growing in the field. During an eight-month growing season non-competing R. obtusifolius grew larger than competing plants although herbivore loads (beetle productivity per unit leaf area) were similar for competing and non-competing plants. However, grazing significantly reduced the growth of R. obtusifolius only in competing plants thus demonstrating a synergism between plant competition and invertebrate herbivory. The relevance of these findings for the distribution of plant species is discussed.     

Neighbours matter: natural selection on plant size depends on the identity and diversity of the surrounding community

Plant diversity can affect ecological processes such as competition and herbivory, and these ecological processes can act as drivers of evolutionary change. However, surprisingly little is known about how ecological variation in plant diversity can alter selective regimes on members of the community. Here, we examine how plant diversity at two different scales (genotypic and species diversity) impacts natural selection on a focal plant species, the common evening primrose (Oenothera biennis). Because competition is frequently relaxed in both genotypically and species rich plant communities, we hypothesized that increasing diversity would weaken selection on competitive ability. Changes in plant diversity can also affect associated arthropod communities. Therefore, we hypothesized that diversity would alter selection on plant traits mediating these interactions, such as herbivory related traits. We grew 24 focal O. biennis genotypes within four different neighbourhoods: genotypic monocultures or polycultures of O. biennis, and species monocultures or polycultures of old-field species that commonly co-occur with O. biennis. We then measured genotypic selection on nine plant traits known to be ecologically important for competition and herbivory. Focal O. biennis plants were smaller, flowered for shorter periods of time, had lower fitness, and experienced greater attack from specialist predispersal seed predators when grown with conspecifics versus heterospecifics. While neither conspecific nor heterospecific diversity altered trait means, both types of diversity altered the strength of selection on focal O. biennis plants. Specifically, selection on plant biomass was stronger in conspecific monocultures versus polycultures, but weaker in heterospecific monocultures versus polycultures. We found no evidence of selection on plant traits that mediate insect interactions, despite differences in arthropod communities on plants surrounded by conspecifics versus heterospecifics. Our data demonstrate that plant genotypic and species diversity can act as agents of natural selection, potentially driving evolutionary changes in plant communities.     

Effects of pepper (Capsicum chinense) genotypic diversity on insect herbivores

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Effects of plant diversity on invertebrate herbivory in experimental grassland

The rate at which a plant species is attacked by invertebrate herbivores has been hypothesized to depend on plant species richness, yet empirical evidence is scarce. Current theory predicts higher herbivore damage in monocultures than in species-rich mixtures. We quantified herbivore damage by insects and molluscs to plants in experimental plots established in 2002 from a species pool of 60 species of Central European Arrhenatherum grasslands. Plots differed in plant species richness (1, 2, 4, 8, 16, 60 species), number of functional groups (1, 2, 3, 4), functional group and species composition. We estimated herbivore damage by insects and molluscs at the level of transplanted plant individuals (“phytometer” species Plantago lanceolata, Trifolium pratense, Rumex acetosa) and of the entire plant community during 2003 and 2004. In contrast to previous studies, our design allows specific predictions about the relative contributions of functional diversity, plant functional identity, and species richness in relation to herbivory. Additionally, the phytometer approach is new to biodiversity-herbivory studies, allowing estimates of species-specific herbivory rates within the larger biodiversity-ecosystem functioning context. Herbivory in phytometers and experimental communities tended to increase with plant species richness and the number of plant functional groups, but the effects were rarely significant. Herbivory in phytometers was in some cases positively correlated with community biomass or leaf area index. The most important factor influencing invertebrate herbivory was the presence of particular plant functional groups. Legume (grass) presence strongly increased (decreased) herbivory at the community level. The opposite pattern was found for herbivory in T. pratense phytometers. We conclude that (1) plant species richness is much less important than previously thought and (2) plant functional identity is a much better predictor of invertebrate herbivory in temperate grassland ecosystems.     

Herbivory on coral reefs: algal susceptibility to herbivorous fishes

Summary The susceptibility of several tropical algal species to fish grazing was studied on the Belizean barrier reef off the Caribbean coast of Central America. Short-term transplant experiments indicate that plant species vary markedly in their rates of biomass loss to grazing by a shallow-water guild of herbivorous fishes. Algal species transplanted from habitats with low grazing pressure are highly susceptible to grazing, while species occurring in habitats with high herbivore densities are highly resistant to grazing. Algal species show differential susceptibility to grazing by two major components of the tropical herbivore guild, Acanthurus (surgeonfishes) and Sparisoma (parrotfishes).Variability in plant susceptibility to grazing by herbivorous fishes was not clearly correlated with morphological or chemical characteristics that have been previously suggested as plant defenses against herbivory. Plants found to be highly resistant to fish grazing, such as Halimeda, exhibit both morphological characteristics and secondary chemical compounds which do appear to reduce herbivory. In contrast, species of Caulerpa, Sargassum, Turbinaria, and Padina, which also possess alleged morphological and/or chemical defenses, are nevertheless highly susceptible to fish grazing.     

Effects of marine algal proximate composition on methane yields

The suitability of differentGracilaria spp. and twoSargassum species for bioconversion to methane was determined through bioassays of methane yield.Gracilaria species and strains were excellent feedstocks for high methane yields, ranging from 0.28 to 0.40 m³ kg^–1 volatile solids added. These yields ranged from 58 to 95% of theoretical stoichiometric yields. Methane yields were highly correlated with acid soluble carbohydrate components of theGracilaria spp. BothSargassum fluitans andS. pteropleuron were poor feedstocks, with methane yields ranging from 0.12 to 0.19 m³ kg^–1 volatile solids added, 27 to 46% of theoretical stoichiometric yields, respectively. The various tissue types of theseSargassum species were also poor feedstocks for anaerobic digestion to methane. While there is no clear explanation for the low methane yields, the twoSargassum spp. appear to contain a high proportion of an insoluble, non-extractable component which may not be available as a substrate for bioconversion to methane.     

Tree species diversity alters plant defense investment in an experimental forest plantation in southern Mexico

How plant species diversity affects traits conferring herbivore resistance (e.g., chemical defenses), as well as the mechanisms underlying such effects, has received little attention. One potential mechanism for the effect of diversity on plant defenses is that increased plant growth at high diversity could lead to reduced investment in defenses via growth–defense trade‐offs. We measured tree growth (diameter at breast height) and collected leaves to quantify total phenolics in 2.5‐year‐old plants of six tropical tree species (N = 597 plants) in a young experimental plantation in southern Mexico. Selected plants were classified as monocultures or as polycultures represented by mixtures of four of the six species examined. Tree species diversity had a significant negative effect on total phenolics, where polycultures exhibited a 13 percent lower mean concentration than monocultures. However, there was marked variation in the effects of diversity on defenses among tree species, with some species exhibiting strong reductions in phenolic levels in mixtures, whereas others were unresponsive. In addition, tree species diversity had no effect on growth, nor was the negative effect of diversity on chemical defenses mediated by a growth–defense trade‐off. These results demonstrate that tree diversity can alter investment in chemical defenses in long‐lived tree species but that such effect may not always be under strong control by plant endogenous resource allocation trade‐offs. Regardless of the underlying mechanism, these findings have important implications for predicting effects on consumers and ecosystem function.     

Elevated nutrient content of tropical macroalgae increases rates of herbivory in coral,seagrass, and mangrove habitats

We explored the role of food quality in herbivore preference for macroalgae by comparing consumption of Acanthophora spicifera with and without elevated tissue nitrogen and phosphorus concentrations. Algal enrichment effects on herbivory were examined in coral, seagrass, and mangrove habitats along a sparsely populated Honduran island protected from fishing. Nutrient enrichment led to significantly increased grazing by herbivores across habitats. Consumption of enriched algae increased by 91% compared to controls among the mangrove roots, where herbivory rates were generally lowest. In the heavily grazed seagrass and coral habitats, nutrient enrichment increased consumption by 30 and 20%, respectively, with the effect more spatially variable than among the mangrove roots. We suggest that, at least on the local scale, intact herbivore populations may be able to compensate for effects of increased nutrient supply by locating and consuming nutrient-enriched algae, but that the importance of this mechanism varies both among and within habitats.Communicated by: R. C. Carpenter     

Effects of aphid-tending Argentine ants, nitrogen enrichment and early-season herbivory on insects hosted by a coastal shrub

The presence of the exotic Argentine ant, Linepithema humile Mayr (Hymenoptera: Dolichoderinae), nitrogen enrichment, and early-season herbivory by the specialist beetle Trirhabda bacharidis (Coleoptera: Chrysomelidae) have been shown, through separate experiments, to affect the densities of insect herbivores of the coastal shrub Baccharis halimifolia (Asteraceae), in Florida. Using a fully-factorial field experiment, we examined the relative importance of all three of these factors to the six most common insect herbivore species utilizing this host plant in a West Central Florida coastal habitat. The presence of ants affected more herbivore species than either early-season herbivory by larval T. bacharidis or nitrogen enrichment. Experimental reductions of L. humile resulted in reductions of an aphid, its coccinellid predators, and adult T. bacharidis, and increases of two species of leafminers and one species of stemborer. Due to the strong negative effects of stemborer herbivory on host plant survival, the increase in stemborer abundance led to increased host plant mortality. Early-season herbivory by larval T. bacharidis only affected the abundance of aphids and their predators, both of which were more abundant on trees with reduced early-season herbivory. Nitrogen fertilization had the most limited effects and only T. bacharidis larvae achieved higher densities on fertilized trees. Our results indicate that aphid tending by the exotic L. humile affects other insects on B. halimifolia more so than herbivory by the exploitative competitor T. bacharidis or nitrogen as a limiting nutrient.     

Herbivore response to vegetational diversity: spatial interaction of resources and natural enemies

Vegetational diversity in agricultural systems is predicted to reduce herbivore populations, but we observed the opposite effect: higher nymph population densities of a functionally monophagous herbivore, the squash bug, Anasa tristis (Hemiptera: Coreidae) in a vegetationally diverse squash-bean-corn polyculture than in a squash monoculture. We examined spatial and temporal aspects of squash bug and predator populations in relation to vegetational diversity. Average colonization, oviposition, and mortality rates for the herbivore were similar in monocultures and polycultures. In the polyculture, however, we found that squash bugs eggs were highly aggregated on plants on the outer edges of plots. Predation was also lower on plants near the edges, allowing the large aggregations of eggs found in the polyculture to escape predation and ultimately produce more squash bugs. Spatial interactions between herbivores and natural enemies may underlie some of the general effects of vegetational diversity on herbivores.     

Density of Anomoea flavokansiensis on Desmanthus illinoensis in monoculture and polyculture

The effect of plant species diversity on the density of the herbivore, Anomoea flavokansiensis Moldenke (Coleoptera: Chrysomelidae), on Desmanthus illinoensis (Michaux) MacMillan (Mimosaceae), a promising North American legume for exploring the principles of diverse, perennial grain agriculture was examined. From mid-June to early August A. flavokansiensis feeds on young leaves and inflorescences of D. llinoensis. At high density, A. flavokansiensis potentially reduces seed yield and is thus an important consideration for long-term stands that are to be grown without insecticides. The potential to manage this insect via intercropping its host species with other, non-host perennial species by monitoring A. flavokansiensis density on D. illinoensis within experimental monocultures, two-species mixtures with Tripsacum dactyloides, and three-species mixtures with T. dactyloides and Leymus racemosus was examined. Insects were censused 2–3 times weekly from mid-June to early August at two sites from 1991 to 1995. In the first three years, beetle density was generally low (<1 per plant), and did not differ among treatments. In the fourth year, however, beetle density peaked at 15 and 25 insects per plant at the two sites, and was highest within monoculture for most dates. In 1995, density was again low, but tended to remain higher in monoculture at one site. The results suggest that beetle density on D. illinoensis can be reduced in polyculture and may hold promise for the management of this insect herbivore within perennial grain polycultures.     

Can tropical seaweeds reduce herbivory by growing at night? Diel patterns of growth,nitrogen content,herbivory, and chemical versus morphological defenses

Summary Tropical seaweeds in the genus Halimeda reduce losses to grazing by capitalizing on diel patterns of herbivore activity. These seaweeds produce new, more herbivoresusceptible growth at night when herbivorous reef fishes are inactive. Plant portions more than 48 h old are low in food value, well defended morphologically (calcified and high in ash content), and relatively resistant to herbivory. Younger plant portions represent 3–4.5 times the food value (nitrogen or organic content) of older portions but are only moderately more susceptible to herbivores due to their high concentrations of the terpenoid feeding deterrents halimedatrial and halimedatetraacetate. Halimedatrial significantly deters grazing by both parrotfishes (Scaridae) and surgeonfishes (Acanthuridae) and occurs in high concentrations (2–4.5% of plant ash-free dry mass) in plant portions that are 4–12 h old, intermediate concentrations (0.3–2.3%) in portions that are 16–26 h old, and low concentrations (0.3%) in older plant portions. The related compound halimedatetraacetate is absent from the youngest plant portions, shows a rapid increase in concentration (from 0 to 1%) in plant material that is approximately 16 h old, and then rapidly declines to low levels (0.1 to 0.2%) in older plant portions. Thus, newly produced tissues are nutritionally valuable but contain high concentrations of defensive chemicals. As these tissues age, morphological defenses increase, the tissue becomes less valuable as a food for herbivores, and chemical defenses decrease. Additionally, new growth of Halimeda remains unpigmented until just before sunrise. Thus, the valuable, nitrogen-containing molecules associated with photosynthesis are not placed in the new, and more herbivore susceptible, growth until lights is available and they can start producing income for the plant.Experiments in a coral-reef microcosm, where diel patterns of light and water chemistry could be altered, indicated that Halimeda's growth pattern is cued by the timing of light-dark cycles rather than by co-occurring diel changes in water chemistry. Although the growth patterns of Halimeda seem unusual, similar patterns appear to occur in numerous other seaweeds and in microalgae such as diatoms and dinoflagellates.     

Effects of tree species diversity on a community of weaver spiders in a tropical forest plantation

The effects of producer diversity on predators have received little attention in arboreal plant communities, particularly in the tropics. This is particularly true in the case of tree diversity effects on web‐building spiders, one of the most important groups of invertebrate predators in terrestrial plant communities. We evaluated the effects of tree species diversity on the community of weaver spiders associated with big‐leaf mahogany (Swietenia macrophylla) in 19, 21 × 21‐m plots (64 plants/plot) of a tropical forest plantation which were either mahogany monocultures (12 plots) or polycultures (seven plots) that included mahogany and three other tree species. We conducted two surveys of weaver spiders on mahogany trees to evaluate the effects of tree diversity on spider abundance, species richness, diversity, and species composition associated with mahogany. Our results indicated that tree species mixtures exhibited significantly greater spider abundance, species richness, and diversity, as well as differences in spider species composition relative to monocultures. These results could be due to species polycultures providing a broader range of microhabitat conditions favoring spider species with different habitat requirements, a greater availability of web‐building sites, or due to increased diversity or abundance of prey. Accordingly, these results emphasize the importance of mixed forest plantations for boosting predator abundance and diversity and potentially enhancing herbivore pest suppression. Future work is necessary to determine the specific mechanisms underlying these patterns as well as the top‐down effects of increased spider abundance and species richness on herbivore abundance and damage.     

Scales of dispersal among hosts in a herbivorous marine amphipod

Abstract The spatial arrangement of plants may constrain the expression of herbivore preferences due to interaction between the scales of herbivore mobility and vegetation structure. I tested the hypothesis that the presence of the herbivorous marine amphipod Peramphithoe parmerong (Ampithoidae) on a low‐preference, poor‐quality host (the brown alga Padina crassa) was due to limited dispersal and the clumped nature of algal distributions. In contrast to limited dispersal in laboratory assays, amphipods rapidly colonized vacant algae in the field with natural densities being achieved within 1 day. These high rates of colonization did not decline with increasing distance from existing algal beds over the spatial scales separating Pa. crassa from the higher‐quality host algae (Sargassum spp.). Thus, the spatial arrangement of host algae did not interact with herbivore mobility to constrain host choices.     

Strong feeding preference of an exotic generalist herbivore for an exotic forb: a case of invasional antagonism

Many hypotheses dealing with the success of invasive plant species concern plant–herbivore interactions. The invasional meltdown and enemy inversion hypotheses suggest that non-native herbivores may indirectly facilitate the invasion of a non-native plant species by either favorably changing environmental conditions or reducing competition from native plant species. Our objective was to determine the role of herbivory by the non-native snail Otala lactea in structuring California grassland communities. We conducted two experiments to examine the feeding preferences of O. lactea for eight representative grassland species. Overall, O. lactea preferred Brassica nigra, a non-native forb, over all other species tested. Field monocultures of B. nigra supported significantly higher snail densities than monocultures of any of the other species tested. O. lactea also preferred B. nigra over all other species tested in controlled laboratory feeding trials. However, based on trait comparisons of each of the eight grassland species, we cannot pinpoint the preference for B. nigra to a basic nutritional requirement on the part of the herbivore or an allocation to defense on the part of the plants. Our study provides evidence for an antagonistic relationship between a non-native herbivore and a non-native plant species in their invasive range. We term this relationship “invasional antagonism”.     

Evidence for rapid evolutionary change in an invasive plant in response to biological control

We present evidence that populations of an invasive plant species that have become re‐associated with a specialist herbivore in the exotic range through biological control have rapidly evolved increased antiherbivore defences compared to populations not exposed to biocontrol. We grew half‐sib families of the invasive plant Lythrum salicaria sourced from 17 populations near Ottawa, Canada, that differed in their history of exposure to a biocontrol agent, the specialist beetle Neogalerucella calmariensis. In a glasshouse experiment, we manipulated larval and adult herbivory to examine whether a population's history of biocontrol influenced plant defence and growth. Plants sourced from populations with a history of biocontrol suffered lower defoliation than naïve, previously unexposed populations, strongly suggesting they had evolved higher resistance. Plants from biocontrol‐exposed populations were also larger and produced more branches in response to herbivory, regrew faster even in the absence of herbivory and were better at compensating for the impacts of herbivory on growth (i.e. they exhibited increased tolerance). Furthermore, resistance and tolerance were positively correlated among genotypes with a history of biocontrol but not among naïve genotypes. Our findings suggest that biocontrol can rapidly select for increased defences in an invasive plant and may favour a mixed defence strategy of resistance and tolerance without an obvious cost to plant vigour. Although rarely studied, such evolutionary responses in the target species have important implications for the long‐term efficacy of biocontrol programmes.