Plant choice,herbivory and weight gain of wood crickets under the risk of predation

Predators can indirectly reduce herbivory by killing herbivores. In addition, predation risk can influence the feeding rate and feeding location of herbivores. Herbivores are expected to avoid plants currently occupied by a predator. Consequently, less herbivory is expected on plants bearing fresh predator cues. We examined whether wood crickets, Nemobius sylvestris Bosc (Orthoptera: Gryllidae), avoided plants bearing the chemical cues of nursery web spiders, Pisaura mirabilis Clerck (Araneae: Pisauridae), or red wood ants, Formica rufa L. (Hymenoptera: Formicidae). We conducted a series of behavioural experiments, in which crickets had the choice between a plant with spider or ant cues vs. a control plant, a plant with spider cues vs. a plant with ant cues, or two control plants. For all plants, we quantified leaf damage and the position and weight change in the crickets. Crickets avoided plants with spider cues. In contrast, ant cues did not significantly deter crickets. The herbivory pattern among the plants reflected the plant choice of the crickets. However, net herbivory was not affected by the presence of predator cues. Thus, our results suggest that spider cues affect feeding location rather than the total amount of herbivory.     

Plant-attracted ants affect arthropod community structure but not necessarily herbivory

Abstract.  1. The effectiveness of ants as plant defenders is equivocal for plants that attract ants via extrafloral nectaries (EFNs).
2. This study focused on the myrmecophilic savannah tree Pseudocedrela kotschyi that attracts ants to EFNs and on the arthropod fauna associated with P. kotschyi . Herbivory and arthropod community composition were compared between trees that were dominated by one of three congeneric ant species, Camponotus acvapimensis , C. rufoglaucus , and C. sericeus , and between trees where ants were experimentally excluded and untreated control trees.
3. Short-term ant-exclusion experiments failed to demonstrate a consistent effect of ants on herbivory.
4. Plants dominated by different ant species differed significantly in leaf damage caused by herbivorous insects. The relative ranking of herbivory levels of the trees dominated by different ant species was persistent in three consecutive years.
5. Ants significantly reduced the abundance of different arthropod groups (Araneae, Blattodea, Coleoptera, Homoptera, non-ant Hymenoptera). Other groups, including important herbivores, seemed not to be affected (Lepidoptera, Orthoptera, Thysanoptera, Heteroptera).
6. The study suggests that the presence of ants only benefits plants when specific ant species are attracted, and protection by these ants is not counterbalanced by their negative effect on other beneficial arthropods.     

Herbivory in canopy gaps created by a typhoon varies by understory plant leaf phenology

1. Availabilities of light and soil nitrogen for understory plants vary by extent of canopy gap formation through typhoon disturbance. We predicted that variation in resource availability and herbivore abundance in canopy gaps would affect herbivory through variation in leaf traits among plant species. We studied six understory species that expand their leaves before or after canopy closure in deciduous forests. We measured the availabilities of light, soil nitrogen, soil water content, and herbivore abundance in 20 canopy gaps (28.3–607.6 m²) formed by a typhoon and in four undisturbed stands. We also measured leaf traits and herbivory on understory plants. 2. The availabilities of light and soil nitrogen increased with increasing gap size. However, soil water content did not. The abundance of herbivorous insects (such as Lepidoptera and Orthoptera) increased with increasing gap size. 3. Concentrations of condensed tannins, total phenolics, and nitrogen in leaves and the leaf mass per area increased in late leaf expansion species with increasing gap size, whereas none of the leaf traits varied by gap size in early leaf expansion species. 4. Herbivory increased on early leaf expansion species with increasing gap size, but decreased on late leaf expansion species. In these late leaf expansion species, total phenolics and C : N ratio had negative relationships with herbivory. 5. These results suggested that after typhoon disturbance, increased herbivory on early leaf expansion species can be explained by increased herbivore abundance, whereas decreased herbivory on late leaf expansion species can be explained by variation in leaf traits.     

The Influence of Soil Type on Rain Forest Insect Herbivore Communities

Insect herbivores were collected from five species of dipterocarp tree seedling within a large‐scale reciprocal transplant experiment in Sabah, Malaysia, on alluvial and sandstone soils in both gap and understory plots. The aim was to determine whether the location and ecological specialization of seedlings influenced the herbivore communities found on and around them. Three major groups of folivores were collected: Coleoptera, Orthoptera, and larval Lepidoptera. Herbivory of all species was confirmed through laboratory trials. Herbivore abundance in the understory plots was extremely low relative to the gaps. Rank‐abundance curves were similar on both soil types, differing only within the Lepidoptera. Coleoptera and Orthoptera communities were numerically dominated by a small suite of species capable of feeding on all dipterocarp species tested, whereas lepidopteran communities had both greater species richness and diversity. When corrected for leaf area surveyed, the abundance of Coleoptera was similar on both soil types, while larval Lepidoptera were more abundant in sandstone plots and Orthoptera were more abundant in alluvial plots. Estimated species richness of all three taxa was greater in alluvial forest, but there were contrasting patterns in Simpson diversity and evenness between groups. Species richness of Lepidoptera was greatest on seedlings when grown in their native soil type, providing partial evidence for possible escape effects, although this was not matched by differences in folivore abundance. The link between herbivore communities and herbivory rates on rain forest tree seedlings is complex and is unlikely to be detected through simplistic measures of abundance, species richness, or diversity.     

Phytophagous insect-woody sprout interactions in tropical eucalypt forest. II. Insect community structure

Abstract Sucking insects constituted 79% of all phytophagous insects collected from woody sprouts in the ground layer of a tropical eucalypt forest. Mobile insect groups such as non-psyllid Hemiptera and Orthoptera were relatively frequent in this environment compared to temperate, Eucalyptus-dominated vegetation. The high fire frequency of the tropical eucalypt forest may favour mobile insect groups. The capture of sucking insects and caterpillars peaked in dry season samples. Other patterns of abundance of phytophagous insect groups showed little consistency in their seasonal trends between host species or between vegetation types within host species. Disparities between chewing insect abundance in daytime samples and the damage chewing insects cause, may result from disproportionate consumption by large, mainly nocturnal insects, such as members of the Orthoptera. In this study, 21% of insect species were specialists on single plant species. This study suggested that insect abundance reflected the growth patterns of woody sprouts after regular burning, rather than that plant growth and development were tuned to the pressures of insect herbivory.     

The alkaloidal responses of wild tobacco to real and simulated herbivory

Summary I compared the induced alkaloidal response in undamaged leaves of plants subjected to herbivory by the larvae of Manduca sexta and to different simulations of this herbivory; all herbivory treatments removed similar amounts of leaf mass. Although larval feeding induced a significant increase (2.2x) in alkaloid concentrations compared to undamaged plants, the alkaloid responses to larval feeding were significantly lower than the responses to an herbivory simulation (4x controls) which involved removing the same amount of leaf area from the same positions on the leaf, over a similar time period. Moreover, another herbivory simulation, identical in amount of leaf mass removed and duration of damage to the larval feeding, but without regard to spatial array of leaf damage, resulted in an alkaloidal response (5.5x controls) higher still than the previous herbivory simulation. In a second experiment the importance of leaf vein damage on the induced alkaloidal response was examined. Here, leaf removal that involved cutting leaf tissues from between secondary veins before removing the midrib, resulted in alkaloidal responses that were significantly lower (1.7x controls) than responses from leaf removal that involved cutting both veins and midribs along with the intervein tissues (2.6x controls). Vein damage alone did not produce a significant response. These results indicate that herbivory is difficult to simulate: that how a leaf is damaged can be as important as the magnitude of leaf damage in determining a plant's response to damage.     

Seasonal changes in arthropod diversity patterns along an Alpine elevation gradient

1. Deciphering patterns in species distributions and species interactions along ecological gradients are fundamental topics in ecology. Theory holds that species diversity is greater and interactions are stronger under warmer and more stable environments, such as low elevations and latitudes. However, recent findings have shown conflicting evidence, potentially due to seasonal effects.
2. We aimed to address this gap by studying seasonal changes in arthropod communities over an elevation gradient in the Swiss Alps, as well as herbivore-predator interactions and their resulting consequences on plant herbivory levels.
3. Overall, we found hump-shaped patterns in arthropod abundance, richness and diversity with increasing elevation, with all factors peaking below the tree line. However, these patterns varied seasonally, with strong mid-elevation peaks at the beginning of the summer, shifting to a pattern of linear decrease at the beginning of the fall. In searching for mechanisms explaining these changes, we found that shifts in arthropod communities over elevation and seasons usually followed shifts in vegetation productivity estimates. Other factors, such as top-down control by natural enemies, which was generally stronger at low elevations, and plant species-specific resistance rates along elevation gradients were also implicated as drivers of diversity and herbivory rates.
4. These results highlight the complexity of arthropod communities' responses to environmental gradients, which vary during the season in response to relative changes in both bottom-up and top-down forces.

     

Prioritising potential guilds of specialist herbivores as biological control agents for prickly acacia through simulated herbivory

Understanding plant response to herbivory facilitates the prioritisation of guilds of specialist herbivores as biological control agents based on their potential impacts. Prickly acacia ( Acacia nilotica ssp. indica ) is a weed of national significance in Australia and is a target for biological control. Information on the susceptibility of prickly acacia to herbivory is limited, and there is no information available on the plant organ (i.e. leaf, shoot and root in isolation or in combination) most susceptible to herbivory. We evaluated the ability of prickly acacia seedlings, to respond to different types of simulated herbivory (defoliation, shoot damage, root damage and combinations), at varying frequencies (no herbivory, single, two and three events of herbivory) to identify the type and frequency of herbivory that will be required to reduce the growth and vigour. Defoliation and shoot damage, individually, had a significant negative impact on prickly acacia seedlings. For the defoliation to be effective, more than two defoliation events were required, whereas a single bout of shoot damage was enough to cause a significant reduction in plant vigour. A combination of defoliation + shoot damage had the greatest negative impact. The study highlights the need to prioritise specialist leaf and shoot herbivores as potential biological control agents for prickly acacia.     

Rapid response of Orthoptera to restoration of montane heathland

Understanding how to restore threatened ecosystems is of special relevance for nature conservation. The aim of this study was to use Orthoptera as ecological indicators for the effects of montane heathland restoration in Central Europe. We analysed the three following treatments: (i) montane heathlands (MONHEATH) (N = 7), (ii) restoration sites (RESSITE) (N = 3) and (iii) clear-cuts of spruce forests as unprocessed and ungrazed control sites (CONTROL) (N = 3). Vegetation structure and microclimate differed considerably between MONHEATH on the one hand and RESSITE and CONTROL on the other hand. Orthoptera species richness and density did so too. MONHEATH was characterised by a high-growing dense dwarf-shrub and moss layer having a cool microclimate and high soil moisture. In contrast, RESSITE and CONTROL had sparse vegetation and a warm microclimate; Orthoptera species richness and density was highest on these sites. Our study clearly showed that heathland Orthoptera responded rapidly to restoration measures, while Ericaceae dwarf shrubs slowly established. The vast majority of Orthoptera species found on the restoration sites are early and mid-successional species. The colonization of the sites by late-successional Orthoptera species in the future will depend on the further development of the heathland vegetation; that is, if Ericaceae will expand to the sites. We conclude that the realised restoration measures are suitable to promote heathland Orthoptera of early and mid-successional stages. However, the current management of montane heathlands is insufficient and needs to be intensified in order to provide structurally diverse habitats with their characteristic orthopteran assemblages.     

Assessing density–damage relationships between water hyacinth and its grasshopper herbivore

Plants are variable in their responses to insect herbivory. Experimental increases in densities of phytophagous insects can reveal the type of plant response to herbivory in terms of impact and compensatory ability. The relationship between insect density and plant damage of a grasshopper, Cornops aquaticum Brüner (Orthoptera: Acrididae: Tetrataeniini), a candidate biological control agent, and an invasive aquatic plant, water hyacinth, Eichhornia crassipes Mart. Solms‐Laubach (Pontederiaceae), was investigated to assess potential damage to the weed. The impact of different densities of male and female grasshoppers on E. crassipes growth parameters was determined in a quarantine glasshouse experiment. Damage curves indicated that the relationship between plant biomass reduction and insect density was curvilinear whereas leaf production was linear. Female C. aquaticum were more damaging than males, causing high rates of plant mortality before the end of the trial at densities of three and four per plant. Feeding by C. aquaticum significantly reduced the total plant biomass and the number of leaves produced, and female grasshoppers caused a greater reduction in the number of leaves produced by water hyacinth plants than males. Grasshopper herbivory suppressed vegetative reproduction in E. crassipes, suggesting C. aquaticum could contribute to a reduction in the density and spread of E. crassipes infestations. The results showed that E. crassipes vigour and productivity decreases with an increase in feeding intensity by the grasshopper. Cornops aquaticum should therefore be considered for release in South Africa based on its host specificity and potential impact on E. crassipes.     

Role of sediment and internal loading of phosphorus in shallow lakes

Current velocity is a pervasive feature of lotic systems, yet this defining environmental variable is rarely examined as a factor for regulating stream herbivory. To investigate how current modifies herbivory in the upper Colorado River, U.S.A., loops of electrified fencing wire were used to reduce in situ grazer densities on 30 × 30 cm tile substrates. After 45 d, electrified tiles had significantly fewer grazers (P = 0.03) and >2X more algal biomass than controls (P = 0.0002). Reduced grazing on electrified tiles yielded periphytic assemblages having more diatoms and chlorophytes, as well as greater algal species richness. Current velocity effects alone did not significantly regulate algal abundance; however, the interaction between current velocity and grazer exclusion resulted in more algae in slow vs. fast current (P = 0.02). Grazer abundances were similar between fast and slow current velocities, suggesting that grazers in the Colorado River differ in their ability to regulate algae across the current velocity gradient. Our results indicate that stream current-mediated herbivory in streams may be more important than is generally recognized.     

Responses of assemblages of Orthoptera to management and use of ski slopes on upper sub-alpine meadows in the Austrian Alps

The Orthoptera assemblages occurring on sub-alpine ski slopes were compared with those found on neighbouring unskied meadows by making frequent transect counts at two pairs of sites in the Gastein valley in the Austrian Central Alps. On one of the ski slopes no Orthoptera were present, although two species were abundant on the control meadow a few meters away. On the second ski slope, the Orthoptera assemblage exhibited reduced species richness, lower densities of individuals and a generally accelerated rate of nymphal development compared to the control meadow populations. These results may be explained in terms of the changed habitat conditions on the ski slopes and the known biologies of the species concerned. The implications of the findings for winter tourism management in high altitude ecosystems are briefly discussed.     

Pre-release evaluation of the efficacy of the leaf-sucking bug Carvalhotingis visenda (Heteroptera: Tingidae) as a biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae)

In classical weed biological control, assessing weed response to simulated herbivory is one option to assist in the prioritization of available agents and prediction of their potential efficacy. Previously reported simulated herbivory studies suggested that a specialist herbivore in the leaf-feeding guild is desirable as an effective biological control agent for cat's claw creeper Macfadyena unguis-cati (Bignoniaceae), an environmental weed that is currently a target for biological control. In this study, we tested (i) whether the results from glasshouse-based simulated herbivory can be used to prioritise potential biological control agents by evaluating the impact of a leaf-sucking tingid bug Carvalhotingis visenda (Drake & Hambleton) (Hemiptera: Tingidae) in quarantine; and (ii) the likely effectiveness of low- and high-densities of the leaf-sucking tingid after its release in the field. The results suggest that a single generation of C. visenda has the potential to reduce leaf chlorophyll content significantly, resulting in reduced plant height and leaf biomass. However, the impact of one generation of tingid herbivory on below-ground plant components, including the roots and tuber size and biomass, were not significant. These findings are consistent with results obtained from a simulated herbivory trial, highlighting the potential role of simulated herbivory studies in agent prioritisation.     

Effect of herbivory on growth and biomass allocation of Brazilian peppertree (Sapindales: Anacardiaceae) seedlings in the laboratory

Brazilian peppertree (Schinus terebinthifolius Raddi), native to South America, is invading many ecosystems in south and central Florida. The defoliating tortricid moth Episimus unguiculus Clarke was selected as a potential biocontrol agent of Brazilian peppertree in Florida. The objective of this study was to examine the effect of different levels of herbivore damage on growth and biomass allocation of Brazilian peppertree seedlings in the laboratory. Three treatments were established: (1) no herbivory (control), (2) low herbivory (~4 larvae/plant), and (3) high herbivory (~12 larvae/plant). High levels of herbivory significantly reduced the number of leaflets, plant height, foliar biomass, foliar relative growth rate (RGR) and shoot: root ratio of Brazilian peppertree seedlings. Moreover, plants were not able to recover from herbivory after 2 months. The performance of Brazilian peppertree subjected to low herbivory levels did not differ from the control plants (no herbivory). The potential effectiveness of the biocontrol agent E. unguiculus to suppress this noxious weed is examined.     

The impact of grazing management on Orthoptera abundance varies over the season in Mediterranean steppe-like grassland

As semi-natural grassland has a high level of biological diversity, understanding the effects of grazing and its variation over time is important in order to identify sustainable grazing practices. We measured temporal variation in Orthoptera abundance and spatial vegetation structure during seasonal grazing in an extensive sheep-farming system. We studied five grazed pasture areas (pre-grazing and post-grazing) and two adjacent ungrazed grasslands. We recorded the total abundance of Orthoptera and described the vegetation structure of 175 replicate plots (25 per pasture/grassland) during six field sampling sessions. We demonstrated that the impact of grazing on Orthoptera abundance is species-specific and greatly varies over the grazing season. The decrease of phytovolume is significant after 4–7 weeks of sheep grazing. Total Orthoptera abundance was higher in pre-grazed plots than in ungrazed plots, and higher in ungrazed plots than in post-grazed plots. These differences were particularly high during the peak of adult abundance. No difference in species richness was observed between grazing intensities. Total Orthoptera abundance positively correlated to phytovolume only when grazing pressure was high. However, the relationship between abundance and phytovolume differed between species. Extensive grazing by sheep tends to homogenize spatial vegetation structure and to temporarily reduce total Orthoptera abundance at pasture scale. However, rotational grazing allows spatial and temporal heterogeneity in vegetation structure to be maintained at farm scale, heterogeneity that is beneficial for Orthoptera. In contrast, absence of grazing has a negative impact on Orthoptera abundance as it favours the accumulation of litter, which is detrimental for a high proportion of xerothermophilic Orthoptera associated with bare ground and short vegetation.     

Effects of insect attack to stems on plant survival,growth, reproduction and photosynthesis

Studies of insect herbivory have mostly focused on leaf‐feeding even though most woody plant biomass is stem tissue. Attack to stems has the potential to be more detrimental to plant performance than attack to leaves. Here we asked how severe is the impact of insect stem herbivory on plant performance. We quantify the effect of insect stem herbivory via a meta‐analysis of 119 papers in 100 studies (papers by the same authors were treated as the same study). These studies involved 92 plant species and 70 species of insect herbivore (including simulated herbivory). Attack to plant stems reduced plant performance by an average of approximately 22%. Stem herbivory had greatest impacts on plant and branch survival, which was reduced by 63%. Measures of plant reproduction and vegetative biomass were reduced by 33% and 16% respectively, while measurements of photosynthetic rate were not significantly different between plants with and without stem herbivore attack. Stem herbivory led to a decline in leader performance but an increase in performance of laterals, highlighting the importance of plant compensation. Juvenile plants were more severely affected by stem herbivory than adult plants, and studies conducted in greenhouses found more severe effects than studies conducted in the field. Stem herbivory did not have a significant effects on any of the non‐performance responses measured (defence compounds, SLA, root:shoot, phenology and plant carbon and nitrogen). We compare our results with results from various meta‐analyses considering herbivory on other plant parts. The impact of insect herbivory to stems on plant performance appears at least as severe as insect herbivory to roots and leaves, if not more.     

Direct and indirect effects of ski run management on alpine Orthoptera

Alpine landscapes are heavily influenced by ski run management, which can have severe impacts on alpine biodiversity. To assess these impacts on alpine Orthoptera, we compared species richness and species abundance in 41 plot pairs on ski runs and adjacent off-slope control plots in three ski resorts in Austria and Germany. A mixed modelling approach was used to assess the impacts of ski run preparation, artificial snow-making and environmental variables such as altitude, cover of dwarf shrubs and the application of fertilizer. Ski run plots showed a significantly lower species richness and number of individuals than control plots. Moreover, artificial snow led to a further decrease in species number. Hierarchical variance partitioning revealed that Orthoptera community composition is best predicted by environmental variables indirectly related to ski run management (fertilization, cover of dwarf shrubs) and to altitude. Only one out of five species significantly decreased in abundance after artificial snow-making. Other species were more sensitive to fertilizing and altitude. Dwarf shrubs were negatively associated with ski run management but positively associated with abundance of three species and species richness. Our data provide evidence for both direct and indirect consequences of ski runs and artificial snow-making on alpine Orthoptera. Overall, Orthoptera communities are suitable indicators for human-induced changes in alpine environments. In particular, a shift towards generalist species such as Chorthippus parallelus along with a decrease in typical alpine species gives cause for concern as this implies a homogenisation of biodiversity owing to ski run management.     

Oak-insect herbivore interactions along a temperature and precipitation gradient

The interactions between herbivorous insects and their host plants are expected to be influenced by changing climates. Modern oaks provide an excellent system to examine this assumption because their interactions with herbivores occur over broad climatic and spatial scales, they vary in their defensive and nutritional investment in leaves by being deciduous or evergreen, and their insect herbivores range from generalists to highly specialized feeders. In this study, we surveyed leaf-litter samples of four oak species along an elevation gradient, from coastal northern California, USA, to the upper montane woodlands of the Sierra Nevada, to examine the relationship between climatic factors (mean annual temperature and precipitation) and oak herbivory levels at multiple scales; across all oak species pooled, between evergreen and deciduous species and within species.Overall, temperature and precipitation did not appear to have a significant effect on most measures of total herbivore damage (percent leaves damaged per tree, percent leaf area removed and average number of feeding damage marks per leaf) and the strongest predictor of herbivore damage overall was the identity of the host species. However, increases in precipitation were correlated with an increase in the actual leaf area removed, and specialized insects, such as those that make leaf mines and galls, were the most sensitive to differences in precipitation levels. This suggests that the effects of changing climate on some plant–insect interactions is less likely to result in broad scale increases in damage with increasing temperatures or changing precipitation levels, but is rather more likely to be dependent on the type of herbivore (specialist vs. generalist) and the scale (species vs. community) over which the effect is examined.