Development, growth, and egg production of Ageneotettix deorum (Orthoptera: Acrididae) in response to spider predation risk and elevated resource quality

Abstract.  1. Predation risk to insects is often size- or stage-selective and usually decreases as prey grow. Any factor, such as food quality, that accelerates developmental and growth rates is likely to reduce the period over which prey are susceptible to size-dependent predation.
2. Using field experiments, several hypotheses that assess growth, development, and egg production rates of the rangeland grasshopper Ageneotettix deorum (Scudder) were tested in response to combinations of food quality and predation risk from wolf spiders to investigate performance variation manifested through a behaviourally mediated path affecting food ingestion rates.
3. Grasshoppers with nutritionally superior food completed development ≈ 8–18% faster and grew 15–45% larger in the absence of spiders, in comparison with those subjected to low quality food exposed to spider predators. Growth and development did not differ for grasshoppers feeding on high quality food when predators were present in comparison with lower quality food unimpeded by predators. Responses indicated a compensatory relationship between resource quality and predation risk.
4. Surviving grasshoppers produced fewer eggs compared with individuals not exposed to spiders. Because no differences were found in daily egg production rate regardless of predation treatment, lower egg production was attributed to delayed age of first reproduction. Results compare favourably with responses observed in natural populations.
5. Risk of predation from spiders greatly reduced growth, development, and ultimately egg production. Increased food quality counteracts the impact of predation risk on grasshoppers through compensatory responses, suggesting that bottom-up factors mediate effects of spiders.     

Resource-mediated impact of spider predation risk on performance in the grasshopper Ageneotettix deorum (Orthoptera: Acrididae)

In response to increased exposure to predators when searching for food, many prey increase the frequency of antipredator behaviors, potentially reducing foraging rate and food intake. Such direct, nonlethal interactions between predators and prey resulting in reduced food intake can indirectly influence lifecycle development through effects on growth, developmental rate, and survival. We investigated the general hypothesis that individual performance of a herbivorous insect can be negatively affected when exposed to nonlethal predation risk, and that the response can be mediated by food quality. This hypothesis was tested using the common rangeland grasshopper Ageneotettix deorum with and without exposure to common wolf spider predators (Lycosidae, Schizocosa spp.) on both untreated natural and fertilized vegetation. All spiders were rendered temporarily incapable of direct feeding by restricting function of the chelicerae with beeswax. Detectable responses by grasshoppers to spiders indicate indirect consequences for lifecycle development. Grasshopper performance was measured as hind femur growth, duration of nymphal lifecycle stages, and survivorship in a caged field experiment conducted over 2 years. Grasshoppers developed faster and grew 3–5% larger when allowed to forage on fertilized vegetation in the absence of risk from a spider predator. Failure-time analysis illustrated enhanced survival probability in response to elevated food quality and the negative effects of grasshopper susceptibility to nonlethal predation risk. Performance on food of relatively low, ambient quality with no predation risk equaled that of grasshoppers caged with high quality vegetation in the presence of a modified spider. Increased resource quality can clearly moderate the negative life history responses caused by the behavioral modification of grasshoppers when exposed to spider predation risk, a compensatory response.     

Populations of the northern grasshopper, Melanoplus borealis (Orthoptera: Acrididae), in Alaska are rarely food limited

In some systems, grasshoppers appear to be food limited in most years, whereas in other systems top down forces, for example, predators, are more often implicated in population regulation. Sustainable strategies to manage grasshopper populations through habitat management require knowledge of the forces that regulate grasshopper populations. This experiment was undertaken to determine whether populations of Melanoplus borealis (Fieber), a common pest species in Alaska, are food-limited in Alaska. Cages were set up in a fallow field near Delta Junction, AK, in 3 yr (2007-2009). In 2007 and 2008, fertilizer was added to half the plots to increase primary production, and, in all years, cages within each plot were stocked with 0, 5, 9, or 13 fourth-instar M. borealis (equivalent to 0, 20, 36, or 52 grasshoppers/m(2)). Grasshoppers in each cage were counted weekly. Near the end of the growing season, surviving female grasshoppers (≈40% of the original number) were collected. Femur length was taken as a measure of adult size, and functional ovarioles were counted as a measure of current fecundity. If the grasshoppers were food limited, we expected to see significant effects of either density or fertilizer on grasshopper survival, size, or fecundity. The fertilizer treatment greatly increased primary production in both years. Neither fertilizer treatment nor grasshopper density had consistent effects on survival, size, or potential fecundity, leading us to conclude that food is seldom limiting to populations in the interior of Alaska at densities <50 m(-2).     

Nest and foraging‐site selection in Yellowhammers Emberiza citrinella: implications for chick provisioning

Capsule Vegetation structure and invertebrate abundance interact to influence both foraging sites and nestling provisioning rate; when invertebrate availability is low, adults may take greater risks to provide food for their young.

Aims To investigate nesting and foraging ecology in a declining farmland bird whose fledging success is influenced by the availability of invertebrate prey suitable for feeding to offspring, and where perceived predation risk during foraging can be mediated by vegetation structure.

Methods Provisioning rates of adult Yellowhammers feeding nestlings were measured at nests on arable farmland. Foraging sites were compared with control sites of both the same and different microhabitats; provisioning rate was related to habitat features of foraging‐sites.

Results Foraging sites had low vegetation density, probably enhancing detection of predators, or high invertebrate abundance at high vegetation density. Parental provisioning rate decreased with increasing vegetation cover at foraging sites with high invertebrate abundance; conversely, where invertebrate abundance was low, provisioning rate increased with increasing vegetation cover.

Conclusions Vegetation structure at foraging sites suggests that a trade‐off between predator detection and prey availability influences foraging site selection in Yellowhammers. Associations between parental provisioning rate and vegetation variables suggest that where invertebrate abundance is high birds increase time spent scanning for predators at higher vegetation densities; however, when prey are scarce, adults may take more risks to provide food for their young.     

Armed Rollers: Does Nestling’s Vomit Function as a Defence against Predators?

Chemical defences against predators are widespread in the animal kingdom although have been seldom reported in birds. Here, we investigate the possibility that the orange liquid that nestlings of an insectivorous bird, the Eurasian roller (Coracias garrulus), expel when scared at their nests acts as a chemical defence against predators. We studied the diet of nestling rollers and vomit origin, its chemical composition and deterrent effect on a mammal generalist predator. We also hypothesized that nestling rollers, as their main prey (i.e. grasshoppers) do from plants, could sequester chemicals from their prey for their use. Grasshoppers, that also regurgitate when facing to a threat, store the harmful substances used by plants to defend themselves against herbivores. We found that nestling rollers only vomit after being grasped and moved. The production of vomit depended on food consumption and the vomit contained two deterrent chemicals (hydroxycinnamic and hydroxybenzoic acids) stored by grasshoppers and used by plants to diminish herbivory, suggesting that they originate from the rollers’ prey. Finally, we showed for the first time that the oral secretion of a vertebrate had a deterrent effect on a model predator because vomit of nestling rollers made meat distasteful to dogs. These results support the idea that the vomit of nestling rollers is a chemical defence against predators.     

Grasshoppers affect grassland ecosystem functioning: Spatial and temporal variation

Using experiments and monitoring, we find that grasshoppers in a grassland ecosystem impact ecosystem functioning (nutrient cycling and primary production) in different ways among sites in the ecosystem. Experiments conducted over many years at two sites (21 and 15 years, respectively) with the same grasshopper and plant species demonstrated that grasshoppers increased nitrogen availability (N) and consequently annual plant production (ANPP) at one site, and decreased N and consequently ANPP at the other site. Comparing the two sites, N increased on average by 8% and up to 21.6%, and resulting ANPP increased on average by 18.6% and up to 33.3%. Grasshoppers increase N and ANPP by preferentially feeding on slower decomposing plants, and the opposite occurs by preferentially feeding on faster decomposing plants. Monitoring 20 random sites in the ecosystem, grasshoppers consistently increased N and ANPP over 3 years at 40% of sites, consistently decreased N and ANPP at 35% of sites, and sometimes increased and decreased N and ANPP at 25% of sites. Therefore, grassland grasshoppers, and insects in many ecosystems, may strongly affect ecosystem functioning.     

Effects of Predation Risk and Food Availability on the Activity,Habitat Use,Feeding Behavior and Mating Behavior of a Pond Water Strider,Gerris marginatus (Hemiptera)

By experimentally manipulating predation risk and food availability, we explored the effects of these factors on refuge use, general activity, feeding activity, and mating activity of adults of the pond water strider Gerris marginatus. Predation risk (from adult backswimmers Notonecta undulata) profoundly influenced the behavior of G. marginatus, reducing mating activity, general activity in males and females, decreasing feeding activity in females and increasing the use of refuge by both sexes. Surprisingly, variations in food availability failed to produce significant effects on any of these behavior patterns. Risk x food interactions were also not statistically significant for any gerrid behavior. The significant responses to predators all changed in directions that should decrease the risk of predation for G. marginatus.     

Stage-Specific Behavioral Responses of Ageneotettix deorum (Orthoptera: Acrididae) in the Presence of Lycosid Spider Predators

Grasshoppers must gather food while avoiding size-selective predation from other arthropods, especially spiders, potentially leading to a trade-off between foraging and defensive behaviors. This trade-off becomes less intense as prey grow larger and are less susceptible to arthropod predation. Activity budgets were constructed for three nymphal (third- to fifth- instar) and adult life cycle stages of Ageneotettix deorum, a common rangeland grasshopper, for three conditions of predation risk by lycosid spiders (spider absence, spider presence, and presence of a nonlethal, chelicerae-modified spider). In third and fourth instars, exposure to predators resulted in reduced feeding activity, increased time spent in antipredator and defensive behaviors, and reduced general activity compared to individuals not exposed to spiders. No significant shifts in behaviors were observed for fifth-instar nymphs and adult A. deorum in response to spider presence. Activity levels in functional spiders and chelicerae-modified spiders were statistically indistinguishable.     

Temporal pattern of resource use and variation in diets of individual grasshoppers (Orthoptera: Acrididae)

Patterns of resource fidelity, switching, and variation in individual diet were examined in the polyphagous grasshopper Taeniopoda equesforaging in experimental cages over an 11-day period. Ten novel dicots were provided in the cages, but grasshoppers fed primarily on kale (Brassica oleraceavar. acephala),lobelia (Lobelia erinus),pansy (Viola x wittrockiana),and dry bermuda grass (Cynodon dactylon).Grasshoppers showed significant fidelity to a single plant within a meal but tended to eat less of the same plant in the next meal and, in the case of pansy, on the next day. Grasshoppers switched nonrandomly, suggesting that compensatory feeding on complementary resources may have occurred. There was no evidence that behavioral interactions among individuals increased switching rate or lowered fidelity. Overall patterns of resource use varied significantly among grasshoppers during the study, despite frequent switching among resources and a significant tendency to aggregate while feeding. All individuals were polyphagous but differed in relative consumption of available plants. The results suggest that individual grasshoppers express different feeding patterns that are consistent over time and that variation in diet among individual observed in the field may be more than simple sampling error.     

Choice of feeding station in Atlantic salmon, Salmo salar, parr: effects of predation risk, season and life history strategy

The current speed at which underyearling salmon parr held feeding stations was examined from late summer to early winter in laboratory flume tanks that offered a choice between (a) areas with high water flow, high food availability but high predation risk and (b) areas with low flow, little food but shelter from predators. In August, those fish that would become smolts aged 1 + (and which by late winter formed the upper modal group, UMG, of the bimodal size distribution) adopted positions in faster currents than did the fish which would take a further year to reach the smolt stage (the lower modal group, LMG). However, the chosen current speed of UMG fish decreased through the period of study, so that by December all fish were found in areas of low flow, and hence little food. Both date and water temperature had independent effects on the chosen current speed of UMG fish.
The effect of predation risk was investigated using a model trout. A brief sight of this predator caused 47% of fish that had been in the main, exposed currents to move to slacker, sheltered areas; they took 1 h, on average, to return to their previous position. The fish that remained in position upon seeing the predator reduced their rate of tail beating, presumably increasing crypticity. Eventual UMG parr were less likely than were LMG fish to move away upon seeing a predator. The fish moved to faster currents than normal 2–3 h after seeing the predator, possibly compensating for the earlier reduction in feeding rate.     

Arsenic hyperaccumulation in the Chinese brake fern (Pteris vittata) deters grasshopper (Schistocerca americana) herbivory

Brake fern, Pteris vittata, not only tolerates arsenic but also hyperaccumulates it in the frond. The hypothesis that arsenic hyperaccumulation in this fern could function as a defense against insect herbivory was tested. Fronds from control and arsenic-treated ferns were presented to nymphs of the grasshopper Schistocerca americana. Feeding damage was recorded by visual observation and quantification of the fresh weight of frond left uneaten and number of fecal pellets produced over a 2-d period. Grasshopper weight was determined before and after 5 d of feeding. Grasshoppers consumed significantly greater amounts of the frond tissue, produced more fecal pellets and had increased body weight on control plants compared with grasshoppers fed arsenic-treated ferns. Very little or none of the arsenic-treated ferns were consumed indicating feeding deterrence. In a feeding deterrent experiment with lettuce, sodium arsenite at 1.0 mm deterred grasshoppers from feeding whereas 0.1 mm did not. In a choice experiment, grasshoppers preferred to feed on lettuce dipped in water compared with lettuce dipped in 1.0 mm sodium arsenite. Our results show that arsenic hyperaccumulation in brake fern is an elemental defense against grasshopper herbivory.     

Seeing without being seen: a removal experiment with mixed flocks of Willow and Crested Tits Parus montanus and cristatus

This paper tests the hypothesis that foraging site selection reflects a trade-off between the various needs for concealment from predators, to find food, and for the individual to maintain some view of its surroundings. After removal of Crested Tits Parus cristatus (the dominant species in mixed flocks), Willow Tits P. montanus did not decrease their foraging heights as expected but remained in the most exposed parts of young pines. In contrast, after removal of Willow Tits, Crested Tits increased their foraging height from the sheltered lower canopy to sites previously occupied by Willow Tits. When flock size was reduced, the birds maintained the same high levels of vigilance without concealing themselves in dense vegetation. I suggest that flock members may benefit from foraging in sites that afford good anti-predator vigilance.     

Interspecific competition among grasshoppers and their effect on plant abundance in experimental field environments

Summary We tested whether grasshoppers in experimental field environments, i.e. cages (40×40 cm) placed on existing old field vegetation, (1) were limited in density by plant abundance and/or nitrogen content, (2) exhibited interspecific competition, and (3) altered the relative abundance of different plant species. We examined interactions among a pair of early season grasshopper species (May–June; Arphia conspersa and Pardalophora apiculata) and a late season pair (July–August; Melanoplus femur-rubrum and Melanoplus bivittatus). Each grasshopper species was placed in cages by itself and with another grasshopper species. Grasshoppers generally survived at higher density in fertilized cages and they reduced plant abundance relative to empty cages, suggesting that grasshoppers may be food limited at these densities. In unfertilized plots, early season grasshoppers preferred grasses (Schizachyrium scoparium and Poa pratensis) and favored the growth of forbs (especially Solidago spp.). However, late in summer, Melanoplus spp. preferred Solidago spp. and favored the growth of grasses.The pattern of grasshopper survivorship and plant reduction within these experimental environments provide preliminary support for some of the predictions of resource competition theory. Grasshoppers exhibited interspecific competition only if they significantly reduced plant biomass. If two species competed, a grasshopper species was eliminated only if the superior competitor, when living by itself, could reduce plant biomass to a significantly lower level than the inferior competitor. Competitors persisted only if they did not differ in their ability to reduce plant biomass or reduced the abundance of different plant species.     

Response of grasshoppers (Orthoptera: Acrididae) to livestock grazing in southeastern Arizona: differences between seasons and subfamilies

Summary Grasshopper densities were compared between grazed and ungrazed semidesert grassland sites in southeastern Arizona. Bouteloua-dominated perennial grass cover was about 1.5 times greater on the livestock exclosure. Grasshoppers were 3.7 times more abundant on the protected area in the summers of 1983 and 1984, when dominant species were grass-feeding members of the subfamily Gomphocerinae. In fall 1984, grasshoppers were 3.8 times more common on the grazed site, when dominants were mainly herb-feeders in the subfamily Melanoplinae. These results indicate important seasonal and taxonomic differences in the responses of grasshoppers to the activities of vertebrate grazers.     

Loss of body mass under predation risk: cost of antipredatory behaviour or adaptive fit-for-escape?

Predation risk may compromise the ability of animals to acquire and maintain body reserves by hindering foraging efficiency and increasing physiological stress. Locomotor performance may depend on body mass, so losing mass under predation risk could be an adaptive response of prey to improve escape ability. We studied individual variation in antipredatory behaviour, feeding rate, body mass and escape performance in the lacertid lizard Psammodromus algirus. Individuals were experimentally exposed to different levels of food availability (limited or abundant) and predation risk, represented by reduced refuge availability and simulated predator attacks. Predation risk induced lizards to reduce conspicuousness behaviourally and to avoid feeding in the presence of predators. If food was abundant, alarmed lizards reduced feeding rate, losing mass. Lizards supplied with limited food fed at near-maximum rates independently of predation risk but lost more mass when alarmed; thus, mass losses experienced under predation risk were higher than those expected from feeding interruption alone. Although body mass of lizards varied between treatments, no component of escape performance measured during predator attacks (endurance, speed, escape strategy) was affected by treatments or by variations in body mass. Thus, the body mass changes were consistent with a trade-off between gaining resources and avoiding predators, mediated by hampered foraging efficiency and physiological stress. However, improved escape efficiency is not required to explain mass reduction upon predator encounters beyond that expected from feeding interruption or predation-related stress. Therefore, the idea that animals may regulate body reserves in relation to performance demands should be reconsidered.     

The effects of natural enemies,competition, and host plant water availability on an aphid population

Summary I used a factorial experiment repeated in two years to assess the relative effects of natural enemy attack, interspecific competition, and water availability to the host plant, and of interactions among these factors, on the population dynamics of the aphid Aphis varians feeding on fireweed (Epilobium angustifolium). The impact of a suite of coccinellid and syrphid predators emerged as the predominant factor affecting the success of aphid colonies: colonies protected from natural enemies grew in size at a rate of ten percent per day, were only one tenth as likely to go extinct, and produced over ten times more dispersing alates. In contrast, I found only minor effects of removing flea beetles, the most abundant herbivore with which A. varians colonies cohabit fireweed stems, and of supplementing water availability to fireweed host plants, in spite of a significant effect of watering frequency on aphid growth in the green-house. There was no evidence of significant two- or three-way interactions among factors. Hence, despite the potential complexity of the food web in which it is embedded, the dynamics of A. varians appears to be driven predominantly by a single factor, i.e. interactions with natural enemies.     

A quantitative analysis of habitat preferences of Woodcock Scolopax rusticola in the breeding season

In Britain, prior to breeding, Woodcock Scolopax rusticola switch from feeding on pastures by night to feeding in woodland by day. Their major food is earthworms (Lumbricidae). In April and May 1985, vegetation structure and composition and soil parameters including earthworm numbers (30 habitat variables) were recorded for quadrats containing 50 feeding locations of six radio-tagged birds, seven nests and 50 random locations in Whitwell Wood, northeast Derbyshire.
Significantly different mean values between feeding and random sites were found for 11 habitat variables. Feeding sites were in younger stands, with a higher percentage ground cover of dog's mercury Mercurialisperennis and consistently high values for pH. Earthworm biomass was on average 82% greater than in random plots. Areas of beech Fagus sylvaticus were avoided. A discriminant function based on six habitat variables correctly classified 85-5% of the feeding and random sites; 70% of the feeding locations were correctly classified by a jack-knife procedure. The important factors determining where feeding occurs are probably safety from avian predators and high availability of earthworms.
Discriminant analysis of the habitats used for feeding by solitary birds, broods and for nesting produced significant functions which correctly classified 84% of sites. Nests were in areas with a high percentage cover of brambles Rubus spp. and more open ground vegetation. Broods and solitary birds used similar areas characterized by denser ground vegetation than nest sites.
The results of this study indicate that both habitat structure and food availability influence the distribution of Woodcock in the breeding season.     

A comparison of two mountain gorilla habitats in Bwindi Impenetrable National Park, Uganda

Bwindi Impenetrable National Park, Uganda, is a small protected area (331 km²) within which there is large climatic and altitudinal variation. Therefore we compared habitat types and forest composition between two locations to investigate differences that may influence ecological conditions for large mammals, including endangered mountain gorillas. We demonstrate that there is considerable intersite variation in habitat types and spatial and temporal availability of vegetation resources of which the most apparent are the differences in species composition of plants and fruit availability. Buhoma (the lower altitude site) has a greater diversity of plant species and higher tree and shrub densities, yet has a much lower density of herbaceous vegetation eaten by gorillas compared with Ruhija. Fruit availability is significantly higher throughout the year in Buhoma compared with Ruhija. Both sites exhibit seasonal variation in fruit availability although they do not follow the same seasonal pattern, perhaps because of inherent asynchronous phenological differences amongst individual tree species. The results of this study are important because distribution and abundance of both terrestrial herbaceous vegetation and fruit resources are believed to influence aspects of feeding ecology for large mammals.     

Living and dying in a multi‐predator landscape of fear: roe deer are squeezed by contrasting pattern of predation risk imposed by lynx and humans

The theory of predation risk effects predicts behavioral responses in prey when risk of predation is not homogenous in space and time. Prey species are often faced with a tradeoff between food and safety in situations where food availability and predation risk peak in the same habitat type. Determining the optimal strategy becomes more complex if predators with different hunting mode create contrasting landscapes of risk, but this has rarely been documented in vertebrates. Roe deer in southeastern Norway face predation risk from lynx, as well as hunting by humans. These two predators differ greatly in their hunting methods. The predation risk from lynx, an efficient stalk‐and‐ambush predator is expected to be higher in areas with dense understory vegetation, while predation risk from human hunters is expected to be higher where visual sight lines are longer. Based on field observations and airborne LiDAR data from 71 lynx predation sites, 53 human hunting sites, 132 locations from 15 GPS‐marked roe deer, and 36 roe deer pellet locations from a regional survey, we investigated how predation risk was related to terrain attributes and vegetation classes/structure. As predicted, we found that increasing cover resulted in a contrasting lower predation risk from humans and higher predation risk from lynx. Greater terrain ruggedness increased the predation risk from both predators. Hence, multiple predators may create areas of contrasting risk as well as double risk in the same landscape. Our study highlights the complexity of predator–prey relationship in a multiple predator setting. Synthesis In this study of risk effects in a multi‐predator context, LiDAR data were used to quantify cover in the habitat and relate it to vulnerability to predation in a boreal forest. We found that lynx and human hunters superimpose generally contrasting landscapes of fear on a common prey species, but also identified double‐risk zones. Since the benefit of anti‐predator responses depends on the combined risk from all predators, it is necessary to consider complete predator assemblages to understand the potential for and occurrence of risk effects across study systems.     

The worm re-turns: hiding behavior of a tube-dwelling marine polychaete, Serpula vermicularis

The polychaete worm Serpula vermicularis (Serpulidae) filterfeeds at the mouth of its calcareous tube, but retreats intothe tube when startled by mechanical stimuli likely to be associatedwith predators. While in its tube, a worm is safe but cannotfeed. Thus, hiding has a lost-opportunity cost. We show thatthis cost can be substantial, given that food in the naturalhabitat appears in pulses, and good feeding conditions may notlast long or recur frequently. We expect that a worm's hidingtime will be sensitive to the lost-opportunity cost, and wepresent data from a series of experiments that support thisprediction. The worms seem able to track relatively short-termchanges in food availability, and some evidence suggests thatthey assess food availability on a relative basis, comparingcurrent feeding conditions to those recently experienced. Hidingand other types of cryptic behavior are common antipredatortactics, and animals may commonly adjust the durations of suchbehaviors to current benefits and costs (including lost opportunity),as they perceive them