Foraging patterns and dietary mixing in the field by the generalist grasshopperBrachystola magna (Orthoptera: Acrididae)

The foraging behavior of a cryptic romaleine grasshopper was observed in its natural habitat in southern Arizona. Individual polyphagy, dietary mixing, and the pattern of feeding on different substrates were monitored. Individuals were found to be extreme generalists with surprising levels of feeding on other insects. Differences in the foraging behavior between grasshopper species are discussed in relation to the habitat, the season, and the different defensive strategies.     

Ontogeny of territoriality in the desert clickerLigurotettix coquilletti (Orthoptera: Acrididae)

Field observations and experiments showed that settlement prior to reproductive maturation strongly influenced territorial success in the desert clicker(Ligurotettix coquilletti), an acridid grasshopper in which males defend individual host plants (Larrea tridentata bushes) as mating arenas. High tendencies to move and reposition among the host plants were displayed in two distinct episodes. The first occurred during early nymphal development, and it resulted in many individuals settling on high-qualityLarrea bushes. Prior work showed thatL. coquilletti that fed on foliage from these specific bushes exhibited higher relative growth rates and therefore were expected to eclose sooner. Early-eclosing males enjoyed a priority advantage in defending high-quality bushes as mating territories, and they consequently encountered more females and mated more frequently. Despite the distribution of most insects on high-quality bushes at the time of eclosion, though, a second movement episode occurred shortly thereafter. This reshuffling contrasts markedly with the site fidelity of mature adult males, most of which settle on mating territories, and it may function as a means of examining an enlarged sample of potential sites. The above results imply that territorial success of maleL. coquilletti does not result from retaining sites defended by the parents. This point is also supported by the finding that females do not oviposit particularly close to theLarrea bushes in which they reside regardless of their quality; most egg pods are deposited in bare soil midway between the bushes. Nonetheless, certain oviposition sites may be conducive to earlier hatch, and this can lead to earlier eclosion and ultimately to defense of a valuable territory. Therefore, the parental generation, through selective oviposition, may yet influence the success of their male offspring.     

Foraging behavior ofPachydiplax longipennis (Odonata: Libellulidae)

Food intake, prey availability, and prey capture behavior at feeding areas were quantified in the dragonflyPachydiplax longipennis by observing focal individuals on artificial perches, where they exhibited marked short-term site fidelity. Prey capture success was high and relatively constant, but the frequency and duration of feeding flights depended on the time of day and season (at least in part because of associated variation in temperature and other physical factors), study site, sex, density of other dragonflies, and prey density. Individuals rapidly responded temporally and spatially to changes in prey availability, particularly to localized prey concentrations.     

Dietary mixing in grasshoppers: switching induced by nutritional imbalances in foods

Dietary mixing by nymphs of Schistocerca americana was studied in the laboratory using artificial diet cakes. Individuals were given either two different inadequate but complementary diet cakes or two adequate and identical ones. When unique flavors (coumarin or NHT) were added to the diet cakes, insects given the inadequate diet treatment switched between cakes more than insects exposed to the adequate diets. This was not the case when no identifying flavors were added.     

Genetics and polymorphism of a duplicated malate dehydrogenase (MDH) locus in the grasshopperOxya japonica japonica (Orthoptera:Acrididae:Oxyinae)

A biochemical genetic study of the enzyme malate dehydrogenase (MDH) was conducted in the grasshopperOxya j. japonica. Analysis of MDH electrophoretic variation in this species of grasshopper shows that one of the two autosomal loci for MDH in grasshoppers, the Mdh-2 locus, controlling the anodal set of MDH isozymes, is duplicated. Results of breeding studies confirm this and the observed polymorphism at theMdh-2 locus in the two populations ofOxya j. japonica studied can be attributed to three forms of linked alleles at the duplicated locus in equilibrium in both populations. In this respect, all individuals of this species possess heterozygous allelic combinations at the duplicatedMdh-2 locus, which may account for the spread of the duplicated locus in the populations of this species of grasshopper.This research was supported by a grant (Vote F) from the University of Malaya, Kuala Lumpur.     

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.     

The importance of palpation in food selection by a polyphagous grasshopper (Orthoptera: Acrididae)

The behavioral responses of final-instar nymphs of Schistocerca americanato a variety of acceptable and unacceptable plants were recorded. Palpation occurred on all plants and the palps are involved in both acceptance and rejection. On most unacceptable plants, rejection was at first dependent on biting the leaf, but subsequently on Lantana, Machaeranthera, Moms,and Physalis,rejection often occurred after palpation of the surface alone. This is consistent with the suggestion that associative learning occurs. This response did not wane even when the insects had been without food for over 2 h. There was also some evidence of an innate response to the surface characteristics of Physalis.The features of the surfaces that produced these responses were not determined.     

The Effect of Food Size and Dispersion Pattern on Retrieval Rate by the Argentine Ant, Linepithema humile (Hymenoptera: Formicidae)

Food acquisition in central-place foraging animals demands efficient detection and retrieval of resources. Most ant species rely on a mass recruitment foraging strategy, which requires that some potential foragers remain at the nest where they can be recruited to food once resources are found. Because this strategy reduces the number of workers initially looking for food, it may reduce the food detection rate while increasing the postdiscovery food retrieval rate. In previous studies this tradeoff has been analyzed by computer simulation and mathematical models. Both kinds of models show that food acquisition rate is greatly influenced by food distribution and resource patch size: as food is condensed into fewer patches, the maximal acquisition rate is achieved by a shift to fewer initial searchers and more potential recruits. In general, these models show that a mass recruitment strategy is most effective when resources are clumped. We tested this prediction in two experiments by letting laboratory colonies of the Argentine ant (Linepithema humile) forage for resources placed in different distributions. When all prey were small, retrieval rate increased with increasing resource patch size, in support of foraging models. When prey were large, however, the mass of prey returned to the colony over time was much lower than when prey were small and widely distributed. As more ants reached a large prey item, the distance the prey item was transported decreased due to a greater emphasis on feeding rather than transport. Because Argentine ants can transport more biomass externally than they can ingest, food retrieval that depends only on ingestion can depress the biomass retrieval rate. Thus, our results generally support theoretical foraging models, but we show how prey size, through differential prey-handling behavior, can produce an outcome greatly different from that predicted only on the distribution of resources.     

Female preferences for large, heavy mates inSchistocerca americana (Orthoptera: Acrididae)

The significance of female choice in shaping mating systems remains problematic. The following study clearly documents female preferences in a species of grasshopper,Schistocerca americana, over the course of 3 years. Three sets of animals were observed in an outdoor seminatural cage. Females were found to mate more often with large, heavy males. Furthermore, remating by a female within a week was less likely if she mated with a large or heavy male. Because females were found to oviposit 2 days after a mating on average, these large heavy males may also be chosen cryptically (as defined in the literature) and father more offspring.     

Foraging behavior of the communal spider,Philoponella republicana (Araneae: Uloboridae)

The communal orb-weaving spider, Philoponella republicana,was observed in the subtropical moist forest of Southeast Peru. These spiders live in colonies of conspecifics whose individual orbs are connected by silk. The wrapping of a prey prior to feeding is a large component of the prey capture process because P. republicanahas no venom with which to kill an insect. Wrapping time was the only aspect of prey capture that was strongly correlated with the size of the insect captured. Occasionally we observed several individuals working together to wrap a prey item. These joint efforts were more frequent on prey larger than the capturing spider. Although group captures accounted for only 5.5% of captures, they represented 14.7% of the biomass obtained. A comparison of the relationship between wrapping time and prey size for solitary and group efforts suggested that, by working together, the spiders reduced their total handling time. In most cases only one spider fed on the captured prey.     

Foraging Behaviour and Patch Exploitation by Campoplex dubitator (Hymenoptera: Ichneumonidae), a Parasitoid of Bark-mining Larvae

This study examines the foraging behaviour of Campoplex dubitator Horstmann (Hymenoptera: Ichneumonidae), a parasitoid of the highly concealed bark-mining host, Enarmonia formosana Scopoli (Lepidoptera: Tortricidae), and how this foraging behaviour is affected by host density. The ultimate aim was to determine whether foraging behaviour and patch fidelity could explain patterns of parasitism in the field. An ethogram was first constructed to portray C. dubitator behaviour prior to and immediately following host attacks. The pre-attack behavioural sequence was highly structured whereas behavioural transitions became less predictable after a sting event. Females spent more time on patches with higher host densities and host encounters caused a significant reduction in the leaving tendency. The giving up time was only slightly affected by host encounters. Campoplex dubitator did not demonstrate the ability to distinguish between occupied and empty host mines. The results show that C. dubitator forages more efficiently at lower host densities and this behavioural phenomenon may result in the patterns of parasitism observed in the field.     

Consumption rates and the evolution of diet-induced plasticity in the head morphology of Melanoplus femurrubrum (Orthoptera: Acrididae)

Summary Phenotypic plasticity may be an ecologically important evolutionary response to natural selection in multiple environments. I have determined the effect of diet-induced developmental plasticity in the head size of grasshoppers (Melanoplus femurrubrum) onfeeding performance on two types of plants. Full-sib families were divided and raised on either red clover, Trifolium repens, or rye grass, Lolium perenne. In three different stages of ontogeny, grasshoppers raised on rye grass had significantly larger heads, relative to body size, than full-sibs raised on clover. A principal components analysis indicated that two to five relative head size characters covaried as a block in their plastic response to the feeding environment. Regressions of adjusted consumption rates (mg/sec) against relative head size revealed that larger head sizes, induced by the rye grass diet, enhanced consumption rates of rye grass, but not clover. Unexpectedly, a similar positive association was observed between head size and consumption rate for grasshoppers raised on clover when they were feeding on clover. These results support the inference that grasshoppers exhibit adaptive phenotypic plasticity. However, the unexpected influence of head size on consumption rates of clover indicates that the functional relationship between head morphology and feeding performance is complex and that variation in this relationship among plant environments is not sufficient to explain the evolution of diet-induced phenotypic plasticity.     

Transmission and perception of acoustic signals in the desert clicker,Ligurotettix coquilletti (Orthoptera: Acrididae)

Males of the desert clicker, Ligurotettix coquilletti (Acrididae: Orthoptera) defend a femalerequired resource, the creosote bush Larrea tridentata, in desert habitats of the southwestern United States. Males signal acoustically to each other as well as to searching females. The call is produced by tegminal/femoral stridulation where one or both legs are used for sound production. Sound pressure levels, measured laterally, are influenced by the intervening tegmen between the stridulating leg and the microphone. Differences in measured sound pressure levels between sides can vary up to 7 dB. When clicks are produced multiply,these multiple clicks may be 4 dB louder than single clicks. We examine the structure of the call and the effective broadcast area of single males by monitoring acoustic ascending neurons of the ventral nerve cord in the neck. By taking the neurophysiological preparation into the field, we were able to map the broadcast area of isolated males and also of males calling within aggregations. The distance over which the signal of isolated males could be detected was 8–14 m, whereas neural representation of the calls of males within aggregation were detectable within 4–6 m. The sound spectrum of the song, although having a major lower-frequency component around 10 kHz, has extensive power in the ultrasonic range. The tuning characteristics of the ascending auditory neuron matched the overall structure of the male call. The importance of the acoustic cue, as compared to visual cues, is discussed in relation to female attraction.     

The influence of proline on diet selection: sex-specific feeding preferences by the grasshoppers Ageneotettix deorum and Phoetaliotes nebrascensis (Orthoptera: Acrididae)

Diet selection based on the level of proline in an insect's host plant has been observed for a number of phytophagous insects, but few studies have examined potential differences in feeding preferences between males and females. The level of proline among an insect's host plants, particularly in drought-stressed plants, can be highly variable and often is positively correlated with soluble nitrogen levels. Additionally, proline is known to participate in a number of physiological functions in insects. We tested the effect of proline as a feeding stimulant in reproductively active grasshoppers using the graminivorous Ageneotettix deorum and the generalist, but mostly graminivorous, Phoetaliotes nebrascensis. Feeding preference tests using diets with representative free amino acid and sucrose levels but varying proline levels (zero, normal and 3 x normal) were examined. The feeding preference exhibited by both species was sex-specific, although the sex-specific response was more pronounced in P. nebrascensis than in A. deorum. Females of both species displayed preferences for diets high in proline. Males of neither species exhibited a preference for proline when responses were averaged over all treatment levels. However, within specific treatment combinations, male A. deorum preferred diets with high proline over diets with zero proline. These results suggest that diet selection for specific nutrients may vary between males and females because of differences in their physiological status and, possibly, differences in the nutritional requirements associated with reproduction. These results also suggest that subtle shifts in the concentration of individual nutrients within an insect's host plant may greatly influence insect feeding patterns.     

Foraging Behavior and Patch Time Allocation by Fopius arisanus (Hymenoptera: Braconidae), an Egg-Larval Parasitoid of Tephritid Fruit Flies

This study quantitatively describes the host-searching behavior of Fopius arisanus (Sonan) (Hymenoptera: Braconidae), an important egg-larval parasitoid of tephritid fruit fly pests, on coffee berries infested with host eggs of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). We also investigate the parasitoid's response to local variation in host patch quality. The temporal pattern of behavioral organisation was examined by constructing an ethogram. The parasitoid spent over 90% of its foraging time in detecting and locating hosts after arriving on a host-infested fruit, and displayed a relatively fixed behavioral pattern leading to oviposition. Patch residence time increased in the presence of host-associated cues, following successful ovipositions, and with increasing size of host clutches per fruit, but decreased with each successive visit to the same host patch and with increasing availability of alternative host patches. The parasitoid females discriminated against previously parasitized hosts and spent significantly less time and searching effort on patches previously exploited by herself or by conspecific females. The effective host-searching behavior, perfect host discrimination ability, and success-motivated searching strategy shown by F. arisanus ensured a thorough exploitation of host resources by this parasitoid.     

Percussive Foraging: Stimuli for Prey Location by Aye-Ayes (Daubentonia madagascariensis)

Aye-ayes (Daubentonia madagascariensis) use the thin middle finger to tap on wood in search of subsurface cavities containing insect larvae. When a cavity is located, they gnaw away wood until the prey can be extracted. Previous researchers suggested that acoustical cues reveal cavity location. We designed five studies to identify the cavity features that provide acoustical cues. When cavities were backfilled with gelatin or acoustical foam, excavation was still successful, suggesting that the reverberation of sound in air-filled cavities is not necessary for detection. Moreover, when the density of cavity content was varied, there was no difference in excavation frequency. On the other hand, a one-dimensional break in the subsurface wood was an effective stimulus for excavation. These studies suggest that a simple interface beneath the surface is sufficient to elicit excavation and that neither prey nor cavity nor even small air pockets are necessary to elicit the behavior. These results raise provocative questions as to how the aye-aye manages to forage efficiently.     

Alternative schedules of male reproductive diapause in the grasshopperAnacridium aegyptium (L.): Effects of the corpora allata on sexual behavior (Orthoptera: Acrididae)

We studied the ontogeny of male sexual behavior in laboratory cohorts of the grasshopper Anacridium aegyptium to understand how male and female sexual activity is synchronized in this species. Contrary to previous accounts, certain male A. aegyptium undergo an extended period of reproductive diapause and do not show an onset of sexual behavior until several months postfledging. Considerable variation in diapause length existed, however, and some males showed onsets as early as 21 days postfledging, probably a nondiapause condition. A series of allatectomies and implantations of corpora allata (CA) demonstrated that male sexual behavior was dependent upon the presence of active CA and indicated that the periods of natural refractoriness noted above were related to inactive CA. In some cohorts, male longevity was extended in allatectomized individuals or in those exhibiting lengthy reproductive diapause. We speculate that variable diapause periods may represent a bet-hedging strategy of the parental generation selected for by two opposing pressures: the uncertainty of extended survival, selecting for brief periods, and intermale competition in the form of sperm precedence, favoring lengthy diapause.     

The complete mitochondrial genome of Gomphocerus tibetanus Uvarov, 1935 (Orthoptera: Acrididae: Gomphocerinae)

The complete nucleotide sequence of the mitochondrial genome (mitogenome) of Gomphocerus tibetanus Uvarov, 1935 (Orthoptera: Acrididae: Gomphocerinae) was determined. It is 15,571 bp in length and contains 74.8% A + T. All Gomphocerus tibetanus protein-coding sequences start with a typical ATN codon. The usual termination codons (TAA and TAG) were found from 13 PCGs except COI and COII which took incomplete codon T as termination codons. All tRNA genes could be folded into the typical cloverleaf secondary structure, except tRNA^Ser(AGN) lacking of dihydrouridine (D) arm. The sizes of the large and small ribosomal RNA genes are 1313 and 822 bp, respectively. The A + T content of the A + T-rich region is 82.3%. A preliminary analysis on characteristics of Gomphocerinae mitogenome was made by comparision among three Gomphocerinae mitogenomes and Locusta migratoria.