Effect of Paranosema (Nosema) locustae (Microsporidia) on morphological phase transformation of Locusta migratoria manilensis (Orthoptera: Acrididae)

Field and laboratory studies demonstrated that Paranosema (Nosema) locustae had significant effects on the morphological phase transformation of Locusta migratoria manilensis (Meyen 1835). In the field, spraying P. locustae on gregarious locusts caused a substantial population reduction by 16 days after treatment, with most of the surviving locusts being phase solitaria. However, the effects of P. locustae on locust phase transformation began before direct mortality had caused a substantial reduction in locust density: locust numbers were still high at day 10, but locusts had already transformed to phase transiens. Laboratory assays showed that while a low dose of P. locustae had no effect on phase transformation, at a higher dose of 1×10⁵ spores/mL, locusts had F/C ratios that were significantly (P<0.05) more solitaria than untreated locusts, with locusts having ratios that were either phase solitaria or on the solitaria side of phase transiens. In a second laboratory experiment that analysed the effects of locust density on phase transformation by P. locustae, there was no obvious effect of density on female locusts 10 days later as all were solitaria at all locust densities. At day 16, female locusts were transiens at higher densities, but were solitaria at 4/cage. With males there were lesser effects. These results provide new evidence for P. locustae having sub-lethal effects on locust phase transformation at a wide range of locust densities.     

Relationships between food quality and fitness in the desert locust, Schistocerca gregaria, and its distribution over habitats on the Red Sea coastal plain of Sudan

The effect of millet, Pennisetum typhoideum Rich. (Poaceae), leaf nitrogen content on fitness parameters of the desert locust, Schistocerca gregaria Forsk. (Orthoptera: Acrididae), was studied under laboratory conditions. Locusts reared on high‐nitrogen leaves were larger, developed faster, had higher survival, reproduced more and earlier, and showed greater synchronization than those fed on low‐nitrogen leaves. Active and passive cannibalism contributed to mortality when locusts were reared on low‐nitrogen leaves, but not when reared on high‐nitrogen leaves. Elevated leaf nitrogen content of host plants increased net reproduction and intrinsic rate of increase, and lowered generation time. The findings show that nitrogen content of host plants affects the potential for population increase in the desert locust. Leaf samples of common plant species were collected in the Heliotropium arbainense (Fresen.) (Boraginaceae) and Panicum turgidum (Forssk.) (Poaceae) plant communities on the Red Sea coastal plain of Sudan during the winters of 1999 and 2000. The levels of leaf nitrogen in host plants were comparable to those in the laboratory studies and consistently higher in plant samples from the Heliotropium community than in samples from the Panicum community. Both in 1999 and 2000, locust densities were much higher in the Heliotropium than in the Panicum plant community. It should be assessed whether the desert locust would be attracted to sites where host plants have high leaf nitrogen content, as this would not only increase their fitness, but also the likelihood of gregarization and outbreaks.     

Invasion of the pathogenic fungus Metarhizium anisopliae through the guts of germfree desert locusts,Schistocerca gregaria

Less than 1% of an ingested inoculum of the pathogenic fungus Metarhizium anisopliae was retained for long enough (ca. 24 h) in the gut of the desert locust, Schistocerca gregaria, for germination and penetration to have occurred. The residual inoculum did not initiate an infection in guts of fed conventional or axenic locusts. However, symptoms of mycosis (hyphal bodies in the haemolymph, fungal penetration of the hindgut intima and epithelium, tetanic paralysis) were consistently observed in axenic but not conventional locusts which were starved post-inoculation.It is concluded that the antifungal toxin produced by the gut bacteria defends the desert locust against gut invasion by Metarhizium anisopliae during periods of starvation when the physical defences, prominent in fed insects, are less apparent.     

Estimation of density threshold of gregarization of desert locust hoppers from field sampling in Mauritania

For desert locusts, Schistocerca gregaria (Forskål) (Orthoptera: Acrididae), the hopper density threshold of gregarization remains poorly documented. Field sampling was carried out in traditional seasonal breeding areas of Mauritania during two successive years without invasion to approximate the gregarization threshold. Hopper densities were assessed at numerous sampling sites. Vegetation was also sampled to characterize the habitats. Hopper behavior was analyzed in situ with the help of a behavioral circular arena to test our assumptions on empirical locust phases determination based on physical appearance (coloration and behavior) following FAO guidelines. The results provided a critical density value around 2.45 hoppers m^?2, above which gregarious hoppers were expected to be seen more frequently in nature. Hopper density was confirmed as the main factor explaining the presence of gregarious individuals. The level of involvement of vegetation parameters such as plant density, basal area, volume, distance between plants, greenness, or combinations of these indicators was low in explaining the observation of gregarious hoppers compared with hopper density. Vegetation cover and height were the only vegetation characteristics that could enhance the prediction of phase status with hopper density. The hoppers' phase determined from their behavior observed in the arena was similar to that characterized through FAO guidelines phase assessment, making consistent the field sampling method. Additionnally, the use of this arena illustrated that the grouping behavior of hoppers is a gradual response to density. This study can be seen as a step forward in the estimation of hopper density thresholds of gregarization in the field. This should improve the decision making for intervention during preventive control operations.     

Diversity of locust gut bacteria protects against pathogen invasion

Diversity–invasibility relationships were explored in the novel context of the colonization resistance provided by gut bacteria of the desert locust Schistocerca gregaria against pathogenic bacteria. Germ‐free insects were associated with various combinations of one to three species of locust gut bacteria and then fed an inoculum of the pathogenic bacterium Serratia marcescens. There was a significant negative relationship between the resulting density of Serratia marcescens and the number of symbiotic gut bacterial species present. Likewise there was a significant inverse relationship between community diversity and the proportion of locusts that harboured Serratia. Host mortality was not negatively correlated with resistance to gut‐invasion by Serratia marcescens, although there were significantly more deaths among pathogen fed germ‐free insects than tri‐associated gnotobiotes. The outcome is consistent with the predictions of community ecology theory that species‐rich communities are more resistant to invasion than species‐poor communities.     

Effect of Paranosema locustae (Microsporidia) on the behavioural phases of Locusta migratoria (Orthoptera: Acrididae) in the laboratory

The ability of parasites to modify the behaviour of their hosts is a wide spread phenomenon, but the effects of microsporidian parasites on locust behaviour remain unexplored. Here the frequencies of directional changes (ND) and jumping (NJ) per minute of gregarious locusts infected with 2000 spores of the microsporidian parasite Paranosema locustae were significantly different from those of untreated locusts 10 and 16 days after infection, being similar to values for solitary nymphs. In contrast, the behaviour of locusts inoculated with the lower doses of 200 spores/locust was sometimes like that of solitary nymphs. At other times, behaviour was intermediate between solitary and gregarious, i.e. transitional. The rearing density did not affect the turning and jumping behaviour of infected locusts, and their behaviours were similar to those of solitary locusts at 10–16 days after infection. Our study demonstrates that infection with P. locustae may lead gregarious locusts to change some of their behaviour to that typical of solitary locusts.     

Composition and emission dynamics of migratory locust volatiles in response to changes in developmental stages and population density

Chemical communication plays an important role in density‐dependent phase change in locusts. However, the volatile components and emission patterns of the migratory locust, Locusta migratoria, are largely unknown. In this study, we identified the chemical compositions and emission dynamics of locust volatiles from the body and feces and associated them with developmental stages, sexes and phase changes. The migratory locust shares a number of volatile components with the desert locust (Schistocerca gregaria), but the emission dynamics of the two locust species are significantly different. The body odors of the gregarious nymphs in the migratory locust consisted of phenylacetonitrile (PAN), benzaldehyde, guaiacol, phenol, aliphatic acids and 2,3‐butanediol, and PAN was the dominant volatile. Volatiles from the fecal pellets of the nymphs primarily consist of guaiacol and phenol. Principal component analysis (PCA) showed significant differences in the volatile profiles between gregarious and solitary locusts. PAN and 4‐vinylanisole concentrations were significantly higher in gregarious individuals than in solitary locusts. Gregarious mature males released significantly higher amounts of PAN and 4‐vinylanisole during adulthood than mature females and immature adults of both sexes. Furthermore, PAN and 4‐vinylanisole were completely lost in gregarious nymphs during the solitarization process, but were obtained by solitary nymphs during gregarization. The amounts of benzaldehyde, guaiacol and phenol only unidirectionally decreased from solitary to crowded treatment. Aliphatic aldehydes (C7 to C10), which were previously reported as locust volatiles, are now identified as environmental contaminants. Therefore, our results illustrate the precise odor profiles of migratory locusts during developmental stages, sexes and phase change. However, the function and role of PAN and other aromatic compounds during phase transition need further investigation.     

Take-off activity in caged desert locusts,Schistocerca gregaria (Forsk.) (Orthoptera,Acrididae) in relation to meteorological disturbances

Take-off activity of the desert locust,Schistocerca gregaria (Forsk.), phasegregaria, was studied in a temperature-controlled laboratory environment and possible relationships with certain meteorological factors were analysed. A statistically significant relationship between the passage of cold fronts and an increased take-off rate was found. The reaction of the locusts could not be correlated to changes in VLF electromagnetic radiation or to the atmospheric pressure.     

Central nervous processing of behaviourally relevant odours in solitary and gregarious fifth instar locusts, Schistocerca gregaria

Physiological and morphological characteristics of antennal lobe neurons of solitary and gregarious fifth-instar nymphs of the desert locust, Schistocerca gregaria, were studied using intracellular recording and staining techniques. Physiological characteristics of antennal lobe neurons of both locust phases responding to stage-dependent aggregation pheromones, egg-laying attractants, a putative sex pheromone and plant-associated volatiles are described. Antennal lobe neurons showed excitatory, inhibitory, combined excitatory and inhibitory and delayed responses. In addition, one neuron␣showing an initial inhibition followed by an excitation and inhibition response was found. Pheromone-specific-, plant-specific- and pheromone-plant-generalist neurons were found in both locust phases. Antennal lobe neurons displayed stage- and phase-dependent differences in the processing of aggregation pheromone component input. Nymphal antennal lobe neurons showed stage-dependent response characteristics highly correlated with the preferential behavioural attraction to the nymphal aggregation pheromone. Phase-dependent differences were found in the response spectra and the sensitivity of the same neuron types. Neurons of solitary locusts responded significantly more frequently to some of the tested components than neurons of gregarious locusts. Furthermore, antennal lobe neurons of solitary locusts showed a higher sensitivity to most of the tested compounds. Accepted: 4 July 1998     

Behavior of geladas and other endemic wildlife during a desert locust outbreak at Guassa,Ethiopia: ecological and conservation implications

Desert locust (Schistocerca gregaria) outbreaks have occurred repeatedly throughout recorded history in the Horn of Africa region, devastating crops and contributing to famines. In June 2009, a desert locust swarm invaded the Guassa Plateau, Ethiopia, a large and unusually intact Afroalpine tall-grass ecosystem, home to important populations of geladas (Theropithecus gelada), Ethiopian wolves (Canis simensis), thick-billed ravens (Corvus crassirostris), and other Ethiopian or Horn of Africa endemics. During the outbreak and its aftermath, we observed many animals, including geladas, ravens, and a wolf, feeding on locusts in large quantities. These observations suggest surprising flexibility in the normally highly specialized diets of geladas and wolves, including the potential for temporary but intensive insectivory during locust outbreaks. To our knowledge, Guassa is the highest elevation site (3,200–3,600 m) at which desert locusts, which require temperatures >20°C for sustained flight, have been reported. Continued monitoring will be necessary to determine whether the June 2009 outbreak was an isolated incident or part of an emerging pattern in the Ethiopian Highlands linked to global warming. The intensive consumption of desert locusts by geladas, wolves, and ravens during the outbreak at Guassa raises concerns about pesticide-based locust control strategies and potential unintended adverse effects on endemic and endangered wildlife.     

Effect of vegetation on density thresholds of adult desert locust gregarization from survey data in Mauritania

In the desert locust, Schistocerca gregaria (Forskål) (Orthoptera: Acrididae), the threshold density inducing the gregarization phenomenon has never been determined under natural conditions. The influence of environmental factors on this phenomenon has been studied mostly in controlled environments. Based on data collected during several years by the survey teams of the National Center for Locust Control in Mauritania, we analyzed the influence of locust density, vegetation cover, and vegetation status on the probability of observing gregarious locusts. We assumed that a probability to observe gregarious locusts of 0.5 corresponded to the density threshold of gregarization. The results showed in detail the change in the threshold of gregarization according to the cover and status of the vegetation. Low cover and dry vegetation led to a low density threshold of gregarization probably due to high probability of individuals to touch each other. Dense and green vegetation favored a high threshold of gregarization probably due to a dispersion of the individuals and a low probability of individual encounters. These findings should help the management of locusts and decision making during control operations.     

PCR primers for polymorphic microsatellite loci in the desert locust,Schistocerca gregaria (Orthoptera: Acrididae)

Nine pairs of polymerase chain reaction (PCR) primers that amplify polymorphic microsatellite loci in the desert locust, Schistocerca gregaria (Forskal), were developed using a magnetic bead‐based enrichment protocol. A sample of 48 locusts collected during the 1993 and 1995 upsurge periods in Eritrea, East Africa, were genotyped. The number of alleles per locus ranged from six to 20; the average was 12.67. Allelic distributions were significantly different between samples from different localities.     

The influence of environmental microstructure on the behavioural phase state and distribution of the desert locust Schistocerca gregaria

Abstract. In previous studies we used logistic regression analysis to quantify the change in behavioural phase state of Schistocerca gregaria (Forskål) nymphs subjected to variations in population density. Such work involved restricting insects in small containers either alone or in a crowd. In the present paper we have shown that the fine-scale distribution of food plants, perches and favourable microclimatic sites influences the spatial distribution of locusts, both in the laboratory and under semi-field conditions. When multiple resource sites were provided, solitarious locusts tended to disperse and behavioural gregarization was inhibited. However, provision of only a single site promoted congregation, overcoming the tendency of solitarious insects to avoid each other, and led to behavioural gregarization. The time-course and extent of this response was fully consistent with our earlier experiments using enforced crowding. We suggest that such quantitative, experimental studies of the effects of environmental microstructure on behaviour may yield fundamental insights into the dynamics of plague formation in the desert locust.     

Is there an intraspecific role for density-dependent colour change in the desert locust?

Attempts to uncover the adaptive significance of density-dependent colour polyphenism in the desert locust, Schistocerca gregaria (Orthoptera: Acrididae), have been unsuccessful. Desert locust juveniles can change colour as part of a phenotypically plastic response to changes in local population density known as phase polyphenism. They are typically cryptic in colour at low rearing density (solitarious phase), but become conspicuous at high density (gregarious phase). Recent evidence indicates that this colour change functions interspecifically as an aposematic signal. Other recent evidence, however, suggests that previous attempts to demonstrate an intraspecific function of gregarious coloration in mediating group interactions among locusts may have been confounded by the effects of multiple sensory cues. We reinvestigated the intraspecific function of density-dependent colour polyphenism and specifically controlled for potentially confounding olfactory and tactile cues. We found no effect of gregarious phase (yellow and black) coloration as either a gregarizing stimulus to behaviourally solitarious locusts or as a visual aggregation stimulus behaviourally to gregarious locusts. We did, however, find that nonmoving solitarious phase (green) coloration significantly increased the activity levels of behaviourally gregarious locusts. We cannot explain this result and its biological relevance remains unknown. In the absence of support for the intraspecific visual cue hypothesis, we favour an aposematic perspective on the function of density-dependent colour polyphenism in the desert locust. The aposematic perspective parsimoniously accounts for density-dependent changes in both colour and behaviour. Copyright 2000 The Association for the Study of Animal Behaviour.     

Defence tactics cycle with diel microhabitat choice and body temperature in the desert locust,Schistocerca gregaria

We examined environmental factors influencing plasticity in antipredator defences of adult gregarious desert locusts, Schistocerca gregaria, including daily cycles in temperature, light, microhabitat occupied and predator threat. In the Sahara Desert in Mauritania, West Africa, daily temperature fluctuated widely from below locusts’ cold thermal limits for jump‐ and flight‐defence, to above their preferred body temperature. Locusts changed microhabitats throughout the 24‐hr period in synchrony with the daily thermo‐photocycle. They roosted in tall trees and large bushes at night, moved to the ground in the morning, shaded under or in small bushes and annuals at midday, moved back to the ground in the afternoon and then returned to night roosting sites around dusk. Locust antipredator defences varied throughout the 24‐hr period, and these changes were correlated with temperature, photocycle and habitat. Flight escape was associated with daytime, high temperatures and the ground habitat. Dropping escape (= releasing hold of vegetation and dropping to the ground or into vegetation) was associated with cool temperatures and low‐to‐medium sized bushes. Stationary behaviour was associated with the tree microhabitat and height off the ground. Roosting (a primary defence) was associated with cool temperatures at night and early morning, tree habitats and nocturnal ground‐foraging times of endothermic mammals. In summary, we propose that temperature is the key factor in determining both changing microhabitat choice and changing antipredator defence, due to thermal constraints on locust muscle for this ectothermic insect. The thermocycle also influences temporal predator loads, which influence the evolution of locust diel defence strategies. These various environmental factors not only influence one another, they also interact to influence antipredator defence expression in locusts. Overall, our study suggests that plasticity in locust antipredator defences is a complicated matter mediated by the interactions of multiple environmental factors and physiological and ecological constraints and trade‐offs.     

Effect of phase status on responses to AKHI in the desert locust,Schistocerca gregaria gregaria

Hyperlipaemic response to adipokinetic hormone (AKH I) was demonstrated in both solitary and gregarious phases of the desert locust, Schistocerca gregaria gregaria. Time-course studies showed that the gregarious locusts had a faster response to the hormone than their solitary counterparts. At peak response time (90 min), the gregarious locusts were more sensitive to AKH I doses below 2 pmol while the solitary locusts had a higher response above this dose. Upon injection of the hormone, lipoprotein conversion occurred, resulting in the formation of the low density lipoprotein (LDLp). The LDLp formed in the gregarious locusts was much larger than that of the solitary locusts. The fat body lipid reserve (expressed as % fat body dry weight) was significantly (P < 0.01) higher in the gregarious (79.02 ± 2.77%) than in the solitary locusts (65.23 ± 2.55%). Triacylglycerol was the major lipid class representing 83.9 and 73.9% of the total lipids in gregarious and solitary locusts, respectively. The higher fat body lipid reserves and efficient LDLp formation in response to AKH in gregarious locusts compared to solitary locusts suggests a physiological adaptation for prolonged flights. © 1996 Wiley-Liss, Inc.     

Does landscape composition alter the spatiotemporal distribution of the pine processionary moth in a pine plantation forest?

Landscape composition and physiognomy affect community structure and species distribution across space and time. The pine processionary moth (PPM) (Thaumetopoea pityocampa Den. & Schiff., Lepidoptera, Notodontidae) is a common pine defoliator throughout southern Europe and Mediterranean countries. We surveyed the spatiotemporal distribution of the PPM in a pine plantation forest in southwestern France and used the density of the winter nests as a proxy for population density. The study spanned 4 years (2005–2008) and showed a high temporal variability in nest density. We found a strong edge effect with nest densities at stand edges more than twice as large as within-stand densities. At the landscape scale, the spatial distribution of the moth exhibited a significant spatial autocorrelation in 3 out of 4 years of our study. The spatial scales of the autocorrelation ranged from ca. 2 km to more than 22 km. We found a positive correlation between spatial distributions corresponding to certain sampling years, but the relationship was not systematic. Landscape configuration appeared to be an important driver of the PPM spatial pattern. Bivariate Moran’s I correlograms showed that patch richness density as well as the percentage of local landscape covered by various land uses were correlated with population density. The study showed that accounting for landscape characteristics may be important in order to understand forest insect pest distribution, even in cases where the host species is abundant and homogeneously distributed throughout the study area, e.g., pure plantation forests.     

Functional Characterization of the Short Neuropeptide F Receptor in the Desert Locust,Schistocerca gregaria

Whereas short neuropeptide F (sNPF) has already been reported to stimulate feeding behaviour in a variety of insect species, the opposite effect was observed in the desert locust. In the present study, we cloned a G protein-coupled receptor (GPCR) cDNA from the desert locust, Schistocerca gregaria. Cell-based functional analysis of this receptor indicated that it is activated by both known isoforms of Schgr-sNPF in a concentration dependent manner, with EC₅₀ values in the nanomolar range. This Schgr-sNPF receptor constitutes the first functionally characterized peptide GPCR in locusts. The in vivo effects of the sNPF signalling pathway on the regulation of feeding in locusts were further studied by knocking down the newly identified Schgr-sNPF receptor by means of RNA interference, as well as by means of peptide injection studies. While injection of sNPF caused an inhibitory effect on food uptake in the desert locust, knocking down the corresponding peptide receptor resulted in an increase of total food uptake when compared to control animals. This is the first comprehensive study in which a clearly negative correlation is described between the sNPF signalling pathway and feeding, prompting a reconsideration of the diverse roles of sNPFs in the physiology of insects.     

The locust foraging gene

Our knowledge of how genes act on the nervous system in response to the environment to generate behavioral plasticity is limited. A number of recent advancements in this area concern food‐related behaviors and a specific gene family called foraging (for), which encodes a cGMP‐dependent protein kinase (PKG). The desert locust (Schistocerca gregaria) is notorious for its destructive feeding and long‐term migratory behavior. Locust phase polyphenism is an extreme example of environmentally induced behavioral plasticity. In response to changes in population density, locusts dramatically alter their behavior, from solitary and relatively sedentary behavior to active aggregation and swarming. Very little is known about the molecular and genetic basis of this striking behavioral phenomenon. Here we initiated studies into the locust for gene by identifying, cloning, and studying expression of the gene in the locust brain. We determined the phylogenetic relationships between the locust PKG and other known PKG proteins in insects. FOR expression was found to be confined to neurons of the anterior midline of the brain, the pars intercerebralis. Our results suggest that differences in PKG enzyme activity are correlated to well‐established phase‐related behavioral differences. These results lay the groundwork for functional studies of the locust for gene and its possible relations to locust phase polyphenism. © 2010 Wiley Periodicals, Inc.     

Contrasting scales of oviposition and parasitism in praying mantids

We report on spatial patterns of parasitism of oothecae (egg cases) of praying mantises (Stagmomantis limbata) by torymid wasps, Podagrion spp. Using collections of mapped mantid oothecae from Riparian sites in the Sonoran desert and Grassland sites in the Chiricahua Mountains (both in Arizona, USA), we characterized the spatial distributions of oothecae and parasitism. The likelihood of an egg case suffering some parasitism was higher at Grassland sites, which had high oothecal densities, than at low-density Riparian sites. However, experimental isolation of Grassland oothecae to densities comparable to Riparian sites reduced parasitism rates. At Riparian sites, parasitized oothecae exhibited on average the same extent of parasitism as parasitized oothecae at high densities but with much greater variation. Indeed, large fractions of Riparian oothecae suffered both severe (>50%) and light (<20%) parasitism, whereas most parasitized Grassland oothecae suffered intermediate levels of parasitism. Analysis of first nearest neighbor distances indicated that the parasite load of an ootheca did not depend on its immediate isolation. However, extending the analysis to include subsequent nearest neighbors (using a technique from spatial statistics called the R(K) function), we found that even though oothecae of S. limbata were spatially clustered, some oothecae in a (statistically defined) cluster escaped parasitism when overall oothecal densities were low. This pattern suggests that when oothecae are sparsely distributed, Podagrion wasps exploit only a fraction of the oothecae available locally, even though the oothecae are strongly aggregated relative to their overall density. We suggest this lack of congruency in the scales of oothecal deposition and parasitism at low densities (which is absent when oothecae are at high densities) may be explained in part by behavioral aspects of the parasite's reproduction, including increased host fidelity by relatively sedentary female parasites. Received: June 13, 2000 / Accepted: October 16, 2000