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.     

Phase polyphenism and preventative locust management

The ecology of phase polyphenism plays a major role in locust swarm formation. We describe how recent advances in the understanding of phase polyphenism can be combined with existing management approaches as part of a preventative Desert locust management strategy. We start with a brief overview of phase polyphenism with particular emphasis on the role that resource distribution patterns play in the process of locust phase change. We then review current perspective on preventative locust management, and conclude by proposing a framework for quantitatively assessing the risk that phase change will occur in local locust populations. Importantly, the data required to implement this framework can be readily collected with little additional effort or cost just by slightly modifying locust habitat survey protocols that are already in operation. Incorporating gregarization risk assessment into existing preventative management strategies stands to make a considerable contribution toward realizing sustainable goals of reductions in the pesticide, manpower and financial support necessary to combat Desert locust upsurges, outbreaks and ultimately plagues.     

Small-scale processes in desert locust swarm formation: how vegetation patterns influence gregarization

Desert locusts ( Schistocerca gregaria ) change phase in response to population density: 'solitarious' insects avoid one another, but when crowded they shift to the gregarious phase and aggregate. This individual-level process is the basis for population-level responses that may ultimately include swarm formation. We have recently developed an individual-based model of locust behavior in which contagious resource distribution leads to phase change. This model shows how population gregarization can result from simple processes operating at the individual level. In the present study, we performed a series of laboratory experiments in which vegetation pattern and locust phase state were assigned quantitative, measurable indices. The pattern of distribution of the resource was represented via fractal dimension; the phase state was evaluated using a behavioral assay based on logistic regression analysis. Locusts were exposed to different patterns of food resource in an artificial arena, after which their behavioral phase state was assayed. These experiments showed that when the distribution of the vegetation was patchy, locusts were more active, experienced higher levels of crowding, and became more gregarious. These results are consistent with simulation predictions and field observations, and demonstrate that small-scale vegetation distribution influences individual behavior and phase state and plays a role in population-level responses.     

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.     

Relating rainfall and vegetation greenness to the biology of spur‐throated and Australian plague locusts

• 1 Rainfall and vegetation greenness are widely claimed to influence fat content and egg development in locusts. Body size, abdominal fat and egg length of spur‐throated Austracris guttulosa and Australian plague Chortoicetes terminifera locusts were related to the normalized difference vegetation index (NDVI; a greenness indicator) to test these assumptions and quantify biological responses.
• 2 Cumulative rainfall was highly correlated with NDVI values of an area (25 × 25 km) surrounding the trap to which locusts were attracted. Catches of locusts were greater in hotter periods preceding rain, associated with an increase in NDVI indicative of brown (0.298) versus green (0.465) vegetation.
• 3 Pre‐overwintering descendent A. guttulosa were larger and fattier than post‐overwintering antecedents. Egg maturation of antecedents occurred coincident with increasing NDVI. Higher abdominal fat in pre‐overwintering locusts was positively associated with higher NDVI.
• 4 Male C. terminifera of two descendent generations were larger than their antecedents but only pre‐overwintering locusts were significantly fattier than their predecessors. Only pre‐overwintering females were significantly larger than their predecessors and no generation was significantly fattier than any other. Fat content was negatively correlated with egg maturation and differed significantly with NDVI, although strong and consistent relationships were not obtained.
• 5 The findings suggest that locust–environment interactions are species‐ and habitat‐specific. If A. guttulosa hoppers develop during periods when regional NDVI exceeds 0.4, fatter adults will arise. The elucidation of relationships between greenness, fat and egg development in C. terminifera is unlikely using NDVI values from areas comprising mixtures of monocotyledonous and dicotyledonous vegetation.

     

Landscape structure and locust swarming: a satellite's eye view

Desert locust Schistocerca gregaria outbreaks consistently start in the same places, suggesting that certain landscapes are particularly favourable for outbreaking. Outbreaks are generated by multiplication, concentration and gregarisation of locust populations. Previous research has shown how small-scale vegetation patterns in desert ecosystems influence locust gregarisation; the present study examines the effects of large-scale landscape structure on locust multiplication and concentration. NOAA/AVHRR satellite imagery was used to relate abundance and spatial distribution of resources at the landscape scale to the historical record of locust outbreaks. Threshold NDVI values were investigated to define what constitutes 'resource' for locusts. The first part of the study showed that abundance and spatial distribution of resource were not sufficient to distinguish between outbreak and non-outbreak areas in the western part of the locust distribution area. Thus, outbreak danger zones cannot be identified by landscape structure at this spatial resolution. The second analysis investigated spatio-temporal patterns of vegetation growth in two locust breeding areas with very different landscape structure; in both cases, the patterns differed significantly between outbreaking and non-outbreaking years. In Mauritania, a flat homogeneous desert landscape, both resource abundance and fragmentation were higher in outbreaking years. On the Red Sea coast, a fragmented landscape, resource spatial distribution was consistent between years, and abundance alone was a significant predictor of outbreaking. High resource abundance promotes locust multiplication, and contraction of resource into small patches increases locust concentration; these two mechanisms explain how landscape structure influences locust outbreaking.     

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.     

CSP and takeout genes modulate the switch between attraction and repulsion during behavioral phase change in the migratory locust

Behavioral plasticity is the most striking trait in locust phase transition. However, the genetic basis for behavioral plasticity in locusts is largely unknown. To unravel the molecular mechanisms underlying the behavioral phase change in the migratory locust Locusta migratoria, the gene expression patterns over the time courses of solitarization and gregarization were compared by oligonucleotide microarray analysis. Data analysis revealed that several gene categories relevant to peripheral olfactory perception are strongly regulated in a total of 1,444 differentially expressed genes during both time courses. Among these candidate genes, several CSP (chemosensory protein) genes and one takeout gene, LmigTO1, showed higher expression in gregarious and solitarious locusts, respectively, and displayed opposite expression trends during solitarization and gregarization. qRT-PCR experiments revealed that most CSP members and LmigTO1 exhibited antenna-rich expressions. RNA interference combined with olfactory behavioral experiments confirmed that the CSP gene family and one takeout gene, LmigTO1, are involved in the shift from repulsion to attraction between individuals during gregarization and in the reverse transition during solitarization. These findings suggest that the response to locust-emitted olfactory cues regulated by CSP and takeout genes is involved in the behavioral phase change in the migratory locust and provide a previously undescribed molecular mechanism linked to the formation of locust aggregations.     

Behavioural phase change in the Australian plague locust, Chortoicetes terminifera, is triggered by tactile stimulation of the antennae

Density-dependent phase polyphenism is a defining characteristic of the paraphyletic group of acridid grasshoppers known as locusts. The cues and mechanisms associated with crowding that induce behavioural gregarization are best understood in the desert locust, Schistocerca gregaria, and involve a combination of sensory inputs from the head (visual and olfactory) and mechanostimulation of the hind legs, acting via a transient increase in serotonin in the thoracic ganglia. Since behavioural gregarization has apparently arisen independently multiple times within the Acrididae, the important question arises as to whether the same mechanisms have been recruited each time. Here we explored the roles of visual, olfactory and tactile stimulation in the induction of behavioural gregarization in the Australian plague locust, Chortoicetes terminifera. We show that the primary gregarizing input is tactile stimulation of the antennae, with no evidence for an effect of visual and olfactory stimulation or tactile stimulation of the hind legs. Our results show that convergent behavioural responses to crowding have evolved employing different sites of sensory input in the Australian plague locust and the desert locust.     

Cannibalism can drive the evolution of behavioural phase polyphenism in locusts

During outbreaks, locust swarms can contain millions of insects travelling thousands of kilometers while devastating vegetation and crops. Such large-scale spatial organization is preceded locally by a dramatic density-dependent phenotypic transition in multiple traits. Behaviourally, low-density 'solitarious' individuals avoid contact with one another; above a critical local density, they undergo a rapid behavioural transition to the 'gregarious phase' whereby they exhibit mutual attraction. Although proximate causes of this phase polyphenism have been widely studied, the ultimate driving factors remain unclear. Using an individual-based evolutionary model, we reveal that cannibalism, a striking feature of locust ecology, could lead to the evolution of density-dependent behavioural phase-change in juvenile locusts. We show that this behavioural strategy minimizes risk associated with cannibalistic interactions and may account for the empirically observed persistence of locust groups during outbreaks. Our results provide a parsimonious explanation for the evolution of behavioural plasticity in locusts.     

外来物种刺槐对土壤微生物功能多样性的影响

以延河流域不同植被区内人工刺槐(Robinia pseudoacacia)群落和乡土植物群落的土壤微生物为研究对象,利用Biolog微平板技术对土壤微生物功能多样性进行测定,分析人工引种刺槐在不同环境梯度(3个植被区)下对土壤微生物功能多样性的影响。结果表明:刺槐对不同环境梯度下的土壤微生物的影响明显不同。从草原区到森林区,刺槐林之间土壤微生物群落的平均颜色变化率(AWCD)和土壤微生物功能多样性指数均没有显著变化;但与乡土植物群落比较,草原区、森林草原区和森林区土壤AWCD分别表现为刺槐乡土植物、刺槐乡土植物、刺槐乡土植物;在草原区和森林草原区刺槐林土壤微生物群落的群落丰富度指数(S)、Shannon-Wiener指数(H)、Simpson指数(D)、Mc Intosh指数(U)均大于乡土植物,森林区刺槐林群落丰富度指数(S)、Shannon指数(H)、Simpson指数(D)均小于乡土植物;刺槐林和乡土植物群落下土壤微生物碳源利用存在差异,主要体现在对糖类、氨基酸类的利用上。PCA分析显示主成分1贡献较大的碳源有24种,在主成分分离中起主要贡献作用的是糖类、氨基酸类和羧酸类。土壤碳氮含量能影响土壤微生物功能多样性指数,土壤含水量和温湿度能够影响碳源的利用类型。刺槐对土壤微生物功能多样性的影响存在区域差异,在评价刺槐对土壤生态过程与功能的影响时必须要考虑这种空间差异性。     

Estimation of climatic favourable areas for locust outbreaks in Spain: integrating species' presence records and spatial information on outbreaks

For millennia, locust swarms have recurrently devastated crop productivity across continents. In much of Europe, locust outbreaks have been considerably reduced by human pressure in recent decades, but important foci of outbreaks still exist in Spain. Distribution models are often used to derive spatial hypotheses and risk maps. Because insufficient information is available to include the extreme plasticity of the solitary and gregarious phases of locusts in large‐scale spatial models, we modelled the distribution either of Acrididae species or of outbreaks per se. Confirmed occurrences of Dociostaurus, Calliptamus and Chorthippus species were obtained from a field survey complemented by museum collection data and the published literature. The locations of confirmed or potential outbreaks covering two time periods of 20 years each were obtained from the literature and from Spanish autonomous community reports. Models were built with one topographic and eleven climatic predictors. We evaluated the ability of different models to predict outbreak recurrence and found that models based on Moroccan locust data or outbreak occurrence data performed the best. We generated a predictive map of the climatic favourability for locust outbreaks in Spain and found that the major foci of locust swarms were encompassed by those areas categorized by the models as areas of highest risk. Predictive maps of outbreak favourability can facilitate the more sustainable use of insecticides and more efficient integrated pest management.     

A behavioural analysis of phase change in the desert locust

A programme of research into phase change in the desert locust, Schistocerca gregaria, is described. The ability to change phase between solitarious and gregarious forms in response to population density is a key feature of locusts and is central to their occasional yet catastrophic impact on humans. Phase polymorphism is an extreme form of phenotypic plasticity. The most labile phase characteristic is behaviour. It is argued that a fully integrated study of behavioural phase change provides a powerful tool for understanding both the mechanisms of phase change and locust population dynamics, both of which offer possibilities for improved management and control of desert locust plagues. An assay for measuring behavioural phase-state in individual locusts was derived, based on logistic regression analysis. Experiments are described that used the assay to quantify the time-course of behavioural change, both within the life of individual locusts and across generations. The locust-related stimuli that provoke behavioural gregarization were investigated. Complex interactions were found between tactile, visual and olfactory stimuli, with the former exerting the strongest effect. Behavioural analysis also directed a study of the mechanisms whereby adult females exert an epigenetic influence over the phase-state of their developing offspring. Female locusts use their experience of the extent and recency of being crowded to predict the probability that their offspring will emerge into a high-density population, and alter the development of their embryos accordingly through a gregarizing agent added to the foam that surrounds the eggs at laying. There is also a less pronounced paternal influence on hatchling phase-state. An understanding of the time-course of behavioural phase change led to a study of the effect of the fine-scale distribution of resources in the environment on interactions between individual locusts, and hence on phase change. This, in turn, stimulated an exploration of the implications of individual behavioural phase change for population dynamics. Cellular automata models were derived that explore the relationships between population density, density of food resources and the distribution of resources in the environment. The results of the simulation showed how the extent of gregarization within a population increases with rising population size relative to food abundance and increasing concentration of food resources. Of particular interest was the emergence of critical zones across particular combinations of resource abundance, resource distribution and population size, where a solitarious population would rapidly gregarize. The model provided the basis for further laboratory and field experiments, which are described.     

Adult female desert locusts require contact chemicals and light for progeny gregarization

Crowding causes many organisms to express phenotypic plasticity in various traits. Phase polyphenism in desert locusts represents one extreme example in which a solitary form (solitarious phase) turns into a gregarious form (gregarious phase) in response to crowding. Conspicuous differences in body size and colour occur even in hatchlings. The phase‐specific differences in hatchling characteristics are caused by the tactile stimuli perceived by the antennae of their mother. However, the nature of the tactile stimuli and the mechanism by which the perceived stimuli are processed as a gregarizing signal remain unknown. To explore this problem, the antennae of solitarious adult females of the desert locust Schistocerca gregaria are touched with the bodies of conspecific locusts at different physiological stages and those of other species. The results suggest that a cuticular chemical factor at a specific developmental stage of conspecific locusts causes the solitarious females to produce large eggs that give rise to black hatchlings characteristic of gregarious forms (progeny gregarization), and that this or a similar compound occurs in other acridids, crickets and cockroaches but not in beetles. The involvement of a chemical substance is also supported by hexane extracts of cuticular surfaces of locusts that induce the same effects. Interestingly, crowding induces such gregarizing effects only when the female receives the appropriate stimulus in the presence of light. Solitarious female S. gregaria with their head capsule coated with phosphorescent paint exhibit progeny gregarization in response to crowding and light pulses in darkness, whereas those treated in the same way without light pulses fail to do so.     

Modelling locust foraging: How and why food affects group formation

Locusts are short horned grasshoppers that exhibit two behaviour types depending on their local population density. These are: solitarious, where they will actively avoid other locusts, and gregarious where they will seek them out. It is in this gregarious state that locusts can form massive and destructive flying swarms or plagues. However, these swarms are usually preceded by the aggregation of juvenile wingless locust nymphs. In this paper we attempt to understand how the distribution of food resources affect the group formation process. We do this by introducing a multi-population partial differential equation model that includes non-local locust interactions, local locust and food interactions, and gregarisation. Our results suggest that, food acts to increase the maximum density of locust groups, lowers the percentage of the population that needs to be gregarious for group formation, and decreases both the required density of locusts and time for group formation around an optimal food width. Finally, by looking at foraging efficiency within the numerical experiments we find that there exists a foraging advantage to being gregarious.     

喀斯特关键带植被时空变化及其驱动因素

中国南方喀斯特地区广泛面临着生态问题,植被的保护与恢复倍受关注,对这一区域植被覆盖的进行监测和预测是非常必要的。以MODIS-NDVI为数据源,分析2000—2016年间,研究区不同地质背景,多种土地覆被类型的NDVI时空变化特征及驱动因素。结果表明:(1)从2000—2016年间,研究区植被覆盖整体呈增长趋势;其中喀斯特区域增长情况略优于非喀斯特区域。植被覆盖在空间上呈现东高西低;其中林地的NDVI值最高,耕地次之,依次草地,居民用地,水域,未利用地最低;在林地和耕地中,非喀斯特区域的NDVI值比喀斯特高,其余的土地覆被类型中都比喀斯特区域低。(2)研究区植被覆盖改善的地区占60.19%,退化地区占17.06%;草地,耕地区改善明显,退化主要在水域和建设用地; Hurst指数显示在研究区持续性改善的NDVI大于持续性退化;相比非喀斯特区域,喀斯特区域改善及持续性改善情况更佳。(3)整体而言,海拔对NDVI的空间分布影响力最大,温度次之,依次为降雨,夜间灯光指数;相比而言,非喀斯特区域NDVI空间分布更易受地形因子影响;喀斯特区域NDVI空间分布更易受气候差异及人类活动影响。(4)研究区分别有49%,45%,61%的NDVI与气温,降雨,日照的相关系数通过a=0.05的显著性检验;相比非喀斯特而言,喀斯特区域植被生长更易受气候变化的影响。     

Are recession populations of the desert locust (Schistocerca gregaria) remnants of past swarms?

The desert locust (Schistocerca gregaria) undergoes crowding-induced phase transformation from solitary to gregarious, which involves changes in behaviour, colour, development, morphometry, fecundity and endocrine physiology. During recession, solitary locusts persist in the central, drier part of the species' range in small pocket populations that are prone to extinction. During the intermittent upsurges and the subsequent plagues, gregarious swarms attain huge population size and invade a vast area causing major damage to agriculture. A highly variable nuclear DNA marker, a noncoding 3' end fragment of an antennapedia-class homeobox gene, was screened in locust samples from Eritrea. Despite the homogenizing potential of plague swarms, the last of which was in 1986-89 and originated in this region, the population genetic structure of solitary phase locusts along the Red Sea coast of Eritrea revealed significant divergence. The pattern of divergence indicated that the invasion of the western and northern plains in the summer of 1995 may not, as reported then, have originated in eastern Chad or western Sudan. A number of interrelated hypotheses have been presented to explain the observed genetic heterogeneity between the sampled populations. We conclude, with caution due to the limited sample sizes, that: (i) geographical isolation between breeding sites during plagues and recession; (ii) the marked differences in the flight behaviour of plague swarms and recession populations; (iii) possible failure of gregarious locusts to solitarize and re-establish in recession areas; and (iv) the effect of repeated extinction and recolonization in the meta-population contribute to the maintenance of the genetic structure of recession populations. Potentially productive future research has been identified.     

Migrating locusts can detect polarized reflections to avoid flying over the sea

The desert locust Schistocerca gregaria is a well known migrating insect, travelling long distances in swarms containing millions of individuals. During November 2004, such a locust swarm reached the northern coast of the Gulf of Aqaba, coming from the Sinai desert towards the southeast. Upon reaching the coast, they avoided flying over the water, and instead flew north along the coast. Only after passing the tip of the gulf did they turn east again. Experiments with tethered locusts showed that they avoided flying over a light-reflecting mirror, and when given a choice of a non-polarizing reflecting surface and a surface that reflected linearly polarized light, they preferred to fly over the former. Our results suggest that locusts can detect the polarized reflections of bodies of water and avoid crossing them; at least when flying at low altitudes, they can therefore avoid flying over these dangerous areas.     

Can late summer Landsat data be used for locating Asian migratory locust,Locusta migratoria migratoria,oviposition sites in the Amudarya River delta,Uzbekistan?

Existing survey methods for assessing the Asian migratory locust, Locusta migratoria migratoria L. (Orthoptera: Acrididae), infestation risk in the Amudarya River delta, Uzbekistan, are largely constrained by economic resources and site accessibility. The surveys are restricted to a few easily accessible areas, which leads to a misinterpretation of the threat of locust infestation. This often results in indiscriminate blanket treatments of vast areas of wetlands with broad-spectrum insecticides, which may adversely impact non-target fauna and flora. In order to minimize the bias during surveys, one approach would be to allocate the sampling locations based on the distribution of the primary food and shelter plant of the locusts, the common reed, Phragmites australis (Cav.) Trin. ex Steud (Poaceae). In this study, we evaluated the utility of satellite-based remotely sensed data (Landsat TM) acquired in August 2006 to characterize reed distribution in the delta and identify potential locust oviposition sites. The overall accuracy of the Landsat data to map land cover classes in the delta was 84%. The Landsat TM data identified 90% of the reeds, but it was less useful in identifying areas where other vegetations (shrubs and grasses) were mixed with reeds. During the following summer field survey in June 2007, we identified 37 sites that were infested with early-instar locusts. The low migration capacity of young nymphs in dense reed vegetation allowed us to presume that these sites were used for oviposition in the previous summer. Twenty-eight (74%) of these 37 sites had reeds in the previous year. Results from these studies demonstrate that reed distribution maps derived from satellite data could be used for targeting locust egg-pod survey locations, in order to minimize sampling bias while predicting locust infestation risks for the following season.