Stage-based mortality of grassland grasshoppers (Acrididae) from wandering spider (Lycosidae) predation

Mortality rates in insects, including grasshoppers (Acrididae), are often stage- or size-specific. We estimated stage-specific mortality rates for three common grasshopper species from a Nebraska (USA) sandhills grassland (Ageneotettix deorum, Melanoplus sanguinipes and Phoetaliotes nebrascensis), and partitioned the impact due to wandering spider predation from remaining sources. Survivorship was estimated for multiple developmental stages (3rd instar through adult) under experimental conditions that either prevented or permitted predation from free-living, wandering spiders (primarily Schizocosa species). Total stage-specific mortality, including spider predation, examined over the period of single stages was greatest for the youngest stages (91% for 3rd instar, 73% for 4th instar, 63.5% for 5th instar and 30.4% for adults). For the developmental stages considered and averaged for all species, the contribution to total mortality from spider predation over the 10-d period (approximately the length of a developmental stage) ranged from 17% for 3rd instar nymphs to 23% for 4th and 5th instars, and an undetectable level for adults. While spiders may depress grasshopper numbers, contributions from spider predation to grasshopper population dynamics are uncertain.     

Persistence of <Emphasis Type="Italic">Paranosema</Emphasis> (<Emphasis Type="Italic">Nosema</Emphasis>) <Emphasis Type="Italic">locustae</Emphasis> (Microsporidia: Nosematidae) among grasshopper (Orthoptera: Acrididae) populations in the Inner Mongolia Rangeland,China

Paranosema (Nosema) locustae Canning has persisted in grasshopper populations in the Inner Mongolia Rangeland since spore treatment was applied in 1993. In the treatment year, a 58.32–71.43% reduction of grasshoppers was recorded and 59.26–80.36% of surviving grasshoppers of various species were infected. In the years following application, prevalence of P. locustae varied from 13.65% to 60.71%, demonstrating that P. locustae remains in grasshopper populations for many years. Infection prevalence was more than 60.00% for Dasyhippus barbipes (F.-W.), Bryodema luctuosum luctuosoma (Stoll) and Angaracris barabensis (Pall.) in 1993 but was very variable, ranging from 20.00% to 60.71% over the subsequent nine years. Disease prevalence was 43.72–60.71% in 1993 and 1994 for Oedaleus asiaticus B.-Bienko and Myrmeleotettix palpalis (Zub.) but gradually declined to approximately 20%, where it remained. Persistence of P. locustae in grasshopper populations may be important in keeping densities below economic thresholds for a number of years after application. Handling Editor: Helen Roy.     

New case of long-term persistence of Paranosema locustae (Microsporidia) in melanopline grasshoppers (Orthoptera: Acrididae: Melanoplinae) of Argentina

We report an additional case of long-term persistence of Paranosema locustae in grasshoppers of Argentina. The pathogen was introduced from North America on rangeland at Loncopué, Neuquén province. Microsporidia were not detected in pre-introduction samples whereas infected grasshoppers were found 11 years after introduction. Affected grasshoppers were the melanoplines Dichroplus elongatus, Dichroplus maculipennis, and Scotussa lemniscata, some of them with high spore loads. The case highlights the ability of P. locustae to recycle in local grasshopper communities by parasitizing susceptible species other than the natural hosts.     

Field observations of the effects of fenitrothion and Metarhizium anisopliae var. acridum on non-target ground dwelling arthropods in the Sahel

The effect of the chemical insecticide, fenitrothion, and a mycoinsecticide based on Metarhizium anisopliae var. acridum on the activity of non-target epigeal arthropod scavengers was investigated in areas of open savannah in southeast Niger Republic, West Africa. Both insecticides were applied as full cover sprays to unreplicated 800 ha plots to assess their season-long control of Sahelian grasshoppers. Compared with control plots, fenitrothion caused an immediate but temporary reduction in grasshopper numbers, whereas M. anisopliae var. acridum provided delayed but prolonged control. Scavenging rates of pyrethroid-killed grasshoppers placed along transects in unsprayed plots and those treated with fenitrothion and M. anisopliae var. acridum at various intervals after spraying were assessed. In the fenitrothion plot, an immediate reduction in scavenging activity occurred that was still apparent after 40 days at the plot center, although recovery at the plot edges was more rapid. By contrast scavenging rates remained high over equivalent areas in the M. anisopliae var. acridum and two untreated plots. Concurrent to the scavenging study, counts of grasshopper cadavers resulting from the spray treatments were conducted. These counts revealed that the density of grasshopper cadavers remained low throughout the M. anisopliae var. acridum plot and explained <1% of the reduction in live grasshoppers resulting from treatment, compared with >20% in the fenitrothion plot. This shortfall in grasshopper cadavers resulting from the spray treatment in the M. anisopliae var. acridum plot was unexpected because in a monitoring study, fungus-killed (unlike pyrethroid-killed) grasshoppers were unattractive to scavengers and readily persisted in this plot, and thus should have become apparent. Given we did not observe significant grasshopper dispersal, the scarcity of cadavers generated in the M. anisopliae var. acridum plot, together with unquantified visual observations, suggests that predation of infected but living grasshoppers was high. Our data provide circumstantial evidence that the different effects of chemical and biological grasshopper control on grasshopper natural enemies may influence the efficacy of large-scale treatments.     

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.     

Weather variation and trophic interaction strength: sorting the signal from the noise

Weather can have important consequences for the structure and function of ecological communities by substantially altering the nature and strength of species interactions. We examined the role of intra- and inter-annual weather variability on species interactions in a seasonal old-field community consisting of spider predators, grasshopper herbivores, and grass and herb plants. We experimentally varied the number of trophic levels for 2 consecutive years and tested for inter-annual variation in trophic abundances. Grasshopper emergence varied between years to the extent that the second growing season was 20% shorter than the first one. However, the damage grasshoppers inflicted on plants was greater in the second, shorter growing season. This inter-annual variation in plant abundance could be explained using the foraging-predation risk trade-off displayed by grasshoppers combined with their survival trajectory. Decreased grasshopper survival not only reduced the damage inflicted on plants, it weakened the strength of indirect effects of spiders on grass and herb plants. The most influential abiotic factor affecting grasshopper survival was precipitation. We found a negative association between grasshopper survival and the total yearly precipitation. A finer scale analysis, however, showed that different precipitation modalities, namely, number of rainy days and average precipitation per day, had opposing effects on grasshopper survival, which were inconsistent between years. Furthermore, our results suggest that small changes in these factors should result in changes of up to several orders of magnitude in the mortality rate of grasshoppers. We thus conclude that in this system the foraging-predation risk trade-off displayed by grasshoppers combined with their survival trajectory and relevant weather variability should be incorporated in analytical theory, whose goal is to predict community dynamics.     

The occurrence of the transposable elementpogo inDrosophila melanogaster

We examined the genomic occurrence of the transposable elementpogo in over 120 strains ofDrosophila melanogaster, from around the world and from different eras. All had multiple copies of a 2.1 kilobase (kb)pogo element, and multiple copies of several size classes between 1.0 and 1.8 kb. There were differences between strains in intensities or presences of deletion-derivative size classes, suggesting current or recent mobility in the species. We were unable to find anypogo-hybridization in eight other species in the genus, in three subgenera, or in the relatedScaptomyza pallida. Thepogo element may be a ‘middle-aged’ element in the genome ofD. melanogaster, having entered the species since its divergence from its sibling species, but long before theP andhobo elements.     

黑河上游天然草地蝗虫空间异质性与分布格局

蝗虫空间分布格局是物种长期适应自然的结果,是蝗虫与环境、蝗虫之间关系的反映,有助于理解蝗虫发生学的环境背景。采用地统计学方法,在黑河上游山区天然草地研究了微生境影响下的蝗虫空间格局与异质性。结果表明:该区域5种优势种蝗虫的半变异函数模型均为非线性模型,表现为聚集分布;不同种类蝗虫样点之间空间依赖的范围为2.29—24.59 m;在蝗虫种群总异质性中随机部分引起的空间异质性占15.77%—88.64%。受食性和栖息地选择的影响,蝗虫种群形成了片状和斑块状的分布格局;尺度依赖性使蝗虫分布格局趋于多元化和繁杂化的特点,不同蝗虫种群之间相互交错嵌插,同种蝗虫高低值呈非均匀扩散,整体上形成了均衡性与互补性的斑块镶嵌结构,这种分布格局反映了蝗虫对生境和气候变化的多元适应性结构。     

Parasitism of the grasshopper Melanoplus sanguinipes by a sarcophagid fly, Blaesoxipha atlanis: influence of solitary and gregarious development on host and parasitoid

Blaesoxipha atlanis (Aldrich) (Diptera: Sarcophagidae) is a common parasitoid of the grasshopper Melanoplus sanguinipes (F.) (Orthoptera: Acrididae) in western Canada. We tested the hypothesis that B. atlanis can develop as either a solitary or a gregarious parasitoid, and assessed the influence of parasitism on the growth and survival of infected grasshoppers. Males and females of M. sanguinipes were parasitized manually with one, two, or three first-instar larvae of B. atlanis in the laboratory. Parasitism was more deleterious to males than females of M. sanguinipes; females are larger than males. Host survival and longevity declined with the number of larvae per host in a sex-specific manner. In females, 39%, 24%, and 8% of hosts containing, respectively, one, two, and three sarcophagid larvae survived parasitism. Although 41% of single-parasitized males survived, all males containing more than one larva died. Variations in host quality as measured by dry mass explained much of the response to parasitism in male, but not female, hosts. Parasitoid larvae, apparently, did not cause significant physical damage to host organs and tissues but instead functioned as a metabolic sink. The greater metabolic activity associated with egg production could account for the relatively higher tolerance to parasitism of female, as opposed to male, grasshoppers. Developmental time, adult size, and percentage survival of B. atlanis declined with the intensity of parasitism, especially in parasitoids developing in male hosts. Females developing gregariously contained fewer ovarioles at eclosion than counterparts developing as solitary larvae. The mean body size of field-collected B. atlanis did not differ from that of laboratory-reared parasitoids developing singly in a host. Gregarious development is an alternative strategy to solitary development that may enable B. atlanis to maintain population numbers during periods of grasshopper scarcity.     

Cloned DNA probes distinguish endemic and exotic Entomophaga grylli fungal pathotype infections in grasshopper life stages

Entomophaga grylli is a fungal pathogen of grasshoppers and at least three pathotypes are recognized world-wide. Pathotypes 1 and 2 are endemic to North America while the Australian pathotype 3 had been released into two field sites in North Dakota between 1989 and 1991. Grasshoppers were collected over the summer at the field sites in 1992 and assessed for pathotype infection by cloned DNA probe analysis. The three most predominant grasshopper species that were infected ( Melanoplus sanguinipes, M. bivittatus and Camnula pellucida ) were assessed for pathotype infection with respect to their life stages (nymphal instars and adult males and females). Pathotype 1 predominantly infected grasshoppers in the subfamilies Oedipodinae and Gomphocerinae and pathotype 2 predominantly infected grasshoppers in the subfamily Melanoplinae. Early-instar M. sanguinipes and M. bivittatus had higher pathotype 2 infection frequencies, while late-instar and adult C. pellucida had higher pathotype 1 infection frequencies. Cross-infection by the pathotypes did occur in up to 3% of the individuals, on a per species basis, and primarily in later instar and adult grasshoppers. Pathotype 3 infections occurred in later instar and adults of the three grasshopper species. Infection of grasshoppers by E. grylli pathotypes is discussed with reference to the fungal life cycles.     

Natural Incidence of Metarhizium flavoviride Infection in Two Grasshopper Communities in Northern Benin

The infection of grasshoppers by naturally occurring entomopathogenic fungi was monitored at two sites in Malanville, northern Benin, Africa. Grasshoppers were collected during the monsoon seasons of 1993, 1994 and 1995. At the first site, 982 individuals of 38 grasshopper species were incubated. At the second site, 899 individuals of 33 grasshopper species were incubated. Two hyphomycete fungi (Deuteromycotina: Hyphomycetes), Metarhizium flavoviride Gams and Rozsypal and Sorosporella sp., were found infecting grasshoppers. The mean incidence of M. flavoviride infection was 0.3-1.7% at the first site and 1.2-3.2% at the second site for all grasshopper species. Sorosporella sp. was only collected from two grasshopper individuals in 1993. The majority of infected species had adult quiescence or continuously reproducing life cycles, and were either arboricolous or terricolous. Comparisons are made with a previous study of the two sites performed during 1992.     

Grasshopper assemblage response to a restored national park (Mountain Zebra National Park, South Africa)

Twelve grassland sites were sampled inside and outside the Mountain Zebra National Park (MZNP), South Africa to assess changes in grasshopper assemblages to grazing by indigenous mammals inside the park compared to grazing by domestic cattle outside. The MZNP has been restored from cattle-grazed farmland to indigenous mammal parkland for 62 years. The number of grasshopper species and families inside the park was not significantly different from outside the park, but the number of individuals inside the park was significantly higher. Multivariate statistics did not reveal any strong site groupings based on simple inside/outside comparisons, but there were clear groupings of sites based on vegetation characteristics and other environmental variables. The park boundary, therefore, does not significantly determine grasshopper assemblages, although intensity of grazing does. The indigenous mammals inside the park had the same effect on grasshoppers as the domestic cattle outside, and it was the level of defoliation and trampling that was important rather than the type of mammal. Intensive livestock grazing and trampling leads to bush encroachment and reduction in grass cover and/or disappearance of several grass species. In response to this pressure, grasshopper populations dropped, with localized extirpation of some species. Vegetation composition and structure, particularly grass height and percentage cover, had a significant effect on grasshopper assemblages. The MZNP is thus an area of localized, elevated grasshopper abundance in comparison with the surrounding farms, and presumably represents a situation prior to the current, intensive farming activities. Such elevated grasshopper abundances are important for maintaining soil quality and hence ecological integrity of this landscape which is poor in organics and nitrogen. The MZNP could be viewed as a centre where species with high mobility may seek refuge from anthropogenic pressures. The MZNP also serves as a reference illustrating the differences between restored-through-natural-succession and anthropogenically disturbed habitats, and compares desirable with undesirable ecosystem changes for herbivorous invertebrates such as grasshoppers.     

内蒙古锡林河流域不同生境中蝗虫种类组成的分析

本文对内蒙古锡林河流域不同生境中蝗虫种类发生的情况进行了分析。经过系统取样和大面积扫网调查,共采集到蝗虫21种,这些种类在各生境中出现的情况,与生境的复杂程度存在很大的关系。在复杂的生境中,蝗虫种类最丰富。根据锡林河流域不同生境中蝗虫种类发生的情况,经过比较分析,我们认为在该区域存在的沙带生境是典型草原的“蝗虫库”。“蝗虫库”的存在,对牧业生产构成一定的潜在威胁。经存在度分析表明,一些常见的种类,生态分布幅度宽,同时,它们又是当地的优势种害虫,为害大,是重点防治对象。     

Neurons projecting from the brain to the corpora allata in orthopteroid insects: anatomy and physiology

Retrograde and orthograde labeling of neurons projecting to the corpus allatum was performed in locust, grasshopper, cricket, and cockroach species in order to identify brain neurons that may be involved in the regulation of juvenile hormone production. In the acridid grasshopper Gomphocerus rufus L., and the locusts Locusta migratoria (R.&F.) and Schistocerca gregaria Forskal, the corpora allata are innervated by two morphologically distinguishable types of brain neurons. One group of 9–13 neurons (depending on species) with somata in the pars lateralis extend axons via the nervus corporis cardiaci 2 and nervus corporis allati 1 to the ipsilateral corpus allatum, whereas two cells in each pars lateralis have bilateral projections and innervate both glands. No direct connection between the pars intercerebralis and corpus allatum has been found. In contrast, neurons with paired axons innervating both glands are not present in Periplaneta americana (L.) and Gryllus bimaculatus de Geer. Instead, two cells in each pars lateralis project only to the gland contralateral to their somata. Electrophysiological experiments on acridid grasshoppers have confirmed the existence of a direct conduction pathway between the two glands via the paired axons of four cells that have been identified by neuroanatomy. These cells are not spontaneously active under experimental conditions. Ongoing discharges in the left and right nerves are unrelated, suggesting that the corpora allata receive independent neuronal inputs from the brain.     

Acid phosphatase activity in hemolymph of the migratory grasshopper,Melanoplus sanguinipes, duringBeauveria bassiana infection

The distribution and abundance of acid phosphatase (AP) in hemolymph (HL), plasma (PM) and hemocyte lysate supernatant (HLS) of the migratory grasshopper,Melanoplus sanguinipes infected withBeauveria bassiana (strain GK 2016) has been examined. AP activity was determined at intervals from 30 min to 60 h postinjection of 2 μl of 1×10⁸ conidia/ml per grasshopper. The enzyme was detected with the substrate β-glycerophosphate in sodium acetate acetic acid buffer form hemocytes (HC) and withp-nitrophenol phosphate sodium salt for HL, PM and HLS. In results of experiment 1 proportion of HC showing AP activity increased 1–2 h, then returned to normal after 4 h. However, inB. bassiana-injected grasshoppers, a second increase was noted 24 h later which was not seen in the Tween-80-injected insects. Uninjected controls showed no change with time in the proportion of HC with AP activity. Studies were also made of the distribution of AP activity in the HL, PM, and HLS. AP activity in HL appeared to vary with the sex of the grasshoppers. Females showed increase in AP activity in HL 18–24 h after injection withB. bassiana, whereas males only showed an increase 1 h after injection. Assay of HLS showed that the level of AP activity did not change significantly throughout the experiment. Changes in AP activity in PM, in bothB. bassiana — and Tween-80-injected insects (both sexes) paralleled those of the HL, indicating that the enzyme is released from the HC. The observations are discussed in terms of the possible role of AP in the immune response ofM. sanguinipes.     

Microbial and protozoan pathogens of grasshoppers and locusts as potential biocontrol agents

Worldwide, grasshoppers and locusts destroy several billion dollars (US) worth of crops every year. The known microbial and protozoan pathogens of grasshoppers and locusts are reviewed. They may be valuable control agents, particularly for some of the economically devastating locust species in Africa and Asia, and grasshopper species in North America and Asia. Aspects of the pathogenic process, if known, of fungi, protozoa, viruses and rickettsia towards acridids are discussed. Finally, production, formulation and application of these biocontrol agents are reviewed.     

Effects of large herbivore grazing on grasshopper behaviour and abundance in a meadow steppe

1. Adaptive phenotypic plasticity has been a major subject in evolutionary ecology, but how a species' behaviour may respond to certain environmental change is still not clear. In grasslands worldwide, large herbivores are increasingly used as a tool for grazing management, and occur to interact with grasshoppers that dominate grassland insect communities. Previous studies have been well-documented about grazing effects on diversity and abundance of grasshoppers. Yet, how grazing may alter grasshopper behaviour, and potential effects on their abundance remains elusive. 2. We conducted a field experiment by manipulating grazing using sheep, cattle, and their mix to examine the behavioural responses and abundance of the grasshoppers (Euchorthippus unicolor) to grazing in a Leymus chinensis-dominated grassland. 3. Results showed that the grasshoppers spent less time on feeding and resting on grasses, but more time on switching and resting on forbs under cattle grazing and mixed grazing with cattle and sheep. In contrast, the grasshoppers spent more time on feeding but less time on switching and resting on forbs under sheep grazing. The behavioural changes were also potentially linked to grasshopper abundance in the context of grazing management. 4. The responses of grasshopper behaviour and abundance to grazing may be largely triggered by altered vegetation and microclimates. Such behavioural flexibility of grasshoppers must be considered when large herbivores are recognised as a management tool for influencing grasshopper abundance, and grazer species should be paid more attention both individually and jointly for better grassland conservation.