Competitive interactions between parasitoid larvae and the evolution of gregarious development

We report experiments using two closely related species of alysiine braconids directed at understanding how gregarious development evolved in one subfamily of parasitoid wasps. Theoretical models predict that once siblicide between parasitoid wasps has evolved, it can only be lost under stringent conditions, making the transition from solitary to gregarious development exiguous. Phylogenetic studies indicate, however, that gregariousness has independently arisen on numerous occasions. New theoretical models have demonstrated that if gregarious development involves reductions in larval mobility, rather than a lack of fighting ability (as in the older models), the evolution of gregariousness is much more likely. We tested the predictions of the older tolerance models (gregariousness based on non-fighting larval phenotypes) and the reduced mobility models (gregariousness based on non-searching larval phenotypes) by observing larval movement and the outcome of interspecific competition between Aphaereta genevensis (solitary) and A. pallipes (gregarious) under multiparasitism. Differences in larval mobility matched the prediction of the reduced mobility model of gregarious development, with the solitary A. genevensis having larvae that are much more mobile. The proportion of hosts producing the solitary species significantly declined after subsequent exposure to females of the gregarious species. This contradicts the prediction of the older models (fighting vs non-fighting phenotypes), under which any competitive interactions between solitary and gregarious larvae will result in a highly asymmetrical outcome, as the solitary species should be competitively superior. The observed outcome of interspecific competition offers evidence, with respect to this subfamily, in favour of the new models (searching vs non-searching phenotypes).     

Adipokinetic Hormone and Flight Fuel Related Characteristics of Density-Dependent Locust Phase Polymorphism: A Review

Recent findings on differences between the gregarious and solitary phases of locusts are reviewed in relation to flight fuel utilization, adipokinetic responses, and adipokinetic hormones. Laboratory results obtained with Locusta migratoria migratorioides show that the amount of lipid reserves, resting levels of haemolymph lipids, and hyperlipaemic responses to flight and to injection of corpus cardiacum extract or of synthetic adipokinetic hormones, are higher in crowded than in isolated locusts. No major phase-dependent differences seem to exist in flight-related carbohydrate metabolism. The adipokinetic hormone content of the corpora cardiaca is higher in younger isolated locusts than in crowded ones. Adipokinetic hormone precursor-related peptide content of the corpora cardiaca is also higher in isolated than in crowded locusts. Crowded locusts have higher lipid reserves and higher hyperlipaemic responses to flight than isolated locusts also in Schistocerca gregaria and, following injection of synthetic adipokinetic hormone, the formation of low density lipophorin is higher in crowded than in isolated locusts of this species. The laboratory results obtained with isolated and crowded locusts are extrapolated to understand the ecophysiology of the migrations of solitary and gregarious field populations of L.m. migratorioides according to available information on the differences in the migration of the two phases. It is inferred that in this species solitary locusts have a rather coarse adipokinetic strategy focused on a single prereproductive long-distance migratory flight, whereas gregarious locusts possess a fine adipokinetic balance for reiterative, sometimes unpredictably long-distance, migrations in the prereproductive, as well as reproductive, periods. The differences between the adipokinetic strategies of solitary and gregarious S. gregaria seem to be less dramatic, nevertheless, they indicate a better adaptation of the gregarious phase to prolonged flights.     

Effects of voltinism, interspecific competition and parasitism on the population dynamics of the hemlock scales, Fiorinia externa and Tsugaspidiotus tsugae (Homoptera: Diaspididae)

Abstract.

• 1 Life tables were constructed for solitary and coexisting populations of univoltine Fiorinia externa Ferris and bivoltine Tsugaspidiotus tsugae (Marlatt) (Homoptera: Diaspididae), two exotic scale pests of eastern hemlock, Tsuga canadensis Carriere, in the northeastern United States.
• 2 Solitary and coexisting populations of F. externa had similar survivorship and population growth rates resulting in an annual increase in density of 7–16%. Survivorship of solitary and coexisting populations of T. tsugae also was similar, but growth rates differed substantially. While solitary populations increased their density by 68% annually, populations coexisting with F. extema were reduced 74% each year.
• 3 The annual reduction in T. tsugae density where it coexists with F. externa was due in part to interspecific competition which resulted in higher mortality to nymphs from dispersal and starvation and in lower fecundity of adult females relative to solitary populations. A host shift by the parasitoid, Aspidiotiphagus citrinus (Crawford) (Hymenoptera: Aphenlinidae) from F. externa to T. tsugae in autumn also accounted for 71% of the annual decrease in T. tsugae numbers. Therefore, F. externa adversely affects the growth of T. tsugae populations not only because of its superior competitive ability but also because adult para-sitoids which emerge from it subsequently attack nymphs of T. tsugae in autumn.
• 4 Parasitism and starvation of nymphs resulting from competition were the key mortality factors in the population dynamics of these exotic hemlock scales.

     

Weed seed granivory by carabid beetles and crickets for biological control of weeds in commercial lowbush blueberry fields

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Variations in ecdysteroid levels in 5th instar larvae of Schistocerca gregaria in gregarious and solitary phases

Moulting hormone levels in whole bodies of 5th instar S. gregaria were determined throughout the 5th larval instar using gas chromatography with electron capture detection of the derivatized hormone. No differences were found between hormone levels in insects in the gregarious and solitary phase. No evidence for the existence of polar ecdysteroid conjugates used for storage or transport of these hormones was found. Approximately one tenth of the 20-hydroxyecdysone observed in the gregarious insects was detected in the faeces, almost equally divided between free hormone and polar conjugates of it.     

Variation in hydraulic architecture and gas-exchange in two desert sub-shrubs,<Emphasis Type="Italic"> Hymenoclea salsola</Emphasis> (T. &; G.) and<Emphasis Type="Italic"> Ambrosia dumosa</Emphasis> (Payne)

Adjustment of hydraulic architecture in response to environmental conditions was studied in two warm-desert sub-shrubs, Hymenoclea salsola and Ambrosia dumosa, both at the level of genetic adaptation along a climatic gradient and plastic response to immediate growth conditions. Individuals of both species originating from southern populations developed higher leaf-specific hydraulic conductance in the common greenhouse than individuals from northern populations. Hydraulic conductance was higher in plants grown at high temperature, but did not vary as a function of growth relative humidity. Hydraulic conductance was not correlated within species with individual variation in vessel diameter, cavitation vulnerability, or root:shoot ratio, but was strongly, negatively correlated with the fraction of total plant biomass allocated to leaves. For both species, stomatal conductance (g _s) at high leaf-to-air vapor pressure difference (ν) was tightly correlated with variability in hydraulic conductance, as was the sensitivity of stomatal closure to increasing ν. Experimentally increasing shoot water potential by soil pressurization, under conditions where high ν had already caused stomatal closure, led to substantial stomatal reopening in both species, but recovery was significantly higher in H. salsola. Hydraulic conductance was higher in H. salsola than A. dumosa. H.salsola also differed from A. dumosa by being a representative of a highly specialised group of desert shrubs which use the twigs as a major photosynthetic organ. The southern population of H. salsola produced far fewer leaves and relied much more heavily on twig photosynthesis than the northern population. At the whole-plant level, increased reliance on twig photosynthesis was associated with higher leaf-specific hydraulic conductance, but equivalent whole-plant photosynthesis on either a dry weight (μmol CO₂ g^–1) or nitrogen basis (μmol CO₂ g^–1)). This suggests that twig photosynthesis might be one way of increasing hydraulic conductance per unit photosynthetic canopy by increasing allocation to an organ which simultaneously performs photosynthetic, support, and transport functions. Received: 13 December 1999 / Accepted: 31 March 2000     

A Comparison of Behavioral Interactions in Solitary and Presocial Zetoborinae Cockroaches (Blattaria,Blaberidae)

The social behavior of five species of Zetoborinae cockroaches is compared with respect to inter-individual interactions of nymphs in the laboratory. These species belong to the same Neotropical subfamily and were characterized as gregarious (Lanxoblatta emarginata, Parasphaeria boleiriana, Phortioeca nimbata, Schultesia lampyridiformis) and solitary (Thanatophyllum akinetum) by previous field studies. Our results show that gregarious species accept closer contacts than does the solitary one. The solitary species did not display especially short, infrequent or less diverse behavioral sequences when forced to remain aggregated, but its interactions are characterized by fewer acts promoting contact and more dominance-like acts. The solitary species symmetrically interacts with conspecifics and does not show specific dispersal-promoting behaviors. This suggests that the solitary behavior observed in the field for species of Zetoborinae mainly results from a passive spacing tendency and a lack of attraction for conspecifics. One of the gregarious species, P. boleiriana, was previously described as subsocial with nymphs remaining with the female in a wood chamber after brood birth. This species does not show a peculiar behavioral repertoire but its interactions are characterized by more dominance-like behaviors than are those of the non-subsocial gregarious species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temperature can limit the invasion range of the ambrosia beetle Euwallacea nr. fornicatus

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Do pools impede drift dispersal by stream insects?

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Density-related variation in vertical transmission of a virus in the African armyworm

Larvae of the African armyworm, Spodoptera exempta, are darker and more resistant to baculovirus infection when reared in groups (gregarious form) compared to being reared singly (solitary form). Lepidoptera that survive virus challenge as larvae could potentially retain a sublethal virus infection which is then transmitted vertically to the next generation. Here we examine whether gregarious and solitary forms of the armyworm differ in the costs of surviving virus infection and in their capacity to transmit an active baculovirus infection to their offspring. Pupae of larvae reared gregariously that survived virus challenge weighed significantly less than uninfected individuals, but this was not so for those reared solitarily. This did not, however, translate into differences in fecundity, at least under laboratory conditions. As found in previous studies, pre-oviposition period was shorter for solitary than gregarious insects, and it was also shorter for females that had been challenged with virus as larvae. Both the prevalence of egg batches containing larvae that died from nucleopolyhedrovirus (NPV) infection and the proportion of infected larvae within each egg batch were significantly increased (approximately doubled) when parental moths were previously challenged with the virus during their larval state. This demonstrates that horizontal transmission in one generation can elevate vertical transmission to the next generation. Moreover, prevalence of overt infection in the offspring generation was two to three times greater when parental moths were reared solitarily as larvae than when reared gregariously. Disease prevalence and proportional infection were both independent of the sex of the infected parent and whether or not the egg batch was surface-sterilized to remove potential contaminants. This suggests that the eggs are infected internally (transovarial) rather than externally (transovum). These results help to shed light on the observed temporal pattern of virus epizootics in eastern Africa.     

Natural enemies of natural enemies: the potential top‐down impact of predators on entomopathogenic nematode populations

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Foraging advantages of mixed-species association between solitary and colonial orb-weaving spiders

This study investigated association between solitary orb-weaving spiders and a colonial orb-weaving spider, Metepeira incrassata (Araneae: Araneidae). Spiders were sampled along transects and an index of species association showed that two of the species were associated more frequently than expected based on a null hipothesis of random co-occurrence. The potential advantages of mixed-species association were investigated by comparing prey-capture success of one of these associates, Nephila clavipes (Araneae: Tetragnathidae), when it occurs alone, in single-species groups, and when associated with M. incrassata colonies. Field observations of prey-capture success by all three of these categories of N. clavipes revealed that individuals in M. incrassata colonies captured significantly more prey than solitaries or individuals in single-species groups. The increase in prey capture by N. clavipes in M. incrassata colonies may result from utilization of a foraging niche which intercepts a diffirent spectrum of prey than that available to single-species groups or solitaries. Related to this enhanced prey consumption is greater fecundity of spiders in association with M. incrassata as compared to solitaries or individuals in single-species groups.     

Small-scale vegetation patterns in the parental environment influence the phase state of hatchlings of the desert locust

Desert locusts (Schistocerca gregaria Forskål (Orthoptera: Acrididae)) change phase in response to population density. Solitarious insects avoid one another; when crowded, they shift to the gregarious phase and aggregate. Laboratory experiments and individual‐based modelling have shown that small‐scale resource distribution can affect locust phase state via an influence on crowding. Laboratory work has also shown that parental phase state is transmitted to offspring via maternal inheritance. These effects had not been investigated in the field previously. We maintained small populations of adult desert locusts in semi‐field enclosures with different distribution patterns of a single plant species (Hyoscyamus muticus L. (Solanaceae)). The offspring of locusts exposed to more clumped patterns of vegetation exhibited more gregarious behaviour when tested in a behavioural phase assay than did progeny from parents left in enclosures with more scattered vegetation. These effects on nymphal behaviour appeared to be mediated by influences of resource distribution on adult phase state. Phase state in small semi‐field populations was influenced by small‐scale vegetation distribution. Phase differences engendered by environmental structure were maintained in time and transmitted to progeny.     

Inter- and intrapopulational variation in time to metamorphosis in a freshwater copepod

• 1 Adjacent populations of the copepod Cyclops scutifer Sars living in lakes in southern Norway exhibit remarkably different life cycles. A series of laboratory-common environment experiments were used to partition variance in one life cycle trait—time to metamorphosis—among and within five populations of C. scutifer, to examine the extent to which variation in this trait has a generic basis and to test whether populations are polymorphic for development rates. The experimental populations exhibit a variety of life cycles in the field and occupy environments that represent a broad range of conditions.
• 2 Populations with different life cycles in the field continued to express differences in time to metamorphosis in the laboratory, indicating a genetic contribution to life cycle variation. Differences in developmental rates in the laboratory were smaller, however, than differences observed in the field, suggesting that environmental conditions also contribute to observed life cycle variation.
• 3 In addition to interpopulation variation, each population maintained substantial intrapopulation variation in time to metamorphosis; differences between individuals from the same population were often as large as or larger than differences between populations.
• 4 Individual females in most populations produced highly variable offspring, and there often was little difference in time to metamorphosis among families within a population.
• 5 Cyclops scutifer exhibits a hierarchy of variation in time to metamorphosis, with a major portion of this variation expressed among siblings. Intrapopulation life history variation important to natural selection may be maintained by different processes among the major groups of freshwater zooplankton.

     

群居型和散居型东亚飞蝗雌成虫飞行肌的超微结构

应用电子显微镜对群居型和散居型东亚飞蝗Locusta migratoria manilensis（Meyen）雌成虫背纵肌进行了比较观察。结果表明：群居型和散居型成虫背纵肌具有类似的亚细胞结构,飞行肌的肌原纤维具有1∶3粗细丝比例,每根粗丝由6根细丝环绕排列成六角形结构。飞行肌的发育和线粒体的形成均是渐进的过程,在不同日龄成虫间存在差异。肌节长度为2.1～3.4 μm;7和10日龄时群居型成虫肌节长度小于散居型;7日龄群居型肌原纤维直径显著大于散居型。背纵肌内线粒体含量约占肌纤维的20%～43%,两型飞蝗之间存在着显著的差异,7日龄时群居型线粒体占肌原纤维的比例高达42.96%,而散居型的只有22.45%;10日龄群居型线粒体含量为41.32%,散居型线粒体29.98%。上述差异可能是东亚飞蝗群居型成虫飞行能力显著强于散居型成虫的重要原因之一。     

Effects of colony size on larval performance in a processionary moth

1. Some lepidopteran species have larvae that live gregariously, especially in early instars. Colony‐living species may benefit from improved protection from predators, thermoregulation, and feeding facilitation, for example. 2. While many studies have compared solitary and gregarious life styles, few data exist as to the relationship between size of the larval colony and larval performance in gregarious species. The present study was aimed at understanding the importance of colony size for growth and survival of the northern pine processionary moth (Thaumetopoea pinivora) larvae. 3. Field studies, comparing three different sizes of colonies of T. pinivora larvae, showed that individuals in larger colonies had a higher survival rate compared with those living in smaller colonies and also a faster growth rate. 4. The higher survival rate of large colonies was attributed to improved protection from predacious arthropods. 5. In early spring, the young larvae bask in the sun to increase their body temperature. In field experiments the thermal gain was higher in large colonies, and individuals in such colonies also grew faster. As growth rate was not affected by colony size when the ability to bask was experimentally removed in a laboratory experiment, the higher growth rate of the larger colonies was probably due to improved thermoregulation rather than feeding facilitation. 6. The size of larval colonies of gregarious insects depends on natural mortality events as well as on female oviposition strategy. Our results show that decreasing colony size can lead to a reduction in growth rate and survival. It is therefore important to understand whether or not small colonies will benefit equally from the gregarious behaviour.     

Facultative gregarious development in a solitary koinobiont parasitoid,Ephedrus californicus: implications of larval ecology and host constraints

In solitary parasitoids, the mandibulate first instars behave aggressively towards potential competitors so that generally only one larva survives per host. A ‘failure of competition’ may result in facultative gregarious development, however. We used Ephedrus californicus Baker (Hymenoptera: Braconidae: Aphidiinae), a solitary koinobiont parasitoid of aphids, to test two hypotheses in the laboratory that could explain facultative gregarious development. Gregarious development increased with the intensity of parasitism, with two (rarely three) parasitoids successfully developing in a single aphid. In heavily superparasitized hosts, interference between surviving larvae often caused abnormal pupation behaviour and inability to emerge from the mummy. The hypothesis that the survival of more than one larva per host is dependent on differences in larval age was not supported. The total body size in terms of dry mass of two males or two females developing together in the same host was higher than that of same‐sex counterparts developing singly. Females were larger than males with which they shared a host. Hypotheses about the evolutionary transition from a solitary to a gregarious lifestyle in parasitoid Hymenoptera have focused on lethal fighting between first instars but have ignored other constraints including immature mortality during later development and limiting host resources. Especially in species that pupate inside the dead host, specific requirements for pupation and emergence may determine whether one or several offspring per host can develop to adult.     

Gregariousness and repellent defences in the survival of phytophagous insects

Group living has both costs and benefits for plant‐feeding insects, but defence against predators is the most widely acknowledged benefit. Gregarious folivores typically have warning coloration and elaborate anti‐predator defences. Do these defences protect these species from predation? To see if protection from predators generally results from gregariousness, I compared the shapes of published survivorship curves of externally feeding, gregarious and solitary Lepidoptera and Symphyta. Gregarious species are less likely than solitary species to die in the larval stages. However, solitary species that have anti‐predator defences do not have higher larval survival compared to gregarious species. This result, along with evidence from experimental manipulations of group size, suggests that repellent defences per se do not increase survival of gregarious larvae. Group behaviour is undoubtedly important in affecting the higher larval survival of gregarious species, but we currently cannot determine whether predator learning, dilution of risk, or rapid development contribute most to increasing survival.     

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.