Differences in competitive ability for the occupancy of shelters in triatomines

Triatomines (Hemiptera: Reduviidae: Triatominae) are nocturnal blood‐sucking insects. During daylight hours they remain in an akinetic state inside their shelters, whereas at dusk they become active and move outside. When they are outside their shelters during the photophase, triatomines are vulnerable to diurnal predators and the period just before dawn is critical to their survival. This work analyses the existence of competitive interactions involved in the occupancy of shelters by triatomines. Behavioural assays were performed in which nymphs of different stages, nutritional status or species were released in an experimental arena containing a space‐limited artificial shelter. The proportions of individuals occupying the shelter during the photophase were quantified to estimate the competitive abilities of each stage and species. Intraspecific comparisons showed higher levels of shelter occupancy for fourth over fifth instars and fed over unfed nymphs of Triatoma infestans. Interspecific comparisons showed higher rates of shelter occupancy for Triatoma sordida in comparison with T. infestans, and for T. infestans over Rhodnius prolixus. Arrival order was also relevant to determining shelter occupancy levels: early arrival was advantageous in comparison with later arrival. The study of intra‐ and interspecific competitive interactions for shelter occupancy provides relevant information about colonization and recolonization processes in the natural environments of triatomines.     

Dislodgement effect of natural semiochemicals released by disturbed triatomines: a possible alternative monitoring tool

The quick detection of domestic and peridomestic triatomines in their environments becomes difficult without the use of dislodgement substances that flush them out from their shelters. At present, tetramethrin 0.2% is being widely used in control programs. Although it is an efficient dislodging agent, its toxicity might affect the health of captured triatomines, of other insects and, to a lesser extent, of other animals, including humans. Here, we tested if semiochemicals released by disturbed adults of Triatoma infestans and/or Rhodnius prolixus can make larvae of the same species exit from their refuges. In a walking olfactometer we found that: 1) larvae of T. infestans were repelled by the odors released by disturbed adults of their own species and of R. prolixus, 2) larvae of R. prolixus did not change their behavior in the presence of odors released by adults of both species, and 3) activity levels were not modulated by these odors in any of both species. Besides, in pseudo‐natural conditions we found an increased flushing‐out activity of larvae of T. infestans when their shelters were sprayed with isobutyric acid or 3‐pentanol, and of larvae of R. prolixus when sprayed with 3‐methyl1butanol. We succeeded in this work to dislodge larvae of triatomines from artificial shelters using natural volatile compounds, allowing the capture of live bugs for further investigations (e.g., xenodiagnosis or genetic studies) and favoring ecological aspects (e.g., minimizing environmental insecticide‐contamination and non‐targeted mortality).     

Social aggregation of the marine isopod Cirolana harfordi does not rely on the availability of light‐reducing shelters

Social aggregation under shelters can afford benefits to animals such as protection from predators. Many isopods and insects are negatively phototactic and this may help them gravitate towards shelter. Previous studies show that, when placed in an arena with two red shelters, specimens of the marine isopod Cirolana harfordi and the terrestrial isopod Porcelio scaber pick one of the two shelters at random and aggregate under it, demonstrating social aggregation under light‐reducing shelters. In the present study, an arena with two clear shelters was used to determine whether group sizes of 4, 8, 12 and 16 C. harfordi specimens display social aggregation when the shelters do not accommodate negative phototaxis. In all group sizes, C. harfordi specimens picked one of the shelters at random and significantly more animals aggregated under this shelter compared with the other. Cirolana harfordi also displayed aggregation in an arena with no shelters. Accordingly, C. harfordi specimens do not require shelters that reduce light to display social aggregation. The ability to locate suitable shelter under which there is no substantial reduction in light could benefit the animal in a natural environment comprising heavily shaded areas, as well as at night.     

Gregarious aggregative behavior in the marine isopod Cirolana harfordi

Gregarious huddling behavior where animals seek the close company of conspecifics is thought to have facilitated the evolutionary transition from sea to land in some species of isopods, the theory being that closely apposed bodies decrease surface area and reduce desiccation. In this study, specimens of the marine isopod Cirolana harfordi were investigated to determine whether they displayed gregarious aggregative behavior. Animals were placed in a 20‐cm‐diameter acrylic cylinder arena that contained two 5‐cm‐diameter circular acrylic shelters that were tinted red (to reduce light transmittance), one at either end of the arena. Specimens of C. harfordi chose one of the two identical shelters at random, and the majority of animals congregated under this winning shelter. When an arena with one shelter tinted red and one clear shelter was used, the majority of animals congregated under the red shelter. These data indicate that the aggregative behavior displayed by this marine isopod species is driven by gregarious, conspecific attraction as well as reactions to environmental heterogeneity and negative phototaxis. Gregarious behavior in this marine isopod may afford the species advantages other than avoiding desiccation.     

Meiotic differences between three triatomine species (Hemiptera,Reduviidae)

We have found the following differences in the male meiosis among three triatomine species: (1) The three largest autosomal bivalents ofTriatoma infestans are heterochromatic.Rhodnius prolixus has two autosomal bivalents with heterochromatic blocks.Triatoma rubrovaria does not show any heteropycnotic autosomes. (2) Sex chromosomes inT. infestans form a chromocenter. At early prophase terminal associations are seen between sex chromosomes inT. rubrovaria, and they maintain a close association until diakinesis. An intimate association between the X and Y chromosomes is observed during early prophase inR. prolixus, but a distant association is maintained by the sex chromosomes at diffuse and diplotene stages in this species. (3) Polyploid nuclei of the nutritive cells are quite distinct. Numerous chromocenters of different shapes and sized are seen in those ofT. infestans. InT. rubrovaria one chromocenter having two positively heteropycnotic elements is observed surrounded by homogeneous chromatin. Only one compact chromocenter is found amongst unevenly distributed chromatin, inR. prolixus.     

Aggregation behavior of Harmonia axyridis under non‐wintering conditions

The invasive multicolored Asian ladybeetle, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), aggregates inside dwellings during winter to avoid cold weather. This adaptive behavior disturbs homeowners, because of the large numbers of individuals that aggregate, which induces allergic reactions. The migratory flight patterns of this species have been well documented, with individuals preferentially moving toward prominent and high color contrast elements. However, the factors involved in the selection of aggregation sites by this species have yet to be elucidated. Here, we evaluated the influence of (i) the density of individuals and (ii) the type of available shelters on decisions by H. axyridis to settle and aggregate under shelters. A dual choice bioassay conducted in the laboratory demonstrated the presence of mutual attraction to conspecifics. We also found that individuals preferentially settled under red covered shelters compared to transparent shelters, and that the type of shelter outweighed the effect of social interactions among conspecifics. Moreover, this experiment was performed under non‐wintering conditions, providing the first evidence that aggregative behavior in this species can also occur under those specific conditions.     

Cues For Shelter Use in a Phytophagous Insect

Many insects spend a large proportion of their life inactive, often hiding in shelters. The presence of shelters may, therefore, influence where insects feed. This study examines stimuli affecting the use of shelters by adults of the pine weevil, Hylobius abietis (L.) (Coleoptera, Curculionidae). This species is an economically important forest pest in Europe since the adults feed on the stem bark of newly planted conifer seedlings. When there are hiding or burrowing places present in close proximity to a seedling, pine weevils may hide there and repeatedly return to feed on the same seedling. Experiments were conducted in a laboratory arena with above-ground or below-ground shelters and in the presence or absence of wind. Pine weevils were highly attracted to shelters both above and below ground. Weevils in shelters were often observed assuming a characteristic “resting” posture. Experiments with opaque and transparent shelters showed that visual stimuli are used for orientation towards shelters and also increase the probability of an individual remaining in a shelter. The presence of wind increased the weevils’ propensity to use shelters both above and below ground. The present study indicates that shelters have a major influence on the behavior of the pine weevil and possible implications of the results are discussed.     

Spatio‐temporal analysis of the role of faecal depositions in aggregation behaviour of the triatomine Rhodnius prolixus

Several triatomine species aggregate over conspecific faeces. In some domestic species, this behaviour might help individuals to find their refuges because faeces are usually deposited around such places. In the present study, the defaecation behaviour of the triatomine bug Rhodnius prolixus Stål (Heteroptera: Reduviidae) is described in detail, including the temporal pattern of aggregation of nymphs with different nutritional status to faeces of different post‐deposition times. First, using an experimental arena, the spatial distribution of faecal deposition spots released by nymphs, in relation to the position of a refuge, is recorded. The results show that a high density of faeces is deposited inside the refuge, and few and distant spots, in relation to the position of the refuge, are dropped outside. Second, whether the aggregation behaviour of nymphs is altered by the time elapsed from deposition of the faeces, and by the nutritional status of the insects, is determined. The post‐deposition time of the faeces strongly modulates the aggregation response of the insects. Differences in aggregation between fed and unfed insects are only marginally significant; however, different temporal windows of aggregation according to the post‐deposition times of the faeces are demonstrated between bugs of different nutritional status. The results are discussed with respect to the relevance of the aggregation signal in relation to the characteristics of the natural refuges used by R. prolixus.     

Insecticide resistance in Brazilian Triatoma infestans and Venezuelan Rhodnius prolixus

Resistance levels to insecticides used in control of Chagas Disease vectors were assessed in two species of bugs (Hemiptera: Reduviidae): Triatoma infestans (Klug) from Brazil and Rhodnius prolixus Stål from Venezuela. The resistance ratios, compared to susceptible laboratory strains, were determined by topical application bioassays. The T. infestans PA strain exhibited resistance ratios of 7× to deltamethrin, 3.6× to β‐cyfluthrin and 3.3× to cypermethrin, but was susceptible to β‐cypermethrin and lambda‐cyhalothrin. Rhodnius prolixus CA strain showed resistance to all the pyrethroids evaluated, the resistance ratios ranging between 4.5× to lambda‐cyhalothrin and 12.4× to cypermethrin. Deltamethrin resistance in both strains was decreased by piperonyl butoxide, suggesting oxidative metabolism as cause of resistance.     

Species composition and life histories of shelter-building caterpillars on Salix miyabeana

We studied the species composition and life history patterns of shelter‐building microlepidoptera on the willow Salix miyabeana in Hokkaido, northern Japan. We identified 23 microlepidopteran species across seven families that constructed leaf shelters. Species in Tortricidae and Pyralidae comprised approximately 90% of the total number of sampled shelter‐building microlepidoptera that reached adult eclosion in the laboratory. Seasonal changes in the density of leaf shelters showed two peaks: early June and mid‐August. In June, caterpillars of Gypsonoma bifasciata, Gypsonoma ephoropa, Acleris issikii and Saliciphage acharis were the principal shelter builders, while in August shelters were constructed primarily by caterpillars of Nephopterix adelphella, A. issikii and S. acharis. Approximately 90% of leaf shelters were constructed on the top portions of shoots, suggesting that most shelter‐building caterpillars prefer to build leaf shelters here.     

Aggregation in woodlice: social interaction and density effects

Terrestrial isopods are known to be sensitive to humidity, brightness or temperature. Until now, aggregation was assumed to depend on these sensitivities as a result of individual preferences. In this paper, we show that the social component is also important in the isopod aggregation phenomenon. In experimental arenas with two identical shelters up to nearly 90% of woodlice aggregated under shelters. This aggregation was quick as in 10 minutes most of the animals aggregated, irrespective of their density. Nonetheless, 10-15% of the animals walked around the arena, rarely forming very small and short-lasting aggregates outside shelters. Woodlice aggregated preferably under one of the shelters in 77% of experiments. Indeed, almost 80% of the animals out of 40, 60 or 80 animals in the arena aggregated under one shelter. In arenas with 100 individuals the aggregations were proportionally smaller (70%). Our results revealed that 70 animals was a maximum number of woodlice in an aggregate. We concluded that the location of aggregates is strongly governed by individual preferences but the dynamics of aggregation and collective choice are controlled by social interaction between congeners. The tested densities of the animals in the arena did not impact the aggregation patterns.     

Evaluation of a CO2‐free commercial mosquito attractant to capture triatomines in the laboratory

Efforts have been made to develop vertebrate odor‐based attractants to lure hematophagous triatomines into traps. However, more work is needed to reach a practical, cheap, and efficient odor lure. We carried out attraction and capture tests in a dual‐choice olfactometer and a pitfall trap. Here we report that a three‐component, CO₂‐free, synthetic blend of vertebrate odor (consisting of ammonia, L(+) lactic acid and hexanoic acid, and known as Sweetscent®) significantly induces 3^rd‐instar Rhodnius prolixus and Triatoma infestans nymphs to fall into the test capture‐tube of the olfactometer. Blend constituents presented singly or in two‐component blends did not evoke a response and, therefore, we propose that the insects respond specifically to the three‐component blend in a synergistic way. When tested in a pitfall trap in an experimental arena, this blend induced capture in 37.5% of the lured traps, whereas 9% of the nymphs tested were captured in a single night. No insects were captured in control traps. Our work represents a proof‐of‐concept regarding capture of triatomines using host odor‐based, CO₂‐free synthetic mixtures as lures for pitfall traps. CO₂‐free lures are more practical for field work than natural or CO₂‐containing synthetic blends.     

An enemy of your enemy is your friend: Impact of predators on aggregation behavior of gammarids

Predation pressure may affect many aspects of prey behavior, including forming groups and changes in social interactions. We studied the aggregation behavior of competing gammarids Dikerogammarus villosus and Pontogammarus robustoides (Amphipoda, Crustacea) to check whether they modify their preferences for conspecifics or heterospecifics in response to predator (the racer goby Babka gymnotrachelus) kairomones in the presence or absence of stone shelters (alternative protection source). Both species exhibited preferences toward shelters occupied by conspecifics over empty shelters and conspecifics apart from shelters, suggesting that their aggregation depends not only on habitat heterogeneity, but also on their social interactions. Moreover, gammarids in the presence of shelters (safer conditions) preferred conspecifics over heterospecifics, but predator kairomones made them form aggregations irrespective of species. In the predator presence, P. robustoides increased its aggregation level only in the sheltered conditions, whereas D. villosus exhibited this response only in the absence of shelters, suggesting that this behavior can protect it against predators. Therefore, we tested the antipredator effectiveness of D. villosus aggregations by exposing them to fish predation. Gobies foraged most effectively on immobile single gammarids compared to moving and aggregated individuals. Fish also avoided aggregated prey, confirming the protective character of aggregations. We have demonstrated that the predator presence increases aggregation level of prey gammarids and affects their social behavior by reducing antagonistic interactions and avoidance between competing species. This is likely to affect their distribution and functioning in the wild, where predator pressure is a standard situation.     

Circadian control of photonegative sensitivity in the haematophagous bug Triatoma infestans

The negative phototactic response of Triatoma infestans (Hemiptera) was studied in an arena, half of which was kept dark and the other half illuminated with different light intensitites. For each intensity we measured the time the insects spent in the dark half, the time to reach the side opposite to that where they were released, and the number of passages through the middle line of the arena. T. infestans displayed a photonegative behaviour that was enhanced by high light intensities. Bugs maintained in 12:12 light-dark cycles responded differently to the same illumination levels when tested in their photophase and scotophase: sensitivity to light was higher during the latter. Bugs entrained to light-dark cycles, kept afterwards either in constant darkness or in constant light, and tested in their subjective night and day, showed the same responses as bugs from the light-dark group tested in their corresponding photophase and scotophase. Thus, phototactic sensitivity is under endogenous control. The behaviour shown by T. infestans may be understood as being composed of at least two different drives: an exploratory one, and a negative phototactic response that is under endogenous control and is particularly sensitive to light during the scotophase, when activity peaks occur. Accepted: 16 July 1998     

Aggregation behaviour and interspecific relationships in Dermaptera

The rate of gregariousness and intra- and inter-specific attractivity were evaluated under laboratory conditions in five species of Dermaptera. The three species of the genus Forficula (Forficula auricularia L., F. decipiens Gené and F. pubescens Gené) showed distinctly gregarious behaviour in old larvae and adults in the pre-reproductive phase. This gregariousness was seen at the intraspecific level and also among individuals of different species. The other two species, Labidura riparia Pallas and Euborellia moesta Gené, were randomly distributed over the available shelters, with no distinct tendency for aggregation or isolation. In individual tests, insects of all five species were attracted by shelters previously visited by conspecifics in preference to control shelters. For the two non-gregarious species, this phenomenon might function as a burrow marker. The same tests showed an interattractivity for four of the species studied. Only L. riparia did not respond to the trails of the four other species, although its trails were attractive to the other species. The pheromonal nature of the substances involved in the interspecific relationships and the involvement of behaviour in the biology and speciation of Dermaptera are discussed.     

Sheltering behaviour of<Emphasis Type="Italic"> Macrobrachium nobilii</Emphasis> (Henderson and Matthai, 1910)

Shelter acquisition seems to be one of the main causes for agonistic interactions in the communal culturing of decapod crustaceans, leading to reduction in survival and growth-rate values. Understanding how to reduce aggressive behaviour among individuals by providing suitable shelters would promote production efficiency and welfare in such aquaculture environments. Factors influencing the sheltering behaviour of a freshwater prawn, Macrobrachium nobilii, were studied in laboratory conditions. Prior ownership significantly increased the ability to retain a shelter; males were significantly more likely to acquire and retain a shelter than females, except females carrying eggs. Various movements of the prawn while acquiring the shelter and the behaviour pattern involved in evicting an occupant are described. The size of the shelter selected by an animal is directly related to its body size. Regarding the choice of the colour of the shelter, juveniles and adults preferred dark shelters over light-coloured shelters and never chose a transparent shelter. Communicated by R.F. Oliveira     

Invasion of the shelter snatchers: behavioural plasticity in invasive red swamp crayfish,Procambarus clarkii

1. In many freshwater systems, competition for shelter plays an important role in determining the persistence of both native and alien species. The red swamp crayfish, Procambarus clarkii, is currently invading the native habitat of the signal crayfish, Pacifastacus leniusculus, in southern Oregon, and interspecific competition for shelter may be driving the species replacement in this region. 2. We designed a 2 × 3 factorial mesocosm experiment, with shelter density and species combination as factors, to investigate shelter occupancy and resource competition. Contrary to our predictions, the two crayfish species are equal competitors for shelter. Further, the invasive P. clarkii modified its shelter occupancy behaviour in the presence of the native P. leniusculus and has broader microhabitat preferences. 3. Specifically, we found that P. clarkii alters shelter occupancy and space use patterns when the two species occurred together, such that shelter use was identical between P. clarkii and P. leniusculus in mixed‐species treatments. In such treatments, both species increased their use of shelters when shelter density increased. When P. clarkii was alone, however, individuals did not alter shelter use as a function of shelter density, whereas P. leniusculus exhibited similar density‐dependent behaviour in both mixed‐ and single‐species treatments. 4. In a complementary field survey, we employed an ‘epicentre‐based’ design to sample two field sites. We observed patterns of microhabitat use and breadth for each species similar to those in our mesocosm experiment: the invasive P. clarkii was more abundant across different habitats and used a broader range of microhabitats than the native P. leniusculus. As such, we found that P. clarkii was more abundant across both field sites than the P. leniusculus, occupying microhabitats within and beyond the preferred range of P. leniusculus. Both field sites were affected by urban development and agriculture. 5. The use of microhabitats by both species was similar in the laboratory and the field. This study confirms that P. clarkii individuals can, and do, successfully occupy microhabitats preferred by P. leniusculus in the Willamette Valley. The results from our study may be relevant to other freshwater systems inhabited by P. clarkii and contribute to the understanding of ‘niche opportunity’, a concept which defines the environmental conditions that promote biological invasions.     

Ecological consequences of shelter sharing by leaf-tying caterpillars

A wide variety of insect herbivores construct and inhabit leaf shelters (ties, rolls, folds, and webs). Shelter construction can lead to a high rate of secondary occupation by other arthropods, including other species of constructors. The consequences for the inhabitants of secondarily occupying these shelters are currently unknown. In this study, we conducted field experiments to examine the fitness consequences (survival and attack by natural enemies) for caterpillars that (i) occupy a shelter with conspecifics vs. occur singly; and (ii) establish a new shelter vs. colonize a pre‐existing one. In addition, we conducted factorial laboratory experiments to test the hypothesis that caterpillars sharing shelters with conspecifics might have reduced construction costs (a potential benefit of shelter‐sharing or secondary occupation). Larvae of Psilocorsis quercicella Clemens (Lepidoptera: Oecophoridae) placed in white oak [Quercus alba L. (Fagaceae)] leaf ties alone or in groups of three had equal likelihood of survival from natural enemies. This same caterpillar species, however, had a higher disappearance rate when placed in pre‐existing leaf ties than when placed in newly formed ones, suggesting a potential cost of secondary colonization. A similar experiment with a closely related species [Psilocorsis cryptolechiella (Chambers)], however, failed to detect a cost of secondarily occupying shelters made on beech, Fagus grandifolia Ehrh. (Fagaceae). In the laboratory experiment, we found no evidence of shelter‐sharing benefits; rather larvae reared in shelters in groups of three had lower pupal mass (and thus lower potential fecundity) than larvae reared singly, suggesting a cost of shelter sharing. Moreover, groups of larvae forced to repeatedly construct new shelters tended to have reduced survival relative to the other treatment, suggesting that energetic constraints are more likely to reduce fitness when larvae cohabit shelters. Taken together, these results indicate that the common phenomenon of shelter sharing by leaf‐tying caterpillars has either neutral or negative effects for the occupants. The fact that these leaf‐tying caterpillars actually share shelters may simply reflect limited availability of oviposition sites.     

Inoculation of susceptible and resistant potato plants with the late blight pathogen Phytophthora infestans: effects on an aphid and its parasitoid

Plants are exposed to microbial pathogens as well as herbivorous insects and their natural enemies. Here, we examined the effects of inoculation of potato plants, Solanum tuberosum L. (Solanaceae), with the late blight pathogen Phytophthora infestans (Mont.) de Bary (Peronosporales: Pythiaceae) on an aphid species commonly infesting potato crops and one of the aphid's major parasitoids. We observed the peach‐potato aphid, Myzus persicae Sulzer (Hemiptera: Aphididae), and its natural enemy, the biocontrol agent Aphidius colemani Viereck (Hymenoptera: Braconidae), on potato either inoculated with water or P. infestans. Population growth of the aphid, parasitism rate of its natural enemy, and other insect life‐history traits were compared on several potato genotypes, the susceptible cultivar Désirée and genetically modified (GM) isogenic lines carrying genes conferring resistance to P. infestans. Effects of P. infestans inoculation on the intrinsic rate of aphid population increase and the performance of the parasitoid were only found on the susceptible cultivar. Insect traits were similar when comparing inoculated with non‐inoculated resistant GM genotypes. We also tested how GM‐plant characteristics such as location of gene insertion and number of R genes could influence non‐target insects by comparing insect performance among GM events. Different transformation events leading to different positions of R‐gene insertion in the genome influenced aphids either with or without P. infestans infection, whereas effects of position of R‐gene insertion on the parasitoid A. colemani were evident only in the presence of inoculation with P. infestans. We conclude that it is important to study different transformation events before continuing with further stages of risk assessment of this GM crop. This provides important information on the effects of plant resistance to a phytopathogen on non‐target insects at various trophic levels.     

Sulfated glycosaminoglycans in two hematophagous arthropod vectors of Chagas disease, Triatoma brasiliensis and Rhodnius prolixus (Hemiptera: Reduviidae)

The characterization of sulfated glycosaminoglycans (GAGs) in hematophagous arthropod vectors in general has been limited, with the exception of the studies in the triatomine Rhodnius prolixus. Heparan sulfate (HS) and chondroitin sulfate (CS) were previously identified and structurally characterized in extracts of whole bodies of fourth instar larvae of R. prolixus. Recently, we showed the expression of these two sulfated GAGs in specific body tissues of adult males and females and in embryos of R. prolixus. In the present work, we identified and compared the sulfated GAG composition in specific tissues of adult insects and in embryos of another triatomine species, Triatoma brasiliensis. Sulfated GAGs were isolated from the fat body, intestinal tract, and the reproductive tracts of adult insects and from embryos. Only HS and CS were found in the tissues analyzed. The present results extend the initial observations on the sulfated GAG composition in R. prolixus by showing that these molecules are widely distributed among internal organs of triatomines. These observations may be useful for future investigations aiming to evaluate the possible implication of these compounds in physiological events that take place in a specific organ(s) in these insects.