Behavioural responses of diverse insect groups to electric stimuli

Anecdotal evidence suggests that cockroaches respond to electrical appliances or outlets. Our objectives were to determine the effect of field‐inducing sources and field attributes on attraction of German cockroaches, Blattella germanica (L.) (Blattodea: Blattellidae), and to test those parameters found effective for attraction of B. germanica for attraction of other groups of insects. In two‐choice, large‐arena experiments, significantly more female, but not nymphal, B. germanica settled in or near electrified coils with static or fluctuating electromagnetic fields produced by low‐level direct current (DC) or alternating current (AC) sources than in control coils without current. Electromagnetic fields with the magnetic, but not the electric, component of the field nulled still attracted B. germanica, suggesting that the electric component of the field may contribute to the attraction or arrestment response of B. germanica. DC‐powered coils with static electromagnetic fields also attracted/arrested brown‐banded cockroaches, Supella longipalpa (Fabricius) (Blattodea: Blattellidae), common silverfish, Lepisma saccharina (L.), firebrats, Thermobia domestica (Packard) (both Thysanura: Lepismatidae), and European earwigs, Forficula auricularia (L.) (Dermaptera: Forficulidae), but they repelled American cockroaches, Periplaneta americana (L.) (Blattodea: Blattidae). If proven in field experiments, electrified coils as trap baits may offer non‐toxic alternatives to pesticides for selective insect control in urban environments.     

Pattern recognition and call preferences in treefrogs (Anura: Hylidae): a quantitative analysis using a no-choice paradigm

Studies of mate attraction have traditionally employed one of two experimental methods: choice tests in which female preferences from among two or more signals are tabulated, or single-stimulus tests in which the female response to one signal is quantified. Choice tests have long been preferred for examining mate attraction in anurans. Inspired by Wagner's (1998, Animal Behaviour,55, 1029-1042) discussion of the strengths and weaknesses of the two experimental approaches, we used a single-stimulus design to examine mate attraction in two sibling species of treefrogs (Hyla versicolor and H. chrysoscelis). We quantified female responses based upon the relative time required to approach signals varying in pulse rate, pulse rise time and pulse number. The data were used to generate response functions providing a quantitative measure of female attraction to the stimuli. Comparisons with data from choice experiments reveal broad similarities, as well as properties of female responses that had not been detected with choice tests. The results are discussed with regard to female selectivity for call parameters that are likely to mediate sexual selection and homospecific pairing.     

Evidence for cocladogenesis between diverse dictyopteran lineages and their intracellular endosymbionts

Bacteria of the genus Blattabacterium are intracellular symbionts that reside in specialized cells of cockroaches and the termite Mastotermes darwiniensis. They appear to be obligate mutualists, and are transmitted vertically in the eggs. Such characteristics are expected to lead to equivalent phylogenies for host and symbiont, and we tested this hypothesis using recently accumulated data on relationships among termites and cockroaches and their Blattabacterium spp. Host and symbiont topologies were found to be highly similar, and various tests indicated that they were not statistically different. A close relationship between endosymbionts from termites and members of the wood-feeding cockroach genus Cryptocercus was found, supporting the hypothesis that the former evolved from subsocial, wood-dwelling cockroaches. The majority of the Blattabacterium spp. sequences appear to have undergone similar rates of evolution since their divergence from a common ancestor, and an estimate of this rate was determined based on early Cretaceous host fossils. The results support the idea that the stem group of modern cockroaches radiated sometime between the late Jurassic and early Cretaceous-not the Carboniferous, as has been suggested on the basis of roach-like fossils from this epoch.     

Conspecific attraction and shelter selection in gregarious insects

During habitat selection, the presence of conspecifics can frequently drive a nonuniform distribution of animals across habitats of equivalent quality. In group-living species, subgroups of individuals might display mutual attraction while differing in their preferences for environmental resources. The final decision to settle requires individuals to integrate both environmental and social cues. This raises the question of the relative importance of sociality and resources preferences in determining habitat choice. In this study, we examined the interactive influence of conspecific attraction on individual resource preferences on refuge choice in groups of cockroaches. Shelters scaled to the sizes of nymphs and adult males were offered to groups of only nymphs and only males and to mixed groups. The choices of males were consistent across social conditions. Conversely, the preferences of nymphs shifted depending on the social context; the presence of males overrode the affinity nymphs had for scaled-size shelters. We developed a numerical model implementing parameters derived from these experiments to test whether the final spatial distribution of individuals originated from a differential attraction between nymphs and males that was associated with their relative body size. Finally, we propose a general framework for understanding how similar mechanisms can promote the skewed distribution of organisms at different spatial scales.     

Biological response of German cockroaches fed diets containing allopurinol

The acceptability of dietary allopurinol to German cockroaches, Blattella germanica (L.), was assessed. In diet choice tests between rat chow with or without 0.1% w/w allopurinol there were no significant differences in nymphal development, feeding duration, visits or consumption. The LT50 of cockroaches fed a choice of diets was 3 weeks greater (7.88) than those fed non-choice a 0.1% allopurinol diet. Female cockroaches provided a choice of diets aborted a significantly greater percentage of their oothecae (98.6%) than those fed the untreated diet (1.7%). Choice arena tests of 2% allopurinol in rat chow significantly reduced cockroach populations compared with untreated controls. After 6 weeks, populations were reduced by approximately 50%, and 97% after 14 weeks. These results indicate that allopurinol is acceptable to B. germanica as a dietary supplement which could be used in baits for cockroach control.     

SEXUAL SELECTION AND FEMALE CHOICE IN MOLE CRICKETS (SCAPTERISCUS: GRYLLOTALPIDAE): MODELLING THE EFFECTS OF INTENSITY AND MALE SPACING

ABSTRACT

Attraction of flying mole crickets to individual males calling in an outdoor arena was influenced significantly by the relative intensity of the males calling in the arena. Louder males attracted more individuals than males whose calling songs were less intense. Making simple assumptions about the acoustic output from calling males and about the flight pattern and response of flying females, the differential attraction can be explained by a mathematical model. Computer simulation of the model was used to examine the importance of a male's intensity relative to others and the effect of distance between males on the attraction of females. The model and its relation to active female choice and passive attraction are discussed. The model makes predictions about differences in spacing behaviour of males that maximize attraction relative to other males.     

Testing decreasing variabililty of cockroach forewings through time using four Recent species: Blattella germanica,Polyphaga aegyptiaca,Shelfordella lateralis and Blaberus craniifer,with implications for the study of fossil cockroach forewings

Abstract Vr?anský (2000) considered there is decreasing variability in the forewings of cockroaches through time, based on a study of fossil and Recent species. This study tests his theory, based on a study of four Recent species of cockroaches: Blattella germanica (Blattellidae), Polyphaga aegyptiaca (Polyphagidae), Shelfordella lateralis (Blattidae) and Blaberus craniifer (Blaberidae). The new results, based on the coefficient of variation (CV) of the number of veins, indicate that Recent species can be more variable than fossil species and therefore do not support the theory. The results also show that at least 30 complete wings of the same species are required for a reliable CV value that is comparable between species.     

Attraction of wild‐like and colony‐reared Bactrocera cucurbitae (Diptera:Tephritidae) to cuelure in the field

The attraction of wild tephritids to semiochemical‐based lures is the ideal basis for trap network design in detection programmes, but in practice, mass‐reared colony insects are usually used to determine trap efficiency. For Bactrocera cucurbitae Coquillett, a lower response by wild males compared with colony‐derived individuals, usually used to estimate attraction parameters, could mean that the sensitivity of detection networks targeting this pest is reduced. We describe the results of mark–release–recapture experiments with wild‐ and colony‐derived B. cucurbitae males in a grid of cuelure‐baited traps within a macadamia nut orchard in Hawaii Island designed to quantify the attraction of cuelure to each fly type. For colony males, we estimate a 65% probability of capture at 27 m, intermediate with previous estimates on the attraction of methyl eugenol to Bactrocera dorsalis Hendel (36 m) and trimedlure to Ceratitis capitata Wiedemann (14 m) at the same site. Results suggest similar response over distance by wild‐derived B. cucurbitae compared with colony in the field, but there may be qualitative differences in response between wild and colony based on very low response of wild males in a standard bioassay of attraction. For both fly types, the estimates of attraction in the smaller of two grid sizes tested were lower than for the larger spacing, suggesting trap competition was a factor at an intertrap distance of 75 m. Dispersal patterns within the grid were generally to the south for the colony‐derived males and more variable for the wild‐derived males. In neither case was the direction of recapture correlated with the prevailing direction of the wind.     

Evaluation of three methods for detecting permethrin resistance in adult and nymphal Blattella germanica (Dictyoptera: Blattellidae).

The insecticide resistance status of the German cockroach, Blattella germanica (L.), against permethrin was evaluated with three different bioassay methods. The LT50 and KT50 in adults were measured by glass jar test, the LT50 and KT50 in nymphs were assessed by glass petri dish-test, and the LD50 in adults and nymphs was measured by topical application. The results indicate that in all three bioassays, cockroaches showed different degrees of resistance to permethrin. Resistance ratios of 6.7-, 12.8-, and 7.5-fold for KT50, LD50, and LT50, respectively, were obtained for adult cockroaches compared with those from a susceptible strain. Similarly, resistance ratio values of 17-fold for KT50, 8-fold for LD50, and 6.4-fold for LT50 were obtained for nymphs compared with those from a susceptible strain. Although both adults and nymphs exhibited a similar level of susceptibility to permethrin, adults showed a higher degree of susceptibility to topical application than to the other exposure method. The order of sensitivity (based on resistance ratio values) was topical application (LD50) > LT50 > KT50 in adults. For nymphs, the resistance ratio in the knockdown test was higher than in other tests (resistance ratio = 17-fold). Therefore, the order of test sensitivity based on comparisons of resistance ratio values was KT50 > topical application (LD50) > LT50 in nymphs. Topical application exhibited a higher sensitivity than the LT and KT methods in adults. Thus, the results suggest that topical application and knockdown testing methods are the most sensitive assays for measuring resistance of adult and nymphal German cockroaches, respectively, to permethrin.     

Diet specialization in an extreme omnivore: nutritional regulation in glucose‐averse German cockroaches

Organisms have diverse adaptations for balancing dietary nutrients, but often face trade‐offs between ingesting nutrients and toxins in food. While extremely omnivorous cockroaches would seem excluded from such dietary trade‐offs, German cockroaches (Blattella germanica) in multiple populations have rapidly evolved a unique dietary specialization – an aversion to glucose, the phagostimulant in toxic baits used for pest control. We used factorial feeding experiments within the geometric framework to test whether glucose‐averse (GA) cockroaches with limited access to this critical metabolic fuel have compensatory behavioural and physiological strategies for meeting nutritional requirements. GA cockroaches had severely constrained intake, fat and N mass, and performance on glucose‐based diets relative to wild‐type (WT) cockroaches and did not appear to exhibit digestive strategies for retaining undereaten nutrients. However, a GA × WT ‘hybrid’ had lower glucose aversion than GA and greater access to macronutrients within glucose‐based diets – while still having lower intake and survival than WT. Given these intermediate foraging constraints, hybrids may be a reservoir for this maladaptive trait in the absence of positive selection and may account for the rapid evolution of this trait following bait application.     

Density dependence,boundedness, and attraction: detecting stability in stochastic systems

Summary By analogy with deterministic stability, the stability of stochastic ecological systems can be viewed as a tendency for population densities to avoid dynamic boundaries (i.e. boundedness) or to approach a dynamic attractor (i.e. attraction). At the population level, these two views generate predictions consistent with density dependence. I therefore devised two new statistical tests of attraction, the random-walk attraction test and the randomized attraction test; I then used them successfully, along with randomization techniques that detect boundedness and two autocorrelation methods, to test for density dependence in published sequences of population densities. The attraction tests identify the apparent attractor, the band of densities toward which density tends to shift between generations. Locating the apparent attractor can generate a prediction of the next direction of density change; for data from a dragonfly assemblage, about 80% of these predictions were correct. From the single-population tests, I also developed two multispecies tests of attraction (the multispecies random-walk and randomized attraction tests) and two multispecies tests of boundedness (the multispecies permutation and randomization tests). These detected attraction and boundedness in the dragonfly assemblage and attraction in a collection of laboratory fruitfly populations. An evaluation of the statistical power of the new density attraction tests indicates a strong dependence on the sequence length n and on the number of populations m: power increases with n and particularly with m. Nevertheless, detecting attraction becomes likely even in populations with strong linear density-dependence only with n>30 or for shorter sequences in multispecies assemblages.     

Newly emerged Batesian mimicry protects only unfamiliar prey

The evolution of Batesian mimicry was tested experimentally using avian predators. We investigated the effect of a search image on the protection effectiveness of a newly emerged Batesian mimic. The two groups of predators (adult great tits, Parus major) differed in prior experience with prey from which the mimic evolved. The Guyana spotted roach (Blaptica dubia) was used as a palatable prey from which the mimic emerged, and red firebug (Pyrrhocoris apterus) was used as a model. Optical signalization of the insect prey was modified by a paper sticker placed on its back. The cockroaches with the firebug pattern sticker were significantly better protected against tits with no prior experience with cockroaches. The protection of the firebug sticker was equally effective on cockroaches as it was on firebugs. The cockroaches with firebug stickers were not protected against attacks of tits, which were familiar with unmodified cockroaches better than cockroaches with a cockroach sticker. We suppose that pre-trained tits acquired the search image of a cockroach, which helped them to reveal the “fake” Batesian mimic. Such a constraint of Batesian mimicry effectiveness could substantially decrease the probability of evolution of pure Batesian mimic systems.     

Behaviours promoting grouping or dispersal of mother and neonates in ovoviviparous cockroaches

Summary. The nature of early relationships between mother and nymphs and among siblings was compared in four cockroach species belonging to the same ovoviviparous family (Zetoborinae) in order to characterise the behavioural interactions favouring dispersal or maintenance of the group of neonates. Behavioural interactions between mothers and their new-born nymphs and between two sibling neonates were video recorded and analysed with flow charts on factorial maps. In the solitary species Thanatophyllum akinetum, nymphs dispersed a few hours after birth without aggressiveness between siblings or between mother and offspring. In contrast females of Schultesia lampyridiformis displayed a behaviour never previously observed in cockroaches: aggression towards their own neonate nymphs, which could contribute to their dispersal. In Phortioeca nimbata and Lanxoblatta emarginata, nymphs stayed with their mother for 10 days, but social interactions differed between the two sp ecies: P. nimbata mothers actively searched for their nymphs, this behaviour being favoured by an active search for mutual contact by the nymphs themselves, while L. emarginata nymphs sought more actively the proximity of their mother but less contact between themselves. Thus, different species of Zetoborinae presented two types of dispersal of the young and two types of maintenance of the birth group, both achieved by specific behavioural interactions. The relevance of behavioural interactions for the characterisation of early gregarism and parental care is discussed.     

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.     

Local structure of DNA toroids reveals curvature-dependent intermolecular forces

In viruses and cells, DNA is closely packed and tightly curved thanks to polyvalent cations inducing an effective attraction between its negatively charged filaments. Our understanding of this effective attraction remains very incomplete, partly because experimental data is limited to bulk measurements on large samples of mostly uncurved DNA helices. Here we use cryo electron microscopy to shed light on the interaction between highly curved helices. We find that the spacing between DNA helices in spermine-induced DNA toroidal condensates depends on their location within the torus, consistent with a mathematical model based on the competition between electrostatic interactions and the bending rigidity of DNA. We use our model to infer the characteristics of the interaction potential, and find that its equilibrium spacing strongly depends on the curvature of the filaments. In addition, the interaction is much softer than previously reported in bulk samples using different salt conditions. Beyond viruses and cells, our characterization of the interactions governing DNA-based dense structures could help develop robust designs in DNA nanotechnologies.     

The antennal system and cockroach evasive behavior. II. Stimulus identification and localization are separable antennal functions

Cockroaches ( Periplaneta americana) orient their antennae toward moving objects based on visual cues. Presumably, this allows exploration of novel objects by the antennal flagellum. We used videographic and electrophysiological methods to determine if receptors on the flagellum are essential for triggering escape, or if they enable cockroaches to discriminate threatening from non-threatening objects that are encountered. When a flagellum was removed, and replaced with a plastic fiber, deflection of a "prosthetic flagellum" still activated the descending mechanosensory interneurons associated with escape and produced typical escape responses. However, escape was essentially eliminated by constraining the movement of the scape and pedicel at the antennal base. When cockroaches approached and briefly explored the surface of a spider or another cockroach with the flagellum, they produced escape significantly more often in response to subsequent controlled contact from a spider than from a cockroach. This discrimination did not depend on visual or wind-sensory input, but required flagellar palpation of the surface. The crucial sensory cues appear to involve texture rather than surface chemicals. These results indicate that cockroaches acquire basic information on stimulus identity during exploration of surfaces with flagellar receptors, but that basal receptors are triggers for escape behavior.     

Models of female choice in acoustic communication

In this article we present generalized theoretical models ofmate attraction in acoustic communication.The models use simpledecision rules for females and show theoretical differencesbetween the rules in relative attraction of females to soundsources (calling males). The important variables of the modelsare the relative powers of the sources, the distance betweenthe sources and the number of competing signalers.Relative attractionamong sources is a function of their relative power output withmore powerful signals attracting more females. Using differentassumptions about the spacing of sources with respect to femalesensitivity, data from phonotaxis studies can be explained wellby either random pick or active choice models. Bioacousticiansmust reexamine their use of, and conclusions based on, the two-choiceplayback paradigm in studying female mate choice.     

Comparisons of toxic baits for controlling the cockroach, Blattella germanica: attractiveness and feeding stimulation

Female German cockroaches, Blattella germanica (L.) (Dictyoptera: Blattellidae) and nymphs (stages 1, 3 and 6) were used to test the attractiveness and the feeding stimulation of several commercial gel formulations of insecticide food baits in laboratory binary choice tests. The four toxic gel baits tested were: 'Avert' (0.05% abamectin=avermectin B1), 'Drax' (33.3% boric acid), 'Goliath' (0.05% fipronil) and 'Maxforce' (2.15% hydramethylnon). The attractiveness of a bait was estimated by the number of cockroaches that chose it first, and by their tendency to change to it secondarily. Feeding stimulation was estimated by the mean duration of feeding bouts and total duration of food intake. Variability of these factors was assessed in relation to age of cockroaches and to age of baits. With a few exceptions, cockroach age did not affect performance in any of these tests. Attractiveness to B. germanica ranked 'Goliath' gel higher than 'Avert', 'Drax' and 'Maxforce' gels, whether or not the active ingredient fipronil was present in 'Goliath' gel. Feeding stimulation ranked 'Avert' and 'Maxforce' gels higher than 'Goliath' gel, and 'Drax' gel was inferior. With ageing up to 3 months, the feeding stimulation power of 'Goliath' and 'Maxforce' gels increased, whereas that of 'Avert' fluctuated. Assuming that all types of bait effectively kill cockroaches once ingested, these results illustrate the importance of behavioural observations on bait efficiency and show that ranking of these different food baits varies in relation to the parameter analysed.     

Toxicity of abamectin to cockroaches (Dictyoptera: Blattellidae, Blattidae).

Abamectin was fed to German cockroaches, Blattella germanica (L.), in non-choice tests. LT50s and LC50s were estimated by probit analysis. The LT50s for the German cockroach ranged from 4.4 to 1.7 d for males, from 9.0 to 2.4 d for females, and from 4.4 to 1.6 d for nymphs for bait concentrations of abamectin between 0.0025 and 0.0500%. The LC50s of abamectin were 0.0110 and 0.0040% from males, 0.0240 and 0.0090% for females, and 0.0200 and 0.0080% for nymphs at 3 and 6 d, respectively. The LT50 values of 0.0550% abamectin bait were 3.4, 3.4, 2.4, 7.5, 2.9, and 4.5 d for Periplaneta americana (L.), P. fuliginosa (Serville), P. brunnea Burmeister, P. australasiae (F.), Blatta orientalis L., and Supella longipalpa (Serville). Although the bait was effective against various cockroach species, time to death for the larger species was longer than for the German cockroach. In preference tests in which male German cockroaches were allowed to feed on rat chow or abamectin bait, all died within 5 d of exposure to abamectin bait. Abamectin bait consumption was not significantly lower than that of untreated rat chow. Arena tests with 0.0550% abamectin bait resulted in 31-75% mortality of German cockroaches after 9 d, with most control being achieved by treating harborages with the bait. The hydramethylnon standard resulted in 65% mortality after 9 d.     

Seasonality of peridomestic cockroaches (Blattoidea: Blattidae): mobility, winter reduction, and effect of traps and baits.

Populations of smokybrown cockroaches, Periplaneta fuliginosa (Serville), and Eurycotis floridana (Walker) were studied to compare mobility and population size between fall 1985 and spring 1986 seasons at site A, and to assess the effect of a combination of traps and baits at site B in northcentral Florida. Adult smokybrown cockroaches were more mobile in the spring than in the fall; a similar trend for E. floridana adults was not significant. Overall population levels were reduced after winter by approximately 73%. No significant shift in spatial distributions of nymphs occurred, but adult distribution changed from a predominance in palms (fall) to hardwood trees with tree holes (spring), suggesting that this habitat offers better survival during winter. Removal of cockroaches by trapping at site B, followed by a single application of dry distiller's grain containing 0.5 chlorpyrifos, reduced expected population levels (based on site A data) by 90% measured at 6 mo after treatment. These data refute the hypothesis that an increase in numbers of cockroaches infesting homes in the fall is due to an increase in mobility. Instead, the results suggest that population size and proximity of principal habitats to the home are probably responsible for domestic infestations in the fall. Use of traps, baits, or both at these foci significantly reduce populations over an extended period.