Notes on paternal care and sibling cannibalism in the giant water bug, Lethocerus deyrolli (Heteroptera: Belostomatidae)

Males of the giant water bug, Lethocerus deyrolli, care for egg masses on vegetation above the water surface. They supply the developing eggs with water and guard them against predators. In the present study, mechanisms by which paternal care is extended were found. Males were found situated just below the water on the natal substrate (usually a stick), and the first instar nymphs were aggregated around the substrate. When disturbed, the males showed aggressive behavior, threatening the intruder with their forelegs. Nymphs up to 12 h old did not attack the offered sibling nymphs or anuran larvae, which are common prey in the field. The 24 h‐old nymphs attacked both prey animals; however, they preferred anuran larvae. Cannibalistic behavior in the nymphs was well developed 72 h after hatching, when the nymphs had already dispersed from the natal substrate. The suppression of sibling cannibalism in younger nymphs would promote the maintenance of tight nymphal aggregations and consequently extend male care in this predatory species.     

Absence of chemical alarm in a primitively eusocial wasp (Belonogaster petiolata,Hymenoptera: Vespidae)

Summary The hypothesis thatBelonogaster petiolata (fam. Vespidae) is able to communicate alarm chemically, using odours released with the venom, was tested in bioassays involving presentation of artificial targets to a wasp colony, simultaneously with crushed venom apparatuses. The odour of venom did not lower the threshold of attack and visual stimuli alone (particularly a black, moving object) were sufficient to release attack. Venom odour on a previously stung target probably does not play a role in focusing further attacks on such a target. The results therefore support the null hypothesis that a venom-based alarm pheromone is absent in this species.     

The seabird tick, <Emphasis Type="Italic">Ixodes</Emphasis><Emphasis Type="Italic">uriae</Emphasis>, uses uric acid in penguin guano as a kairomone and guanine in tick feces as an assembly pheromone on the Antarctic Peninsula

In the vicinity of Palmer Station, Antarctica, the seabird tick, Ixodes uriae, forms large aggregations under rocks at the periphery of Adelie penguin rookeries. When the adult penguins return to the rookeries at the beginning of the nesting season the ticks leave their off-host aggregation site, attach to the penguins for a period of feeding, and then subsequently return to the aggregation site. In this study, we searched for chemical cues that may be used by the ticks to locate their aggregation sites as well as cues involved in finding penguins. Tick excreta and soil extracts from beneath tick aggregations contained a pheromone that elicited assembly behavior in unfed larvae, non-fed nymphs and non-fed adults. Guanine, the major excretory product of ticks, elicited assembly behavior, thus, guanine is likely an active component involved in assembly. Non-fed stages also responded positively to penguin guano and uric acid, the primary excretory product of penguins, suggesting that uric acid and other components of penguin guano function as a kairomone used by the non-fed ticks to locate their host. After feeding, the immature ticks’ response to both the assembly and kairomones is switched off for several days, and the ticks regain responsiveness only after they have molted. Fed adult females lay eggs and die without ever regaining responsiveness. Thus, I. uriae relies on two closely related chemicals to regulate two critical aspects of its life: assembly and host-finding. Guanine and other components of tick excreta function as an assembly pheromone in promoting the formation of off-host aggregations, while uric acid and other components of penguin guano function as a kairomone used by the tick to locate its host.     

Use of Alternative Food Plants Exclusively by Adult Male Froghoppers (Homoptera: Cercopidae)1

Male-specific associations of adult froghoppers (Homoptera: Cercopidae) with alternative food plant hosts is reported. Four accounts from the field and one from the literature are presented: Iphirhina quota on Bourreria costaricensis (Boraginaceae), Huaina inca on Solanaceae sp., Prosapia simulans and Prosapia sp. (near P. bicincta) on Ilex haberi (Aquifoliaceae), and Prosapia bicincta on Ilex cornuta and Ilex opaca. In three cases host use is limited to adults while in two cases use is limited to only adult males. In four cases adults feed on plants that differ taxonomically, structurally and ecologically from hosts normally utilized by all life stages. These associations are not considered aggregations per se because froghoppers are not clustered and because these associations cannot be explained by the usual criteria supporting the adaptiveness of insect aggregations. Indirect evidence suggests that these unusual hosts may be unavailable to nymphs and not sought by females. This passive lack of opportunity by nymphs and females, in tandem with an active selection/attraction by males, is a plausible explanation for the existence of male-specific hosts.     

Horizontal and vertical movements of host‐seeking Ixodes pacificus (Acari: Ixodidae) nymphs in a hardwood forest

The nymph of the western black‐legged tick (Ixodes pacificus) is an important bridging vector of the Lyme disease spirochete (Borrelia burgdorferi) to humans in the far‐western United States. The previously unknown dispersal capabilities of this life stage were studied in relation to logs, tree trunks, and adjacent leaf‐litter areas in a mixed hardwood forest using mark‐release‐recapture methods. In two spatially and temporally well‐spaced trials involving logs, the estimated mean distances that nymphs dispersed ranged from ≈0.04 to 0.20 m/day on logs vs 0.11 to 0.72 m/day in litter. Prior to recapture in either trial and within the confines of the sampling grids, the greatest estimated dispersal distances by individual nymphs released on logs, and in litter 0.5 m or 1.5 m from logs, were 2.4, 3.0, and 3.0 m, respectively. Nymphs released on logs or litter tended to remain within the same biotopes in which they were freed while host‐seeking. In two simultaneous trials involving trunks spaced close‐at‐hand, nymphs released at the trunk/litter interface on all four aspects collectively dispersed a mean of 0.353 m/day on trunks vs 0.175 m/day in litter. In either trial, the greatest distances that recaptured nymphs climbed trunks, or dispersed in litter in an encircling 3‐m grid, were 1.55 m and 2.97 m, respectively. Nymphs ascending trunks did not exhibit a preference for any one aspect, and the B. burgdorferi‐infection prevalences in nymphs that climbed trunks (3.2–4.0%) did not differ significantly from those that moved horizontally into litter (10.5–17.6%). We conclude that I. pacificus nymphs use an ambush host‐seeking strategy; that they disperse slowly in all biotopes studied; that they usually continue to host‐seek in or on whatever substratum they access initially; and that B. burgdorferi‐infected nymphs are as likely to move horizontally as vertically when offered a choice.     

Predator detection and avoidance by lotic mayfly nymphs of different size

We studied antipredatory responses of lotic mayfly (Baetis) nymphs in a factorial experiment with four levels of fish presence: (1) a freely foraging fish (the European minnow,Phoxinus phoxinus), (2) a constrained fish, (3) water from a fish stream, (4) water from a fishless stream. LargeBaetis nymphs drifted mainly during night-time in treatments involving either the chemical or actual presence of fish, whereas no diel periodicity was observed when the water was not conditioned with fish odour. The response was strongest when the fish was uncaged, which suggests that visual or hydrodynamic cues are needed in addition to chemical ones for an accurate assessment of predation risk. Fish presence had no effect on the drift rates of small nymphs. Instead, they increased their refuge use in the presence of a live fish. Chemical cues alone did not have any effect on the refuge use of any of theBaetis size classes. Our results indicate active drift entry by mayfly nymphs. Because predation pressure is spatially and temporally variable, nymphs must sample the environment in order to locate predator-free areas or areas with low predation risk. Drifting should be the most energy-saving way to do this. To avoid the risk from visually feeding fish, large individuals can sample safely (i.e. enter drift) only at night-time, while the small ones can also do this safely during the day. We suggest that, contrary to some earlier assumptions, mayfly drift is not a fixed prey response. Instead,Baetis nymphs are able to assess the prevailing predation pressure, and they adjust their foraging behaviour accordingly.     

Alarm signals in Daphnia?

Daphnia magna can respond to chemical cues from freshly crushed conspecifics with various behavioural reactions. A shift in vertical distribution towards the bottom, the formation of aggregations and direct escape responses can all be induced by water-borne signals released from crushed Daphnia. The pattern and strength of the first two behavioural responses (i.e. the persistent tendency to occupy deeper strata in the experimental columns and to stay within patches) indicate that Daphnia perceive the signal from crushed conspecifics as nonspecific information, not necessarily associated with any particular kind of danger from either vertebrate or invertebrate predators. The adaptive value and possible costs associated with performing these two behavioural reactions are discussed. The adaptive value of the induced escape response was directly tested: Daphnia which had experienced the presence of a cue from crushed conspecifics avoided attacks by common bream more efficiently than naive Daphnia. The recognition of the signal originating from crushed conspecifics can be especially adaptive in encounters with unfamiliar predators and with predators that undergo ontogenetic shifts in their diet. Under natural conditions, the combination of such a signal with a predator cue can, very reliably, advertise the local scale of the predatory impact. Received: 10 October 1996 / Accepted: 20 May 1997     

Colonization of natural substrata of different roughness and colour by Ephemeroptera nymphs using retrieval and direct observation techniques

We investigated the colonization of stream substrata by Ephemeroptera nymphs in a Canadian Rocky Mountain stream. Stream substrata used in our study were different textures (smooth and rough) and colours (dark and light); the taxonomic groups studied were Drunella coloradensis Dodds (Ephemerellidae), Heptageniidae (Cinygmula and Epeorus) and a Baetis Ameletus group (Baetidae and Siphlonuridae). We tested the hypothesis that as D. coloradensis nymphs and other abundant Ephemeroptera nymphs increase in size they choose a dark substratum to remain cryptic and a substratum of greater texture to reduce the risk of being swept from the substratum by the water flow. Two approaches were used to test this hypothesis, a substratum basket experiment and direct observations.For the substratum basket experiment, there were no statistical differences in abundance of nymphs on the substratum types. But there was a general tendency that the D. coloradensis nymphs were least abundant on the rough dark-coloured substratum and developmental stage three nymphs were most abundant on the smooth white-coloured substratum.Colonization was investigated for other Ephemeroptera nymphs by making direct observations on the substratum baskets using a glass-bottomed box. For the taxonomic groups and two colonization periods, nymphs were statistically at greatest density on the smooth white-coloured substratum and lowest on the smooth black-coloured substratum. The Baetis-Ameletus groups were also found at greatest densities on the rough dark-coloured substratum.     

Behavioral ecology of Strongylocentrotus droebachiensis (Muller) (Echinodermata: Echinoidea)

Summary Field observation indicated that extensive aggregation behavior occurs in Strongylocentrotus droebachiensis, an important echinoid herbivore in rocky subtidal zones of northern New England. The relationship of this behavior to the behavioral ecology of this species was investigated. Laboratory studies indicated that urchins are chemically sensitive to the stimuli of food and other urchins. Quantitative field experiments with tagged animals and transects showed the existence of two types of aggregations: feeding and nonfeeding, which appear to be pervasive phenomena in the biology of S. droebachiensis.It is concluded that, depending upon their hunger state, urchins shuttle between feeding aggregations in exposed locations and non-feeding aggregations located in relatively sheltered areas (where they spend most of their time). Individuals of S. droebachiensis aggregate actively through response to other urchins in the form of chemotaxis. The aggregative behavior may be maintained for its selective value in terms of defense from predation and, to lesser extents, facilitation of feeding and breeding.     

The function of sound in male spacing behaviour in bush-crickets (Tettigoniidae,Orthoptera)

This paper examines the spatial dynamics of an aggregation of singing male bush-crickets (Tettigoniidae) in respect to the degree of regularity between males. Our results establish that singing males of the genus Mygalopsis are regularly spaced within aggregations and that sound is involved in spacing. Attention is focussed on the male strategy of regular spacing within a widely dispersed aggregation in order to attract females reaching sexual maturity within the confines of that population. This is seen as an alternate to male aggregations attracting females from outside their boundary.     

Sexual dimorphism of arrestment and gregariousness in the bed bug (Cimex lectularius) in response to cuticular extracts from nymphal exuviae

Releaser pheromones have direct behavioural effects to arrest, attract or disperse insects, whereas interactions within groups of social insects are often influenced by primer pheromones. The behaviour of insects displaying intermediate levels of sociality is largely unexplored in this context. In the present study, both the gregariousness and arrestment (settling near the odour source) of bed bugs Cimex lectularius L. (Hemiptera: Cimicidae) in response to conspecific exuvial extracts are described. Adult males are arrested on filter papers with extracts derived from exuviae of fifth‐instar nymphs. Adult females and nymphs display no significant evidence for such behaviour. Adults of both sexes show no preference for extracts of male versus female fifth‐instar exuviae. Arrestment of adult males does not occur on papers treated with fourth‐instar exuvial extracts. Because the insects are assayed behaviourally in groups, an index is calculated describing how far bugs are away from being located independently of one another, as a measure of gregariousness. Adult males have lower values for this index (i.e. locations are closer to independence). Adult females, nymph cohorts and mixed age groups all have higher values for this index, which tend to increase over time. Females exhibit a clear increasing dose‐dependent relationship for this index. It is concluded that the extracts of fifth‐instar nymphal exuvia arrest males on refuges that possess the odour source. However, gregariousness is induced in females, without evidence of a tendency to assemble near the odour source.     

Endocrine manipulations and ontogenesis of male sexual behaviour in Locusta: studies on azadirachtin-induced over-aged nymphs

ABSTRACT. Dose-response relations of the moult-inhibitory and lethal effects of azadirachtin (AZA) injected into fifth and fourth instar hoppers of Locusta migratoria migratorioides are presented. The optimal dose for obtaining maximum number of long-surviving over-aged nymphs is 1.6 and 1.0 μg of AZA per insect for young fifth and fourth instar nymphs, respectively. Over-aged male nymphs are capable of exhibiting sexual behaviour. The mating behaviour of fifth instar over-aged male nymphs, and of chemically allatectomized but otherwise similar over-aged male nymphs (obtained by combined precocene treatment and injection of AZA), with and without injections of exogenous Juvenile Hormone III (JH), is investigated quantitatively; the results are compared with those obtained for normal, chemically allatectomized, and chemically allatectomized plus JH-injected adult males. Over-aged fifth instar male nymphs show a low intensity of mating behaviour and in chemically allatectomized ones this intensity is even lower. Injections of JH intensify the mating behaviour of chemically allatectomized fifth instar over-aged male nymphs in a dose-dependent way, but less markedly so than in chemically allatectomized adult males. We conclude that the intensity of male mating behaviour of the over-aged nymphs is subnormal and their response to the mating behaviour intensifying effect of the JH is weaker than that of adult males. The relations between endocrine factors and ethological ontogenesis of male sexual behaviour in locusts are discussed.     

Sampling and interpretation of psyllid nymph counts in potatoes

Development of effective management practices for insect pests relies heavily on sampling methods to accurately detect and quantify emerging populations. Herein we describe a novel method to extract and count nymphs of potato psyllid, Bactericera cockerelli (?ulc) (Hemiptera: Triozidae), from leaves in commercial fields of potato, Solanum tuberosum L. (Solanaceae). The proposed sampling method (referred to as the leaf washing method, LWM) consists of: (1) immersing samples of infested leaves in cold water to remove dust and sand; (2) immersing leaves in >85 °C water for 5 s; (3) extracting psyllid nymphs from heated water by passing it through a sampling unit composed of a funnel with a fine mesh‐organza fabric inside and attached to a vacuum pump; and (4) removing organza fabric and count psyllid nymphs under a stereoscope. With five sampling units operating simultaneously, one person is able to process and count immatures from about 400 leaf samples in two normal work days. As examples of applications, the LWM was used to characterize: (1) the vertical distribution of potato psyllid nymphs in the potato canopy; and (2) the spatio‐temporal distribution of potato psyllid nymphs in three potato varieties. Using LWM, we showed that psyllid nymphs were most predominant in the middle portion of the canopy and that spatio‐temporal distributions of nymphs varied among potato varieties.     

The role of amphibian prey in the diet and growth of giant water bug nymphs in Japanese rice fields

Predatory insects that depend upon particular prey animals are commonly regulated by the prey animal’s abundance. Nymphs of the giant water bug Kirkaldyia (=Lethocerus) deyrolli (Heteroptera: Belostomatidae) are predators regarded as specialists in feeding on tadpoles. We studied the ontogenetic diet shift of aquatic nymphs by quantifying instar abundance and by analyzing captured prey and prey relative abundance during the period of rice irrigation in three localities. We also evaluated the contribution of major prey items (tadpoles, frogs, and Odonata nymphs) on specific growth rates of each nymphal stage in a rearing experiment. First to third-instar nymphs of K. deyrolli fed mainly on tadpoles, regardless of differences in prey availability. Nymphs of subsequent fourth and fifth instar stages shifted from tadpoles to other prey animals within each rice field. A rearing experiment demonstrated that giant water bug nymphs provided with tadpoles had greater specific growth rates at all nymphal stages, except for the final stage, than nymphs fed other prey (frogs and Odonata nymphs). The emergence of young K. deyrolli nymphs seemed to coincide with the period during which tadpoles became abundant in the rice fields. Consumption of tadpoles seems important to allow the nymph to complete its larval development in an unstable temporary habitat. An erratum to this article can be found at     

Sex differentiation studies relating to releaser aggregation pheromones of the desert locust,Schistocerca gregaria

Behavioural responses of nymphs and adults in the gregarious phase of the desert locust,Schistocerca gregaria (Forskal) (Orthoptera: Acrididae) were investigated in a single-chamber bioassay system to a choice of two columns of air, one permeated with airborne volatiles emanating from either sex of nymphs or adults and the other untreated. There was no sexual differentiation in the production of or response to nymphal volatiles. Young adults of either sex did not produce a stimulus with significant activity. Of the older adults, only the males produced the aggregation stimulus to which both sexes were equally responsive. Charcoal-trapped volatiles from the two sexes of nymphs and adults evoked similar aggregation responses. Antennae of the older adults showed significantly higher EAG responses than those of fifth instar nymphs to all four volatile collections, of which volatiles from older adult males were the most stimulatory and evoked the highest EAG amplitudes.     

Influence of diet on growth,food retention time,and gill ventilation rate of nymphs of Cloeon sp. (Ephemeroptera: Baetidae)

Early instar nymphs of the mayfly, Cloeon sp. were kept on an algal or a detrital diet at 20 ± 1 °C in the laboratory. A control group was not given any food. Nymphs kept on algae showed significantly (P = 0.001) higher growth both in terms of length and biomass than for those kept on detritus. None of the nymphs given algae or detritus died during the experiment, and 40 per cent of those fed algae and 20 per cent fed detritus reached maturity by the termination of the experiment. Food retention time as well as gill ventilation rate of nymphs feeding on algae were significantly (P = 0.001) higher than those feeding on detritus. It is probable that these two factors largely enabled the nymphs to survive as well as to grow and reach maturity, though relatively slowly, solely on a detrital diet. However, algae was clearly shown to be a superior food resource for Cloeon sp. nymphs.     

Possible Regulation of Feeding Behavior in Cockroach Nymphs by the Neurotransmitter Octopamine

Insects, including cockroaches, have the ability to select a proper diet from different nutrient choices. We have showed previously that various neurotransmitters and neuromodulators appear to regulate certain aspects of feeding in the cockroach, Rhyparobia madera. In the current study, we examined the role of octopamine in feeding behavior of cockroach nymphs. By either injection or direct incorporation into the diet blocks, an octopamine agonist (octopamine or synephrine) or antagonist (phentolamine) was effective in altering feeding in R. madera nymphs. Both octopamine and synephrine increased feeding slightly, while phentolamine decreased feeding dramatically. Phentolamine was able to decrease feeding, but not motor activity, when injected directly into the nymphs. Octopamine appears to cause increased feeding in the cockroach.     

Prolonged maternal care inElasmucha dorsalis (Heteroptera: Acanthosomatidae)

Female parents ofElasmucha dorsalis attend their offspring and show specific sequential defensive behavior when disturbed. Intense responses of brooding females such as wing fanning were usually triggered by exposure to crushed nymphs. Maternal care continued for a long time, sometimes into the fourth or last nymphal instar. Female removal experiments showed that maternal defense is highly effective under field conditions. High egg mortality observed in the field was probably due to the predation by an ant,Myrmica ruginodis, which occurred at high densities on the host plant,Aruncus dioicus. Most of the egg masses were found on the leaves subtending axillary inflorescence of the female plants, and nymphs usually aggregated and fed on fruits in the inflorescence. Because the food of nymphs is temporally limited, the female can produce only a single brood. Thus, temporality of food resources as well as a heavy predation pressure may have led to prolonged maternal care.     

Understanding the toad code: Behavioural responses of cane toad (Chaunus marinus) larvae and metamorphs to chemical cues

Abstract Larvae of many anuran taxa display strong behavioural responses to chemical cues, including alarm signals from injured conspecific tadpoles. We exposed tadpoles and metamorphs from an Australian population of the invasive cane toad (Chaunus[Bufo] marinus) to a range of chemical stimuli and quantified their responses both in the laboratory and in the field. Filtered fluids containing scent cues from crushed conspecifics elicited strong avoidance from tadpoles, whereas other cues (e.g. scent of food, of native‐range fish or urodele predators, and thermal stimuli) did not. Apparent aggregation of tadpoles in response to scent cues proved to be an artifact of tank design, and was an indirect consequence of avoidance of those cues. Field trials confirmed that free‐ranging toad tadpoles and metamorphs avoided chemical cues from crushed conspecifics, suggesting that the chemicals inducing this response might provide an opportunity to develop targeted control methods for this invasive species.     

The response of grass-cutting ants to natural and synthetic versions of their alarm pheromone

The responses of the grass‐cutting ants Atta bisphaerica (Forel) and Atta capiguara (Gonçalves) to the main components of their alarm pheromones were examined in simple field bioassays. Both species react most strongly to 4‐methyl‐3‐heptanone, which causes the full range of alarm behaviour and a large increase in the number of individuals near the sources. In later experiments with A. capiguara, this increase was found to be due primarily to attraction, with some arrestment also occurring. The ant response to 4‐methyl‐3‐heptanone was compared with that to crushed heads and to that with whole ants with crushed heads. The pheromone 4‐methyl‐3‐heptanone by itself stimulates the same level of attraction as crushed heads, but results in far less alarm behaviour and arrests fewer ants. Whole ants with crushed heads attract a greater number of ants than the other sources and also cause more alarm behaviour. Bodies alone attract ants, but do not result in alarm behaviour. The main component in both species is the same, supporting the view that alarm pheromones lack species specificity. However, it appears that other components may also be important either as synergists of the main compound, or by stimulating behaviours that would not be observed in its absence.