Predatory behavior of Polybia sericea (Olivier), a tropical social wasp (Hymenoptera: Vespidae)

Summary Polybia sericea (Olivier) (Hymenoptera: Vespidae) foragers were trained to visit experimental foraging plots and tests were conducted to determine the role of visual, olfactory, and chemotactile cues in prey location. Foragers approached prey from downwind and hovered downwind of visual and olfactory stimuli. Olfactory cues were more likely to elicit landing than were visual cues. Biting of potential prey was most consistently elicited by a combination of visual, tactile, and chemotactile cues. Foragers encountering large prey carried a piece back to the nest and returned for prey remains. These returning foragers used visual cues to direct intensive aerial search; olfactory prey cues elicited landing.     

Responses of Metaphycussp. nr. flavus to semiochemicals released from a scale host, Coccus hesperidum

Summary. Metaphycus sp. nr. flavus (Encyrtidae: Hymenoptera) is a parasitoid species collected from the Mediterranean region which lays its eggs in the immature stages of several economically important soft scale insects (Hemiptera: Coccidae), including brown soft scale, Coccus hesperidum L. (= host insect). Preliminary tests suggested that the parasitoid is most successful in producing offspring when it oviposits in the younger stages of brown soft scale. In Y-olfactometer bioassays measuring wasp choices and residence times, naïve parasitoids were significantly more attracted to yucca leaves infested with 26, 27, or 28 d-old scale than to uninfested leaves, whereas leaves with older (29-30 d-old) scale were no more attractive than uninfested leaves. Parasitoids also spent significantly more time in the arm with yucca leaves infested with 26 d-old scale than in the arm with uninfested leaves. These results are consistent with observations of the parasitoids reproductive success on scale of different ages, whereby older scale are more likely to encapsulate the developing eggs of M. sp. nr. flavusfemales than are younger scale. Further bioassays determined that yucca leaves that had been infested with 26 d-old scale but from which the scale had been removed were as attractive as infested leaves. In contrast, infested yucca leaves from which scale had been removed and the leaves subsequently washed with distilled water were less attractive than infested leaves. Furthermore, the wash water containing scale residues was attractive to female wasps. In total, these results suggest that Metaphycussp. nr. flavus females utilize volatile, water soluble compounds produced by brown soft scale as cues to locate suitable hosts.     

“Armpit effect” in an African cichlid fish: self-referent kin recognition in mating decisions of male Pelvicachromis taeniatus

Kin discrimination in nepotistic as well as in sexual contexts is widespread in animals including humans. However, the underlying mechanisms of kin discrimination are assumed to vary between species and—within species—between contexts. During solitary life stages, kin recognition based on social learning is assumed to be less reliable because kin templates that are not continuously updated may get lost. Therefore, self-referent phenotype matching (“armpit effect”), i.e., the comparison of unknown phenotypes with own characteristics, should be particularly relevant when no social cues are available. However, experimental evidence for this mechanism is scarce. Here, we examine self-referent kin recognition in a mate-choice context in adult male Pelvicachromis taeniatus, a socially monogamous cichlid fish from West Africa with biparental brood care and pronounced kin-mating preferences. Juvenile P. taeniatus live in groups, whereas adult males compete for access to breeding sites which they aggressively defend against rivals. Using computer-animated females as standardized visual stimuli in combination with olfactory cues of related and unrelated females, we show that adult males reared isolated from kin since egg stage were able to discriminate sisters from unrelated females. As males could have learned kin cues only from themselves, our study provides evidence for self-referent kin recognition and indicates that the observed inbreeding preferences are mediated by self-derived olfactory cues. Male preferences for sisters were correlated with male body size. We discuss the implications of quality-related mate choice for the evolution of kin-mating preferences.     

Attraction and learning in mate-finding by solitary bees,Lasioglossum (Dialictus) figueresi Wcislo and Nomia triangulifera Vachal (Hymenoptera : Halictidae)

Summary Males of the solitary sweat bee, Nomia triangulifera, patrol over large areas where thousands of females emerge, searching for receptive females. The daily operational sex ratio is strongly male-biased. Males contact dead, frozen (untreated) females more frequently than they contact females which were washed in hexane, showing that olfactory cues are utilized in mate-finding. A major source of female sex pheromone is in the head. Male pouncing on females is temporally non-random, indicative of group stimulation. Bioassays show that newly emerged females are more attractive to males than are older pollen-collecting females. Female odors are individually distinctive, based on male responses, and there is much variation among females in their attractive properties. Male responses to female odors suggest that learning is important for mating in natural populations. In contrast, the following hypotheses are unlikely to account for the observed behavior: (1) dissipation of female odors; (2) site learning and avoidance behavior by males; (3) decay of male motivation; or (4) male-produced repellents effective against other males. Laboratory and field studies show that female Lasioglossum figueresi produce individually-distinctive odors, which are attractive to males. There is considerable inter-individual variation among females in their attractiveness to males among sexually immature females. Male responses to female odors decay over the course of the presentation, suggesting the importance of learning in natural populations, although several alternatives could not be tested.     

Ubiquitous eight-carbon volatiles of fungi are infochemicals for a specialist fungivore

An enormous variety of volatile substances are released in distinctive blends by fungal substrates that should be recognisable for fungivores. Certain compounds dominate in most of the fungal species. Fungal oxylipins as the eight-carbon volatiles are the most prominent. This raises the question whether such are specific enough to qualify as appropriate host cues for a fungivore. We could demonstrate differentiated responses of the fungivorous beetle Bolitophagus reticulatus to eight-carbon volatiles: Nine eight-carbon volatiles were identified with GC–MS from its host fungus Fomes fomentarius. 1-Octen-3-ol, 3-octanone and 3-octanol induced contrasting behaviour of beetles in olfactometer bioassays. Electroantennographic experiments investigating the beetle olfactory sense revealed distinguishable antennal responses. Moreover, their individual release from F. fomentarius fruiting bodies changes not only considerably, but also independently over successive stages of beetle colonisation. Concentrations of attractive and repellent eight-carbon volatiles correlate to frequency of beetles in the field and further substantiate their relevance as host cues. Our results show that a specialist fungivore is able to differentiate the most common eight-carbon volatiles of fungi to assess host quality. Key roles and marked similarities of fungal to plant oxylipins suggest a comparable importance of eight-carbon volatiles to fungivores as green leaf volatiles have to herbivores.     

Salience of multimodal flower cues manipulates initial responses and facilitates learning performance of bumblebees

Visual and olfactory cues are the first interface between flowers and their visitors and are adaptations to facilitate successful pollination. Initial responses to and associative learning of multimodal cues by flower visitors are based on the perception of colours and volatiles. In this study, we tested how visual, olfactory and multimodal stimuli affect the behaviour of bumblebees (Bombus terrestris) and correlated the properties of 28 stimuli in relation to the insects’ sensory equipment to these behaviours. Initial responses and associative learning were recorded using a radio frequency identification system, tracking the visitation sequences of individual bumblebees to artificial flowers treated with naturally occurring pigments and/or volatiles. The salience of the stimuli was evaluated as the colour contrast to the background and as electroantennogram responses. The main finding was that both initial responses and learning performance were positively correlated to the salience of the stimuli, suggesting that salience is a key feature of flower cues in the interactions with insects. The salience of compound stimuli consisting of two or more pigments and/or volatiles was largely additively determined by the saliences of individual compounds. Potentially, the valence of the stimuli may interfere with the positive relationship between salience and behaviour, which is indicated by our results, too. The salience of multimodal cues depends on the species-specific equipment of visual and olfactory receptors and thus enables flowers to be advertising for some but rather inconspicuous for other flower visitors.     

Cave molly females (Poecilia mexicana, Poeciliidae, Teleostei) like well-fed males

We examined the preference of Atlantic molly females (Poecilia mexicana) to associate with a well-fed or a starved male in simultaneous choice tests. Females from three different populations were tested in three treatments: (1) the females could choose on the basis of multiple cues from the males (visual plus non-visual); (2) only non-visual cues could be perceived in darkness, (3) only visual cues were presented. The three tested populations differ clearly in their ecology: one population occurs in a typical river habitat, the second one in a milky sulfur creek outside a cave, and the third population occurs in a cave habitat (cave molly). In the river-dwelling population, females never showed a preference. In the population from the sulfur creek, females preferred to associate with the well-nourished male when visual cues from the males were available. Only cave molly females exhibited a strong preference for well-nourished males in all treatments. A morphological comparison demonstrated that wild-caught males from river habitats are typically in a good nutritional state. In the sulfur creek, males showed signs of starvation. Cave molly males were in an even worse nutritional state. In the cave population, saturated males probably indicate high fitness, thereby driving the evolution of the preference for good male nutritional state.Communicated by K. Lindström     

Sensory cues associated with host detection in a marine parasitic isopod

Parasite population dynamics and the evolution of life history characteristics are strongly associated with the process of host infection. Parasites with free-living life stages have a narrow window to infect a host and have evolved a number of mechanisms to detect and locate a host. Cymothoa excisa is a parasitic isopod that is commonly found on the Atlantic croaker, Micropogonias undulatus, in the Gulf of Mexico. Here, we determined the infection window that constrains host-searching behavior in C. excisa and tested the behavioral response of the free-swimming larvae (mancae) to visual and chemical cues. Mancae were found to have an infection window of 7 days. Mancae were responsive to both visual and chemical cues. Our findings are the first to show that cymothoid isopods use visual and chemical cues to locate a host and that individuals display a host-locating strategy that maximizes host encounter rate, while reducing energy expenditure.     

Female association preferences based on olfactory cues in the guppy,<Emphasis Type="Italic"> Poecilia reticulata</Emphasis>

Guppies are highly sexually dimorphic. Females have been shown to mate preferentially with males with various visible traits and previous studies investigating mate choice have focussed on these visual cues. However, guppies have been shown to respond behaviourally to odour cues from conspecifics as well as from predators. Therefore, sexual selection in guppies may act on multiple sensory modalities, but so far only visual cues have been investigated. Here, we used a specially designed olfactory choice tank to determine whether females could detect other guppies on the basis of their olfactory cues only, and to determine whether females preferred to associate with males or females and whether they could distinguish between different males. We show that female guppies associated preferentially with other females when given a choice between water containing the odour cues from another fish and water containing no odour cues. When females were presented with olfactory cues from a male and a female, they investigated the female initially, but then most reversed their decision and swam to the male. Furthermore, we demonstrate that females associated preferentially with certain males based upon olfactory cues alone, but that this choice was in direct opposition to that made when the females received only visual cues from the same males.Communicated by: M. Abrahams     

Preventing behavioural interactions with a male facilitates sex change in female bluebanded gobies, Lythrypnus dalli

Sex change in marine teleost fishes is commonly regulated by social factors. In species that exhibit protogynous sex change, such as the bluebanded goby Lythrypnus dalli, the most dominant female typically initiates sex change when a male is removed from the social group. Females can use visual, chemical or tactile cues to assess the presence or absence of a male. The primary goal of our study was to determine whether the olfactory and visual presence of a male versus its behavioural interactions with females were important for mediating sex change. We exposed females to three different treatments: absence of a male, presence of a male that could physically interact with her and presence of a male behind a barrier that allowed visual and olfactory interactions but prohibited physical interactions. Sex change occurred in the absence of a male but not in the presence of a male that could physically interact with the female. The presence of a male behind the barrier did not prevent sex change but affected the timing of sex change. Season appeared to affect the latency to initiate male typical courtship, with a delay at the end of the reproductive season only when the male was present behind the barrier. We discuss the seasonal results in terms of L. dalli life history and the potential benefits and costs of changing sex late in the season in the presence or absence of aggressive reinforcement by the male. Our results identify direct behavioural interactions as an important proximate mechanism in the social regulation of sex change in L. dalli.     

All at sea: aquatic life modifies mate-recognition modalities in sea snakes (Emydocephalus annulatus, Hydrophiidae)

Alternative sensory modalities (e.g. vision, chemoreception) differ in the spatial scale, permanence and reliability of cues they provide to mate-searching males. Males of terrestrial snake species use chemoreception to locate females over large distances, but phylogenetic shifts to aquatic life render such cues unavailable. Do male sea snakes use alternative modalities for identifying potential sexual partners and if so, are the novel systems as effective for mate-finding as the ancestral ones? Observations and experiments show that free-ranging male turtle-headed sea snakes (Emydocephalus annulatus) in shallow-water reef habitats in New Caledonia use visual cues (including size, movement and color pattern) to assess whether snake-shaped objects are potential sexual partners. Skin lipids (pheromones) are important only after the male comes into physical contact with the stimulus. Visual cues provide unreliable information about potential mates, and function over short distances only (generally, <1 m). In consequence, mate-searching male snakes frequently failed to find nearby females, rarely managed to maintain contact with females they did find, and wasted time and energy investigating inappropriate stimuli (e.g. fishes, sea cucumbers, divers). The loss of effective pheromonal mate-location systems means that mate recognition by aquatic snakes functions over smaller distances than in their terrestrial relatives. Phylogenetic transitions among habitat types thus may directly modify central features of the mating system.     

Host preference,site fidelity,and homing behavior of the symbiotically luminous cardinalfish,Siphamia tubifer (Perciformes: Apogonidae)

The sea urchin cardinalfish, Siphamia tubifer (Perciformes: Apogonidae), is unusual among coral reef fishes for its use of bioluminescence, produced by symbiotic bacteria, while foraging at night. As a foundation for understanding the relationship between the symbiosis and the ecology of the fish, this study examined the diel behavior, host urchin preference, site fidelity, and homing of S. tubifer in June and July of 2012 and 2013 at reefs near Sesoko Island, Okinawa, Japan (26°38′N, 127°52′E). After foraging, S. tubifer aggregated in groups among the spines of the longspine sea urchin, Diadema setosum, and the banded sea urchin, Echinothrix calamaris. A preference for D. setosum was evident (P < 0.001), especially by larger individuals (>25 mm standard length, P < 0.01), and choice experiments demonstrated the ability of S. tubifer to recognize and orient to a host urchin and to conspecifics. Tagging studies revealed that S. tubifer exhibits daily fidelity to a host urchin; 43–50 and 26–37 % of tagged individuals were associated with the same urchin after 3 and 7 days. Tagged fish also returned to their site of origin after displacement; by day two, 23–43 and 27–33 % of tagged individuals returned from displacement distances of 1 and 2 km. These results suggest that S. tubifer uses various environmental cues for homing and site fidelity; similar behaviors and cues might be used by larvae for recruitment to settlement sites and for the acquisition of luminous symbiotic bacteria.     

Shelter selection of the spiny lobster <Emphasis Type="Italic">Palinurus elephas</Emphasis> under different levels of <Emphasis Type="Italic">Octopus vulgaris</Emphasis> predation threat

The skill of recognizing and reacting to predators is often based on a learned component. Few studies have examined the role of learning in spiny lobster anti-predator behavior. We investigated whether European spiny lobster (Palinurus elephas) shelter selection is influenced by olfactory stimuli released by one of the most common lobster predators, the common octopus (Octopus vulgaris), and whether the behavioral response to octopus chemical stimuli is innate or influenced by experience. In experimental arenas, we conditioned wild-caught lobsters with three levels of predation threat: no threat, with no predator–prey interaction; medium threat, with odor and visual predator cues only; high threat, active predation risk. We subsequently tested the shelter choice of the conditioned lobster under different experimental conditions: (1) shelter plus seawater; (2) shelter plus seawater plus chemical octopus cue. Our results showed significant differences in mean shelter occupancy with conditioning level. We conclude that P. elephas individuals use chemosensory systems in predator-avoidance mechanisms. Moreover, lobsters subject to a training period of high-level predation threat were able to learn the octopus chemical stimuli and treat its odor as a cue related to predation risk. The findings relative to the spiny lobster learning abilities could be an important tool for future management of lobster populations, e.g., by re-introduction of reared juveniles, which have not yet experienced predation.     

An experimental field test of host-finding mechanisms in a Caribbean gnathiid isopod

Field experiments were conducted from dusk to dawn off St. John (18° 18′ 59.32″ N, 64° 43′ 24.5″ W) and Guana Island (18° 28″ 28.31″ N, 64° 34′ 30.83″ W), Virgin Islands from June through August 2008-2010 to assess the sensory cues used by the nocturnal/crepuscular fish-parasitic gnathiid isopod, Gnathia marleyi, to locate fish hosts. Experimental traps providing both visual and olfactory cues from live French grunts (Haemulon flavioliniatum) attracted significantly more gnathiids than traps providing only visual cues or control traps (empty or with a rock), which were not significantly different from each other. In another experiment, traps providing both cues and only olfactory cues attracted significantly more gnathiids than empty control traps, but were not significantly different from each other. Our findings suggest that during nocturnal and crepuscular periods, visual cues provided by resting or slow-moving fish are not alone sufficient to attract gnathiids, while olfactory cues alone are. The traps designed for this study offer a new method of sampling free-living gnathiid isopods.     

Female sex pheromone in immature insect males—a case of pre-emergence chemical mimicry?

Sexual selection by competition for mates is a formidable force that has led to extraordinary adaptations in males. Here we present results suggesting a novel case of pheromone mimicry in males of Lariophagus distinguendus, a parasitic wasp of beetle larvae that develop in stored grain. Females of L. distinguendus produce a pheromone even before they emerge from a grain. Males are attracted to the parasitised grain and wait for females to emerge. Males emerging later than others are under enormous selection pressure since females mate only once. We show evidence that developing males fool their earlier emerging competitors by mimicking the female pheromone. Males exposed to pupae of either sex exhibit typical courtship behaviour. Searching males are not only arrested by grains containing developing females but spend as much time on grains containing developing males. Hence, by distracting their competitors away from receptive females late males may increase their own chance to mate with these females. After emergence, males decompose the active compounds within 32 h probably to decrease molestation during their own search for mates. Chemical analyses of active pheromone extracts and bioassays using fractions demonstrate that the active compounds are among the cuticular hydrocarbons.     

Weapons or mating signals? Claw shape and mate choice in a fiddler crab

Males often possess traits, such as horns, claws, and tusks, which are used during male–male combat. Studies suggest that selection has fine tuned these weapons to improve their effectiveness and that the shape of these weapons provides cues for males to assess the strength of rivals. Similarly, females might also assess a male’s weaponry to determine his value as a mate. The largest weapon relative to body size is the major claw of male fiddler crabs (genus: Uca). Males use their claws both as a weapon and as a courtship signal, waving it to attract females. We used robotic crabs in two-choice experiments to test female mating preferences based on male claw shape in Uca mjoebergi. First, females did not prefer conspecific claws over any of three alternatively shaped stimuli matched for color and for the rate and pattern of waving. The alternative stimuli were two different heterospecific claws and a plain rectangular shape. Second, females significantly preferred the alternative stimuli when they were presented at a faster wave rate. We conclude that claw shape in U. mjoebergi has not evolved under sexual selection imposed by female mating preferences and is more likely to have evolved under selection for effectiveness during male–male competition.     

Mating expenditures reduced via female sex pheromone modulation in the primitively eusocial halictine bee, Lasioglossum (Evylaeus) malachurum (Hymenoptera: Halictidae)

The present paper reports on behavioral experiments and gas chromatographic analysis of chemical communication in the mating biology of the primitively eusocial sweat bee Lasioglossum (Evylaeus) malachurum. In a dual-choice experiment, a female made odorless was significantly less attractive than an untreated one. Attraction in L. (Evylaeus) malachurum is therefore mediated by a female-produced sex pheromone. Further bioassays showed that unmated gynes are more attractive to males than mated ones. Males are able to differentiate between the two groups of females as little as 3 h after mating. Biotests with different samples obtained from attractive gynes showed surface extracts to be most attractive. Behavioral tests with synthetic copies of the compounds identified as cuticular constituents of virgin gynes were highly attractive to males; the volatile bouquets consisting of n-alkanes, n-alkenes and iso- pentenyl esters of unsaturated fatty acids were the most attractive samples. Isopentenyl esters of unsaturated fatty acids were the key compounds in inducing male inspections as well as stimulating pounces and copulatory attempts. Virgin and nesting gynes differed clearly in the relative and absolute amounts of the volatiles on the cuticle. The total amount of volatiles was significantly higher in virgin gynes and decreased in breeding queens. Hydrocarbons were the dominant group of compounds in both groups of females. The relative amounts of the wax-type ester, hexadecyl oleate, iso- pentenyl esters, and a hitherto unidentified steroid were higher in attractive virgin gynes, while the relative proportions of hydrocarbons and lactones dominated in nesting queens. The site of sex pheromone production in attractive young L. (Evylaeus) malachurum gynes remains unknown. Head glands or Dufour's gland secretions may be involved. Another possible source of the `active principle' found among the cuticular lipids could be glandular cells of the epidermis. The significance of modulation of female sex pheromone composition is discussed in terms of a reduction in mating expenditures. Received: 30 April 1998 / Accepted after revision: 24 July 1998     

Storage and display of odour by male Saccopteryx bilineata (Chiroptera, Emballonuridae)

Males of the sac-winged bat, Saccopteryx bilineata, actively fill their propatagial sacs with secretions from the genital region, the gular gland, urine and saliva. From our observations and those of Starck we deduce that propatagial sacs in S. bilineata do not have a glandular function, but are instead organs for the storage and display of odours. In addition to the already known “salting” and hovering behaviour of male S. bilineata, we describe in detail how odour is fanned to roosting individuals during the complex, stereotypic hovering displays. S. bilineata males also coat the fur of their backs with saliva using the wing tip and might scent-mark territory boundaries. “Yawning” may represent a visual as well as an olfactory cue. Odour seems to play an important role in the social communication of S. bilineata and in other emballonurids, as revealed by the broad distribution of wing sacs in this family. S. bilineata males display odour during energetically costly hovering flights in front of females. We demonstrate that the number of hovering displays increases with harem size. The mating effort of S. bilineata males comprises a multimodal signalling behaviour. Although males defend harem territories in which females gather, females seem to be able to choose the father of their progeny freely among the males of a colony. This may have led to the evolution of the complex mating displays by male S. bilineata. Received: 9 December 1998 / Received in revised form: 6 May 1999 / Accepted: 13 June 1999     

Whitefly wax as a cue for phoresy in the broad mite, <Emphasis Type="Italic">Polyphagotarsonemus latus</Emphasis>(Acari: Tarsonemidae)

Summary. Broad mite, Polyphagotarsonemus latus (Acari: Tarsonemidae) exhibits a specific phoretic relationship with whiteflies. Under field conditions most broad mites, caught in sticky traps, are attached to whiteflies. Under laboratory conditions, attachment occurs equally well in the dark and light. Mites do not differentiate between the sexes of their phoretic host Bemisia tabaci. However, mite attachment to B. tabaci is greatly diminished by washing the host with various organic solvents, chloroform in particular. The effect of whitefly waxy particles on broad mite behavior was studied using wax from the whitefly Aleyrodes singularis and from the mealybug Planococcus citri. Broad mites were not only attracted specifically to the A. singularis waxy particles-treated leaf areas but were also attached to leaf trichomes in this area. The results of this study suggests the importance of olfactory cues from the whitefly waxy particles in the recognition process of the phoretic host and/or the induction of the attachment behavior to whitefly legs or leaf trichomes.     

Olfactory cues,genetic relatedness and female mate choice in the agile antechinus (<Emphasis Type="Italic">Antechinus agilis</Emphasis>)

Females show mate preferences for males that are genetically dissimilar to themselves in a variety of taxa, but how females choose these males is not clearly understood. In this study, we examined the effects of olfactory stimuli and genetic relatedness on female mate choice in a small carnivorous marsupial, the agile antechinus (Antechinus agilis), during two breeding seasons. Captive female antechinus in oestrus were provided with a combination of male urine and body scent from two novel males, one more genetically similar and one more dissimilar to the females, in a Y-maze olfactometer. Genetic relatedness between females and pairs of males was determined using highly polymorphic, species-specific, microsatellite markers. Females consistently chose to visit the scents of males that were genetically dissimilar to themselves first, spent significantly more time near the source of those scents and showed more sexual and non-exploratory behaviours near those scents. These data demonstrate that chemosensory cues are important in mate choice in the agile antechinus and that females prefer males that are genetically dissimilar to themselves.