The cost of sexual coercion and heterospecific sexual harassment on the fecundity of a host-specific, seed-eating insect (Neacoryphus bicrucis)

Two species of seed-eating true bugs, Neacoryphus bicrucis (Heteroptera: Lygaeidae) and Margus obscurator (Heteroptera: Coreidae) co-occur on ragwort, Senecio tomentosus, in southern Georgia, USA. Males of both species sometimes engage in chases and protracted grappling with females that flee initial mountings. Sometimes genital coupling occurs while the wriggling female is restrained in the male's grasp. Chases, grappling, and mounting attempts are misdirected toward heterospecific females, heterospecific males, or conspecific males. In a laboratory study, confinement of mated N. bicrucis females with either conspecific or heterospecific males reduced fecundity by approximately one-half relative to mated females confined only with other females. Perhaps as a consequence of this, N. bicrucis females frequently leave areas of high host plant density, where they prefer to oviposit, when males are abundant. The abundance of each species is positively correlated with host plant density but the two species rarely occur together on the same plants. This may be an effect of heterospecific courtship which induces the flight of N. bicrucis more than the flight of M. obscurator. The laboratory results suggest that copulations following chases and grappling represent sexual harassment, not a mechanism of active female choice for a vigorous mate. As sexual harassment imposes high fitness costs that favor abandonment of host plants, it may, when misdirected, incidentally limit habitat use by ecologically similar species. Received: 3 October 1998 / Received in revised form: 21 January 1999 / Accepted: 14 February 1999     

Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal

Females often base their mating preferences on male sexual secondary traits that are used to settle contests among males. Such traits are likely to be honest indicators of male quality if they are constantly used during costly male–male agonistic interactions. Carotenoid signals have been shown to work as a handicap because they are costly to produce. However, the role of carotenoids as “honest” signals during male contests is less clear, and it is not known whether a carotenoid-based trait can serve in both male–male competition and female choice. In this study, we studied the dual function of a carotenoid feather ornament in the rock sparrow (Petronia petronia), a bird species in which both sexes have a yellow throat patch whose size positively correlates with phenotypic measures. First, we investigated, in a field study, whether the size of a male’s yellow patch correlates with his ability to acquire a territory. Second, we tested the signal function of the yellow patch in two male–male interaction in captivity experiments. Finally, we measured female preference for males differing in throat patch size in a mate choice experiment. Our experiments revealed that the size of a male’s throat patch positively correlated with the number of nest boxes he was able to defend. Moreover, in controlled conditions, males with relatively large yellow patches had earlier access to food than those with small patches. Also, in an experiment in which a dummy rock sparrow with an experimentally manipulated yellow patch was positioned near a feeder, latency to feed by focal birds positively correlated with dummy patch size. Lastly, in a dichotomous mate choice experiment, females showed a proximity preference for males whose patch was experimentally enlarged. Taken together, these results suggest that the same carotenoid feather signal may be used in both male–male competition and female choice in this passerine bird.     

Elytra color as a signal of chemical defense in the Asian ladybird beetle Harmonia axyridis

Carotenoid- and melanin-based colors are valuable indicators of quality in many vertebrates, but their signaling role in invertebrates remains relatively unexplored. The Asian ladybird beetle Harmonia axyridis is an ideal organism for studies of this nature because males and females exhibit highly variable red and black colors on their elytra and are chemically defended with an alkaloid (harmonine). We used digital photography to quantify elytra coloration and absorbance spectrophotometry and gas chromatography mass spectrometry analyses to quantify pigment and alkaloid concentrations, respectively, in wild-caught male and female H. axyridis. We predicted that extensive or intense coloration would be an aposematic signal of high-alkaloid stores. We found that carotenoid pigments largely controlled variation in red elytra coloration. There was no relationship between alkaloid content and either elytra redness or carotenoid pigment concentration in either sex. However, we found a positive correlation between the extent of elytra coloration and alkaloid content. Animals with proportionally more red (or less black) on the elytra were more alkaloid-rich; this relationship was particularly strong in females. We also found that females with lighter black spots had greater amounts of harmonine than those with darker spots. These results suggest that elytra color patterns have the potential to reveal information about chemical defensiveness to mates or predators. Prior studies in this species show that nonmelanic forms are typically less active and yet more sexually attractive than melanic forms, and both results may be explained by the fact that nonmelanics are better chemically defended.     

The honey bee shaking signal: function and design of a modulatory communication signal

This study explores the meaning and functional design of a modulatory communication signal, the honey bee shaking signal, by addressing five questions: (I) who shakes, (II) when do they shake, (III) where do they shake, (IV) how do receivers respond to shaking, and (V) what conditions trigger shaking. Several results confirm the work of Schneider (1987) and Schneider et al. (1986a): (I) most shakers were foragers (at least 83%); (II) shaking exhibited a consistent temporal pattern with bees producing the most signals in the morning (0810–1150 hours) just prior to a peak in waggle dancing activity; and (IV) bees moved faster (by 75%) after receiving a shaking signal. However, this study differs from previous work by providing a long-term, temporal, spatial, and vector analysis of individual shaker behavior. (III) Bees producing shaking signals walked and delivered signals in all areas of the hive, but produced the most shaking signals directly above the waggle dance floor. (IV) Bees responded to the signal by changing their direction of movement. Prior to receiving a signal, bees selected from the waggle dance floor moved, on average, towards the hive exit. After receiving a signal, some bees continued moving towards the exit but others moved directly away from the exit. During equivalent observation periods, non-shaken bees exhibited a strong tendency to move towards the hive exit. (V) Renewed foraging activity after food dearth triggered shaking signals, and, the level of shaking is positively correlated with the duration of food dearth. However, shaking signal levels also increased in the morning before foraging had begun and in the late afternoon after foraging had ceased. This spontaneous afternoon peak has not previously been reported. The shaking signal consequently appears to convey the general message “reallocate labor to different activities” with receiver context specifying a more precise meaning. In the context of foraging, the shaking signal appears to activate (and perhaps deactivate) colony foraging preparations. The generally weak response elicited by modulatory signals such as the shaking signal may result from a high receiver response threshold which allows the receiver to integrate multiple sources of information and which thereby increases the probability that receiver actions will be appropriate to colony needs. Received: 21 March 1997 / Accepted after revision: 30 August 1997     

Egg coloration in ring-billed gulls (<Emphasis Type="Italic">Larus delawarensis</Emphasis>): a test of the sexual signaling hypothesis

Although many avian eggs appear to be cryptically colored, many species also lay vibrant blue green eggs. This seemingly conspicuous coloration has puzzled biologists since Wallace, as natural selection should favor reduced egg visibility to minimize predation pressure. The sexual signaling hypothesis posits that blue green egg coloration serves as a signal of female quality and that males exert post-mating sexual selection on this trait by investing more in the nests of females laying more intensely blue green eggs. This hypothesis has received mixed support to date, and most previous studies have been conducted in cavity-nesting species where male evaluation of his partner’s egg coloration, relative to that of other females, may be somewhat limited. In this study, we test the sexual signaling hypothesis in colonially nesting ring-billed gulls (Larus delawarensis) where males have ample opportunity to assess their mate’s egg coloration relative to that of other females. We used correlational data and an experimental manipulation to test four assumptions and predictions of the sexual signaling hypothesis: (1) blue green pigmentation should be limiting to females; (2) extent of blue green egg coloration should relate to female quality; (3) extent of blue green egg coloration should relate to offspring quality; and (4) males should provide more care to clutches with higher blue green chroma. Our data provide little support for these predictions of the sexual signaling hypothesis in ring-billed gulls. In light of this and other empirical data, we encourage future studies to consider additional hypotheses for the evolution of blue green egg coloration.     

The modulation of worker behavior by the vibration signal during house hunting in swarms of the honeybee, Apis mellifera

During house hunting, honeybee, Apis melli- fera, workers perform the vibration signal, which may function in a modulatory manner to influence several aspects of nestsite selection and colony movement. We examined the role of the vibration signal in the house-hunting process of seven honeybee swarms. The signal was performed by a small proportion of the older bees, and 20% of the vibrating bees also performed waggle dances for nestsites. Compared to non-vibrating controls, vibrating bees exhibited increased rates of locomotion, were more likely to move into the interiors of the swarms, and were more likely to fly from the clusters and perform waggle dances. Recipients responded to the signal with increased locomotion and were more likely than non- vibrated controls to fly from the swarms. Because vibration signals were intermixed with waggle dances by some vibrators, and because they stimulated flight in recipients, the signals may have enhanced nestsite scouting and recruitment early in the house-hunting process. All swarms exhibited increased vibration activity within 0.5–1 h of departure. During these final periods, numerous vibrating bees wove repeatedly in and out of the clusters while signaling and motion on the swarms increased until it culminated in mass flight. The peaks of vibration activity observed at the end of the house-hunting process may therefore have activated the entire swarm for liftoff once a new nestsite had been selected. Thus, the vibration signal may help to integrate the behavior of numerous groups of workers during nestsite selection and colony relocation. Received: 17 January 2000 / Received in revised form: 5 April 2000 / Accepted: 3 May 2000     

The effect of repeated vibration signals on worker behavior in established and newly founded colonies of the honey bee, <Emphasis Type="Italic">Apis mellifera</Emphasis>

Communication signals used in animal social interactions are frequently performed repetitively, but the function of this repetition is often not well understood. We examined the effects of signal repetition by investigating the behavior of worker honey bees that received differing numbers of vibration signals in established and newly founded colonies, which could use signal repetition differently to help adjust task allocations to the labor demands associated with the different stages of colony development. In both colony types, more than half of all monitored workers received more than one vibration signal, and approximately 12% received ≥5 signals during a given 20-min observation period. Vibrated recipients exhibited greater activity and task performance than same-age non-vibrated controls at all levels of signal activity. However, vibrated workers showed similar levels of task performance, movement rates, cell inspection rates, and trophallactic exchanges regardless of the number of signals received. Thus, the repeated performance of vibration signals on individual bees did not cause cumulative increases in the activity of certain workers, but rather may have functioned to maintain relatively constant levels of activity and task performance among groups of recipients. The established and newly founded colonies did not differ in the extent to which individual workers received the different numbers of vibration signals or in the levels of activity stimulated by repeated signals. Previous work has suggested that compared to established colonies, newly founded colonies have a greater number of vibrators that perform signals on a greater proportion of the workers they contact. Taken in concert, these results suggest that vibration signal repetition may help to adjust task allocations to the different stages of colony development by helping to maintain similar levels of activity among a greater total number of recipients, rather than by eliciting cumulative effects that cause certain recipients to work harder than others.     

The role of the vibration signal in the house-hunting process of honey bee (<Emphasis Type="Italic">Apis mellifera</Emphasis>) swarms

The function of the vibration signal of the honey bee (Apis mellifera) during house hunting was investigated by removing vibrating bees from swarms and examining the effects on waggle dancing for nest sites, liftoff preparations and swarm movement. We compared house hunting among three swarm types: (1) test swarms (from which vibrating bees were removed), (2) manipulated control (MC) swarms (from which randomly selected workers and some waggle dancers were removed), and (3) unmanipulated control (UC) swarms (from which no bees were removed). The removal of vibrating bees had pronounced effects on liftoff preparations and swarm movement. Compared to the MC and UC swarms, the test swarms had significantly greater liftoff-preparation periods, were more likely to abort liftoff attempts, and in some cases were unable to move to the chosen site after the swarm became airborne. However, the three swarm types did not differ in overall levels of waggle dance activity, the time required to achieve consensus for a nest site, the rate at which new waggle dancers were recruited for the chosen site, or the ability to maintain levels of worker piping necessary to prepare for flight. The removal of vibrating bees may therefore have altered liftoff behavior because of a direct effect on vibration signal activity. A primary function of the signal during house hunting may be to generate a level of activity in workers that enhances and coordinates responses to other signals that stimulate departure and movement to a new location.Communicated by R. Page     

The habitat saturation hypothesis and sociality in an allodapine bee: cooperative nesting is not “making the best of a bad situation”

Several factors thought to be important for the evolution of cooperative breeding in vertebrates have received little attention in facultatively social insects. One of these, the “habitat saturation hypothesis” of Selander (1964), predicts that colony sizes will be greater in breeding units where dispersal opportunities are limited, suggesting that group living is a secondary option to independent reproduction. The Australian allodapine bee Exoneura bicolor exhibits a number of traits that occur in cooperatively breeding bird species, including long life-span, repeated opportunities for reproduction, and vulnerability to brood predation and parasitism. We experimentally examined the effect of a potentially limiting environmental factor, nesting substrate availability, as an agent influencing sociality in E. bicolor. We manipulated nesting substrate availability in two separate locations during a time when foundress dispersal is common. No significant difference was found between colony sizes in cases where dispersal options were abundant and cases where dispersal options were limited. An increase in opportunities for dispersal did not lead to higher rates of independent nesting, suggesting that cooperative nesting is a preferred strategy regardless of distance-related costs of dispersal. Reproductivity per female and brood survival were examined as factors selecting for group living. Low survival of brood in single-female nests has the potential to select for cooperative nesting in this bee. Received: 29 September 1995/Accepted after revision: 24 June 1996