The sex pheromone of the grain aphid,Sitobion avenae (Fab.) (Homoptera,Aphididae)

Summary The sex pheromone of the grain aphid,Sitobion avenae (Fab.), is identified as (4aS,7S,7aR)-nepetalactone, for which specific olfactory cells are located in the secondary rhinaria of the male antenna. The related (1R,4aS,7S,7aR)-nepetalactol was not produced by females in significant amounts (ca 10,000-fold less than the nepetalactone), although cells in the male antenna were found that responded specifically to this compound. Behavioural activity in the linear track ol-factometer showed attraction of males to the nepetalactone, but not to the nepetalactol.     

Family insurance: kin selection and cooperative breeding in a solitary primate (Microcebus murinus)

Lactation imposes substantial physiological costs on mothers and should therefore not be directed towards foreign offspring. Such allonursing, however, is common in mammal species that share roosts. Hypotheses to explain allonursing among such plural breeders include misdirected parental care, milk evacuation, brood parasitism, reciprocity, and kin selection. The necessary behavioral data, in combination with data on kinship and kin recognition, have rarely been available to distinguish among these explanations, however. In this study, we provide evidence for cooperative nursing and adoption by plural-breeding females in a nocturnal primate, the gray mouse lemur (Microcebus murinus), in which females forage solitarily during the night, but form day-time sleeping groups with one to two other females. We observed 34 resident females in an 8 ha study area in Kirindy Forest, Madagascar, over three consecutive annual breeding seasons and determined genetic relationships among all members of this population. Five sleeping groups of adult females were filmed inside their roosts during one breeding season after females gave birth. The composition of groups changed substantially across years, but they always consisted of close maternal relatives. All females within a group gave birth to one to three infants. They regularly transferred only their own offspring among roosting sites, demonstrating an ability to discriminate between their own and other’s offspring, but they regularly groomed and nursed related offspring other than their own and adopted related dependent young after their mother’s death. Kin selection may therefore be the main selective force behind cooperative breeding among these closely related females with a high mortality risk, providing each of them with family insurance.     

Worker discrimination among queens in newly founded colonies of the fire ant Solenopsis invicta

Although colonies of the fire ant Solenopsis invicta are often founded by small groups of queens, all but one of the queens are soon eliminated due to worker attacks and queen fighting. The elimination of supernumerary queens provides an important context for tests of discrimination by the workers, since the outcome of these interactions strongly affects the workers' inclusive fitness. To test whether workers in newly founded colonies discriminate among nestmate queens, paired cofoundresses were narrowly separated by metal screens that prevented direct fighting, but through which the workers could easily pass. Soon after the first workers completed development, they often attacked one of the queens; these attacks were strongly associated with queen mortality. When one queen's brood was discarded, so that the adult workers were all the daughters of just one queen, the workers were significantly less likely to bite their mother than the unrelated queen; however, this tendency was comparatively weak. Queens kept temporarily at a higher temperature to increase their rate of investment in brood-rearing lost weight more rapidly than paired queens and were subsequently more likely to be attacked and killed by workers. Workers were more likely to bite queens that had been temporarily isolated than queens that remained close to brood and workers. When queens were not separated by screens, the presence of workers stimulated queen fights. These results show that workers discriminate strongly among equally familiar queens and that discrimination is based more on the queens' condition and recent social environment than on kinship. Received: 9 June 1998 / Accepted after revision: 10 October 1998     

Kinship and food availability influence cannibalism tendency in early-instar wolf spiders (Araneae: Lycosidae)

For potentially cannibalistic animals such as spiders, the ability to recognize and avoid kin and/or preferentially cannibalize non-relatives would permit exploiting conspecifics as prey while minimizing loss of inclusive fitness. We investigated the effects of relatedness and availability of alternative food on cannibalism tendency in pairs of juvenile Hogna helluo (Walckenaer), a North American wolf spider (Araneae: Lycosidae). For second-instar spiderlings (dispersing stage), cannibalism was more likely among pairs of non-sibs than pairs of sibs and, interestingly, was also more likely when other prey were available. We found no evidence of increased cannibalism in pairings involving broods of greatest average size disparity, indicating that size differences are unlikely to explain differences in cannibalism tendency. Additionally, the relative number of deaths from cannibalism or other causes did not increase with increasing risk of starvation. For third-instar spiderlings, which had lived independently of their mother and sibs following dispersal, cannibalism rates were very high in all treatments and there were no significant effects of relatedness or food availability. Our results suggest that spiders with predominantly solitary lifestyles may bias cannibalism toward non-kin during the juvenile associative period, and that this effect is lost in the subsequent instar. Results are discussed in the context of several potential mechanisms that might result in differential cannibalism.Communicated by M. Elgar     

Benefits of kin association: related and familiar zebrafish larvae (<Emphasis Type="Italic">Danio rerio</Emphasis>) show improved growth

Zebrafish (Danio rerio) larvae prefer the olfactory cues of kin to non-kin. We examined the potential benefits of kin preference by comparing growth rate, shoaling, and aggressive behavior in juvenile zebrafish housed in groups of either familiar kin or unfamiliar non-kin. Over an observation period of 5 days, the animals grew 33% more in kin groups; however, neither shoaling nor the frequency of aggressive interactions was different in groups of related versus unrelated individuals. Shoaling behavior increased with increasing observation time and increasing age, while aggressive behavior remained the same. We conclude that associating with kin probably creates a less stressful environment that allows for higher growth rates, which can lead to higher direct fitness based on increased survival and earlier reproduction. Kin recognition leading to kin-structured groups may therefore be under positive selection.     

Sex pheromone of the tea aphid,Toxoptera aurantii (Boyer de Fonscolombe) (Hemiptera: Aphididae)

The tea aphid, Toxoptera aurantii, also called the “black citrus aphid”, is one of the most destructive insect pests in commercial tea plantations and gardens in southern China. In autumn, declining day length triggers production of winged T. aurantii sexuparae, which produce both winged males and wingless oviparae. Oviparous females then release sex pheromone that attracts potential mates. GC–MS analysis of volatile headspace extracts of T. aurantii oviparae revealed that they emit (4aS,7S,7aR)-nepetalactone (I) and (1R,4aS,7S,7aR)-nepetalactol (II) in a ratio of 4.3–4.9:1. Field-trapping experiments with synthetic I and II singly or as two-component blends of different doses and ratios showed significant attraction of T. aurantii males, as well as weak attraction of sexuparae. Identification of the T. aurantii sex pheromone provides a new opportunity for developing a pheromone-based monitoring and management strategy for the sexual phase of tea aphids and, possibly, the alate sexparous generation in late summer and fall.     

Paternal kin bias in the agonistic interventions of adult female rhesus macaques (Macaca mulatta)

When agonistic interventions are nepotistic, individuals are expected to side more often with kin but less often against kin in comparison with non-kin. As yet, however, few mammal studies have been in a position to test the validity of this assertion with respect to paternal relatedness. We therefore used molecular genetic kinship testing to assess whether adult female rhesus macaques (Macaca mulatta) from the free-ranging colony of Cayo Santiago (Puerto Rico) bias their interventions in ongoing dyadic aggressive interactions towards maternal and paternal half-sisters compared with unrelated females. It turned out that females supported maternal half-sisters significantly more often than paternal half-sisters or non-kin regardless of the costs associated with such interventions. Similarly, females targeted maternal half-sisters significantly less often than non-kin when this was associated with high costs. Unrelated females provided significantly higher mean rates of both high- and low-cost support to each other than did paternal half-sisters. However, females targeted paternal half-sisters significantly less often than non-kin when targeting was at low cost, suggesting that females refrain from intervening against paternal half-sisters. Our data confirm the general view that coalition formation in female mammals is a function of both the level of maternal relatedness and of the costs of intervention. The patterns of coalition formation among paternal kin were found to be more complex, and may also differ across species, but clear evidence for paternal kin discrimination was observed in female rhesus as predicted by kin selection theory.     

Costs and benefits of communal rearing to female prairie voles (<Emphasis Type="Italic">Microtus ochrogaster</Emphasis>)

Communal rearing of offspring may help mothers maximize their investment in offspring at a reduced cost to their own bodily condition, thus maximizing their potential for reproductive success. The objective of this study was to quantify the costs and benefits of communal rearing to prairie vole (Microtus ochrogaster) pups and mothers. Mothers were assigned to one of three social units: solitary mothers, singularly breeding groups (i.e. one mother and one non-reproductive sister) and plurally breeding groups (i.e. two lactating sisters). For each type of social unit, some replicates were provided with food ad libitum, while others were provided with limited food. The body mass of focal mothers (i.e. the first mother to produce a litter) was a significant predictor of pup growth. Regardless of food availability, litters of focal mothers in plurally breeding groups gained more weight than litters reared by solitary mothers. Pups reared in singularly breeding groups were intermediate in weight gain, but did not gain significantly more weight than solitary offspring. There was no difference in the body mass of focal mothers from each type of social unit, regardless of food availability. Within plurally breeding groups, the weight gain of the two litters and body mass of focal and second mothers did not differ. However, focal mothers from plurally breeding groups nursed fewer pups than solitary mothers and also fewer pups than their nestmates when food was limited. Our results suggest that plural breeding results in greater fitness to mothers than solitary and singular breeding.Communicated by E. Korpimäki     

Individual roosting strategies in a flock-living bird: movement and social cohesion of hooded crows (Corvus corone cornix) from pre-roost gatherings to roost sites

The function of the conspicuous pre-roost gathering in communally roosting birds is poorly known. We studied movement and social cohesion of radio-tagged hooded crows (Corvus corone cornix) from their daytime location via pre-roost to roost in autumn and winter. With increasing snowfall the previous 24 h, the crows attended pre-roosts farther from the territory, and moved longer from pre-roost to roost. The crows became more likely to roost communally as distance to their territory increased. Attending a pre-roost on average almost doubled the travel distance to a communal roost. Crows were much more likely to join the same roost when they had attended the same pre-roost than otherwise. Breeding mates were more likely to keep together from pre-roost to roost than were other assigned pairs of a territorial male and female. For assigned pairs of non-mates, cohesion from pre-roost to roost decreased with increasing distance to roost, and was higher when both crows roosted communally compared to when at least one of them roosted in its territory. When both roosted communally, cohesion decreased with increasing snowfall and increasing number of crows attending the pre-roost, increased with increasing snow depth, and became higher among two females, compared to other combinations of two crows, with increasing number of crows attending the pre-roost. These patterns may be interpreted as supporting several of the current hypotheses on the function of pre-roost gathering. Electronic Publication     

Kin discrimination in the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>): nepotism among siblings

We examined patterns of affiliation, association, and aggression to inquire whether spotted hyenas (Crocuta crocuta) can distinguish among various groups of maternal and paternal siblings. If so, and if these animals conform to predictions of kin selection theory, then behavioral interactions among hyenas should vary with relatedness. We also considered familiarity-based recognition and phenotype matching as mechanisms hyenas might use to recognize kin. Patterns of affiliative behavior indicated that hyenas favored full-sibling littermates over half-sibling littermates or any other group of half-siblings. Rates of dyadic aggression generally did not vary with kinship. Hyenas associated more closely with half-sibling littermates than with non-littermate half-siblings, and hyenas affiliated more with maternal half-siblings than with paternal half-siblings, suggesting that familiarity-based cues might mediate discrimination among these sibling classes. In addition, operation of a phenotype-matching mechanism was suggested by the preference hyenas demonstrated during affiliative interactions for full- over half-sibling littermates, and by their lack of preference in these interactions for half-sibling littermates over non-littermate half-siblings. Phenotype matching was also suggested by our observation that paternal half-siblings cooperated more, and fought less, than did non-kin. Our data indicate that hyenas can discriminate among various types of siblings, that their social behavior conforms to predictions of kin selection theory, and that they recognize kin using mechanisms of both familiarity and phenotype matching.Communicated by S. Alberts     

Provisioning behavior and the estimation of investment ratios in a solitary bee,Calliopsis (Hypomacrotera) persimilis (Cockerell) (Hymenoptera: Andrenidae)

Summary One aspect of behavioral ecology that has received considerable attention, especially by students of social insects, is the relative amount of energy invested by parents in the rearing of male versus female offspring. Sexual selection theory makes predictions about how individuals should allocate their total investment in the sexes. To test these predictions we must accurately quantify the relative cost incurred by a parent in the production of an average individual of either sex. Body weight ratios are the most common estimate of cost ratio, but the correspondence between offspring body weight and energetic investment on the part of the parent has rarely been determined. Calliopsis (Hypomacrotera) persimilis is a solitary, ground-nesting bee whose natural history makes it particularly convenient for studies of investment patterns and foraging behavior. Each day females construct and provision from 1 to 6 cells in linear, closely-spaced series. Each cell is provisioned with pollen from Physalis Wrightii flowers, which is collected on two or three foraging trips. However, the temporal sequence in which two- and three-trip foraging bouts occur is not random. Females invariably begin each day provisioning cells with three trips worth of pollen and usually switch to provisioning the latter cells of the day with just two trips worth of pollen. The sex of the offspring within the same co-linear series of cells also varies non-randomly — female offspring predominate in the first cells of each series and male offspring in the latter cells. The correspondence between the number of foraging trips to provision a cell, the total time spent foraging, and offspring sex was determined for 36 cells. The data indicate a close, though not absolute, relationship between the number of foraging trips and the sex of the offspring: males usually received two trips of pollen, though some received three, whereas female offspring invariably received three trips worth of pollen. A number of potential estimates of the relative cost of female and male offspring production were calculated. Estimates of the cost ratio based on the amount of time spent foraging, adult dry body weight, and pollen ball dry weight all give similar values. Female offspring receive an energetic investment of from 1.3 to 1.5 times that of males. These results support the use of adult dry body weight ratios in the estimation of cost ratios.     

The effects of feeding by the black bean aphidAphis fabae Scop. (Homoptera: Aphididae) on copper and cadmium accumulation in broad bean (Vicia faba L.)

Broad beans (Vicia faba L.), cultured hydroponically were supplied with 100 g mL^–1 copper or 50 g mL^–1 cadmium in nutrient solution. Samples of plant material from both nutrient regimes were analysed before and after infestation by the black bean aphid (Aphis fabae Scop.). Heavy aphid infestation resulted in a significant reduction in copper content of shoots in comparison with uninfested plants. A similar, but less well- defined, situation occurred in the case of cadmium.Further investigations examined the effects of different levels of aphid infestation on the above phenomena. In all cases the presence of feeding aphids reduced elemental accumulation in plant shoots. Long term infestation with population densities as low as three adult aphids showed a reduction in shoot copper and cadmium content.     

Behavioural evidence for the involvement of Dufour’s gland secretion in nestmate recognition in the social wasp Polistes dominulus (Hymenoptera: Vespidae)

The results of recent studies by our group have suggested that in social wasps of the genus Polistes the Dufour’s gland is involved in kin recognition. In fact, the same hydrocarbons occurring on the cuticle are found in the gland secretion, and in P. dominulus the composition of the glandular secretion is more similar in foundresses belonging to the same colonies than in heterocolonial foundresses. In this study, P. dominulus colonies were experimentally presented with previously treated dead conspecific females. These lures had been deprived of their epicuticular lipids and coated with epicuticular or Dufour’s gland secretion extracts from females hetero- or homocolonial with respect to the tested colonies. The behaviour of the colonies towards these lures indicates that, like the epicuticular lipids, the Dufour’s gland secretion is involved in nestmate recognition. Received: 14 August 1995/Accepted after revision: 5 November 1995     

Social relationships among adult female baboons (Papio cynocephalus) II. Variation in the quality and stability of social bonds

A growing body of evidence suggests that social bonds have adaptive value for animals that live in social groups. Although these findings suggest that natural selection may favor the ability to cultivate and sustain social bonds, we know very little about the factors that influence the quality or stability of social bonds. Here, we draw on data derived from a 16-year study of baboons living in seven different social groups in the Amboseli basin of Kenya to evaluate the quality and stability of social bonds among females. Our results extend previous analyses, which demonstrate that females form the strongest bonds with close maternal and paternal kin, age mates (who may be paternal kin), and females who occupy similar ranks but are not maternal relatives. Here we show that the same factors influence the quality and strength of social bonds. Moreover, the results demonstrate that the quality of social bonds directly affects their stability.     

Divergence in female calling song discrimination between sympatric and allopatric populations of the southern wood cricket Gryllus fultoni (Orthoptera: Gryllidae)

Two species of closely related wood cricket, Gryllus fultoni (Orthoptera: Gryllidae) and Gryllus vernalis, occur together in some parts of the eastern United States and have a similar calling song structure, consisting of three-pulse chirps. A previous study revealed that chirp rate in G. fultoni was highest (greatest difference vis-à-vis chirp rate in G. vernalis) in sympatric populations, intermediate in near allopatric populations that were located close to the sympatric zone, and lowest in allopatric populations. A similar trend was observed in pulse rate, but the mean values of this trait showed much more convergence than chirp rate at the low end of the range of calling temperatures. In this study, we investigated the song discrimination of females from sympatric and allopatric populations of G. fultoni at about 23°C, which is near the middle of the normal range of calling temperatures. We used both single-stimulus and two-stimulus playback experiments to learn if geographical differences in song preferences paralleled those in calling songs. Stimuli presented were representative of calling songs in three classes of G. fultoni populations (sympatric, near allopatric, and far allopatric), a calling song of G. vernalis, and three calling songs with parameter values that were intermediate with respect to those of the songs of far allopatric G. fultoni and G. vernalis. In the single-stimulus playbacks, females of all G. fultoni populations responded poorly if at all to the heterospecific stimulus. Females of sympatric and near allopatric populations responded poorly to all intermediate stimuli, but females of far allopatric populations frequently responded to these sounds. In the two-stimulus playbacks, females of sympatric and near allopatric populations generally discriminated against intermediate and heterospecific stimuli. However, females of far allopatric populations often did not discriminate against intermediate stimuli, whose characteristics resembled the calling songs of G. vernalis. The divergent pattern of female phonotactic discrimination between sympatric and far allopatric populations was thus generally congruent with the pattern of divergence in chirp and pulse rates and would be expected to significantly reduce heterospecific mating in sympatry. These geographical patterns of female song discrimination and male calling songs conform to a commonly used definition of reproductive character displacement.     

Intraspecific parasitism and split sex ratios in a monogynous and monandrous ant (Leptothorax nylanderi)

Sex ratios were bimodally distributed in a population of the monogynous and monandrous ant Leptothorax nylanderi during each of 3 study years. The population-wide investment ratios suggested worker control of sex allocation. Nest-level variation in the proportional investment in virgin queens was not affected by the presence or absence of a queen and only slightly by collecting year, but was correlated with nest size, total sexual investment and, unexpectedly, with differences in nestmate relatedness: small, low-investment nests and nests with several worker lineages produced male-biased sex ratios. Colonies containing several worker lineages arise from usurpation of mature colonies by unrelated founding queens and the fusion of unrelated colonies under strong nest site limitation. In contrast to facultatively polygynous and polyandrous species of social insects, where workers can maximize their inclusive fitness by adjusting sex ratios according to the degree of relatedness asymmetry, workers in mixed colonies of L. nylanderi do not benefit from manipulating sex allocation, as here relatedness asymmetries appear to be the same as in homogeneous colonies. Received: 7 December 1999 / Received in revised form: 29 February 2000 / Accepted: 13 March 2000     

Individual variation and prior experience affect the discrimination of a brood-parasite by its subsocial beetle host

In birds, host experience can modify response to parasites but nothing similar is known for insects. We studied two desert tenebrionid beetles, the subsocial host Parastizopus armaticeps and its obligate and ubiquitous brood-(clepto)parasite Eremostibes opacus, which mimics the hosts odour spectrum. Nearly 10% of host burrows in the field (n=214) remain unparasitised even after introducing E. opacus experimentally (n=22). In discrimination tests, 7% of naive host beetles eject the parasite, showing individual variation in olfactory discrimination ability. To test for effects of prior breeding experience, naive (first time) or experienced (second time) breeders with or without prior parasite contact were confined with E. opacus at breeding onset, their behaviour recorded and parasite number counted on breeding completion. More first- than second-time breeding pairs completely excluded the parasite and second-time breeders with no prior contact with E. opacus had fewer parasites than any others. These pairs increased burrow-guarding dramatically, mostly during early reproduction, this being the best predictor of parasite number. Prolonged guarding, however, must be traded-off against other brood-care behaviours. Hence, it might pay breeding pairs to take the risk of accepting some parasites breeding within their burrow, and we expect this effect will be more pronounced if poor feeding and digging conditions are taken into account. This might explain why apparently 90% of the P. armaticeps breeding burrows in the field contain at least one parasitic E. opacus.Communicated by D. Gwynne     

Prey chemical discrimination in ambush foragers: absence in representatives of two additional iguanian lizard families and probable olfactory mediation in a gekkonine gecko

Summary. Lingually mediated prey chemical discrimination in lizards has evolved in active foragers, been lost in taxa that have reverted to ambush foraging, and has not evolved in taxa that have retained the ancestral ambushing. Previous studies have shown that all families of insectivorous ambushers lack prey chemical discrimination, including most families of iguanian lizards and two gekkonid species. I conducted experimental studies of prey chemical discrimination in representatives of two additional iguanian families and a third gekkonid lizard. An oplurid species, Oplurus cuvieri and a corytophanid, Corytophanes cristatus, did not discriminate among prey chemicals and control substances. Prey chemical discrimination is now known to be absent in insectivorous ambush foragers in all but one of the families in Iguania, one of the two major lizard radiations. Hoplocercidae remains unstudied. Like other ambushing gekkonid lizards, Pachydactylus turneri did not exhibit elevated tongue-flick rates in response to prey chemicals. However, after tongue-flicking or being touched on the labial scales by cotton swabs, these lizards bit swabs bearing prey chemicals more frequently than control stimuli. They also exhibited buccal pulsing more frequently in response to prey chemicals than deionized water, suggesting olfactory sampling. The unusually highly developed olfactory organs of gekkonid lizards and their nocturnal habits suggest that olfaction may be more important to foraging than in other lizards. Further studies are needed to determine relative roles of olfaction and vomerolfaction in selective response to prey chemicals and to ascertain whether and to what extent the tongue may be used to locate and identify prey. Received 30 March 1999; accepted 26 July 1999     

Neighbor–stranger discrimination in Audubon's shearwater (Puffinus l. lherminieri) explained by a “real enemy” effect

Neighbor–stranger discrimination (NSD) occurs when animals respond with more aggression to strangers than to territorial neighbors. NSD has been reported in many species that defend multi-purpose territories for breeding and foraging, but it is rare among species that defend other types of territories. For birds that defend only their nest sites, there is no experimental evidence for NSD, and observational studies have provided mixed results. In a colony of Audubon's shearwaters (Puffinus l. lherminieri), I played back the calls of a neighbor and a stranger to males defending nest sites. Subjects responded with longer calls to playbacks of strangers than to those of neighbors. In shearwater colonies, strangers are often birds looking for future breeding sites. In contrast, there is no evidence that established breeders compete with their neighbors for any resources. Shearwaters should benefit from NSD because strangers represent a “real enemy” and established neighbors do not.     

Natural selection and the communal rearing of pups in African wild dogs (Lycaon pictus)

Summary The contributions of adult (>18 months) and yearling (10–18 months) African wild dogs (Lycaon pictus), to raising pups were quantified at eight natal dens, where pups remain for their first three months of life.Adults between 2–6 years of age did most of the hunting, and the dominant male often made the first grab at fleeing prey. Yearlings contributed to the hunting but were reluctant to tackle large prey animals. Yearlings and breeding females had prior access to food. Yearlings and adults regurgitated comparable amounts of food, but in one pack watched at a time of food shortage, the yearlings failed to regurgitate and stole food from the pups.Dominant dogs chased predators, especially spotted hyenas (Crocuta crocuta) from the area of the den more frequently than other adults or yearlings chased predators. Mothers, particularly in the first six weeks post-partum, stayed and protected the pups when the pack hunted.There is a positive, but non-significant correlation between the number of adult helpers and the number of pups raised. However, the sex ratio bias towards males at birth suggests that male helpers (which predominate) usually increase pup survivorship. The roles of direct and indirect selection in the evolution of the helping behavior are discussed. Indirect selection has probably played an important role in the unusual post-reproductive survival of males and their helping behavior.Serengeti Wildlife Research Institute contribution number 284