Sex, intimidation and severed limbs: the effect of simulated predator attack and limb autotomy on calling and emergence behaviour in the field cricket Gryllus bimaculatus

Crickets can autotomize a limb in order to evade predation; however, this autotomy compromises their escape speed and, therefore, their ability to avoid subsequent predation events. We recorded two measures of predator-avoidance behaviour in field crickets (Gryllus bimaculatus) exposed to varying levels of predation threat, with the most extreme level leading to autotomy of a limb. Our first measure of caution was time to emergence from cover, which was affected by perceived predation treatment, with both autotomized males and females being significantly more cautious than intact individuals. For males (but not females), the presence of a calling conspecific encouraged earlier emergence. Our second measure of caution was alteration of male calling behaviour. Autotomized males remained silent for significantly longer time after disturbance than intact males, but there was no difference in call rate once they had resumed calling, suggesting that behavioural changes were not merely a result of injury response but a controlled modification of behaviour as a result of autotomy. These data suggest that autotomy in field crickets results in altered calling behaviour, which was not significantly altered with different acoustic environments.     

The influence of kinship on foraging competition in Siberian jays

Foraging competition in Siberian jay groups was examined in relation to dominance and kinship to determine whether juvenile offspring, by associating with adults, gained in food acquisition relative to juvenile immigrants. Members of the adult pair were dominant over juvenile cohort members and males were dominant to females, although an inter-sexual hierarchy, with male juveniles occasionally overlapping adult females, was suggested. Few competitive asymmetries were found between adults and retained offspring or adults and immigrant juveniles when they were competing for food together, but in kin and non-kin foraging groups, respectively. Male offspring visited the bait site more frequently than adult males, and female immigrants spent less time at the bait site than adult females. Under these circumstances, hoarding activities may limit the ability of alpha members to control resources. In mixed groups containing both juvenile offspring and juvenile immigrants, no difference was found in the number of visits made to the bait site, although load sizes and foraging rates were lower for immigrant birds. Retained juveniles obtained greater load sizes and foraging rates when associating with adults. The social dominance of parents suggests that they control juvenile foraging. Although offspring benefit in the presence of adults, adults may incur a cost to their restraint by spending more time at the bait site when competing with immigrants. These results extend conclusions from previous work describing the role of selective tolerance by adults which relaxes competition with retained offspring in Siberian jay winter groups. The present findings suggest that offspring benefit in both immediate and future energy gains, which may have a direct influence on survival. Received: 18 September 1996 / Accepted after revision: 26 January 1997     

Reproductive biology of a small isopod symbiont living on a large isopod host: from the maternal marsupium to the protective grip of guarding males

Mating systems of many symbiotic crustaceans are characterised by a high degree of mate guarding. A peculiar case of mate guarding has been reported for small symbiotic janirid isopods where males mate with immature females. Field samples of individual hosts and laboratory experiments were conducted to reveal the mating behaviour of the symbiont in a natural environment, that is, on their hosts. Along the coast of the Magellan Strait, Chile, the janirid isopod Iais pubescens was frequently found on the shore-living isopod Exosphaeroma gigas. Symbiont prevalence (percent hosts occupied) was high at eight of the nine sampling sites. Mean symbiont intensity was very low at one site (<<1 individual host^-1), intermediate at two sites (1-10 individuals host^-1) and high at the other sites (10-40 individuals host^-1). The mean sex ratio (males:females) was male biased at most sampling sites (n=7). Females of I. pubescens reached substantially larger sizes (1.5-3.0 mm body length, BL) than males (1.1-1.9 mm BL). The majority of males were carrying small juveniles (66.15%), and males with juveniles were significantly larger than males without juveniles - this suggests that males prefer virgin juveniles to adult females and that they compete for small juveniles. In laboratory observations, males were seen to manipulate the marsupium of adult females that were about to release small juveniles. Males obtained virgin juveniles in this manner. Juveniles were carried for ~7 days, and they moulted shortly before being fertilised and released by males. The high proportion of juveniles carried by males in the field (68.2%) supports previous observations that males initially are not able to distinguish male and female juveniles. It is suggested that the mating system of symbiotic janirid isopods with long-term sperm storage and continuous receptivity in females and male mating with virgin females has evolved in response to highly unpredictable encounter probabilities between the sexes. Mate guarding and manipulation of small virgin juveniles may be favoured on the highly mobile hosts of symbiotic janirid isopods. Furthermore, adult females may gain by leaving their emerging offspring in the protective grip of guarding males, thereby reinforcing the maintenance of this peculiar mating system.     

Variation in behavioral and hormonal responses of adult male gray-cheeked mangabeys (<Emphasis Type="Italic">Lophocebus albigena</Emphasis>) to crowned eagles (<Emphasis Type="Italic">Stephanoaetus coronatus</Emphasis>) in Kibale National Park,Uganda

Intensive study of arboreal forest-dwelling primates and their predators in Africa is increasingly revealing that crowned eagles (Stephanoaetus coronatus) are major predators of primates. Gray-cheeked mangabeys (Lophocebus albigena) are overrepresented in the diets of crowned eagles in Kibale National Park, Uganda, and adult male mangabeys are represented more than females. We focused on the behavior of adult male gray-cheeked mangabeys living in social groups in Kibale National Park (1) to clarify the interactions between mangabeys and eagles that might put adult males at greater risk and (2) to better understand individual variation in behavioral responses to predators. Adult male mangabeys in five groups responded to observer-confirmed presence of crowned eagles 88 times over a 13-month period. While all males gave alarm calls, only the highest-ranking male in each of four groups chased eagles. These males had elevated levels of fecal cortisol metabolites in the days immediately after they engaged in active defense, suggesting that they perceived such behavior as risky. In the one group where male ranks were unstable and there were no infants, no male was observed to chase eagles. We suggest that males pursue the dangerous tactic of chasing eagles only when they are likely to have offspring in the group. Males in larger groups also spent less time alarm calling to crowned eagles (from first to last call in a group), and our observations confirmed that the duration of their alarm calls was related to eagle presence. Thus, eagles spent less time around larger mangabey groups. Alarm calling by adult male mangabeys may signal to this ambush predator that it has been detected and should move on.     

Female brown-headed cowbirds' (<Emphasis Type="Italic">Molothrus ater</Emphasis>) social assortment changes in response to male song: a potential source of public information

In many species, females' behavior appears to be influenced by that of other females, particularly regarding mate choice. Females theoretically can reduce the costs associated with independent male assessment by observing conspecifics. Studies of brown-headed cowbirds suggest that females pay attention to other females' behavior. Group-housed females modify their song preferences, whereas females housed in pairs do not. What information is available to females in a group environment? To address this question, we studied two groups of juvenile (i.e. hatch-year birds) and adult female cowbirds in a naturalistic group setting. We used a longitudinal ABA design, consecutively introducing and removing males that differed in age, amount of song production and stage of song development, to isolate the male characteristics that related to changes in female behavior. Juvenile and adult females assorted by age class when singing adult males were in the aviary, but not when singing juveniles or silent males of any age class were in the aviary. Results from playback tests confirmed that adult male song alone influenced female age class assortment. Videotape analysis from playback tests revealed that females also wing stroked in response to male song. Other females sometimes approached females who wing stroked and observed them. We hypothesize that group-level changes in social organization and individual females' responses can serve as visual signals for other individuals.     

Paternity alone does not predict long-term investment in juveniles by male baboons

Adult male chacma baboons (Papio hamadryas ursinus) form preferential associations, or friendships, with particular lactating females. Males exhibit high levels of affiliative contact with their friends’ infants and defend them from potentially infanticidal attacks (Palombit et al. 1997). Little is known about males’ associations with juveniles once they have passed the period of infanticidal risk. We conducted an observational, experimental, and genetic study of adult male and juvenile chacma baboons in the Moremi Reserve, Botswana. We identified preferential associations between males and juveniles and used behavioral data and a playback experiment to explore whether those associations have potential fitness benefits for juveniles. We determined whether males preferentially invest in care of their own offspring. We also determined how often males invest in care of their former friends’ offspring. The majority of juveniles exhibited preferential associations with one or two males, who had almost always been their mother’s friend during infancy. However, in only a subset of these relationships was the male the actual father, in part because many fathers died or disappeared before their offspring were weaned. Male caretakers intervened on behalf of their juvenile associates in social conflicts more often than they intervened on behalf of unconnected juveniles, and they did not appear to differentiate between genetic offspring and unrelated associates. Playbacks of juveniles’ distress calls elicited a stronger response from their caretakers than from control males. Chacma males may provide care to unrelated offspring of former friends because the costs associated with such care are low compared with the potentially high fitness costs of refusing aid to a juvenile who is a possible offspring.     

Social behavior of free-ranging juvenile sooty mangabeys (Cercocebus torquatus atys)

Sooty mangabeys are terrestrial monkeys exhibiting female philopatry and male dispersal. Studies in captivity as well as in the wild have found that adult females form linear dominance hierarchies. However, while captive studies found no evidence for a matrilineal social system, a previous study in Taï National Park, Ivory Coast, suggested that relatedness could influence both dominance rank and affiliation pattern among adult females. Here I test whether the dominance rank, coalitionary behavior, and affiliative behavior of juveniles in a group of free-ranging mangabeys in the Taï National Park are in accordance with a matrilineal, individual, or age-related dominance system. I found that juvenile females' dominance ranks remained stable over time and were highly correlated with the dominance ranks of their mothers, whereas juvenile males' dominance ranks were initially correlated with the ranks of their mothers, but showed greater instability with increasing age. Moreover, coalitions occurred mainly between juveniles and animals that were close in rank, including their mothers and siblings. Finally, juvenile females associated and groomed preferentially with close-ranking juvenile and adult females. Juvenile males showed similar preferences in affiliation with adult females, but when associating with juvenile males, they preferred peers. The observed social behavior of free-ranging juvenile sooty mangabeys resembled the social behavior described for juveniles of many matrilineal primate species.     

Males are selective too: mating, but not courtship, with sequential females influences choosiness in male field crickets (Gryllus bimaculatus)

Mate choice experiments have generally focused on female choice; few have considered that males can also be selective. We examined courtship in male field crickets sequentially introduced to four females of differing size. Large (L) and small (S) females were introduced in order of either LSLS or SLSL. We demonstrate that naive males invest equally (courtship effort) in the first female they meet, regardless of her size, but show greater courtship effort when they subsequently encounter large compared with small females. Moreover, we demonstrate that males show this discriminatory behavior when they are permitted to mate with females (i.e. use a spermatophore) but are less choosy when the female is removed before spermatophore transfer.     

Female preferences for male calling bout duration in a field cricket

Summary The calls of male field crickets attract sexually receptive females. In Gryllus integer, males differ from one another in their durations of uninterrupted calling (calling bout lengths). Tape recordings of the calls of 50 wild-caught males revealed that 14 males spent most of their calling time in short bouts (Fig. 1A), 18 in both short and long bouts (Fig. 1B), and 18 in long bouts (Fig. 1C). Re-recordings of 32 males after 3 weeks showed that calling bout lengths of individual males are stable with time (age) (Fig. 2). Three phonotaxis experiments investigated whether calling bout lengths of males affect female preferences. They demonstrated that (1) females can discriminate among conspecific males on the basis of calls alone; (2) females are preferentially attracted to males with long calling bout lengths; and (3) calling bout length is the specific factor responsible for preferential attraction. These results precisely identify a criterion that females use to discriminate among potential mates of their own species.     

Latency to resume calling after disturbance in the field cricket,<Emphasis Type="Italic"> Teleogryllus oceanicus</Emphasis>, corresponds to population-level differences in parasitism risk

A possible parasitoid-evasion behavioral adaptation is examined in male field crickets, Teleogryllus oceanicus, from three Hawaiian islands where parasitoid prevalence varies naturally among islands. Ormia ochracea, the parasitoid fly that parasitizes T. oceanicus on these islands, uses male calling song to locate its hosts. We used laboratory-reared males from three Hawaiian islands to determine if there are population differences in the time it takes for calling males to resume calling after a standardized disturbance. Males follow the expected pattern; males from the island with the greatest risk of parasitism have the longest latency to resume calling, and males from the island with the least risk of parasitism have the shortest latency to resume calling. Results are discussed in the context of behavioral adaptations to differing parasitism levels, and trade-offs between natural and sexual selection.Communicated by D. Gwynne     

An attractive male trait and aggressiveness are negatively correlated in wild field crickets, but uncorrelated in lab-reared crickets

It is generally thought that attractive male traits are positively correlated with dominance (aggressiveness). However, growing evidence suggests that this is not necessarily the case. We investigated whether calling song, a male mating display used by females to evaluate potential mates in a field cricket (Gryllus integer), is correlated with aggressiveness. In this species, females prefer males with longer durations of singing time. We measured singing time by measuring song over three continuous days using a custom-designed audio-monitor and assessed aggressiveness by pairing males in agonistic interactions. Our results showed that for males caught in the field, the percentage of time spent singing was negatively correlated with aggressiveness. However, in males that were raised in the laboratory, the percentage of time spent singing was not correlated with aggressiveness. Since calling duration is an attractive male trait in these crickets, our data show that the attractiveness of males can be either negatively related to aggressiveness (field) or unrelated to aggressiveness (lab). Neither of these results fit the traditional view that preferred male mating cues should be positively correlated with dominance (aggressiveness). These results also suggest that measurements of aggressiveness made in the lab do not necessarily correspond with aggressiveness in the field.     

The sex in short supply for matings varies over small spatial scales in a katydid (Kawanaphilanartee, Orthoptera: Tettigoniidae)

In katydids such as Kawanaphilanartee, a female bias in the operational sex ratio (OSR) results in female competition for mates and male choice of mates. Previous work showed that the excess of sexually active females occurs when food availability is low, in part because less food increases the propensity of females to mate as they forage for the large edible spermatophores produced by males. In this study with K.nartee, a pollen-feeding species, we estimate natural variation in numbers of sexually active males and females by assessing male calling activity and the propensity of females to respond to experimental calling males. We found an excess of sexually active males at a site with many flowers and an excess of sexually active females at a site with few flowers about 900 m away. Between-site differences in gut masses of calling males were consistent with the hypothesis that pollen availability controls OSR. Finally, at a third site where flowers were at first scarce, we found that the initial excess in sexually active females changed to an excess of sexually active males after a clump of grass-trees flowered. The mean gut mass of all sampled males from this site increased after flowering. The large variation in OSR that we document for K. nartee highlights the importance of identifying the appropriate spatial and temporal scales over which OSRs are measured in studies of factors controlling sexual selection. Received: 13 May 1997 / Accepted after revision: 27 October 1997     

Forced dispersal of juvenile guanacos (<Emphasis Type="Italic">Lama guanicoe</Emphasis>): causes,variation, and fates of individuals dispersing at different times

We examined adult-juvenile conflict in the guanaco (Lama guanicoe). During spring, territorial males become increasingly aggressive toward all juveniles born the previous year and begin expelling them from family groups. In an apparent effort to reduce aggression, juveniles display submissive crouches when being observed, approached, or attacked by the territorial male. Therefore, we assessed the influence of juvenile submissive behavior on the timing of dispersal and also examined if dispersal time was related to survival and reproductive performance as adults. We also evaluated hypotheses regarding the evolution of juvenile mammalian dispersal in the context of if and how each may favor the forced dispersal of juvenile guanacos by territorial males. Juveniles generally dispersed in late spring and early summer, and a nearly equal proportion of females (n=46; 48%) and males (n=49; 52%) dispersed. More-submissive animals generally dispersed later than less-submissive animals. Juvenile sex and dispersal time were not related to survival. In contrast, juvenile sex and dispersal time were related to reproductive performance. The probability of reproducing was highest when juveniles dispersed early and decreased with increasing time in family groups prior to dispersal. The largest proportion of juveniles was forced to disperse during a 2-week interval following the peak of the breeding season. Competition for food resources is likely very intense at this juncture and territorial males may force older juveniles to disperse in order to divert food resources to younger neonates. Additionally, juveniles may be forced to disperse after territorial males mate their mothers to prevent lost mating opportunities, because females leave territories when their offspring disperse and possibly prior to mating with males. We conclude that the forced dispersal of juvenile guanacos by territorial males is ultimately driven by competition for food resources on territories. The timing of dispersal, however, may be tempered by the chronology of matings between territorial males and particular adult females, and/or genetic relatedness between territorial males and juveniles.     

Aggregation and dispersal based on social cues as a nest-site selection strategy in a resource-defence polygynandry mating system

Many animals must choose a nest site in order to reproduce. However, it is unclear how nest-site selection strategies vary across different mating systems. We must therefore explore nest-site selection strategies in a range of mating systems, including the interaction between resource-defence polygyny and polyandry (i.e. polygynandry). In this study, we imposed a re-settlement event in the terrestrial toadlet Pseudophryne bibronii and measured the influence of the spatial position of each male’s nest site with respect to rival males on the likelihood that it would be abandoned or receive eggs. We captured every calling male in a population, measured their breeding success and released them back into the breeding area. We then recorded the establishment and abandonment of nest sites by males over 26 consecutive nights. Spatial positioning did not have any significant effects on male-breeding success, supporting claims that females show less discrimination between nest sites when they are polyandrous and spread their eggs amongst multiple male nests. However, we found that males consistently selected nest sites according to a site’s spatial position, which suggests that fitness benefits unrelated to male breeding success (e.g. reduced mortality risk) might influence male nesting decisions. Overall, our study provides new evidence that the mating system adopted by a population can influence the cues that individuals respond to when selecting nest sites.     

Effect of pair-bonding with adults on the ontogenetic manifestation of aggressive behavior in a primate,Saguinus fuscicollis

Summary Injurious aggression, scent marking and some visual displays shown in situations analogous to territorial defense were studied in young saddleback tamarins who had lived with nonrelated adult and juvenile social partners from 6 months of age under the experimental conditions outlined below. Twenty male and 20 female tamarins were removed from their natal families when 6 months old, and were used to form pairs and trios of the following composition: (1) one juvenile female and one juvenile male (10 pairs); (2) one juvenile female and one adult male (4 pairs); (3) one juvenile female, one juvenile male, and one adult male (6 trios); (4) one adult female and one juvenile male (4 pairs). These groups cohabited permanently and when the subjects were between 10 and 12 months old, their social interactions with adult conspecific strangers were studied, using series of 10-min social encounter tests. During these tests injurious aggression (attacks, fights, chases), a number of visual hostile displays (threat face, ruffle, arch) and scent marking were recorded.Young males and females living with adults of the opposite sex showed more injurious aggression against strangers and scent marked more frequently than young subjects living with partners of their own age. Males living with adult females also showed more facial threat than males living with young females (Figs. 1 and 2). Young males living in trios with adult males and young females scent marked less frequently than males living only with a young female. Aggression, scent marking and facial threat shown by subjects living with adult sex partners were at the level of socially and sexually experienced, pair-bonded adults (Fig. 3). No differences in ruffling and arching scores were found. The injurious aggression scores of the subjects were correlated with those of their mates. These results show that tamarins can fulfill the behavioral roles of pair bonded adults and also breed (Epple and Katz 1980) at an unexpectedly early age, when they live under the influence of an adult sex partner. Young animals cohabiting with an adult of the same sex, on the other hand, show a trend toward inhibition of adult-like behaviors.     

Alternative mating tactics in the Italian treefrog, <Emphasis Type="Italic">Hyla intermedia</Emphasis>

As in many lekking anurans, Italian treefrog males use two mating tactics: they can attract females by calling vigorously or be satellites, that is, they can remain silent in proximity of a calling male and try to intercept females attracted by their neighbour. We investigated the factors that affected the expression of this mating tactic. Consistent with the conditional mating tactic hypothesis, satellites were smaller than average and smaller than their parasitised calling males. They spent a larger-than-average number of nights at the breeding site, where most of them were also observed calling. Moreover, satellites showed lower call rates and lower mating success than those of males they parasitise but not lower than those of males they did not parasitise. Overall, these results, together with those derived from the analyses of the seasonal and spatial distribution of males, provide evidence for a non-random association between satellites and calling males and are consistent with the hypothesis that satellites have spectral and temporal acoustic preferences that parallel those of females. By adopting the less-successful satellite mating tactic, competitively inferior males can nevertheless maximise their potential reproductive fitness by sexually parasitising the most attractive chorusing males.     

Responses to conspecific scent marks and the ontogeny of territorial marking in immature terrestrial salamanders

We investigated the ontogeny of responses to scent marks in immature terrestrial salamanders (Salamandra lanzai) that inhabit the southwest Alps. In this species, sexual maturity is usually reached at about 8 years, and adults exhibit territoriality. One should expect territoriality to take place largely before the acquisition of sexual maturity if sexual competition is not the main force driving territoriality. However, both the difficulties for inexperienced animals to find a suitable territory and size-related competition may delay the acquisition of territoriality in juveniles. We performed choice tests with juveniles belonging to two age groups (1±1 years old versus 4±1 years old). Each focal animal was offered, in random order, the choice between two shelters, one without scent and the other containing one of the following scents: own, same-aged animal, a juvenile belonging to the other age group, or adult female. We also performed choice tests with adult females for which the scents of two juveniles belonging to a different age group were successively offered in a random order. Older juveniles were strongly attracted toward their own shelter and mostly avoided the shelters that contained the scent of juveniles of about the same age and of adult females. Adult females avoided the scents of older juveniles but not younger juveniles. These results therefore suggest that older juveniles use territorial marking. Conversely, young juveniles behaved randomly with respect to their own scents and to those of a same-aged juvenile, and they were significantly attracted towards the odor of an older animal, especially adult females. Both older juveniles and adult females displayed a random behavior toward the scents of young juveniles. Our results suggest that young juveniles do not defend territories but use spaces occupied by older individuals.Communicated by A. Mathis     

Kinship and social tolerance among female and juvenile wild house mice: kin bias but not kin discrimination

Kin-biased social tolerance among house mice has been interpreted in terms of kin discrimination. However, several lines of evidence suggest it may instead be an incidental artifact of group member discrimination. This leads to very different predictions about the social consequences of relatedness within and between social groups. Social interactions between wild-stock adult female and juvenile house mice (Mus domesticus) established in neighbouring territorial groups within enclosures reveal relatedness to dominant males within groups as the major factor determining social tolerance of juveniles by females. Relatedness to the female herself had no significant independent effect on responses indicating tolerance. Females were generally more aggressive toward neighbouring-group juveniles (all unrelated to females) compared with those from their own group (all related to females), but were most aggressive toward neighbouring juveniles sired by the neighbouring dominant male. They were also more aggressive toward their own-group juveniles that had been sired by the neighbouring dominant but only when encountered in the neighbouring territory and with a greater bias against female juveniles. Females were least aggressive toward own-group juveniles sired by their own-group dominant male. The sire-bias in tolerance among females is similar to that reported among the dominant males themselves in an earlier study. As a result of the combined sire-bias in tolerance by adult males and females, juveniles sired by their own-group dominant males become less likely to intrude into a neighbouring territory with time. Overall, the results suggest that differences in social tolerance reflect discrimination on the basis of social group membership rather than relatedness between interactants and thus provide strong experimental evidence in support of incidental kin bias rather than kin discrimination.     

The timing of premating and mating behavior in a field population of the cricket Gryllus campestris L.

Summary In a field population of Gryllus campestris, 209 males and 175 females were marked individually and their migratory and mating behavior was observed. Many females and males mate more than once and both sexes change burrows frequently. All copulations observed were during the day. Automatic recording of calling in 12 selected males, and walking in 7 females, showed that an individual male may call predominantly during the day, during the night, or both day and night, but the females show locomotory activity predominantly during the day. Thus, female locomotion is significantly temporally correlated with copulation but not with male calling.     

Risk trading in mating behavior: forgoing anti-predator responses reduces the likelihood of missing terminal mating opportunities

Life history theory predicts that organisms make certain adjustments to their current and future reproductive effort such that fitness is maximized. Moreover, these adjustments may be fine tuned in response to risks of attack by natural enemies. Thus, we may predict that as an organism ages it will accept increasing levels of exposure to predators during mating activities, effectively trading the risk of losing terminal mating opportunities for the risk of predation. We tested this prediction in an acoustic moth, Achroia grisella, in which females orient toward and evaluate males based on their ultrasonic calling song, and both sexes may be vulnerable to predation by insectivorous bats while in flight as well as on the substrate. In the latter situation, singing males and orienting females show silence and arrestment responses, respectively, when presented with synthetic bat echolocation signals broadcast above a threshold amplitude. We found that both males and females become less sensitive to these broadcasts over the course of their brief reproductive periods, 7 and 5 days, respectively. Over the same periods, sensitivity to male song in both males and females remains constant, and relatively little senescence in sexual behavior is observed. These results support the risk trading hypothesis, and they indicate that life history principles may apply over a very short lifespan.