Alliances in winter flocks of willow tits; effects of rank on survival and reproductive success in male-female associations

Summary The value of being dominant in winter flocks of willow tits may not only be a matter of personal survival for males, although their survival probability was highest among flock members. Winter flocks of willow tits contain male-female subunits, and the dominant pair may be viewed as an alliance based on mutual benefits. Dominant males provide their mates with protection in winter, and females return the help in reproductive success. Survival was directly related to rank only within sexes. Mates of dominant males survived significantly better than low-ranked males even though these females were subordinate to these males in direct interactions. Mates of dominant males were relatively spared from costs of low rank as they were subject to aggression less often than expected. Presumably dominant males prevented such attacks, as they excluded other flock members from high tree sections where their mates fed. Males should gain from improving the survival probability of their mates since it was not always possible to replace lost mates in spring populations having a male-biased sex ratio. Dominance in avian winter flocks may thus have more indirect effects than merely to improve the personal survival probability through resource priority. The benefits of being a dominant suggest that subordinate willow tits join groups because there is no space available for them to take up a territory as dominants.     

Age-specific survivorship in relation to clutch size and fledging success in California gulls

Summary Data from a natural population of California gulls (Larus californicus) demonstrated that increasing reproductive effort with age was associated with reduced survivorship. Number of offspring fledged but not clutch size was inversely related to adult survivorhip indicating that reproductively induced mortality resulted from the cumulative effects of the entire breeding season. Agerelated increases in fledging success were correlated with increased adult mortality. Young gulls fledged few offspring and had high survival rates. Old gulls tended to fledge more offspring and had low survival rates. However, those old gulls fledging few offspring survived as well as young gulls. Data also invalidate the assumption that survivorship is age-constant in this species.     

Transient ontogenetic expression of hermaphroditic gonad morphology within the Gobiosoma group of the Neotropical seven-spined gobies (Teleostei: Gobiidae)

The Neotropical seven-spined gobies (tribe Gobiosomatini), including the Gobiosoma and Microgobius groups, constitute a speciose, monophyletic gobiid taxon. In particular, member species of the Gobiosoma group exhibit a combination of behavioral diversification and microhabitat specialization that may have played a major role in their collective rapid evolution and high rate of speciation. Functional hermaphroditism, which is common among gobiids, can promote exploitation of patchily distributed micro-niches by countering potential reductions in reproductive opportunities. However, the possible role of functional hermaphroditism in the exploitation of microhabitats and rapid speciation in the Gobiosoma group is unknown. One member species, Tigrigobius multifasciatus, is a functional hermaphrodite. Two other Gobiosoma group species are functionally gonochoric (i.e., constant-sexed), but exhibit transient hermaphroditic ovarian structure among immatures. In this study, ovarian morphology among immature and adult females of several Gobiosoma group species was examined to see if hermaphroditic gonadal features are present. Although no evidence of functional hermaphroditism in the form of precursive accessory gonadal structures (pAGS) associated with the adult ovary was found among newly examined species, all species exhibited the transient expression of hermaphroditic gonadal features associated with the immature ovary. In contrast, among six species of non-Gobiosomatini genera having no record of hermaphroditism, none exhibited similar transient hermaphroditic features associated with the immature ovary. These findings suggest that hermaphroditism may have been an ancestral trait which has been secondarily lost within some Gobiosoma group species. This study also shows that ontogenetic and morphogenic processes within the gobiid reproductive system may provide new insights into the evolution of life history traits and significantly further our understanding of the extraordinary evolutionary success of this group.     

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     

Mass-dependent take-off ability in wintering great tits (Parus major): comparison of top-ranked adult males and subordinate juvenile females

When birds are attacked by predators, initial take-off is crucial for survival. Theoretical studies have predicted that predation risk in terms of impaired flight ability increases with body weight. However, studies in which attacks were simulated, and within-individual daily changes in body weight were used to test mass-dependent take-off outside migration period, have so far failed to show an effect of mass on velocity. In this field study I compared the mass/velocity relationships of alarmed adult male and juvenile female great tits, Parus major. Fattening strategies differ among members of the dominance-structured basic flocks of wintering great tits, and dominant individuals often carry significantly less amount of fat reserves than subordinates. Since the range of body weight gain/loss is the least among dominant males, it was expected that impaired flight ability is more likely in lower-ranked female great tits. The results show that the birds differed significantly in their daily increase of relative body weight. Average daily weight increase of adult males was 6.2%, while it was 12.2% in juvenile females. Males were faster than females at take-off both at dawn and at dusk. Flight velocity of males did not differ significantly between dawn and dusk, whereas females took off at a significantly lower speed at dusk than at dawn. The results suggest that the larger fat reserves of subordinate females needed to increase their chances of overwinter survival probably place them at increased risk of predation. Electronic Publication     

Sympatric corvids

Summary Overlap in winter resource use by four farmland corvid species was insufficient to predict any costly interspecific aggression. Rates of inexpensive displacement aggression were too low to affect intake rates of prey, and were not related to flock size. Eurasian crows also chase-attacked corvids and other species. This more costly form of aggression is interpreted not as interference aggression against food competitors, but as a response to the disturbance of earthworm prey into unavailability by the foraging movements of such flocks (Fig. 1). Rooks and jackdaws were less likely to leave the field following an attack if they were in larger flocks, but such flocks were attacked more frequently (Table 1).     

Song as a pursuit-deterrent signal,and its occurrence relative to other anti-predation behaviours of skylark (Alauda arvensis) on attack by merlins (Falco columbarius)

Skylarks show a range of anti-predation behaviours including flocking, refuge-seeking and song. Responses by skylarks to merlin attack were recorded over three winters on a Scottish estuary to determine the effectiveness of song as a pursuit-deterrent signal, and its use with respect to other anti-predation options such as flocking. Mortality due to merlin predation was high. Skylarks used song as a pursuit-deterrent signal. Merlins chased non- or poorly singing skylarks for longer periods compared to skylarks that sang well. A merlin was more likely to catch a non-singing than a poorly singing than a full-singing skylark. Temperature did not affect chase lengths, song types or success rates of attacks. Larger flocks of skylarks were preferentially attacked so that the individual probability of being attacked within some larger flock sizes was greater than in a smaller flock. Success rate did not vary with flock size. Merlins chose skylarks before any song was heard, so there was no cost for non-singing skylarks in joining flocks. The frequency of capture on merlin attack depended on the escape response used by the skylark. Non-singing skylarks were probably more likely to escape by seeking a refuge or staying on the ground on attack, while singing skylarks were more likely to escape if they flew. The optimal escape option available to a skylark on merlin attack was probably dependent on its condition, as indicated by its ability to sing on attack.     

Survival probability estimates for the endangered loggerhead sea turtle resident in southern Great Barrier Reef waters

Sex- and age-class-specific survival of a loggerhead turtle population resident in southern Great Barrier Reef waters was estimated using a long-term capture-mark-recapture (CMR) study and the Cormack-Jolly-Seber modelling approach. The CMR history profiles for 271 loggerheads tagged over 9 years (1984-1992) were classified into two age classes (adult, immature) based on somatic growth and reproductive traits. The sex and maturity status of each turtle was determined from visual examination of reproductive organs using laparoscopy. A reduced-parameter model accounting for constant survival with sex- and time-specific recapture was adequate for estimating age-class-specific survival probabilities, but inclusion of time-specific transient behaviour was informative for the immature age class. The annual fluctuations in the estimated proportion of transient immatures was not a function of sampling effort, but could be due to anomalous oceanographic conditions affecting dispersal of the immature class. There was no sex-specific difference in survival probabilities for either age class, but females were more likely to be recaptured than males, which might be related to behavioural differences such as sex-biased dispersal. The expected annual survival probability for adults was 0.875 (95% CI: 0.84-0.91). The expected annual survival probability for immatures was 0.859 (95% CI: 0.83-0.89), but when the transients were accounted for, the expected annual survival for the resident immature loggerheads was 0.918 (95% CI: 0.88-0.96). These are the first substantive estimates of annual survival probabilities for any loggerhead sea-turtle stock and provide a basis for developing a better understanding of loggerhead population dynamics.     

Kinship and aggression: do house sparrows spare their relatives?

Kin-selection theory predicts that relatedness may reduce the level of aggression among competing group members, leading to indirect fitness benefits for kin-favoring individuals. To test this hypothesis, we investigated whether relatedness affects aggressive behavior during social activities in captive house sparrow (Passer domesticus) flocks. We found that sparrows did not reduce their aggression towards kin, as neither the frequency nor the intensity of fights differed between close kin and unrelated flock-mates. Fighting success was also unrelated to kinship and the presence of relatives in the flock did not influence the birds’ dominance rank. These results suggest that the pay-offs of reduced aggression towards kin may be low in non-breeding flocks of sparrows, e.g. due to competition among relatives as predicted by a recent refinement of kin-selection theory. Our findings indicate that the significance of kin selection may be restricted in some social systems such as winter aggregations of birds. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Fattening strategies in dominance-structured greenfinch (Carduelis chloris) flocks in winter

Relationships between social status and levels of body reserves stored by members of greenfinch (Carduelis chloris) flocks in winter were investigated. In addition, the adjustment of reserves by birds of different rank to experimental changes in food predictability and to changed weather conditions were examined. Birds with low social status carried overall larger body reserves than high-ranked birds. The results of the experiments suggest that this was mainly because subordinates, due to their low priority to food access, perceived future foraging success as less predictable than dominants. In response to severe weather, which probably increased the risk of starvation for birds, dominants temporarily increased their reserves more than subordinates. This response also indicated that birds with low social status carried larger reserves than high-ranked birds, and demonstrated that dominants could increase food intake when there was a risk of energetic shortfall. The results suggest that fattening strategies in greenfinches depend on social status. In winter, subordinates may be forced to carry larger reserves than dominants to safeguard against social constraints in access to food during critical times. As body reserves may be costly to carry and acquire, this should reduce the probability of surviving the winter for subordinates compared to dominants. Received: 2 March 1995/Accepted after revision: 13 April 1996     

Individual differences and within-flock convergence in chickadee calls

Summary A detailed sound analysis of the Chick-adee call of the black-capped chickadee (Parus atricapillus) was performed in order to determine a basis for individual recognition and for imitation within winter flocks. During the winter of 1978–1979 members of five free-living black-capped chickadee flocks were uniquely marked for individual identification, and their calls were recorded in the field. Nested analysis of variance of temporal call parameters measured from sonagrams and spectral parameters from frequency spectra showed that there were significant differences between individuals within flocks for every parameter measured. There were significant differences between flocks in the frequency ranges 800–2,200 Hz and 4,000–5,300 Hz, in the spectral parameters bandwidth and maximum frequency, and in the duration of the Dee syllable and total call duration.Members of four of the five flocks were captured in December 1978 and held in aviaries segregated by original flock membership. In January 1979 the memberships of the three aviaries were rearranged to form experimental flocks. After one month, there were significant differences among the experimental flocks in the duration of the Dee syllable and total call duration. Convergence, as indicated by a significant decrease in variance among members (F-test), occurred in the experimental flock in Aviary 1 and was concentrated in the frequency ranges 1,300–1,800 and 6,200–6,900 Hz. The members of this experimental flock differed from each other in the number of 100 Hz frequency intervals within which each changed its own call.The Chick-a-dee call contains sufficient information that it can potentially be used by black-capped chickadees for individual recognition. In addition, both field and aviary data suggest that flock members converge in some call characteristics. Possible explanations of the social significance of vocal convergence in chickadee flocks are discussed.     

Parental effort in the California gull: tests of parent-offspring conflict theory

Summary Parental effort was studied among knownaged California gulls (Larus californicus) with a brood size of two. Results supported three of Triver's predictions from parent-offspring conflict theory. First, the amount of time parents withheld food from offspring increased with increasing offspring age. Second, older parents were less likely than younger parents to withhold food from offspring throughout the period of parental care. Third, older parents had a longer period of parental care compared to younger parents. Adult survival to future breeding seasons was inversely related to duration of parental care. Older parents, in association with their greater levels of parental effort, had a lower survival rate compared to that of younger parents.     

Functional aspects of the pair bond in winter in Bewick's swans (Cygnus columbianus bewickii)

Summary The benefits to partners of monogamous pairs of maintaining continual spatial proximity in the non-breeding season were studied in Bewick's swans wintering in Norfolk, UK.When separated from their mates, females were less successful in aggressive encounters, were threatened more frequently and spent less time feeding than when close to them. Males also suffered reduced success in encounters and a higher frequency of threats by other flock members, though the effect was less pronounced than for females. This sex difference in effect of separation may be associated with the greater weight of males and the fact that success in encounters is related to weight.Partners appeared to assist each other by joining in aggressive encounters, as well as by inhibiting other birds from threatening their mates. However, the precise manner in which the female assisted the male is still obscure, since the highest intensity aggressive encounters — physical fights — involved only male partners. It is suggested that the male may fight harder in his mate's presence.Proximity of partners varied with situation and between different pairs. Partners maintained greater proximity in dense flocks than in dispersed flocks and showed a tendency to stay closer when feeding on winter wheat than on waste potatoes. Partners where the male was high-ranking spent more time together than those where the male was low-ranking.     

Chick-a-dee call syntax,social context,and season affect vocal responses of Carolina chickadees (<Emphasis Type="Italic">Poecile carolinensis</Emphasis>)

Chick-a-dee calls in many chickadee (Poecile) species are common vocal signals used in a diversity of social contacts. The calls consist of four notes, A, B, C, and D, which follow simple rules of syntax (note ordering and composition) to generate many unique call types. We used field playbacks with Carolina chickadees, P. carolinensis, to ask whether violations of a syntactical rule affected their vocal responses. We show that chickadee responses to typical calls (e.g. AAAACCCC and CCCCDDDD) differ from responses to atypical calls (e.g. CACACACA and DCDCDCDC) depending on playback note composition, season, and social context (presence of heterospecifics). In the fall/winter, playbacks of typical calls with A and C notes elicited the greatest number of A and B notes in chick-a-dee call responses and typical calls with D notes elicited the greatest number of C notes, when in the presence of heterospecifics. In contrast, the corresponding atypical calls did not elicit similar responses. This suggests communicative significance is lost in calls that violate a rule of syntax in the fall/winter. In the spring, neither chickadee feebeefeebay song rate nor chick-a-dee calls responses differed by playback type. We suggest that call syntax is less salient for mated pairs in the spring than it is for fall/winter flocks that rely more on conspecific communication for foraging success and flock cohesion. This study represents the first experimental evidence that chickadees attend to both note composition and ordering in chick-a-dee calls.Communicated by W.A. Searcy     

Exploitative compensation by subordinate age-sex classes of migrant rufous hummingbirds

Summary The three age-sex classes of rufous hummingbirds (Selasphorus rufus) that directly interact on southward migratory stopovers in our California study system differ in territorial ability and resource use. Immature males are behaviorally dominant to adult and immature females and defend the richest territories. Here, we test the hypothesis that the territorially subordinate age-sex classes compensate exploitatively for their exclusion from rich resources. Our results show that females were able to accumulate energy stores at rates comparable to males despite their subordinate territorial status. Territorial females gained body mass at the same rate and in the same pattern as males, and resumed migration at the same body masses. Moreover, during periods when birds were nonterritorial and used dispersed resources, adult and immature females maintained or gained body mass, whereas immature males lost mass. We suggest that females may be energetically compensated by (1) lower costs of flight incurred during foraging and defense, resulting from their lower wing disc loading, and (2) greater success at robbing nectar from rich male territories, resulting from duller coloration (immature females), experience (adult females), and, possibly, hormonal differences. In the future, experiments will be necessary to distinguish the various hypotheses about the mechanisms involved in compensation. Correspondence to: F.L. Carpenter     

Inter-colony movements,at-sea behaviour and foraging in an immature seabird: results from GPS-PPT tracking,radio-tracking and stable isotope analysis

Seabird populations contain large numbers of immatures––in some instances comprising >50% of the fully grown adults in the population. These birds are significant components of marine food webs and may contribute to compensatory recruitment and dispersal, but remain severely understudied. Here, we use GPS-PTTs, radio-tracking and analysis of stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes to investigate the movements and foraging ecology of immature seabirds. Our study focussed on immature northern gannets Morus bassanus aged 2–4 attending non-breeding aggregations alongside a large breeding colony. GPS-PTT tracking of five birds revealed that immatures have the ability to disperse widely during the breeding season, with some individuals potentially prospecting at other colonies. Overall, however, immatures were faithful to the colony of capture. During returns to the focal colony, immatures acted as central place foragers, conducted looping and commuting flights, and analysis of the variance in first-passage time revealed evidence of area-restricted search (ARS) behaviour. In addition, stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope analyses indicate that immatures were isotopically segregated from breeders. Our findings provide insights into the foraging, prospecting and dispersal behaviour of immature seabirds, which may have important implications for understanding seabird ecology and conservation.     

Group-living in cliff swallows as an advantage in avoiding predators

Summary Cliff swallows (Hirundo pyrrhonota) in SW Nebraska, USA, nest in colonies and associate in groups away from their colonies. The degree to which group-living in this species affords advantages in the avoiding of predators was examined. The distance from the colony at which a snake predator was detected increased with colony size. In flocks away from the colonies, group vigilance increased, but the time that each individual spent vigilant decreased, with flock size. As a result, birds in large flocks had more time for preening and mud-gathering. Cliff swallows did not effectively mob predators and thus were unable to deter predators regardless of group size. Nesting within each colony was highly synchronous, but when the effects of ectoparasites on nesting success were removed, individuals nesting during the peak breeding period were no more successful than those nesting before or after the peak. This suggests that swamping of predators is unlikely in cliff swallow colonies. Nests at the edges of colonies were more likely to be preyed upon than nests nearer the center, suggesting that colonial nesting conferred some selfish herd benefits. Overall reproductive success did not vary with colony size. While cliff swallows receive some anti-predator benefits by living in groups, the avoidance of predators is probably not a major selective force for the evolution of coloniality in this species.     

Dominance, not kinship, determines individual position within the communal roosts of a cooperatively breeding bird

Kin selection has played an important role in the evolution and maintenance of cooperative breeding behaviour in many bird species. However, although relatedness has been shown to affect the investment decisions of helpers in such systems, less is known about the role that kin discrimination plays in other contexts, such as communal roosting. Individuals that roost communally benefit from reduced overnight heat loss, but the exact benefit derived depends on an individual's position in the roost which in turn is likely to be influenced by its position in its flock's dominance hierarchy. We studied the effects of kinship and other factors (sex, age, body size and flock sex ratio) on an individual's roosting position and dominance status in captive flocks of cooperatively breeding long-tailed tits Aegithalos caudatus. We found that overall, kinship had little influence on either variable tested; kinship had no effect on a bird's position in its flock's dominance hierarchy and the effect of kinship on roosting position was dependent on the bird's size. Males were generally dominant over females and birds were more likely to occupy preferred roosting positions if they were male, old and of high status. In this context, the effect of kinship on social interactions appears to be less important than the effects of other factors, possibly due to the complex kin structure of winter flocks compared to breeding groups.     

Interactions between fangblennies (Plagiotremus rhinorhynchus) and their potential victims: fooling the model rather than the client?

Lepidophagous (scale-eating) blue-striped fangblennies (Plagiotremus rhinorhynchus Bleeker 1852) are often found sympatrically with the bluestreak cleaner wrasse (Labroides dimidiatus Valenciennes 1839). They have some resemblance to the juvenile L. dimidiatus and have previously been reported as aggressive cleaner wrasse mimics. We observed 14 P. rhinorhynchus on a small area in the barrier reef near Hoga Island, Indonesia to assess the effects of client size on the behaviour and attack success of fangblennies: our results suggest that fangblennies are selective with regard to victim size; fish avoided by the fangblennies are significantly larger than those not avoided and attack success is significantly higher at intermediate victim size classes. The behaviour of the victims also has a significant direct effect on the foraging success of the fangblennies; where the potential victim posed, 63.6% were ignored by the fangblenny and only 7.4% of attacks were successful on posing fish as opposed to a surprise attack success rate of 71.6%. Overall, victims which exhibited the pose behaviour were significantly smaller in size. It appears likely that the predatory strategy of these fangblennies varies with victim size and that mimicry plays a minor role in attracting potential victims. We suggest that in common with other mimetic fish the resemblance of fangblennies to juvenile bluestreak cleaner wrasse allows them to actively hunt in areas where adult cleaners are common thus, indirectly improving their feeding opportunities.     

Costs of switching social groups for dominant and subordinate dark-eyed juncos (Junco hyemalis)

I quantified the costs of switching from a familiar to an unfamiliar flock for captive dark-eyed juncos (Junco h. hyemalis) by measuring several physiological and behavioral variables before and after flock switching. Birds that were initially dominant dropped in status in unfamiliar flocks, and experienced increased metabolic rates, while subordinate birds appeared to undergo less physiological change when switching flocks. This difference occurred despite a lack of any rank-related differences in the effects of joining a new flock on rates of aggression, weight change, access to food, or plasma corticosterone levels. These results suggest that for dominant, but not subordinate, individuals there is a measurable metabolic cost to joining a new social group, even in the absence of adverse factors such as food limitation. Dominant individuals may be less likely than subordinates to leave familiar flocks because of their higher metabolic costs when joining a new social group.