Variation in foraging effort by lactating Antarctic fur seals: response to simulated increased foraging costs

Seasonally breeding predators, which are limited in the time available for provisioning young at a central location, and by the fasting abilities of the young, are likely to maximize energy delivery to the young by maximizing the rate of energy delivery averaged over the whole period of investment. Reduction in food availability or increased foraging costs will alter the optimal behavior of individuals. This study examined the behavioral adaptations of a diving predator, the Antarctic fur seal, to increased foraging costs during lactation. One group of mothers (n=5, treatment) was fitted with additional drag to increase the cost of transport in comparison with a control group (n=8). At the scales of the individual dives, the treatment group made more shorter, shallower (< 30 m) dives. Compensation for slower swimming speeds was achieved by diving at a steeper angle. Overall, diving behavior conformed to several specific theoretical predictions but there were also departures from theory, particularly concerning swimming speed during diving. Diving behavior appears to be adjusted to maximize the proportion of time spent at the bottom of dives. At the scale of diving bouts, no difference was observed between the treatment and control groups in terms of the frequency and duration of bouts and there was also no difference between the two groups in terms of the proportion of time spent diving. At the scale of complete foraging cycles, time taken to return to the pup was significantly longer in the treatment group but there was no difference in the rate of delivery of energy (measured from pup growth rate) to the pups in each group. Since mothers in the treatment group did not use significantly more body reserves, we conclude that behavioral adjustments at the scale of individual dives allowed mothers in the treatment group to compensate for the additional foraging costs. Pup growth rate appears to be less sensitive to the foraging conditions experienced by mothers than foraging trip duration. Received: 14 June 1996 / Accepted after revision: 16 November 1996     

Relatedness, polyandry and extra-group paternity in the cooperatively-breeding white-browed scrubwren (Sericornis frontalis )

We used DNA fingerprinting to examine the genetic parentage and mating system of the cooperatively breeding white-browed scrubwren, Sericornis frontalis, in Canberra, Australia. Our analyses revealed a remarkable variety of mating tactics and social organization. Scrubwrens bred in pairs or multi-male groups that consisted of a female and two or more males. Females were always unrelated to the pair male or alpha (dominant) male. Among multi-male groups we found three different mating tactics. Firstly, when alpha and beta (subordinate) males were unrelated, they usually shared paternity in the brood. This resulted in both males gaining reproductive benefits directly. Secondly, when beta males were not related to the female but were related to the alpha males, beta males sired offspring in some broods. In this situation, beta males gained reproductive benefits both directly and potentially indirectly (through the related alpha male). Thirdly, when beta males were related to the female or both the female and alpha male, they remained on their natal territory and did not sire any offspring. Thus beta males gained only indirect reproductive benefits. Overall, when group members were related closely, the dominant male monopolized reproductive success, whereas when the members were not related closely the two males shared paternity equally. This positive association between monopolization of reproduction and relatedness is predicted by models of reproductive skew, but has not been reported previously within a single population of birds. Other cooperatively breeding birds with both closely related and unrelated helpers may show a similar variety of mating tactics. Finally, we found that extra-group paternity was more common in pairs (24% of young) than in multi-male groups (6%), and we discuss three possible reasons for this difference. Received: 21 May 1996 / Accepted after revision: 14 December 1996     

Correlates of extra-pair fertilization success in hooded warblers

We examined correlates of extra-pair fertilization (EPF) success in the hooded warbler (Wilsonia citrina), a species where females pursue extra-pair matings. The good genes hypothesis predicts that females choose extra-pair mates with morphological or behavioral traits that reflect differences in male genetic quality. EPFs were common, as 35.3% (42/119) of broods contained extra-pair young (EPY) and 26.7% (95/356) of nestlings were the result of EPFs. There was a strong skew in male EPF success which increased the variance in annual male mating success 2–3 fold. However, male morphology did not predict EPF success, as extra-pair males were not older or larger than the males they cuckolded. Likewise, there were no significant correlations between the proportion of extra-pair young in a brood and male size or age. The good genes hypothesis predicts that high-quality males will be consistently preferred as genetic mates, but the number of young sired by a male with his social mate was not consistent from one year to the next. There was a significant negative correlation between female age and proportion of EPY produced, which could result if older females obtain higher-quality social mates. We found no strong evidence that females choose extra-pair mates for good genes, but females may use behavioral rather than morphological cues to assess relative male quality. Received: 31 May 1996 / Accepted after revision: 17 October 1996     

Lack of detectable nepotism in multiple-queen colonies of the fire ant Solenopsis invicta (Hymenoptera: Formicidae)

Multiple-queen (polygyne) colonies of the introduced fire ant Solenopsis invicta present a paradox for kin selection theory. Egg-laying queens within these societies are, on average, unrelated to one another, and the numbers of queens per colony are high, so that workers appear to raise new sexuals that are no more closely related to them than are random individuals in the population. This paradox could be resolved if workers discriminate between related and unrelated nestmate sexuals in important fitness-related contexts. This study examines the possibility of such nepotism using methods that combine the following features: (1) multiple relevant behavioral assays, (2) colonies with an unmanipulated family structure, (3) multiple genetic markers with no known phenotypic effects, and (4) a statistical technique for distinguishing between nepotism and potentially confounding phenomena. We estimated relatedness between interactants in polygyne S. invicta colonies in two situations, workers tending egg-laying queens and workers feeding maturing winged queens. In neither case did we detect a significant positive value of relatedness that would implicate nepotism. We argue that the non-nepotistic strategies displayed by these ants reflect historical selection pressures experienced by native populations, in which nestmate queens are highly related to one another. The markedly different genetic structure in native populations may favor the operation of stronger higher-level selection that effectively opposes weaker individual-level selection for nepotistic interactions within nests. Received: 28 June 1996 / Accepted after revision: 6 October 1996     

Female mate choice in the non-territorial lizard Ameiva plei (Teiidae)

Mate choice by females has been documented in a variety of taxa. Female mate choice in species lacking male resource control or paternal care might occur if preferred males provide protection from harassment. Female mate choice was investigated in a natural population of the non-territorial lizard Ameiva plei (Teiidae). Consort pairs were allowed to form naturally. Consort males were significantly larger than non-consort males. After removal of consort males, the “abandoned” female's reaction to the first male who approached her was recorded. Females rejected all small males. Female preference for large males was significantly higher than preference for small males. Large males may be better equipped to guard the females from harassment and behavior of large males is less harassing than behavior of small males, thereby affording the female increased foraging time. Received: 21 June 1996 / Accepted after revision: 28 December 1996     

Dominance and the dynamics of phenotype-limited distribution in common cranes

We studied the behavior of 13 radiotagged cranes dispersing from a communal roost over days when they changed their main daily foraging area between consecutive days during two winter seasons. Individuals went to a new foraging zone when on the previous day their morning food intake had fallen below their mean morning food intake measured over the whole winter. Food intake on the day before a change in foraging area was positively correlated with dominance rank. Dominant cranes changed to new zones with higher numbers of birds and food density, while subordinate cranes went to new zones with lower numbers of birds. As a result, all birds increased their food intake over that of the previous day. Dominant cranes remained more faithful to their most preferred foraging zone, where they spent 69% of the mornings, while subordinate birds were more mobile, switching among zones frequently. Dominant birds left the roost later than subordinate birds on the days they changed to a new zone, which could be used to track the main departing flows. The results suggest that the dynamics that led to a truncated phenotype-limited distribution were determined by social dominance and food abundance, with dominant cranes shifting to a new zone to maintain their high intake levels and subordinates changing more frequently whenever their daily intake did not reach the minimum metabolic requirements. Received: 16 December 1996 / Accepted after revision: 22 February 1997     

Why is competition more intense if food is supplied more slowly?

We investigated competition for food among two groups of six clone amagos (salmonids), Oncorhynchusmasoumacrostomus, in a laboratory experiment with different rates of food input. We examined the effect of temporal clumping of food resources on the inequality of food sharing between competitors. Monopolization of food by dominants was greater at a low input rate (one food item per 10 s) than at a high input rate (1 food item per 1 s). Aggressive behavior by dominants was more frequent at the low input rate than at the high input rate; its purpose was presumably to interfere with the feeding behavior of subordinates. We assessed the relative importance of three foraging factors (the number of approaches to food items, the chance per approach and the gain per chance) in enhancing inequality in food gain between individuals. Dominants had a disproportionately high chance per approach and gain per chance at the low input rate, but not at the high input rate. The chance of obtaining a food item per approach depended on how many competitors approached simultaneously. The gain per chance depended on the competitive ability of the approaching fish. There was an interaction between these components, such that the number of approaches affected the chance per approach and gain per chance. We evaluated the independent effect of the chance per approach, and showed that it was higher for dominants than for subordinates at the low input rate, but not at the high input rate. This implies that subordinates changed their behavior and became more likely to avoid approaching food at the same time as dominants at the low input rate. Received: 13 August 1996 / Accepted after revision: 30 November 1996     

Female mallard mating preferences for multiple male ornaments

Mating preferences were studied in captive wild-stock female mallards. Independent observers scored ornament brightness for the bill and eight plumage areas of male mallards on a scale of 1 to 6. Ornament scores were generally positively correlated with each other as indicated by univariate correlations and principal component analysis. Males with higher bill scores were significantly more likely to obtain pairings. None of eight individual plumage ornaments were significant predictors of pairing success, but males with higher average plumage scores had significantly higher pairing success. Males that completed the molt first were also significantly more likely to be paired. Thus females may have paid some attention to plumage brightness. Stepwise multiple regression produced a model that included only bill and molt, both of which contributed a similar degree (r ² = 0.39); plumage ornaments and size measurements did not enter the model. These results are interpreted in light of hypotheses for the evolution of multiple ornaments, and phylogenetic plumage patterns in dabbling ducks. Received: 18 December 1995 / Accepted after revision: 22 September 1996     

Magpies' tails: damage as an indicator of quality

Quality-indicating sexually selected traits may have their honesty maintained by their costs or by an inherent “revealing” nature. Long tails in birds are usually considered to be costly “handicaps”, but may have additional potential as revealing indicators through the incidence of breakage. Magpies Pica pica with unbroken and less abraded tails paired earlier, but did not nest or fledge young earlier than pairs with tails in poorer condition. Pairs mated assortatively by tail quality, and magpies with very broken tails remained unmated. Pairs in which both members had almost undamaged tails fledged more offspring than pairs with poorer tails. Tail quality did not correlate with the extent of any habitat type in the territory. Tail damage thus honestly indicated a magpie's reproductive potential, and the data are consistent with its having a role in mate choice, as a revealing element of tail morphology. Received: 28 March 1996 / Accepted after revision: 9 December 1996     

The structural significance of a vocalization that stimulates gaping in black-capped chickadees

In a variety of avian species, vocalizations used to stimulate gaping in nestlings during feeding share similar features such as broad frequency range, multiple spectral components, and sudden onset. It is currently not understood what functional significance, if any, these acoustic features have for close-range communication. One example of such a vocalization is the “squawk” of the black-capped chickadees (Parus atricapillus). The present study analyzes the significant features of the squawk by observing the response of chickadee nestlings to field playbacks of natural and computer-modified squawks. Of the features tested, frequency range appears most critical, but responses to frequency ranges change with age, such that low frequencies are most stimulating around the time of hatching but cease to elicit gaping after 5 days posthatch. Aperiodic stimuli and computer-modified squawks with gradual onsets tend to be less stimulating than natural squawks at some ages, but these differences are not significant at any age. The functional significance of the broad frequency range of the squawk is discussed, including the possible role of accommodating shifting frequency preferences related to ontogenetic changes in auditory sensitivity and elaboration of the vocal repertoire. Received: 16 January 1996 / Accepted after revision: 28 December 1996     

Breeding synchrony and extra-pair mating in red-winged blackbirds

Using data from a 6-year paternity study of red-winged blackbirds, I tested the hypotheses that increased nesting synchrony should either promote extra-pair mating by increasing the advantage of extra-pair mating to females, or decrease extra-pair mating by constraining males from seeking extra-pair copulations. Contrary to these hypotheses, the occurrence of extra-pair paternity did not vary with nesting synchrony over the breeding season, or vary with the number of synchronous nests within territories or within marshes, or with nesting order on territories. However, for nearly all nests with extra-pair young, there were fewer females synchronous with that nest on the cuckolder's territory than on the territory of the cuckolded male. This “advantage” of a synchrony difference was less pronounced for older males that cuckolded younger males, particularly when the two males were not neighbors. Collectively, these results suggest that breeding synchrony affects extra-pair mating by affecting mate guarding, but that breeding synchrony alone can not be used to predict which females are more likely to engage in extra-pair mating, nor with which extra-pair males they will mate. Understanding why extra-pair mating by older males is less affected by breeding synchrony may explain much about both the proximate and ultimate causes of extra-pair mating in red-winged blackbirds. Received: 7 June 1996 / Accepted after revision: 25 November 1996     

Song rates of dark-eyed juncos do not increase when females are fertile

Frequency of singing by birds may vary with reproductive stage in ways that reflect variation in the functions of song in intersexual and intrasexual communication. In dark-eyed juncos (Junco hyemalis) high-amplitude song is produced only by males. To investigate the function of this song, we tested whether fertility of females affected singing by their mates or by neighboring males. Using focal observations, song censuses, and radiotracking data, we determined whether song production varied between and among periods when females were fertile and non-fertile. Our findings show that males do not increase song production when their mates are fertile, nor do they increase song production when neighboring females are fertile. These results suggest that male juncos do not signal their intent to defend territories (or mates) more when females are fertile and that they do not use song to advertise to specific potential participants in extra-pair fertilizations. Received: 13 February 1997 / Accepted after revison: 2 May 1997     

Control of reproduction in social insect colonies: individual and collective relatedness preferences in the paper wasp, Polistes annularis

Social insect colonies often have one or a few queens. How these queens maintain their reproductive monopoly, when other colony members could gain by sharing in the reproduction, is not generally known. DNA microsatellite genotyping is used to determine reproductive interests of various classes of colony members in the paper wasp, Polistes annularis. The relatedness estimates show that the best outcome for most individuals is to be the reproductive egg-layer. For workers, this depends on the sex of offspring: they should prefer to lay their own male eggs, but are indifferent if the queen lays the female eggs. The next-best choice is usually to support the current queen. As a rule, subordinates and workers should prefer the current queen to reproduce over other candidates (though subordinates have no strong preference for the queen over other subordinates, and workers may prefer other workers as a source of male eggs). This result supports the theory that reproductive monopoly stems from the collective preferences of non-reproductives, who suppress each other in favor of the queen. However, we reject the general hypothesis of collective worker control in this species because its predictions about who should succeed after the death of the present queen are not upheld. The first successor is a subordinate foundress even though workers should generally prefer a worker successor. If all foundresses have died, an older worker succeeds as queen, in spite of a collective worker preference for a young worker. The results support the previous suggestion that age serves as a conventional cue serving to reduce conflict over queen succession. Received: 3 May 1996 / Accepted after revision: 22 September 1996     

Ornamentation predicts reproductive success in female pipefish

In the sex-role reversed pipefish Syngnathus typhle females compete for access to males and males are choosy. Females develop a temporary ornament when competing over mates with other females and when performing nuptial dances with males. This ornament is an amplification of the normal striped pattern in these fishes. We here show experimentally that (1) the contrast of this normal pattern forecasts the extent to which the ornament is shown, (2) contrast and ornamentation honestly signal female quality (egg numbers), (3) contrast and ornamentation accurately predict female mating success, (4) contrast is a phenotypically plastic trait specifically exaggerated under situations of female – female competition, and (5) neither contrast nor ornament are energetically expensive to the females (i.e., they are independent of short-term nutritional status). Hence, as predicted in sex-role reversed species, ornament design is constrained by costs to female fecundity: an energetically demanding ornament would impair on a female's ability to produce eggs. The type of ornament described here is the expected one, costly for reasons other than being energetically expensive to produce. Received: 4 April 1996 / Accepted after revision: 27 October 1996     

A molecular genetic analysis of social structure, dispersal, and interpack relationships of the African wild dog (Lycaon pictus )

The African wild dog is a highly social, pack-living predator of the African woodland and savannah. The archetypal wild dog pack consists of a single dominant breeding pair, their offspring, and non-breeding adults who are either offspring or siblings of one of the breeding pair. Non-breeding adults cooperate in hunting, provisioning and the protection of young. From these observations follows the prediction that the genetic structure of wild dogs packs should resemble that of a multigenerational family, with all same-sexed adults and offspring within a pack related as sibs or half-sibs. Additionally, a higher kinship between females from neighboring packs should be evident if females tend to have small dispersal distances relative to males. We test these predictions through analysis of mitochondrial DNA control region sequences and 14 microsatellite loci in nine wild dog packs from Kruger National Park, Republic of South Africa. We show that as predicted, African wild dog packs generally consist of an unrelated alpha male and female, subdominant close relatives, and offspring of the breeding pair. Sub-dominant wild dogs occasionally reproduce but their offspring rarely survive to 1 year of age. Relatedness influences the timing and location of dispersal events as dispersal events frequently coincide with a change in pack dominance hierarchy and dispersers often move to areas with a high proportion of close relatives. Received: 22 February 1996 / Accepted after revision: 16 November 1996     

Common goldeneyes adjust maternal effort in relation to prior brood success and not current brood size

Parental investment theory predicts that parental effort should be related to the reproductive value of the current brood. This depends on both the number of young and the survival prospects of each of them. Thus parents may provide more care to larger broods either because of (1) the direct effect of brood size per se on reproductive value (the “brood size” hypothesis) or because (2) past mortality, reflected in current brood size, predicts future mortality of the brood and hence its reproductive value (the “brood success” hypothesis). Earlier studies have not attempted to distinguish between these alternatives. We tested the hypotheses in the precocial, nidifugous common goldeneye Bucephala clangula, a species with uniparental female care. Maternal effort was measured as the time spent by the female in rearing the brood. We found that brood size itself is not associated with maternal effort, but that females modify their maternal effort according to the mortality already experienced by the brood, supporting the prediction of the brood success hypothesis. We also found that brood mortality varied considerably between broods and that previous mortality predicts future mortality within broods, basic assumptions of the brood success hypothesis. Received: 30 January 1996 / Accepted after revision: 27 October 1996     

The evolution of non-maternal caretaking among anthropoid primates: do helpers help?

Non-maternal infant care among nonhuman primates has frequently been investigated from the perspective of the caretaker. Here we examine whether allocaretaking behavior provides direct reproductive benefits to mothers. Comparative analyses that control for the effects of allometry and phylogeny reveal that allocaretaking behavior correlates with relatively fast infant growth and reproduction, but is not associated with the production of large infants. These results are consistent with those from studies of other taxa; primate helpers appear to increase the reproductive success of female breeders. In addition, our findings contrast with those derived from traditional allometric analyses and underscore the importance of controlling for the potentially confounding effects of phylogeny in comparative analyses. Received: 28 May 1996 / Accepted after revision: 16 November 1996     

Infanticide in seasonally breeding multimale groups of Hanuman langurs (Presbytis entellus) in Ramnagar (South Nepal)

In the seasonally breeding langur (Presbytis entellus) population of Ramnagar, South Nepal, where multimale groups prevail, 25 attacks on 11 infants (including one actual killing) by seven adult males were witnessed in five groups by six observers between 1990 and 1996. Circumstantial evidence also indicates three additional attempts at infanticide and in seven additional cases infanticide was presumed or likely. Infanticide presumably accounted for 30.8–62.5% of infant mortality in the first 2 years of life. Most attackers (91.4%) were residents of the infants' group and had immigrated after the infants had been born (75.0%) or conceived (25.0%). Thus, they were not related to the victims. The interbirth interval was shortened if an infant died either prior to September of its 1st year of life (mean = 1.2 years), or its 2nd year (mean = 2.0) and even its 3rd year (mean = 2.4). All attackers remained in the group at least until the next mating season; high-ranking males maintained their dominance rank and lower-ranking males rose in rank. Since rank and mating success were correlated and rank and reproductive success might be correlated, all attackers had a good chance of siring the next infant of the victims' mothers and could thus have benefited by their action. Infanticide seems to be a male reproductive strategy at Ramnagar. Infanticide has never before been reported among seasonally breeding langurs living at such low densities. This is also the first detailed report of infanticide as a male reproductive strategy in a seasonally breeding primate population. Received: 19 December 1996 / Accepted after revision: 7 June 1997     

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     

Group-specific dialects and geographical variation in coda repertoire in South Pacific sperm whales

Codas, which are patterned series of clicks, were recorded from female and immature sperm whales (Physeter macrocephalus) in a number of locations around the South Pacific Ocean and in the Caribbean Sea. Using K-means cluster analysis, 3,644 codas were categorized based on the number of clicks and their patterning. There were 30 resulting types of coda. The numbers of codas of the different types recorded were used to construct repertoires for each recording session, day, group of whales, place, area, and ocean. Strong group-specific dialects, which seem to persist over periods of years, were apparent, overlaid on weaker geographical variation. Significant differences in repertoire were found between the Caribbean and the Pacific Ocean. Sperm whales now join killer whales (Orcinus orca) as the only cetacean species in which dialects (differences in vocal repertoire among neighboring, potentially interacting groups) have been found. Received: 13 June 1996 / Accepted after revision: 19 January 1997