Graded recruitment and hunting strategies linked to prey weight and size in the ponerine ant Ectatomma ruidum

According to the weight and size of their prey, Ectatomma ruidum workers can employ different recruitment systems (solitary hunting, cooperative hunting and group hunting with recruitment) when mastering and retrieving prey items from short distances from the nest. Prey size determined the backwards entry typically adopted by this species, while prey weight determined the predatory strategy selected. After a common initial sequence (search for prey, detection, localization), predatory sequences varied in terms of the type of approach, the site of seizure, the reaction after stinging and the type of transport. Nevertheless, irrespective of prey weight and size, seizure was preferentially oriented towards the head and prey were always stung. Short-range recruitment and mass recruitment without trail laying were elicited by a large range of heavy prey (> 2.5 times the weight of an individual worker). According to the mortality risk associated with each prey, hunters exhibited a “prudent” stinging posture associated with an increase in the duration of the subsequent phase of waiting for prey immobilization. The overall time of capture was positively correlated with the weight of the prey. When collective hunting strategies were involved, E. ruidum colonies matched the number of recruited hunters to the size and weight of the prey. Compared to solitary hunting strategies, for short food–nest distances, this graded recruitment appeared to enhance the energetic benefits derived by this species from the use of recruitment systems: the higher the number of workers involved in the recruitment process, the greater the energetic benefits obtained. The exhibition or absence of trail laying behavior in the recruitment responses displayed by E. ruidum workers is discussed in relation to their involvement in scavenging or predatory behavior. Received: 27 June 1996 / Accepted after revision: 3 March 1997     

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     

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     

The role of turbidity as a constraint on predator-prey interactions in aquatic environments

Many of the world's most productive aquatic ecosystems usually contain turbid water. Paradoxically, many fish species that live in these habitats are also those that often rely on vision to detect their predators and their prey. For these fish, turbidity will reduce the distance at which predator-prey interactions occur, and there should be a reduction in the opportunity for behavioural modification to control the risk of predation. Under these conditions, we predict that most antipredator behaviour will become ineffective and that predator-prey interactions in turbid water will be primarily characterised by the direct effects of predator consumption of prey, rather than behavioural modification reducing the growth rates of prey. This hypothesis was tested in two laboratory experiments. The first experiment investigated how water turbidity, risk of predation, and their interaction affect habitat choice decisions by fathead minnows (Pimephales promelas). These data demonstrate that fathead minnows reduce their use of dangerous habitats, but that this effect is diminished in turbid water. A second experiment examined mortality patterns when these fish were preyed upon by yellow perch (Perca flavescens) in clear and turbid water. Absolute mortality rates were unaffected by visibility, but patterns of mortality were random in turbid water and skewed towards smaller individuals in clear water. Combined, these results support our hypothesis and suggest that the impact of predation risk will be reduced in turbid aquatic ecosystems. Received: 23 May 1996 / Accepted after revision: 18 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     

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     

Reproductive dominance and differential ovicide in the communally breeding burying beetle Nicrophorus tomentosus

Communal breeding can be characterized by the division of reproduction among cooperating individuals and, if the distribution of reproduction is inequitable, by the mechanisms for achieving skewed reproductive success. The burying beetle (Nicrophorus tomentosus) is a facultative communal breeder. Unrelated adults, especially females, provide extensive parental care to broods of mixed parentage. The frequency and degree of reproductive skew between two females were examined experimentally. On medium-size carcasses, the proportion of eggs attributed to each female was not significantly different from random in 42% of the broods, skewed in 42% and not shared in 16%. Although reproduction was usually skewed in favor of the larger female, the relative sizes of the two females did not predict the degree of skew. On large carcasses, the proportion of eggs attributed to each female was not different from random in 87% of the broods and weakly skewed in 13%. Several mechanisms for biasing reproductive success were investigated. Females increase the proportion of their offspring in the brood by committing differential ovicide. Secondly, burying and preparing a carcass cooperatively stimulates ovarian development of the larger female and slows it for the smaller female, reducing or delaying oviposition by the subordinate. Thirdly, larger females are more likely to be dominant and are more fecund than smaller females. Received: 20 July 1996 / Accepted after revision: 30 November 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     

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     

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     

Weapon size versus body size as a predictor of winning in fights between shore crabs, Carcinus maenas (L.)

Relative body size (carapace width) and weapon size (chela length) were used as indicators of resource holding potential (RHP) in the agonistic behaviour of male shore crabs, Carcinus maenas (L.). Weapon size was found to be a more reliable predictor of the outcome of pairwise fights than body size. Crabs with longer chelae than their opponents were more likely to win fights than crabs with relatively larger bodies. Body size had less influence on the outcome of fights. Relative body and weapon size did not influence initiation of contests but did affect the likelihood of winning; however, this was significant only for weapon size. Winning crabs had heavier claws with greater surface area than losing crabs. There was no relationship between relative size and fight duration. The frequency of cheliped display increased with chela length and win- ners performed significantly more displays than losers. Received: 5 February 1997 / Accepted after revision: 20 May 1997     

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     

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     

Testosterone stress does not increase asymmetry of a hormonally mediated sexual ornament in a lizard

The theory that fluctuating asymmetry is sensitive to both environmental and genetic stress is gaining acceptance among evolutionary biologists. Most empirical work has focused on ornamental traits on the assumption that they are more susceptible to stress. In tegumentary coloration is a common ornament in nature, and frequently has a hormonal basis. Earlier studies in the lizard Psammodromus algirus indicate that testosterone induces the development of head nuptial coloration in large males and, at the same time, produces an increase of the ectoparasite load and higher mortality. Hence, the manipulation of testosterone levels may be a way to increase ornament expression and simultaneously create conditions that may make symmetrical development difficult. This positive covariation between character size and symmetry is opposite to that expected in theory for sexually selected traits, so the predicted elevation of asymmetry due to the treatment cannot be confounded by any intrinsic association between symmetry and character size. We firstly consider the effect that testosterone supplementation has on two variables that reflect the symmetry of bilateral throat nuptial coloration in large P. algirus males. Also, we examine whether ornament symmetry is positively associated with reproductive success, a prediction of theory of sexually selected symmetry. Testosterone treatment did not increase the fluctuating asymmetry of throat coloration. Size asymmetry increased with character size in individuals with a fragmented colour pattern, but changed suddenly to a highly symmetric pattern in individuals with non-fragmented coloration. Mirror asymmetry decreased steadily with character size. These results suggest that the development of coloration on both sides of the throat midline follows a random pattern. Asymmetry did not correlate with variables that estimated reproductive success, suggesting that asymmetry is not affected by physiological stress and that this trait is not a sexually selected signal in P. algirus. Received: 1 July 1996 / Accepted after revision: 4 May 1997     

Experimental evidence for food limitation and sex-specific strategies of American kestrels (<Emphasis Type="Italic">Falco sparverius</Emphasis>) provisioning offspring

For birds with altricial young, the brood-rearing period is one of the most energetically expensive periods in their lives and may be the bottleneck for fitness. Because parents are expected to be prudent in allocating resources between reproduction and self-maintenance, food supply should be an important factor determining reproductive decisions during brood rearing. Parents with abundant food are expected to have enhanced fitness because they are able to reduce their work rates and increase their own survival chances, because their offspring may be of higher quality, or some combination of these. However, few studies have simultaneously documented all of these variables in a single investigation. We performed a food supplementation experiment to test how food supply influenced provisioning decisions by parent American kestrels (Falco sparverius). Female kestrels showed a strong response to extra food and reduced their provisioning rates. As a result, supplemented females had higher return rates than control females, suggesting significant effects of food on female survival. Because females used extra food to increase their fitness, our results suggest that kestrels raising offspring are limited by food. Male kestrels whose nests were supplemented also responded to extra food by reducing provisioning, but to a much lesser extent than their mates. Male parents did not appear to benefit from supplementation, as their return rates were similar to control males. The total amount of food received on a daily basis by nestlings was similar between supplemented and control nests. Supplemented offspring therefore did not fledge in better condition or have higher survival rates than control nestlings; the only significant factor consistently affecting offspring condition and survival was weather.     

Adjustment of parental effort in the puffin; the roles of adult body condition and chick size

 We examined the adjustment of parental effort of puffins by switching 20-day-old chicks randomly between parents of known body condition. Among unmanipulated birds mass gain (5–20 days) and mass of 20-day-old chicks was positively correlated with the body condition of parents at day 6. During the first 5 days after chick switching 28% (n = 55) of the parents deserted their foster chick. Parents which deserted their foster chick originally had a chick of their own that was smaller than that of those which did not desert their foster chick. Whether parents deserted their foster chick was also negatively related to the size of the foster chick. The mass of the foster chick was more important than the size of the parents' own chick in determining the desertion rate of chicks. The mass gain of the foster chick during the first 5 days after switching was positively related to the body condition of foster parents and also positively related to the mass of the foster parents' own chick, but negatively related to the size of the foster chick. The results suggest that puffins adjust their parental effort according to both their own body condition and the size of the chick. The latter may indicate the chick's prospect of survival and recruitment to the population. Received: 20 January 1996 / Accepted after revision: 27 October 1996     

Brood reduction in response to manipulated brood sizes in the common swift (Apus apus)

Following a brood size manipulation experiment on the common swift (Apus apus) in which different levels of parental effort were created, brood reduction occurred in all five nests manipulated to four chicks and in two of the five manipulated to three young. This provided an opportunity for looking in detail at the parental investment decisions concerning allocation strategies between parent and young during the process of brood reduction. The data recorded here were analysed on a visit-by-visit basis regarding changes in parental and chick body masses, the mass of prey delivered and the estimated mass of parental self-feeding. The results show that food delivery rates did not increase in proportion to the number of chicks in the broods. This reduction in delivery rates per chick resulted in lower chick masses and ultimately in the death of one or more chicks in the larger broods. The resulting low parental body masses for adult birds feeding larger broods suggested that these parents could not have raised all their young without risking their own survival. There was a tendency for parents from nests that experienced brood reduction to feed themselves instead of allocating most of the food gathered to the young. After brood reduction, parents regained lost body mass and their young fledged at masses only slightly lower than normal. The differential allocation decisions regarding parental self-feeding which resulted in brood reduction were largely responsible for keeping the parents in good enough condition to care for the surviving nestlings. Therefore, this study clearly demonstrates that brood reduction can operate through the differential allocation of food between parent and young in a way that can have adaptive consequences for the survival of parents and their young. Correspondence to: T.L.F. Martins     

Sexual cohabitation as mate-guarding in the leaf-curling spider Phonognatha graeffei Keyserling (Araneoidea, Araneae)

The leaf-curling spider Phonognatha graeffei incorporates a twisted leaf into the central hub of its orb-web that is used as a retreat. This species is unusual among orb-weaving spiders because males cohabit in the leaf retreat with both immature and mature females, mating with the former shortly after the female molts. Cohabitation appears to be a form of mate-guarding because cohabiting males respond agonistically to rival males that venture onto the web, and their behaviour depends upon the reproductive status of the female; males defending immature females are more aggressive than those defending virgin, adult females. Males copulate with previously mated females for significantly longer than with virgin females. Females may cannibalise cohabiting males, which occurs independently of whether the female has been deprived of food. Females that cannibalise a single male do not have a higher fecundity than non-cannibalistic females. Received: 2 February 1996 / Accepted after revision: 27 October 1996     

Food access, brood size and filial cannibalism in the fantail darter, Etheostoma flabellare

We compared the occurrence of filial cannibalism in fed and starved male fantail darters (Etheostoma flabellare). All males in the experiment consumed eggs, and 56% ate all of their eggs. A male's initial body condition did not explain the number of eggs that he ate. Neither did non-fed males eat more eggs than fed males. Fed males were able to maintain better body condition during the experiment, but the change in body condition also depended on the number of eggs eaten. Thus, males who ate more eggs were able to maintain better body condition.The most important determinant of whether or not a male ate all of his eggs was his initial egg number. Males with small egg masses ate all of their eggs whereas males with large egg masses were only partial cannibals. There was, however, no difference in the total number of eggs eaten by total and partial cannibals. We conclude that eggs are only partially eaten for energetic reasons. We also suggest that small egg masses are completely consumed because the costs of caring for a small egg mass may exceed the expected reproductive benefits of a small egg mass. Received: 26 January 1996 / Accepted after revision: 2 November 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