Are communal pukeko caught in the prisoner's dilemma?

Summary The prisoner's dilemma of game theory provides a possible explanation for communal breeding in the pukeko (Porphyrio porphyrio, Aves) where group breeding birds are at a reproductive disavantag compared to pairs. Territorial defence is largely the realm of males. A male who defect on neighbours by taking in an extra male defender and so becoming communal stands to gain territorial area and possibly higher reproductive success. Neighbours are forced to follow suit if additional males are available or lose their territory and therefore their breeding opportunities. If neighbours also admit an extra defender, the initial defector loses his advantages and all males end up with the lower breeding success of communal groups. Such defection to a communal habit appears to be the only stable solution.     

Components of lifetime reproductive success in communally and solitarily nursing house mice — a laboratory study

Under laboratory conditions, communal nursing among familiar and closely related female house mice (Mus domesticus) improved lifetime reproductive success compared to females rearing litters alone or females living with a previously unfamiliar, unrelated partner (reproductive success was measured within an experimental lifespan of 6 months, standardized as 120 days after mating at the age of about 2 months). An analysis of the contribution of three multiplicatively combined components to variation in reproductive success among breeding females revealed that, in all three social groups, survival of young until weaning contributed most to differences in lifetime reproduction (46–64% of the total variance). Females living with a sister had a significantly higher probability of reproducing successfully than females in the other groups, and also reared significantly more litters communally than females sharing nests with an unrelated partner. Weaning probabilities of young were highest in litters cared for by sisters and lowest in nests of unrelated females. Young were found dead either directly after birth (within the first 2 days of lactation) or after they had been cared for and nursed for at least 1 day. The loss of an entire litter typically occurred directly after birth. In monogamous females rearing litters alone the death of almost all young coincided with such early entire-litter mortality. In polygynous groups, however, offspring died at an older age and more litters suffered the loss of some young. Still, rearing young with a sister improved survival directly after birth and fewer litters were lost entirely in comparison with females in the other groups. In polygynous groups, pregnant females were observed to kill some of their partner's dependent young shortly before they gave birth themselves. As a consequence, individual young had reduced survival when they were firstborn in a communal nest (another litter was born within 16 days). Analyzed over a lifetime, communal care among familiar and closely related female house mice seems to be an adaptation to maximize the survival of offspring until weaning.     

Parental investment and pair-bond behavior among canvasback ducks (Aythya valisineria,Anatidae)

Summary Hypothetical patterns of male and female cumulative reproductive effort in canvasback ducks were used to predict the efforts of males and females in pair-bond reinforcing behavior through a breeding sequence. The predictions were that, at any point in time in a monogamous breeding system, given certain assumptions, (a) the sex investing less should be more active in reinforcing the pair bond, and (b) this relationship should shift within and between breeding attempts with the pattern of cumulative reproductive effort per brood by the male and female. Data on the behavior of newly-formed pairs were obtained by observing apparently paired but unmarked canvasbacks during spring migration. Later, 11 individuallymarked pairs were followed through breeding sequences, including repeated nesting attempts. Observed affiliative behavior fit the predictions reasonably well. That sex whose cumulative reproductive effort was less tended to be more active in maintaining proximity to its mate, initiating bouts of mutual display, and coordinating pair activities. Predictions were met in 14 of 24 cases, results were contrary in 3 cases, and in 7 cases trends were not statistically different (P>0.05). Shifts in relative male and female reinforcement efforts that coincided with major changes in investment were apparent. These results support the proposition that factors other than simple parental investment-related desertion costs might be important to an individual's behavioral decisions regarding pairbond reinforcement and ultimately mate abandonment. Discrepancies in a few predicted outcomes further suggest that males and females may use and balance various pair-reinforcement tactics differently through a breeding season.     

Dynamics of space use and male vigour amongst wood mice,Apodemus sylvaticus,in the cereal ecosystem

The spatial organisation of male and female wood mice,Apodemus sylvaticus, was investigated in a large-scale radio-tracking study on arable farmland near Oxford, United Kingdom, during the breeding season. Both sexes had significantly larger home ranges in the breeding season than at other times, and the breeding season home ranges of male (X = 1.44 ha) were significantly larger than those of females (X = 0.49 ha). Home range overlap was significantly greater between males, and between males and females, than it was between females. Overlap between males tended to be greatest in heavily utilised areas. Except during sexual consortship, there was minimal evidence of dynamic interaction among individuals. Home range sizes of breeding males varied widely, as did their body weights. There was no relationship between male body weight and home range size or any other movement parameter. However, males with the largest home ranges had the highest scores on all other movement parameters, indicating that they expended more energy in movement. These more vigorous males had access to the home ranges of more females than did males with small home ranges.     

Passing the buck: resource defence,lek breeding and mate choice in fallow deer

Summary Lek breeding systems, where males defend small, clustered mating territories, are thought to occur where the distribution of females is heavily clumped but males are unable to defend resources used by females. In this paper, we describe a breeding system in fallow deer where males are able to defend resources used by females but the most successful bucks instead defend small territories on a traditional mating ground; where the lek is sited in an area not heavily used by females at other times of year and is visited primarily by females in or close to oestrus; and where mating success on the lek is related to territory position and to male phenotype but not to the resources available on different lek territories. Comparisons with other ungulates suggest that lek breeding species fall into two groups: those where leks are regularly visited by herds of females many of which are not in oestrus and those, like fallow deer, where leks are visited primarily by oestrous females. In the latter species, it is unlikely that females visit the lek for ecological reasons.     

What do pairing patterns in common tern, Sterna hirundo, recruits reveal about the significance of sex and breeding experience?

In general, reproductive output in long-lived bird species increases in older compared to younger individuals. Therefore, experienced mates should be advantageous for first-time breeders. To examine requirements and consequences of experienced pair mates we investigated the first pair bonds of common terns, Sterna hirundo, recruiting to their natal colony. We found that male recruits were usually the same age as their mates, whereas female recruits were usually the younger member of the pair. In order to acquire experienced mates, it was necessary for males to arrive early in the year of recruitment. Mates with 2 or more years of breeding experience were only attainable by male recruits characterised by greater body mass and age. Female recruits arrived more than 1 week later than their experienced mates and significantly later in the season than male recruits paired with experienced females. In general, females first bred at a younger age than males, and neither the female recruits body mass nor their age was related to the level of experience of their first mate. These sex-specific differences in obtaining an experienced mate did not result in different levels of reproductive success between the sexes. Male and female recruits with mates with 2 or more years breeding experience benefited from having experienced mates: they had greater reproductive success. First-time breeders paired with mates with only 1 year of breeding experience did not differ from pairs where both members were breeding for the first time in terms of reproductive success, but clutches were larger. Our results illustrate not only different prerequisites for males and females, but also males need for experienced mates. Delayed male breeding (postponing breeding for another year), supposed to be a negative trait, and high body mass, supposed to be a trait of superior individual quality, can be combined in some individuals, improving reproductive success and showing that breeding common terns use a range of tactics to begin reproduction.Communicated by F. Trillmich     

Arrival timing and seasonal reproductive performance in a long-distance migratory landbird

The date when a landbird migrant arrives on its breeding grounds may have reproductive consequences. Generally, early arriving individuals begin breeding earlier and consequently experience greater seasonal reproductive performance. Here, we describe relationships between arrival timing and seasonal reproductive performance in the American redstart (Setophaga ruticilla), a long-distance passerine migrant, arriving at northerly breeding grounds in Michigans eastern Upper Peninsula. Evidence suggests that both males and females benefited from early arrival at the breeding grounds. Early males appeared to settle on higher quality territories and hatched nestlings sooner than later arrivals. Early females began their clutches early, produced heavier nestlings and possibly laid more eggs than later arrivals. Larger clutches and heavier offspring increase the likelihood of offspring recruiting into the breeding population. The findings of this study point to fitness consequences arising from when a bird arrives at its breeding grounds. These results also have implications for understanding how events occurring during spring migration influence reproductive performance as migratory delays likely influence arrival timing.Communicated by: W. Wiltschko     

Reproductive effort in Adélie penguins

Summary We estimated reproductive effort (energy expenditures for reproduction, as opposed to maintenance) in Adélie penguins breeding at Palmer Station, Antarctica. Data on body composition changes and metabolic rate were obtained using isotopic methods. Adelie breeding behavior consists of an initial courtship stage (during which both sexes fast), incubation, the guard stage (when chicks are 1 to 18–28 days old), and the creche stage (from the end of guarding until chicks are 28–45 days old). Both males and females lost considerable mass during the initial stages of the reproductive season, but males fasted longer and lost more mass. Mass losses of both sexes consisted of 66% depot fat and 34% lean tissue. Mass and body composition remained constant once birds resumed feeding. The metabolic expenditure for the foraging necessary to accumulate the mass lost while fasting — one component of reproductive effort —was about 63 MJ in males and 39 MJ in females. Field metabolic rates (FMR) were low during courtship and while incubating, increasing more than 2-fold when birds resumed foraging. Although mean FMR increased between incubation and the creche stage, differences between stages were small and not significant. We used FMR data and an energy balance model to estimate the cost of feeding chicks. Results suggest a maintenance FMR of about 2.7 × basal metabolism (BMR), increasing to 3.4–3.6 × BMR during the creche stage. The reproductive effort (as metabolic expenditures) associated with feeding chicks is 31 MJ (males) to 36 MJ (females). Cumulative reproductive effort is 94 MJ in males and 75 MJ in females, or 5.3–6.2% of the annual energy budget. The reproductive effort devoted to chick care does not appear to be constrained by physiological or time limitations. Instead, selection to reduce the risk of predation may prevent the evolution of increased parental care. Correspondence to: M.A. Chappell     

Operational sex ratios and behavioural sex differences in a pipefish population

In the pipefish Syngnathus typhle, only males brood embryos in specially developed brood pouches, supplying oxygen and nutrients. Laboratory studies have shown that this elaborate paternal care has led to sex-role reversal in this species: males limit female reproductive rate, females are the primary competitors for mates and males exercise greater selectivity in accepting mates. In the first field study of this pipefish, we describe mating behaviour in the wild and test the hypothesis that temporal variations in the operational sex ratio (OSR) determine sex differences in mating behaviour. Our study comprised two reproductive seasons of two sequential mating periods each, the latter separated by a lengthy interval of male brooding. During mating periods, females displayed to all males without wandering and males moved about searching for females, without reacting to all females. The OSR was least female-biased (or even male-biased) at the onset of the breeding season, when most pipefish were simultaneously available to mate, but became strikingly female-biased as males' pouches were filled. The OSR remained substantially female-biased during the second mating period, because few males became available to remate at any one time. As hypothesised, female-biased OSRs resulted in more female-female meetings. As well, females were above the eelgrass more often than brooding males, thus exposing themselves to conspecifics and/ or predators. In the second year, males arrived earlier than females on the breeding site and male pregnancies were shorter, because of higher water temperatures, so rematings occurred earlier. Males met more often during that year than the previous one, but male competitive interactions were still not observed. The field results support laboratory studies and demonstrate that behaviours associated with female-female competition are more prominent when the OSR is more female-biased. Correspondence to: A. Vincent     

Sex differences in the energy allocation strategies of house mice

Summary Sex differences in energy allocation were studied in wild stock house mice (Mus domesticus). Peripubertal animals of both sexes, either intact or gonadectomized, were subjected for 8 weeks to one of two feeding requirements by using a caging system in which a pellet dispenser was controlled by activity on a running wheel. Recently weaned animals were required to run 200 vs 300 or more wheel revolutions to obtain a pellet of food. The 200 revolution requirement allowed normal body growth and reproductive development; the 300+ requirement was adjusted weekly to maintain food intake at a level that allowed survival but did not permit normal body growth. Reproductive development was completely inhibited in intact females at 300+ revolutions whereas intact males at 300+ revolutions, despite stunted growth, all experienced normal sexual development. At both feeding requirements, however, females exhibited more total locomotor activity and consumed more food than males, regardless of gonadectomy. Furthermore, at the 200 revolution requirement we often observed extensive running activity beyond that needed to generate the amount of food actually eaten, especially among females. This extra locomotor activity was gonad-dependent in males, but not so in females. These results suggest that male and female house mice employ different strategies when relating their behavior and reproductive development to existing foraging conditions. Females appear more resource-dependent than males.     

Aggression in female Red-Winged Blackbirds: A strategy to ensure male parental investment

Summary Female Red-winged Blackbirds (Agelaius phoeniceus) are often aggressive towards conspecific females during the breeding season. We hypothesize that the function of female-female aggression in this species is to guard the nonshareable portion of the male's parental investment.The investment-guarding hypothesis predicts that a female should be more aggressive toward another female evincing interest in mating with the territory-owning male than toward a female simply perching within the male's territory. Results of mount presentations to females with active nests confirmed this prediction. Nesting females attacked a stuffed conspecific female mounted in a precopulatory, soliciting posture significantly more often than a mount in a normal, perched posture.The male's nonshereable parental care consists of provisioning his young, and most of this care is invested in the brood of his primary (first-to-nest) female. It is therefore predicted that primary females should be more aggressive than secondary (later-nesting) females. Female mount presentations also confirmed this prediction. Primary females attacked the soliciting mount significantly more often than secondary females.     

The effect of intrauterine position on the survival,reproduction and home range size of female house mice (Mus musculus)

Summary In laboratory studies using albino house mice, a female's prior intrauterine position can affect many postnatal physiological, morphological and behavioral characteristics. Females flanked by males in utero (2M females) exhibit more aggressive dominance than females flanked by females (OM females). Thus, wild 2M females may be most successful during peak population densities when their aggressive nature would allow them to displace other females from limited resources. 2M and 0M females and males delivered by cesarean section were individually marked and released as young adults on two occasions onto a highway island (the area enclosed by exit and entrance ramps at an interchange) to determine whether 2 M females have a competitive advantage over 0 M females in the field. Males were included to create realistic population structure; their intrauterine position was not a treatment. Feeding stations afforded individuals an opportunity to exhibit their dominance by maintaining home-ranges at or near the stations. The populations were monitored by periodic live-trapping and reproductive success was determined using field body weights and by post-mortem examination for uterine implantation scars. Survival and capture rates were estimated, using a modified Jolly-Seber markrecapture program, for each of four intervals between trapping occasions over the course of 7 weeks. There were no overall differences in survivorship between 2M and 0M females, neither type of female was caught more frequently at feeding stations and they did not differ in measures of reproductive success. However, 2M females had significantly larger home-range sizes than 0M females and thus space use may be a trait masculinized by prior intrauterine position. Although there are a number of life-history characteristics that differ between 0M and 2M females in the laboratory that we did not test specifically in the field, our findings and other features of wild house mouse biology suggest that prior intrauterine position does not have a strong effect on survival and reproduction in the wild. Offprint requests to: W.J. Zielinski at the present address     

Mate guarding,copulation strategies and paternity in the sex-role reversed,socially polyandrous red-necked phalarope<Emphasis Type="Italic"> Phalaropus lobatus</Emphasis>

In a recent review, Westneat and Stewart (2003) compiled evidence that extra-pair paternity results from a three-player interaction in which sexual conflict is a potent force. Sequentially polyandrous species of birds appear to fit this idea well. Earlier breeding males may attempt to use sperm storage by females to obtain paternity in their mates subsequent clutches. Later-breeding males may consequently attempt to avoid sperm competition by preferring to pair with previously unmated females. Females may bias events one way or the other. We examined the applicability of these hypotheses by studying mating behavior and paternity in red-necked phalaropes (Phalaropus lobatus), a sex-role reversed, socially polyandrous shorebird. Male red-necked phalaropes guarded mates more strongly than other shorebirds. Males increased within-pair copulation attempts during their mates fertile period, and maintained or further increased attempts towards the end of laying, suggesting an attempt to fertilize the females next clutch; these attempts were usually thwarted by the female. Paired males sought extra-pair copulations with females about to re-enter the breeding pool. Multilocus DNA fingerprinting showed that 6% of clutches (4/63) each contained one chick sired by a male other than the incubator, producing a population rate of these events of 1.7% (n=226 chicks). Male mates had full paternity in all first clutches (n=25) and 15 of 16 monogamous replacement clutches. In contrast, 3 of 6 clutches of second males contained extra-pair young likely fathered by the females previous mate. Previously mated female phalaropes may employ counter-strategies that prevent later mating males from discriminating against them. The stability of this polyandrous system, in which males provide all parental care, ultimately may depend on females providing males with eggs containing primarily genes of the incubating male, and not a previous mate.Communicated by M. Webster     

Genetic reconstruction of breeding patterns in gundis (Rodentia: Ctenodactylidae)

Current mating system theory predicts that the number of females breeding in a group will depend on the number of females in the group and the accessibility of unrelated males, whereas the number of males breeding in a group will depend on the ability of males to control access to reproductive females. By combining information on group composition with genetic data, I determined whether breeding patterns in a rock-dwelling rodent, Ctenodactylus gundi, were concordant with these expectations. C. gundi breeding patterns varied from facultative monogamy to uni-male polygyny to multi-male polygyny. The number of reproductive individuals of each sex in a group increased with group size. Whereas communal breeding among related females tended to increase female reproductive success, males breeding in the same group were unrelated to other group members and seemed to compete over access to matings. While some males were assigned offspring from neighboring social groups, most group-living males successfully monopolized the reproduction of females within their group. There was no evidence that females had multiply sired litters, although some bred with alternative males in separate breeding attempts. Although numerous individuals were not assigned as parents or offspring, genetic information enabled me to determine that most unassigned individuals were philopatric group members, whereas only a few were unrelated immigrants into their current social group. Together, these results provide evidence that C. gundi social groups represent fairly distinct reproductive units whose breeding patterns are dependent on group size and composition in accordance with theoretical predictions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Foraging decisions of American robins: somatic and reproductive tradeoffs

Summary Breeding units (occupants of a nest including at least one reproductive female) within two free-living populations of the prairie vole, Microtus ochrogaster, were monitored by live-trapping at nest during two 28-h periods each week from October 1980 to March 1984. Data are presented for 281 breeding units from all seasons, at high and low population densities and during breeding and nonbreeding periods. Fifty percent of the breeding units were monogamous (single resident reproductive male and female), 27% consisted of a single reproductive female with no resident adult male and 23% included more than one resident adult male and/or female (complex units). Monogamous units were present in the same proportions during breeding and nonbreeding periods. The number of monogamous units was significantly greater at low population densities than at high densities. During winter there were relatively more complex units and fewer single female units than during the rest of the year. Monogamous pairs remained together for an average of 42 days. Seventy-eight percent of these pairs were disbanded by the death of one or both members. There were few overlaps of the home ranges of adjacent breeding units. Significantly more nests were visited by nonresident males than by females, and the intervals between visits by males were significantly shorter than those for visits by females. Males visited single female units significantly more often than units with one or more resident males. Survival of juveniles was generally very low; 38% and 34% of young males and females, respectively, that were trapped survived until 30 days of age. Of young females remaining at the natal nest at low population densities, only 17.6% were reproductively activated; 77.1% of such females became reproductively activated at high densities. All young females that dispersed from the natal nest became reproductive.     

Variation in copulation duration in Mnais pruinosa pruinosa Selys (Odonata: Calopterygidae)

Summary The Japanese calopterygid damselfly Mnais pruinosa pruinosa shows three distinct copulation durations (Siva-Jothy and Tsubaki 1989). A variety of factors which might influence copulation duration were investigated. Results indicate that the temperature in the vicinity of copulating pairs, the rate at which males encountered receptive females, territory quality, the rate of agonistic interactions between males, female gravidity and female willingness to oviposit have no effect on copulation duration. The most significant factor was the location of the site at which males captured their mates. We suggest that under natural conditions the location of the capture site provides males with reliable, indirect information about a female's intention to oviposit: it is important for males of this species to acquire such information since sperm precedence, and therefore reproductive success, is in part dependant on the interval experienced by females between copulation and subsequent oviposition. The results are discussed and it is suggested that the three observed mate-securing tactics of M.p. pruinosa are facultative and information-dependant.     

Group size in wedge-capped capuchin monkeys Cebus olivaceus and the reproductive success of males and females

Summary The effect of variation in group size on age-specific survivorship and fecundity rates were examined in a population of wedge-capped capuchin monkeys Cebus olivaceus during a 10 year study. Life tables were constructed separately for four large (15 individuals) and four small groups (<15 individuals). Female reproductive success, and its relative contribution to population growth, was much higher in large groups, primarily through higher age-specific fecundity. Age-specific survivorship was similar in groups of different sizes. The reproductive success of the single breeding male in a group was much higher in large than small groups. Compared to small groups, breeding males in large groups had a longer breeding tenure, and access to greater numbers of reproductive females with a higher average fecundity. Differences in female reproductive success apparently resulted from variation in access to monopolizable fruit trees. Large groups predictably displaced small groups during intergroup encounters. Group rank depended on the number of males resident in groups. The large number of non-breeding males in large groups results from their longer average residency time. I explain the longer residency of males in large groups by the higher average reproductive success of breeding males in these groups.     

Male mating history and female fecundity in the Lepidoptera: do male virgins make better partners?

In insects, large ejaculate and associated materials, including spermatophores, appear to have evolved via sexual selection acting on males to either delay female remating or to increase the rate of egg-laying. It is also possible, however, that females use nutrients transferred during mating to increase their lifetime fecundity. If so, male ejaculate size may also have evolved under natural selection as a form of paternal investment. In Lepidoptera, males with a greater number of prior matings tend to produce smaller spermatophores. However, the reported effects of male mating history on female fecundity vary widely among species. We therefore performed a meta-analysis using data from 29 studies of 25 species. Overall, the reproductive output of females mated to virgin males was significantly higher than that of females mated to sexually experienced males (Hedges d=0.33, P<0.01). A sample size of around 145 females per male mating type is required to detect an effect of this size with 80% statistical power at =0.05 (two-tailed). There was no difference in mean effect size between butterflies/skippers and moths. After controlling for any effect of taxonomic group, however, the mean effect size for polyandrous species was significantly greater than that for monandrous species (Hedges d=0.45 vs 0.25, P=0.01). We then discuss possible reasons why male mating history, presumably acting through its effect on spermatophore size, might have a stronger effect in polyandrous than monandrous species.Communicated by A. Cockburn     

Why do female mice mate with multiple males?

Females often show multi-male mating (MMM), but the adaptive functions are unclear. We tested whether female house mice (Mus musculus musculus) show MMM when they can choose their mates without male coercion. We released 32 females into separate enclosures where they could choose to mate with two neighboring males that were restricted to their own territories. We also tested whether females increase MMM when the available males appeared unable to exclude intruders from their territories. To manipulate territorial intrusion, we introduced scent-marked tiles from the neighboring males into males' territories, or we rearranged tiles within males' own territories as a control. Each female was tested in treatment and control conditions and we conducted paternity analyses on the 57 litters produced. We found that 46 % of litters were multiply sired, indicating that multiple paternity is common when females can choose their mates. Intrusion did not increase multiple paternity, though multiple paternity was significantly greater in the first trial when the males were virgins compared to the second trial. Since virgin male mice are highly infanticidal, this finding is consistent with the infanticide avoidance hypothesis. We also found that multiple paternity was higher when competing males showed small differences in their amount of scent marking, suggesting that females reduce MMM when they can detect differences in males' quality. Finally, multiple paternity was associated with increased litter size but only in the intrusion treatment, which suggests that the effect of multiple paternity on offspring number is dependent on male–male interactions.     

Divorce in common murres (Uria aalge): relationship to parental quality

Behavioral precursors of 12 divorces were examined in 30 color-banded pairs of common murres (Uria aalge) over six breeding seasons. Common murres are long-lived seabirds that typically return each year to the same mate and nest site in dense colonies. At least one parent is present continuously from egg lay to chick fledging. Murres, therefore, have considerable opportunities to compare their mates parental behavior with that of several familiar neighbors. Previous reproductive success was lower for divorcing birds than for reuniting pairs. As predicted by the better option hypothesis, there were clear choosers (seven females and five males) that initiated divorce by moving to a new birds site or by courting a new partner at their current site. Choosers improved their reproductive success after the divorce, whereas their previous partners, the victims did not. Yearly divorce rates (average 8.2% per year) were significantly correlated with yearly mortality rates. Divorces appear to be opportunistic: pairs divorced after varying numbers of reproductive failures with the immediate precursor usually being the disappearance (death) of a murre from a successful neighboring site. In contrast to the delays experienced by victims, choosers formed new pairs quickly and laid their eggs no later than reuniting pairs. Prior to the divorce, victims fed their chicks less often than choosers, and some engaged in other behaviors that compromised egg or chick survival. These observations suggest that deficiencies in parental behavior were precursors to the divorce. This report is one of the first cases where reproductive failure of divorcing pairs has been linked to deficits in the parental behavior of the subsequent divorce victim.Communicated by C. Brown