Embryonic and parental preferences for incubation temperature in herring gulls: implications for parent-offspring conflict

According to Trivers (1974), parent-offspring (P-O) conflict arises because offspring are selected to solicit more care than parents are selected to provide. However, should benefits fail to increase with increasing care, the offspring optimum can be reduced to the point where predicted P-O conflict vanishes. We examined offspring demand and parental care in such a benefit-limited system in herring gulls (Larus argentatus). In this species, parents typically neglect their lasthatched (C-) egg during the final hours of hatching (pipped-egg stage), allowing mean temperature to drop by about 4°C, to near 33°C. Other studies indicate that no increased offspring benefit arises from increasing pipped egg incubation temperature above that level, but embryo damage occurs if temperature drops lower. In such a system, P-O conflict over preferred incubation temperature is predicted to be minimal or absent. We assessed phenotypic manifestations of conflict by determining incubation temperature preferences of parent and offspring independently. Temperature provided solely by parental initiative was 33.9°C (artificial eggs, corrected for embryonic heat production). Preferred incubation temperature of pipped embryos was measured by exposing them to moderate chilling (20°C) punctuated by 4-min periods of rewarming when they called. Temperature of vocally thermoregulating embryos stabilized around a mean of 32.9–33.4°C, about 0.5–1.0°C below parental preference. Acting independently, parents and embryos each maintained egg temperature at or near minimum developmentally safe levels. Results provided no evidence for phenotypic conflict, as predicted by a benefit-limited version of Trivers' P-O conflict model. Benefit limitation may also be relevant to P-O conflict in other contexts such as feeding of newly-hatched young.     

Yolk hormones and sexual conflict over parental investment in the pied flycatcher

Female birds might be able to manipulate the parental effort of their male partner through elevated transfer of hormones to the eggs, since these hormones affect many chick traits that males might use as cues for adjusting the level of their investment. We experimentally studied whether female pied flycatchers Ficedula hypoleuca could manipulate male investment via yolk androgens. There is much more variation in yolk androgen levels between females than within clutches, and in order to change the androgen levels of the eggs, we swapped whole clutches between nests. To estimate the androgen levels of the clutch, we measured the androgen content of a single egg per clutch. Females did not succeed in manipulating male effort using yolk androgens, since there was no relationship between the division of parental care within a pair and either original or foster egg androgen levels. One of these relationships should have occurred if females were manipulating males. The proportion of feeding visits by the male was higher when the male was old (55%) than when he was young (45%) and females laid eggs with higher androgen levels when mated with a young male. Young males did not exhibit any responses to yolk androgen levels either, which indicates that females cannot exploit their effort more than that of old males. We suggest that females may allocate yolk androgens to adjust the growth trajectories of the chicks to poor growing conditions when mated with young males that are poor providers or occupying a poor territory.     

Biased parental investment and reproductive success in Gabbra pastoralists

Demographic data from 848 Gabbra households are used to examine the relationships between herd size and reproductive success in relation to sex, in a traditional, pastoralist population. The number of camels in the household herd has a significant positive effect on the reproductive success of both men and women, although the effect of wealth is greater for men, as predicted from evolutionary theory. The greater the number of elder brothers a man has, the lower his reproductive success, as a result of a smaller initial herd and a later age at marriage. This is not true for women –number of elder sisters does not have a measurable effect on a woman’s fertility, although it does have a small, negative effect on the size of her dowry. These results are interpreted as competition between same-sex siblings for parental investment, in the form of their father’s herd, which is more intense between sons than daughters as parental investments are greatest in males. Received: 30 June 1995/Accepted after revision: 23 October 1995     

Intra-group relatedness affects parental and helper investment rules in offspring care

In any system where multiple individuals jointly contribute to rearing offspring, conflict is expected to arise over the relative contributions of each carer. Existing theoretical work on the conflict over care has: (a) rarely considered the influence of tactical investment during offspring production on later contributions to offspring rearing; (b) concentrated mainly on biparental care, rather than cooperatively caring groups comprising both parents and helpers; and (c) typically ignored relatedness between carers as a potential influence on investment behavior. We use a game-theoretical approach to explore the effects of female production tactics and differing group relatedness structures on the expected rearing investment contributed by breeding females, breeding males, and helpers in cooperative groups. Our results suggest that the breeding female should pay higher costs overall when helpful helpers are present, as she produces additional offspring to take advantage of the available care. We find that helpers related to offspring through the breeding female rather than the breeding male should contribute less to care, and decrease their contribution as group size increases, because the female refrains from producing additional offspring to exploit them. Finally, within-group variation in helper relatedness also affects individual helper investment rules by inflating the differences between the contributions to care of dissimilar helpers. Our findings underline the importance of considering maternal investment decisions during offspring production to understand investment across the entire breeding attempt, and provide empirically testable predictions concerning the interplay between maternal, paternal and helper investment and how these are modified by different relatedness structures.     

Red-winged blackbird parental investment following brood parasitism by brown-headed cowbirds: is parentage important?

Parental investment by red-winged blackbirds (Agelaius phoeniceus) in response to natural and experi‐mental parasitism by brown-headed cowbirds (Molothrus ater), and in response to freeze-dried, female cowbird mounts presented near redwing nests during the egg-laying period was measured. Two measures of redwing parental investment were used: nest defense effort toward a model predator, and rate of feeding nestlings. There were no significant differences in levels of parental investment among unparasitized nests, naturally parasitized nests, or experimentally parasitized nests. Similarly, parental investment did not differ between redwings that were exposed to the cowbird mount and those that were not exposed to the mount, or among redwings exposed to the cowbird mount at different distances from the nest. This suggests that red-winged blackbirds do not recognize when they have been parasitized, and hence do not associate parasitism with a decrease in their parentage, or that parentage is not an important predictor of parental investment in this species. Received: 24 January 1997 / Accepted after revision: 7 June 1997     

The relationship between presumed gamete contribution and parental investment in a communally breeding bird

Summary Measures of relative gamete contribution were related to degree of parental investment for both males and females from six groups of communally breeding pukeko (Porphyrio porphyrio). If there is a fitness cost to participation in parental duties, then for males parental investment should relate to degree of confidence of paternity while for females investment should relate to the number of eggs laid. Data were analysed in two ways: (1) by looking at individuals of the same sex within the same group to see if males that had a greater probability of paternity and females that had greater number of eggs in a nest subsequently invested relatively more in parental duties and (2) by pooling the data from the six groups and determining if there was a positive correlation between measures of presumed gamete contribution and amount of parental investment. The results from both types of analyses are variable and give no clear support for a positive relationship between presumed gamete contribution and parental investment even though a possible fitness cost is suggested. Factors which might influence investment are discussed.     

Within- and between-individual (co)variance partitioning reveals limited pleiotropic effects of testosterone on immune function,sexual signaling,and parental investment

Behavioral Ecology and Sociobiology - An individual’s home range, or how much space it requires to obtain resources and meet its needs for survival and reproduction, affects the scale of many...     

High paternal investment in unrelated young: extra-pair paternity and male parental care in house martins

The response of males to reduced paternity has important consequences for the evolution and maintenance of a mixed reproductive strategy. Paternity is predicted to affect directly the level of male parental care in some cases but not in others. The response of males to reduced paternity will be influenced by their ability to assess their paternity, the predictability of cuckoldry and the costs and benefits of parental care. Although male house martins (Delichon urbica) provide among the highest levels of male parental care known in passerines (incubation, brooding and feeding nestlings), there was no evidence that cuckolded males substantially reduced their level of parental care, and, as a result, all young fledged successfully. Thus, extra-pair fertilizations enhanced the reproductive success of some males because they were able to parasitize the parental care of cuckolded males. We discuss several conditions which may favor extensive male parental care even when the male's paternity is very low.     

Cap-and-trade, taxes, and distributional conflict

Enacting market-based environmental regulation, such as emissions taxes and cap-and-trade programs, often create rents that are contested by agents. In this paper, we create a framework that compares social welfare from alternative market-based environmental policy instruments under the presence of rent seeking. We show that, contrary to the commonly held view, non-revenue-raising instruments (NRRIs) are in many cases preferable over revenue-raising instruments (RRIs). We find that the choice of instrument depends on the size of a potential revenue-recycling effect and the level of preassigned rents.     

Optimality of parental investment: The influence of nursing on reproductive success of mother and female young house mice

Summary The amount of parental milk investment determines not only the number of young the parent can produce, but also affects the offspring's fitness. The antagonism between quantity and quality of offspring was investigated in laboratory mice.In nursing first litters containing 2,4, ..., 14 pups, mothers invested increasing amounts of milk. This mainly extended the intervals between their first and second litters and slightly increased the size of their second litters, whilst third litters were not affected. In the female young, the decreasing amount each individual received as a result of increasing litter size led to delayed birth and reduced the size of their first litters. The intervals between their first and second litters and the size of the second litters were also affected, although to a lesser extent.Taking these results into consideration on a standard lifetime pattern of reproduction, the effects were calculated on net reproductive rates as an expression of fitness. Death rates were assumed that referred to population equilibrium or population growth. In both situations, the fitness curves of mothers and young showed that the actual level of milk investment and the level yielding optimal reproduction were in agreement. There was no significant deviation towards increased investment, as might result from parent-offspring conflict.     

Measurement of parental investment and sex allocation in the European beewolf Philanthus triangulum F. (Hymenoptera: Sphecidae)

Fisher’s 1930 theory of sex allocation predicts a population-wide 1:1 ratio of parental investment. We tested this prediction in the European beewolf, a sphecid wasp that hunts for honeybees as larval food. Because the method to quantify parental investment is of crucial importance, we compared the suitability of several different investment measures. Female/male cost ratios were determined from a sample and the total investment in sons and daughters was calculated. In addition, the actual number of prey items for sons and daughters was directly determined by excavating nests and counting the cuticle remains of the prey. Though mortality was high (70%), it had only a weak effect on the estimate of the investment ratio. Based on commonly used measures like fresh and dry weight of emerged adults, the investment ratio did not deviate from Fisher’s prediction of equal investment. However, progeny weight considerably underestimates investment in males and investment in large progeny. Measures that reflect the allocation of resources more directly (amount of provisions, brood cell volume) revealed a significant male bias and thus contradicted Fisher’s theory. Three kinds of explanation are discussed. First, non-adaptive explanations are unlikely. Second, from the spectrum of alternative adaptive theories, only models that assume a non-linear relationship between amount of investment and progeny fitness seem to be relevant for the study species. Third, though the number of prey in a brood cell seems to be a rather good measure of parental investment in European beewolves, some problems in measuring parental investment remain. These problems are of broad significance. Received: 17 June 1999 / Received in revised form: 6 July 1999 / Accepted: 11 July 1999     

The effects of nesting stage,sex, and type of predator on parental defense by killdeer (Charadrius vociferous): testing models of avian parental defense

Summary Two models predicting the temporal patterns of parental investment in offspring defense over the nesting cycle were tested. The first is based on offspring age, the other on the vulnerability of offspring to predation. Both models make very similar predictions for altricial species after eggs have hatched, i.e., increases in intensity of parental defense until fledging. For precocial species, however, the post-hatching predictions of each model are different: the offspring age model predicts a continued increase in defense intensity, while the vulnerability model predicts a decline. I examined the temporal patterns of parental defense of a precocial shorebird, the killdeer (Charadrius vociferus), and determined which model was supported. Killdeer responses to human and natural predators were observed. Killdeer were less willing to leave the nest, responded most intensely, and displayed closest to a potential predator around hatching. Defense intensity increased from early to late incubation as predicted by the offspring age model. However, after hatching killdeer parental defense declined for both males and females, thus supporting the vulnerability model for this stage. Males and females responded significantly differently to all types of predators. Males took greater risks, remained on the nest longer, defended offspring more intensely, and displayed closer to the predator than females at the approach of a potential predator. Responses to natural predators depended on the type of predator and the approach made by the predator; a greater range of defense behavior was used for predators approaching on the ground compared to aerial predators. In general, killdeer responses to humans were more intense and less variable than their responses to natural predators. This was most likely because the human intruder approached nests and chicks more directly and closer than natural predators.     

Parental conflict and brood desertion by females in blue-headed vireos

We investigated sexual conflict over parental care in blue-headed vireos (Vireo solitarius) and documented the first example of unvarying unisexual brood desertion in passerines. Females at all nests (N = 24) that were monitored closely near fledgling, deserted their broods on or near the day of fledging leaving males alone to complete parental care of young. No males deserted. This observational evidence was confirmed with radiotracking of females (2004, 2007) and both pair members (2008). Radiotracked females began visiting distant males 1–4 days before young left the nest, subsequently paired with males 355–802 m away, and laid first eggs in new nests less than 5 days after deserting. In contrast, females suffering nest predation did not desert and renested with the same male. We suggest equal parental care (nest building, incubation, feeding) in the sexes, genetic monogamy, and an adult sex ratio biased towards males has led to female control of brood desertion in this species. Unisexual desertion may be more important in altricial birds than generally realized and we discuss prerequisites to predict its occurrence. One is genetic monogamy, which may be a female tactic that reduces the likelihood of males evolving counter-adaptations to female desertion.     

Intergroup conflict,ostracism, and the evolution of cooperation under free migration

Theoretical investigations and quasi-experimental evidence from modern conflict areas suggest that intergroup conflict and ostracism play a pivotal role in the evolution of cooperation. However, there is little direct evidence about the influence of intergroup conflict on human social behavior in the presence of endogenous group formation and unrestricted migration. This study introduces an experiment to examine the impact of intergroup conflict and ostracism on group formation, and human cooperation in a dynamic environment where group size, the occurrence of intergroup hostility and the threat of ostracism are endogenously determined. Here, we show that intergroup conflict may inhibit merging into single large human coalitions. The threat of ostracism is shown to increase the average group size and cooperation within the society. In addition, we find that competitive pressure between groups decreases the likelihood of social exclusion. Our results suggest that free migration between groups suppresses the impact of between-group competition on within-group cooperation. Moreover, our results stress the role of ostracism as a means to repress competition within groups and enhance group success in competition against other groups.     

Independence, not conflict, characterizes dart-shooting and sperm exchange in a hermaphroditic snail

Although the sexes are united in hermaphrodites, conflict can still occur because the male and female functions have separate interests. We examined the evidence for conflict in the mating system of the terrestrial snail Cantareus aspersus (formerly Helix aspersa) where sharp, calcareous darts are ‘shot’ during courtship. We predicted that the use of the dart would either reflect or create conflict and this would be evident in either the courtship behavior or the transference of sperm. Previous studies demonstrated that the dart functions after sperm transfer to increase sperm survival. Using detailed observations of mating snails, we examined the factors that determine dart shooting order, the behavioral responses after being hit by a dart, the accuracy of dart shooting, and the allocation of sperm resources. We found that each dart was shot independently, and each animal appeared to be interested only in getting off the best possible shot, probably one that penetrates deeply near the genital pore. There is no evidence of mating conflict. Every snail transfers sperm to its partner, and the size of the donation does not depend on the success or failure of either snail’s dart shot. Although the receipt of a dart does not appear to cause harm, it may produce indirect costs due to the partial loss of control over fertilization. We conclude that mating in C. aspersus is a partnership in which independent actors demonstrate unconditional reciprocity during courtship and sperm transfer.     

Bridging the generation gap in plants: pollination, parental fecundity, and offspring demography

Despite extensive study of pollination and plant reproduction on the one hand, and of plant demography on the other, we know remarkably little about links between seed production in successive generations, and hence about long-term population consequences of variation in pollination success. We bridged this "generation gap" in Ipomopsis aggregata, a long-lived semelparous wildflower that is pollinator limited, by adding varying densities of seeds to natural populations and following resulting plants through their entire life histories. To determine whether pollen limitation of seed production constrains rate of population growth in this species, we sowed seeds into replicated plots at a density that mimics typical pollination success and spacing of flowering plants in nature, and at twice that density to mimic full pollination. Per capita offspring survival, flower production, and contribution to population increase (lambda) did not decline with sowing density in this experiment, suggesting that typical I. aggregata populations freed from pollen limitation will grow over the short term. In a second experiment we addressed whether density dependence would eventually erase the growth benefits of full pollination, by sowing a 10-fold range of seed densities that falls within extremes estimated for the natural "seed rain" that reaches the soil surface. Per capita survival to flowering and age at flowering were again unaffected by sowing density, but offspring size, per capita flower production, and lambda declined with density. Such density dependence complicates efforts to predict population dynamics over the longer term, because it changes components of the life history (in this case fecundity) as a population grows. A complete understanding of how constraints on seed production affect long-term population growth will hinge on following offspring fates at least through flowering of the first offspring generation, and doing so for a realistic range of population densities.     

Dominance orders,worker reproduction,and queen-worker conflict in the slave-making ant Harpagoxenus sublaevis

Summary In a queenright colony of the monogynous slave-making ant Harpagoxenus sublaevis, a subset of workers formed a linear dominance order in which dominance was corrlated with ovarian development, frequency of trophallaxis, length of time spent in the nest, but not body size. Identical dominance orders occurred in queenless colonies. Experiments in which the top-ranking workers were removed from queenless colonies demonstrated that worker dominance behaviour inhibits egg-laying in subordinates. A similar removal experiment showed queens restrict dominance behaviour and egg-laying in workers, probably pheromonally. Observations of slave raids indicated ovary-developed workers spent significantly less time scouting for slaves, and tended to participate less in slave raids, than workers without ovarian development. These findings suggest that potentially fertile H. sublaevis workers aggressively compete for egg-laying rights, consume extra food for egg development, and safeguard their reproductive futures by avoiding risks outside the nest. Hence worker reproduction in this species strongly influences the colony's social structure, nutrient flow, and division of labour, even though all workers in a colony are full sisters. I hypothesize that worker reproduction was formerly even more prevalent in H. sublaevis, with workers following the strategy of raising sisters and producing sons predicted by kinship theory. Its continued existence despite queen opposition conceivably results from selection on orphaned workers to reproduce, and the inability of slave-maker workers to raise female-biased broods. The social organization of H. sublaevis therefore highlights the importance both of worker reproduction and of the concomitant queen-worker conflict over male parentage in Hymenopteran social evolution.     

Maternal investment, sex-differential prospects, and the sex ratio in wild house mice

In a population of first-generation offspring from wild-caught house mice (Mus musculus domesticus), previous evidence suggested that male fitness is more strongly affected by an increase in body weight than female fitness. This paper shows that in these mice the young are weaned at heavier weights the smaller the litter and the better the maternal body condition. These effects persisted into adulthood and were less pronounced in female young. However, contrary to expectation from conventional sex ratio theory, maternal condition and litter size had no detectable effect on sex ratios. Also, litter size did not affect sex ratios in two populations of laboratory-kept, wild-caught western (M. m. domesticus) and eastern house mice (M. m. musculus). Wild house mice, therefore, appear not to adaptively manipulate the sex ratio of offspring. It is argued that this absence of sex ratio trends might not be maladaptive, but rather that models currently used to predict sex ratio trends in rodents may not be valid. Received: 13 March 1997 / Accepted after revision: 9 August 1997     

Philopatry,reproductive success of females,and maternal investment in the red-necked wallaby

Summary Female red-necked wallabies settle within their mothers' home ranges, apparently for life, while males disperse at about two years of age. However, sons spend much more time with their mothers before dispersing than do daughters of similar ages. Females who associate regularly with their subadult offspring are less likely to reproduce successfully at their next breeding attempt than are females who spend little time with their subadults, and sons therefore impose greater short-term reproductive costs on their mothers than do daughters. Females who are generally gregarious also suffer reduced reproductive success, even though reproductive success is independent of local density. It is suggested that the reproductive costs to females of associating with their subadult offspring, and other relatives, are incurred through tolerance of ecological competition from those kin, and therefore reflect a form of prolonged maternal investment, which is initially heaviest in sons but is sustained for longer periods in daughters. Females produce equal numbers of male and female offspring, and spend equal amounts of time suckling them in infancy.