Mate guarding in the swallowHirundo rustica

Summary The importance of mate guarding by males in the monogamous swallowHirundo rustica was studied by temporarily detaining the males. Mate guarding reduced the frequency of extra-pair copulations and of sexual chases involving female mates. Males participated in sexual chases more frequently if they had a non-fertile female. Neighbouring males of ‘widowed’ females increased their own mate guarding presumably in response to the experimentally increased rate of sexual chases. Neighbouring males with a fertile female increased their mate guarding more than did males with a non-fertile female. Addition of eggs to swallow nests in the post-fledging period of the first brood induced mate guarding by male nest owners. These males also copulated more frequently with their mates than did control males. Neighbouring male swallows responded to the increased mate guarding by showing sexual interest in the guarded females. removal of eggs from swallow nests during the laying period, leaving only one egg in the nest, resulted in reduced nest attendance by females. Male mates responded by increasing their mate guarding intensity as compared to controls, and neighbouring males showed an increased sexual interest in these females.     

Parentage and paternal care: consequences of intersexual selection in Savannah sparrows?

Empirical relationships between parentage and male parental care are commonly interpreted in the context of life-history models that consider increased offspring survivorship as the only benefit of paternal effort. However, indirect benefits associated with male care can also influence a male's response to cuckoldry: if females allocate paternity according to their prior experience with male parental care, it may pay for males to provision extra-pair young in early broods. Here, I assess the relationship between first-brood parentage and paternal care in a population of Savannah sparrows (Passerculussandwichensis) where a male's fertilization success in the second brood appears to be influenced by his prior parental performance. Based on the multi-locus DNA fingerprinting of 17 first broods, male feeding effort was influenced by parentage (percent of brood resulting from within-pair fertilizations) but not by brood size, male mating status (monogamous versus polygynous), timing of breeding (hatching date), structural size (wing length) or condition (mass). Males provided more care to broods that contained few within-pair young. This result supports the idea that males provision young to increase their future mating success, but alternative hypotheses involving male quality and timing of breeding cannot be excluded. Received: 13 August 1996 / Accepted after revision: 22 February 1997     

Equality of feeding roles and the maintenance of monogamy in tree swallows

Summary We investigated the division of labor in male and female tree swallows by measuring the rate of food delivery to nestlings at 36 nests. By observing natural nests and performing brood manipulations we found that males and females divided the feeding duties about equally and responded similarly to changes in brood size and age. Feeding rate was most highly correlated with brood mass. Manipulation and removal experiments showed that increased feeding rates could be elicited, but only for limited periods of time. Male and female tree swallows could only partially compensate in feeding nestlings when mates were removed. This, along with the higher mortality in enlarged broods and in those raised by single parents, indicates that both male and female are required to raise an entire brood to fledging. We argue that this requirement contributes to the absence of mate guarding and the maintenance of monogamy in the tree swallow.     

Do males and females differ in the feeding of large and small siblings? An experiment with the bluethroat

Males and females have been reported to differ in their feeding of large and small siblings in several species of birds. According to recent hypotheses, this phenomenon may be related to a sexual conflict over avian hatching patterns. We designed an experiment to test for the existence of such a sex difference by manipulating nestling size hierarchies of the bluethroat (Luscinia s. svecica) in two directions; half the broods were “asynchronized” to yield large size-differences within broods and the other half were “synchronized” to yield small size-differences. In all broods, nestlings were categorized as being either large or small according to body mass. We recorded male and female food distribution by video early (day 4 after hatching) and late (day 8) in the nestling period. Males and females did not differ in their distribution of food among different-sized nestlings. With large size-differences, both males and females fed large nestlings nearly twice as often as small ones. In contrast, when the size-differences were small, food was more evenly distributed among nestlings. Early in the nestling period, males fed more nestlings during each feeding visit than did females. Our finding that male and female bluethroats do not differ in the feeding of large and small siblings is in contrast to most previous studies. Variation in costs and benefits to males and females from feeding different-sized nestlings, and restrictions to parental choice due to nestling interactions, may explain interspecific variation. Received: 27 June 1997 / Accepted after revision: 26 January 1998     

Male mating success and paternal care in Nicrophorus vespilloides (Coleoptera: Silphidae)

Summary If no female is present, male burying beetles Nicrophorus vespilloides (Coleoptera: Silphidae) co-operate in the burial of a corpse. Once a female has arrived, the males fight with one another. The defeated male stays near the corpse and to copulate with the female. Laboratory experiments using sterilised males showed that the defeated male was able sometimes to father some of the offspring raised on the corpse. Male N. vespilloides almost always participate in defence and feeding of the brood. This is not affected by the size of the male. Males quickly leave or are driven from 5 g corpses. Males feed the larvae as often as females do, and larvae raised by males alone are not significantly different in weight from larvae raised by females alone or by both parents. Males which cared for a succession of broods in the laboratory did not differ significantly in median lifespan from males which were removed from their corpses after eggs had been laid. Non-caring males weighed significantly more than caring males over a sequence of corpses, but the caring males did not differ significantly in weight from non-breeding controls.     

Reproductive allocation in female house wrens is not influenced by experimentally altered male attractiveness

The differential allocation hypothesis proposes that females mated to attractive males should invest more resources in their offspring than those mated to less-attractive males, whereas the compensation hypothesis posits that females mated to less-attractive males should invest more resources in their offspring to compensate for lower-quality young. We tested these hypotheses by manipulating attractiveness of male house wrens (Troglodytes aedon) prior to female arrival by adding extra nest sites to territories of some males while leaving control males with only a single nest site. Females laid their eggs sooner in the nests of attractive males, and attractive males were more likely to retain their territory over successive broods and were marginally more likely to obtain a mate for a second brood later in the season than were control males, thereby confirming the effect of our manipulation on male attractiveness. Experimentally enhanced attractiveness also led to increased hematocrit in males. However, there were no consistent differences in the number, size, or quality of eggs laid by females mated to attractive and control males, nor were there any differences in the size, health state, or immune function of nestlings produced from these eggs. There was also no effect of treatment on the number of nestlings surviving to fledging. Collectively, these results are inconsistent with both the differential allocation hypothesis and the compensation hypothesis. Future studies should consider the possibility that the criteria used by females in selecting a mate may vary temporally and be more flexible than generally thought.     

The dynamics of brood desertion among communally breeding females in the treehopper, Publilia concava

In species that exhibit extended parental care, females sometimes lay eggs communally in order to redistribute the costs of offspring care. Communal egg-laying often involves redundancy in female effort, such that the number of females contributing to reproduction is more than is needed to provide adequate parental care. As a result, a subset of females will often abandon the communal brood, with the time of departure ranging from immediate abandonment after egg-laying (brood parasitism) to delayed abandonment with prolonged care (cooperative breeding). In this paper I approach the parental care dynamics of female-female broods as a desertion game similar to that of mate desertion in species with bi-parental care. I describe a field study of the insect Publilia concava (Hemiptera: Membracidae), a species that exhibits communal oviposition and a full range of egg guarding. This species exhibits full redundancy in female care, with no difference in survival between singly and doubly guarded broods. I find that double guarding is extremely rare in the population, with most communal broods having only one female guard. While this guard was usually the female that initiated the brood, these same females were more likely to abandon when secondary females arrived and when secondary females exhibited longer guarding durations. Paradoxically, the secondary females usually abandoned the broods they visited, resulting in up to 50% of broods with double abandonment. These unguarded broods suffered a 50% reduction in hatching success, reflecting an important risk for primary females that abandon egg masses to secondary females. Overall, P. concava exhibits desertion dynamics similar to mate desertion in vertebrates and it is likely that the theoretical work in this area will be useful for future work that addresses the allocation of parental care among communal breeders.     

Costs and benefits of brood desertion in female kentish plovers,Charadrius alexandrinus

Female kentish plovers Charadrius alexandrinus typically desert their broods after the chicks hatch, i.e. 1–4 weeks before the chicks fledge or become independent. In this paper we investigate the costs and benefits of desertion for females. Desertion incurs a cost for females: following desertion chick survival in broods is lower (0.95 +- 0.02 day^–1) than before the female deserts (0.98 +- 0.01 day^–1). We investigated several possible causes for reduced brood survival by comparing characteristics of broods before and after desertion (controlling for differences in brood age). After desertion males increased the time they spent foraging and they tended to reduce time spent brooding chicks. Increased mortality of chicks may occur in deserted broods because following desertion (1) males spend less time alert in vigilance behaviour than before desertion, (2) they attend the chicks from greater distances, and (3) they show greater distraction display distances (in response to human intruders). Growth or development of chicks, measured by weight gain and tarsus length, was not different before and after desertion. Females gain two potential benefits from desertion: (1) they may remate and produce a second brood within the same breeding season or (2) they may enhance their probability of surviving to breed in a subsequent season. At least 27% of female kentish plovers that deserted remated and renested in the same season in this study. In contrast, we found no evidence that brood desertion increased the survival of females: there was no difference in local survival rate (return rate) for females deserting before or after 6 days brood age. These results clearly demonstrate that female kentish plovers that desert their offspring prior to fledging incur costs, but we suggest that there is a trade-off with the potential benefits gained by remating and making a second breeding attempt in the same season.     

Search costs influence the spatial distribution,but not the level,of extra-pair mating in tree swallows

Costs of searching for a mate are an important component of models of sexual selection, yet they have rarely been examined in wild populations of vertebrates. In this paper, we report an experiment in which we handicapped female tree swallows by clipping some flight feathers. This manipulation increased the costs of flight and searching for extra-pair mates. Despite these costs, handicapped females had the same level of extra-pair mating (percentage of extra-pair young, percentage of broods with extra-pair young, and the number of extra-pair sires per brood) as control females. However, handicapped females were more likely to have young sired by extra-pair males that lived closer to her nest than control females. This change in the distribution of extra-pair mating was most likely due to female choice rather than male coercion, and it suggests that extra-pair mating has significant benefits to females. One important implication of our study is that ecological and social factors that influence search costs could affect the spatial distribution of extra-pair sires and, consequently, the intensity of sexual selection. These effects may have been overlooked in previous studies that did not identify extra-pair sires.     

Filial cannibalism in a nest-guarding fish: females prefer to spawn in nests with few eggs over many

In fish, fecundity correlates with female body size and egg-tending males often eat small broods. Therefore, small females may prefer to spawn in nests that already contain many eggs, to ensure the brood is as large as possible. In contrast, large females may prefer nests with few eggs, if high egg number or density has a negative effect on egg survival, or if there are drawbacks of spawning last in a nest. To test the hypothesis that female body size affects nest (and male mate) choice, using the sand goby (Pomatoschistus minutus), we allowed small and large females to choose between two males that were matched in size — one guarding a small clutch and the other a large clutch, respectively. We recorded where females spawned (measure of female preference), the combined brood size, male courtship, egg care and nest building. We also quantified the effect of brood size and egg density on egg survival in a separate data set. Although the combined broods did not exceed the small brood sizes that are at risk of being eaten, both small and large females preferred to spawn in nests with smaller clutch sizes. This preference could not be explained by more courtship or male parental effort, nor by reduced survival of larger or denser broods. Instead, our result might be explained by females avoiding the danger of cannibalism of young eggs by males or the risk of reduced egg health associated with being near the nest periphery.     

Mating opportunities,paternity, and sexual conflict: paternal care in northern and southern temperate house wrens

Males are generally predicted to care less for their young when they have more additional mating opportunities, lower paternity, or when their mates care more. We tested these predictions using male provisioning as a proxy for paternal care in two temperate populations of house wrens (Troglodytes aedon) with divergent life histories. Males in the migratory, occasionally socially polygynous New York, USA (northern) population provisioned less when more local females were fertile. A similar relationship was only weakly supported in the resident, socially monogamous Buenos Aires Province, Argentina (southern) population, possibly due to the higher density of house wrens there. A relationship between male provisioning and level of paternity within the brood was supported in both populations, but in opposite directions: while males in the southern population provisioned less at broods containing more extra-pair young, males in the northern population provisioned such broods more, contradicting predictions. Males provisioned less when their mates provisioned more in both populations, in agreement with sexual conflict theory. Additionally, the populations both exhibited a positive relationship between male provisioning and nestling age, but differed in the direction of the relationships of male provisioning with date and brood size. Our results suggest that even within a species, life history differences may be accompanied by differences in the determinants of behavior such as paternal care.     

The presence of eggs in the nest and female choice in common freshwater gobies (Rhinogobius brunneus)

In many species of fishes with paternal care, females prefer to spawn with males who are already guarding eggs. We studied the effect of egg presence on female mate choice in common freshwater gobies, Rhinogobius brunneus sp. OR. In our tests, females did not prefer males with eggs, suggesting egg presence per se may not act as a cue to attract females. We also examined the effects of brood size on paternal care and offspring survival to look for possible benefits females could obtain when choosing males with eggs. Both fanning by egg-guarding males and egg survival increased with brood size. The presence of neighboring males did have a significantly negative effect on males' parental activity, which subsequently results in a lower level of egg survival than in solitary egg-guarding males. This result provides a partial explanation for the result of eggs not attracting females to mate.     

Active and effective nest defence by males in a social apoid wasp

Altruism in the social Hymenoptera is generally considered to be a feature of females rather than males. A popular explanation for this is that in the solitary ancestors of today's social species, males provided little brood care. Males might therefore lack the preadaptations necessary to evolve altruism in social contexts. While anecdotal observations of male contributions to colony life have been reported, there are few reports of male participation in nest defence. In apoid wasps, there have been several reports of male nest-guarding behaviour in solitary species, potentially setting the evolutionary stage for similar behaviours in social lineages. Here, we present evidence of active and effective nest defence in males of the social apoid wasp Microstigmus nigrophthalmus. Males were observed chasing intruders away from the nest, and the presence of males had a significant effect on nest survival when females were removed. Males potentially obtained direct benefits through defence, so that defence may not represent male altruism. However, our results do show that males can perform acts that benefit their colony.     

The costs of male parental care and its evolution in a neotropical frog

Summary Parental care is practiced exclusively by males of the Puerto Rican frog, Eleutherodactylus coqui. Males brood clutches of direct-developing eggs in non-aquatic nest sites and defend eggs against cannibalistic nest intruders. Here, I report on energetic and mating costs incurred by males that provide parental care, and suggest how these proximate costs affect male fitness and the evolution of male parental care in this species. Energetic costs are small for brooding males in comparison to non-brooding, calling males. Brooding males had a higher frequency of empty stomachs and lost small, but significant, fractions of their initial body mass during parental care. Abdominal fat bodies of brooding males during the middle third of parental care were significantly smaller than those of calling males; those of males brooding eggs in earlier or later stages were not different. The mating cost of parental care is greater. Most brooding males cease calling during parental care. However, gravid females are available (i.e., known to mate) on most nights during the principal breeding season; hence non-calling males miss potential opportunities to mate. A mating cost was estimated by calculating nightly mating probabilities for calling males in a plot where nightly calling male densities and daily oviposition schedules were known. On average, a male exhibiting normal calling behavior would be expected to obtain a new mate once every 35.7 days. Hence a brooding male that ceased calling for a 20-day parental care period would miss, on average, 0.56 additional mates. Males that were more successful than average in attracting mates could miss up to 1.63 matings. A marginal value model (Fig. 1) is used to analyze the net effect on male fitness of parental care benefits and costs in E. coqui (Fig. 3). The model indicates that males garner the highest reproductive success by providing care from oviposition through hatching. There is no stage during the pre-hatching period at which a desertion strategy would yield higher reproductive success. In fact, the model suggests that males should provide full parental care even in the face of much higher mating costs than currently obtain in the system.     

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.     

Brood sex ratio is dependent on female mating status in polygynous great reed warblers

Females capable of adjusting the sex ratio of their offspring should be more fit than females lacking such an ability. In polygynous birds where breeding success in males is more strongly influenced by body size and/or attractiveness than in females, females might produce more sons when predicting good conditions or when mating with attractive males. Polygynous great reed warbler, Acrocephalusarundinaceus, males direct most of their feeding effort to the primary (first-hatching) nest and in these nests increase their feeding effort in relation to the brood sex ratio (proportion of sons). Therefore, with the expectation of well-nourished sons, we would predict that females which start breeding first within harems might produce more sons than those which start breeding later, and in anticipation of sons with good genes, that females mated to polygynous males might produce more sons than females mated to monogamous males. I took blood samples from hatchlings and determined the sex using DNA markers. The sex ratio of primary (monogamous and polygynous primary) broods is more male-biased (mean 0.58 males, n = 50) than that of secondary (polygynous secondary and tertiary) broods (mean 0.46, n = 25). Moreover, in the secondary broods with the largest clutch (five eggs), in which offspring are most likely to suffer food shortage, the sex ratio was distinctively female biased (mean 0.33, n = 10). In the primary broods, sex ratio was correlated to harem size. The results suggest that great reed warbler females modify the brood sex ratio to produce both well-nourished sons and sons with good genes, but the former effect is probably stronger than the latter factor. Received: 11 March 1998 / Accepted after revision: 23 May 1998     

Offspring protection by merlin Falco columbarius females; the importance of brood size and expected offspring survival for defense of young

Summary Nest predation was simulated by presenting a stuffed raven close to nests of merlins. This was done to examine the influence of brood size related factors on female defence intensity. The original clutch size did not affect nest defense after hatching, but brood size was important. It determined the attack frequency and, for each brood size level, the proportion of attacking females. When brood sizes were manipulated, the defense intensity increased and decreased, respectively, in relation to the size of the brood. In addition, broods with high future survival (first broods) were defended more vigorously than broods with low future survival (replacement broods). Hence, expected benefits in terms of fledgling production and chick survival seem to be important determinants of female investment in offspring protection. The lower predation rate among females responding with overt aggression to the raven compared to that of less aggressive females suggests that defence of young is beneficial.     

Sex-specific conditional mating preferences in a cichlid fish: implications for sexual conflict

Conditional mating strategies enable individuals to modulate their mating behaviour depending on ‘individual status’ to maximise fitness. Theory predicts that variation in individual quality can lead to differences in mating preferences. However, empirical evidence is scarce particular in terms of variation in male and female strategies. Here, we experimentally investigated quality-dependent variation in mating preferences concerning reliable quality indicators in Pelvicachromis taeniatus, a colourful cichlid fish with mutual mate choice and ornamentation. Males as well as females were artificially manipulated in phenotypic quality by different feeding regimes. Ornamentation was connected to individual quality in both sexes. Males and females showed conditional mating strategies in different directions. Males showed prudent choice by preferring females of similar quality. In contrast to males, low-quality females preferred highly ornamented males, whereas high-quality females showed neither preferences for high- nor for low-quality males. The results suggest that individuals aim for specific benefits depending on individual quality. Furthermore, the conflicting conditional mating preferences of males and females might lead to sexual conflict, implicating a highly dynamical mating system that evolves even in absence of environmental changes.     

Female choice and the quality of parental care in the great tit Parus major

Summary Previous work on great tits suggested that female mate choice was based on the characteristics of the male rather than the quality of his territory. The aim of this paper was to see whether females were deriving any direct benefits as a result of such choice by comparing male plumage colouration (the size of the central black breast-stripe) with the quality of parental care provided by both members of the pair. Males with large stripes were more likely to defend their broods because their nest attentiveness was higher increasing the chances of predator detection. Males with large stripes produced heavier fledglings primarily as a result of faster growth early in the nestling period. Previous work on great tits has shown that heavier fledglings have a higher survival probability than lighter ones. The results of this study can more easily be explained by differences in parental quality between males rather than as a result of variation in female or territory quality. Therefore, female great tits could increase their reproductive success by pairing with males with a large stripe because such males appear to be better quality parents. This suggests that female choice in the great tit may concern, at least in part, the quality of male parental care.     

Resource allocation varies with parental sex and brood size in the asynchronously hatching green-rumped parrotlet (<Emphasis Type="Italic">Forpus passerinus</Emphasis>)

When eggs hatch asynchronously, offspring arising from last-hatched eggs often exhibit a competitive disadvantage compared with their older, larger nestmates. Strong sibling competition might result in a pattern of resource allocation favoring larger nestlings, but active food allocation towards smaller offspring may compensate for the negative effects of asynchronous hatching. We examined patterns of resource allocation by green-rumped parrotlet parents to small and large broods under control and food-supplemented conditions. There was no difference between parents and among brood sizes in visit rate or number of feeds delivered, although females spent marginally more time in the nest than males. Both male and female parents preferentially fed offspring that had a higher begging effort than the remainder of the brood. Mean begging levels did not differ between small and large broods, but smaller offspring begged more than their older nestmates in large broods. Male parents fed small offspring less often in both brood sizes. Female parents fed offspring evenly in small broods, while in large broods they fed smaller offspring more frequently, with the exception of the very last hatched individual. These data suggest male parrotlets exhibit a feeding preference for larger offspring—possibly arising from the outcome of sibling competition—but that females practice active food allocation, particularly in larger brood sizes. These differential patterns of resource allocation between the sexes are consistent with other studies of parrots and may reflect some level of female compensation for the limitations imposed on smaller offspring by hatching asynchrony.