Mate acquisition tactics in polyandrous spotted sandpipers (Actitis macularia): the role of age and experience

We consider resource-defense polyandry and mate-access polyandryas female mating tactics in spotted sandpipers (Actitis macularia).These tactics can be distinguished by the resource females defend,female interclutch movement, expected reproductive success aftermoving, and male and female dispersion. We examine these characteristicsrelative to patterns observed in a 17-year study of spottedsandpipers, a species traditionally considered resource-defensepolyandrous. On average, 26% of spotted sandpiper females eachyear were monogamous. Older females were more likely to be polyandrous,and polyandrous females of each age employed different matingtactics. Yearlings were typically sequentially resourcedefensepolyandrous. Two-year-olds were primarily simultaneously polyandrous,exhibiting equivalent proportions of resource-defense and mate-accesspolyandry. Older females were primarily simultaneously resource-defensepolyandrous. Females tended to stay on territories where theyand/or their mates had greater breeding experience (i.e., manyclutches laid for females, many clutches diat hatched for males);females that moved went to territories where their mates hada history of breeding success. Location changes between clutchesby polyandrous females were better described by breeding experienceon a territory than by age.     

Competing Females and Caring Males. Polyandry and Sex-Role Reversal in African Black Coucals, Centropus grillii

Most species of birds show bi‐parental or female‐only care. However, a minority of species is polyandrous and expresses male‐only care. So far, such reversals in sex roles have been demonstrated only in precocial bird species, but there was suggestive evidence that such a mating system may occur in one altricial bird species, the black coucal, Centropus grillii. In a field study in Tanzania we investigated whether black coucals are sex‐role reversed and polyandrous. We found that males were mated to one female, rarely vocalized and provided all parental care from incubation of eggs to feeding of young. In contrast, female black coucals were about 69% heavier and 39% larger than males and polyandrous. They spent a large proportion of time calling from conspicuous perches, defended breeding territories, did not help in provisioning young and had a higher potential reproductive rate than males. We conclude that the black coucal currently represents the only altricial bird species with sole male parental care and a classical polyandrous mating system. High nest predation pressure and small territory sizes due to high food abundance may have been important factors in the evolution of sex‐role reversal and polyandry in this species.     

Behavioral and Endocrine Responses in Male Marmosets to the Establishment of Multimale Breeding Groups: Evidence for Non-monopolizing Facultative Polyandry

Studies of wild callitrichids provide conflicting evidence regarding polyandrous groups. One perspective supports a monopolizing breeding strategy on the part of one male, while the alternative perspective suggests that polyandry does not lead to a breeding monopoly. We tested the hypotheses in male marmosets (Callithrix kuhlii) with 5 polyandrous groups composed of related/familiar males as our attempts to establish polyandrous groups of unrelated/familiar males failed. We monitored male social and sexual behavior and urinary testosterone (T) and cortisol (CORT) across the first 80 days of group formation and contrasted them with similar measures in males housed in monogamous groups. We also examined the same measures across the females' ovulatory cycles for polyandrous males. We found little evidence that males in polyandrous groups exercised a mating monopoly over the female and no evidence for overt competition between polyandrous males. We found 2 behavioral differences: polyandrous males were less often in proximity and copulated more often with the female than monogamous males did. Our findings suggest that the males in newly-formed groups of marmosets do not monopolize breeding and social behavior with the female. This appears to be the case for 3 reasons. First, males may use sperm competition rather than overt competition. Secondly, it may take longer for relationships to develop between the female and the males in polyandrous groups versus in monogamous groups. Thirdly, the cost of infant care is sufficiently high to demand that group members get along when groups are small and reproductive benefits are shared.     

Breeding Biology and Variable Mating System of a Population of Introduced Dunnocks (Prunella modularis) in New Zealand

Species with variable mating systems provide a unique opportunity to investigate whether females receive direct fitness benefits from additional male partners. The direct benefits provide an obvious explanation for why females would breed polyandrously, in a situation where males clearly do not attain their optimal reproductive success. Evidence for these direct benefits is, however, mixed. Here, we present a detailed study of the breeding biology of the dunnock, Prunella modularis, which inform an investigation into the effects of the social mating system on the reproductive success in a population of dunnocks in Southern New Zealand. We studied 80 different social groups over the course of three breeding seasons. Dunnocks in our population presented a variable mating system, with socially monogamous (45%), socially polyandrous (54%) and socially polygynandrous (1%) groups being observed in the same breeding season. We did not observe any polygynous social units in our study period although polygyny exists in the population. We found little difference in the numbers of eggs laid, and egg volume between monogamous and polyandrous nests. However, polyandrous groups had better hatching and fledging success than monogamous groups (composite d = 0.385, 95% CI: 0.307 to 0.463). Overall our results support the notion that polyandry is beneficial for females.     

Breeding biology and mating system of Brown Accentors Prunella fulvescens on the Tibet Plateau

ABSTRACT

Capsule: The mating system of the Brown Accentor Prunella fulvescens in the high altitudes of the Tibeta plateau is mainly monogamous but polyandry was recording in 9.7% of breeding groups.

Aims: To describe the poorly known breeding biology of Brown Accentors in the high-altitude environment of the northeastern Tibet plateau.

Methods: Individually colour-ringed birds were observed during two breeding seasons. Biometrics were recorded for breeders, and reproductive variables were quantified.

Results: Brown Accentors were territorial during the breeding season and were resident year-round. Most groups (90.3%, n?=?31 groups) were socially monogamous and the remainder (9.7%) were polyandrous, with two males sharing a female’s territory. Mean group size was 2.1 adults and the population adult sex ratio was 52% male. The birds raised a single brood during the breeding season. Nests were invariably located in Sea Buckthorn Hippophae rhamnoides. Eggs were laid between late June and late July, and mean clutch size was 3.4?±?0.6 (SD) eggs. Incubation was undertaken by the female and lasted 12.9?±?0.9 (SD) days. Both sexes provisioned young in the nest for 13.9?±?1.2 (SD) days. Females provided significantly more feeding visits than did males. Brood size at estimated fledging dates averaged 2.2?±?0.7 (SD) nestlings. Forty-five per cent of nesting attempts (n?=?37) fledged at least one nestling.

Conclusions: Like other species in the family Prunellidae, the Brown Accentor exhibits remarkable variation in its mating system.     

Reproductive success and survivorship of the Galapagos Hawk Buteo galapagoensis: potential costs and benefits of cooperative polyandry

The Galapagos Hawk Buteo galapagoensis is one of few species showing cooperative polyandry. As many as four males may mate with one female and share a territory and nest, Such groups, on average, produce more young than monogamous pairs, but on a per male basis (assuming that each male of a group has an equal chance to fertilize each young) polyandrous males produce fewer young per year than monogamous males. Territorial breeding birds show higher yearly survivorship (90%) than non-territorial, non-breeding birds (50% or less). All territorial birds seem to remain on their territories for life, and monogamous pairs actually constitute the remnants of polyandrous groups. Some potential factors producing such an unusual system are discussed. Limited breeding space and high mortality among non-territorial birds seem to work in favour of group breeding in this and other cooperative species. Polyandry may be favoured because of the reduced value of extra females in hawk breeding behaviour. Equal-status males may occur because of the lack of kinship among cooperating males and the high potential genetic cost of being a helper in a long-lived, low fecundity species. Other possible factors favouring group breeding are territory acquisition and the variable nature of the Galapagos climate.     

Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor

The sexes’ share in parental care and the social mating system in a marked population of the single‐brooded Lesser Spotted Woodpecker Dendrocopos minor were studied in 17 woodpecker territories in southern Sweden during 10 years. The birds showed a very strong mate fidelity between years; the divorce rate was 3.4%. In monogamous pairs, the male provided more parental care than the female. The male did most of the nest building and all incubation and brooding at night. Daytime incubation and brooding were shared equally by the sexes, and biparental care at these early breeding stages is probably necessary for successful breeding. In 42% of the nests, however, though still alive the female deserted the brood the last week of the nestling period, whereas the male invariably fed until fledging and fully compensated for the absent female. Post‐fledging care could not be quantified, but was likely shared by both parents. Females who ceased feeding at the late nestling stage resumed care after fledging. We argue that the high premium on breeding with the same mate for consecutive years and the overall lower survival of females have shaped this male‐biased organisation of parental care. In the six years with best data, most social matings were monogamous, but 8.5% of the females (N=59) exhibited simultaneous multi‐nest (classical) polyandry and 2.9% of the males (N=68) exhibited multi‐nest polygyny. Polyandrous females raised 39% more young than monogamous pairs. These females invested equal amounts of parental care at all their nests, but their investment at each nest was lower than that of monogamous females. The polyandrously mated males fully compensated for this lower female investment. Polygynous males invested mainly in their primary nest and appeared to be less successful than polyandrous females. Polyandry and polygyny occurred only when the population sex ratio was biased, and due to strong intra‐sexual competition this is likely a prerequisite for polygamous mating in Lesser Spotted Woodpeckers.     

Plasticity in the mating system of the longnose filefish,Oxymonacanthus longirostris, in relation to mate availability

The mating system of the longnose filefish,Oxymonacanthus longirostris, was examined on coral reefs of Okinawa, Japan. This species has been shown previously to be monogamous. Fish were usually found swimming together in heterosexual pairs with the male and female sharing the same feeding territory. However, both monogamous and polygynous (bigamous) males were found in the present study. Polygynous males, which were larger than monogamous males, visited and stayed several minutes in turn with each female within the territories. Although most males were monogamous in the early breeding season, over 20% of males mated polygynously in the late breeding season. The adult sex ratio in the former was unbiased, but became slightly female-biased toward the end of the breeding season because of the higher disappearance rate of males. The higher disappearance rate may be due mainly to a higher mortality rate of males resulting from a greater deterioration of physical condition during the breeding season. Thus, the mating system varied with the change of the adult sex ratio. Plasticity in the mating system of this species may be the outcome of male mating tactic depending on local mate availability.     

The relative influence of prey abundance and co-breeders on the reproductive performance of polyandrous Pale Chanting-goshawks

This study investigates if the reproductive performance of polyandrous Pale Chanting-goshawks, Melierax canorus, is governed by the abundance of dominant rodent-prey species or a co-breeding male participating fully in prey being delivered to the female and young. Polyandrous trios in prey-rich habitat, the only habitat where these trios occurred, failed to produce more offspring than monogamous pairs, but attempted more second broods in years of high prey abundance. The higher the prey-delivering rates at the nesting sites by either one monogamous male or two polyandrous males (combined effort), the higher the proportion of time their females spent in close proximity to the nest. Since there were no significant differences between the body masses of nestlings fledged by polyandrous and monogamous groups, higher delivery rates relieved females from away-from-the-nest hunting duties and permitted them to feed on prey not fed to offspring. Although breeding in polyandrous trios held fitness benefits for females, e.g. to accumulate the necessary body reserves for laying a second clutch in years of high prey abundance, it is still not clear how males benefited. The reproductive performance of each group was strongly linked to the abundance of the dominant, otomyine rodent prey (Otomys unisulcatus and Parotomys brantsii) and it was therefore prey abundance, and not the number of males delivering prey to the female and nestlings at each nesting site, that controlled the reproductive performance of cooperative-breeding Pale Chanting-goshawks.     

The effect of mating frequency and mating pattern on female reproductive fitness in cabbage beetle,Colaphellus bowringi

It is generally thought that females can receive more of the material benefits from males by increasing mating frequency and polyandry can lead to greater reproductive success. The cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), is a highly promiscuous species, in which females or males can readily mate repeatedly with a given partner or multiple partners at a very high frequency. In the present study, the effect of mating frequency (number of matings) and mating pattern (polyandry vs. monogamy) on female reproductive fitness was investigated by measuring fecundity, fertility, and female longevity. The results indicated that increased female mating frequency with the same male did not result in variation in lifetime fecundity, but significantly increased fertility and decreased female longevity. Moreover, five copulations were sufficient to acquire maximal reproductive potential. Female lifetime fecundity also did not differ between polyandrous and monogamous treatments. However, monogamous females exhibited a significant increase in fertility and significant prolongation of longevity compared with polyandrous females, further demonstrating that monogamy is superior to polyandry in this beetle.     

High rates of infidelity in the Grey Fantail Rhipidura albiscapa suggest that testis size may be a better correlate of extra‐pair paternity than sexual dimorphism

The Grey Fantail Rhipidura albiscapa is a socially monogamous passerine endemic to Australia. Behavioural and morphological clues point to opposing conclusions as to its breeding system; sexual monomorphism and monochrome colorations suggest monogamy, whereas relatively large testes and a prominent cloacal protruberance are more indicative of multiple mating and sperm competition. We used five highly variable microsatellite loci to investigate the genetic breeding system of this species. Paternity was assigned to 49 of 69 (71%) offspring tested and the overall rate of partner infidelity was high, with 55% of offspring being sired by an extra‐pair male and 64% of all clutches containing extra‐pair young. This puts the Grey Fantail amongst the most promiscuous socially monogamous species yet studied. Where extra‐pair fathers were identified, these were invariably in neighbouring territories, and although larger males did not gain more paternities overall, extra‐pair offspring tended to be fathered by larger males than expected by chance. We interpret our findings in light of some of the potential costs and benefits associated with extra‐pair paternity.     

The diet and foraging behaviour of the Dunnock Prunella modularis in a hedgerow habitat

The diet and foraging behaviour of a population of Dunnocks inhabiting hedgerows was studied between November 1981 and December 1983 at Nedge Hill, Shropshire. Invertebrates were most abundant in hedgerows and adjacent vegetation during the summer when the diet of the Dunnock was largely insectivorous. Beetles (Coleoptera) were the main taxa in the diet, with weevils (Curculionidae) and rove beetles (Staphylinidae) predominating. Arachnids were important, and snails (Gastropoda), earthworms (Oligochaeta), springtails (Collembola) and flies (Diptera) were taken. Seeds were available in the study area from July and featured in the diet from August to April. Nettle Urtica dioica comprised the main seed in the diet. Other important seeds included Yorkshire fog Holcus lanatus , elder Sambucus nigra and dock Rumex obtusifolius. Stomach stones were most common in the faeces in winter. The Dunnocks were predominantly ground feeders and mainly used a hop-picking technique. Some seeds and invertebrates were picked directly off plants by Dunnocks above the ground. Most feeding occurred in hedgerows, which formed a small proportion of the study area. Overgrown hedgerows were the most important of the five hedgerow types. Nettle patches were an important feeding habitat. Dunnocks usually foraged alone or in pairs and rarely fed in association with other bird species. The highest proportion of time spent feeding was in January and the lowest in May.     

Polyandry enhances offspring viability with survival costs to mothers only when mating exclusively with virgin males in Drosophila melanogaster

A prominent hypothesis for polyandry says that male–male competitive drivers induce males to coerce already‐mated females to copulate, suggesting that females are more likely to be harassed in the presence of multiple males. This early sociobiological idea of male competitive drive seemed to explain why sperm‐storing females mate multiply. Here, we describe an experiment eliminating all opportunities for male–male behavioral competition, while varying females’ opportunities to mate or not with the same male many times, or with many other males only one time each. We limited each female subject's exposure to no more than one male per day over her entire lifespan starting at the age at which copulations usually commence. We tested a priori predictions about relative lifespan and daily components of RS of female Drosophila melanogaster in experimental social situations producing lifelong virgins, once‐mated females, lifelong monogamous, and lifelong polyandrous females, using a matched‐treatments design. Results included that (1) a single copulation enhanced female survival compared to survival of lifelong virgins, (2) multiple copulations enhanced the number of offspring for both monogamous and polyandrous females, (3) compared to females in lifelong monogamy, polyandrous females paired daily with a novel, age‐matched experienced male produced offspring of enhanced viability, and (4) female survival was unchallenged when monogamous and polyandrous females could re‐mate with age‐ and experienced‐matched males. (5) Polyandrous females daily paired with novel virgin males had significantly reduced lifespans compared to polyandrous females with novel, age‐matched, and experienced males. (6) Polyandrous mating enhanced offspring viability and thereby weakened support for the random mating hypothesis for female multiple mating. Analyzes of nonequivalence of variances revealed opportunities for within‐sex selection among females. Results support the idea that females able to avoid constraints on their behavior from simultaneous exposure to multiple males can affect both RS and survival of females and offspring.     

Resident female Northern Lapwings Vanellus vanellus fight to exclude settlement by secondary females

In systems with bi‐parental care, sexual conflict theory states that a transition from monogamy to polygyny should increase fitness prospects for males and reduce fitness prospects of resident females. Behavioural observations of Northern Lapwings Vanellus vanellus suggested that resident females try to monopolize their breeding resources by attacking females prospecting for secondary status, and males commonly respond by intervening on behalf of the prospecting females. In four territories the resident females successfully defended their monogamous status, in nine territories the resident females made at least one successful eviction before being forced to accept polygyny and in nine territories observed eviction attempts were unsuccessful. These observations indicate that mating status in Northern Lapwings is partly determined by the competitive ability of resident females.     

Sexual selection and assortative mating: an experimental test

Mate choice and mate competition can both influence the evolution of sexual isolation between populations. Assortative mating may arise if traits and preferences diverge in step, and, alternatively, mate competition may counteract mating preferences and decrease assortative mating. Here, we examine potential assortative mating between populations of Drosophila pseudoobscura that have experimentally evolved under either increased (‘polyandry’) or decreased (‘monogamy’) sexual selection intensity for 100 generations. These populations have evolved differences in numerous traits, including a male signal and female preference traits. We use a two males: one female design, allowing both mate choice and competition to influence mating outcomes, to test for assortative mating between our populations. Mating latency shows subtle effects of male and female interactions, with females from the monogamous populations appearing reluctant to mate with males from the polyandrous populations. However, males from the polyandrous populations have a significantly higher probability of mating regardless of the female's population. Our results suggest that if populations differ in the intensity of sexual selection, effects on mate competition may overcome mate choice.     

EXTRAPAIR MATE CHOICE AND HONEST SIGNALING IN COOPERATIVELY BREEDING SUPERB FAIRY-WRENS

In many species of monogamous birds females copulate with males other than their social mates, resulting in extrapair fertilizations. Little is known about how females choose extrapair mates and whether the traits used to choose them are reliable indicators of male quality. Here we identify a novel male trait associated with extra-group mating success in the superb fairy-wren (Malurus cyaneus), a cooperatively breeding bird with one of the highest known frequencies of extra-group mating. Female fairy-wrens chose extra-group mates that molted earlier into breeding plumage. Males molted up to five months before the breeding season began, and only males that molted at least one month prior to its onset gained any extra-group fertilizations. This conclusion held after controlling statistically for the effect of age and social status on molt date. Once males acquired breeding plumage, they began courtship display to females on other territories. Thus, some males were displaying to females for several months before the breeding season began. This extraordinarily long period of advertisement by males may be facilitated by the long-term ownership of territories. We suggest that early acquisition of breeding plumage or the subsequent display behavior can be reliable cues for mate choice because they are costly to acquire or maintain.     

Variable Social Mating System in the Sedge Warbler,Acrocephalus schoenobaenus

We studied the mating system in a local population of colour-ringed sedge warblers in south Central Sweden in 1990–92. Of 58 territorial males, 59% were socially monogamous, 14% socially polygynous and 27% unpaired. Socially polygynous males in general paired with two females: the only exception was one male that formed pair bonds with three females. Among the males that formed a pair bond, 38% resumed singing after their first female had started egg-laying or incubation and 50% of the paired males that resumed singing succeeded in attracting a second female. Hence, despite a consistently male-biased sex ratio in the population a large proportion of the males tried to become polygynous and they were often successful. The frequency of extra-pair matings did not differ between monogamous and polygynous males. Of 47 breeding females, 6.4% were sequentially socially polyandrous. In two out of three cases, the females fed the young of their first broods until independence before initiating the second brood. In the third case the female deserted her newly fledged young and these were instead cared for by a neighbouring male. DNA fingerprinting revealed that this male had not sired any of these young. Each of the sequentially polyandrous females successfully raised both their broods, and their annual reproductive success was slightly higher than the average for the polygynous males. When the sequentially socially polyandrous females initiated their second brood, their primary male (in all cases polygynous males), cared for young in their secondary female's nest. In all cases, the sequentially polyandrous females formed second pair bonds with unpaired males that were close neighbors. This suggests that females switched pair male for their second brood to obtain a mate that was more likely to provide them with direct benefits (e.g. parental care).     

The evolution of egg size in socially polyandrous shorebirds

In classical and multi-clutch polyandry, females lay multiple clutches during a breeding season for more than one mate. The production of multiple clutches may be energetically demanding. We used comparative analyses to investigate three possible ways of reducing such egg-laying costs in polyandrous shorebirds: (1) reduction in egg size, (2) reduction in clutch size, and (3) evolutionary increase in female size. Paired comparisons of polyandrous and non-polyandrous taxa showed that females of polyandrous shorebirds lay smaller eggs than females of closely related monogamous and polygynous species. Directional analyses corroborated this result by indicating a significant decrease in egg size after phylogenetically independent origins of polyandry. The comparative analyses uniformly rejected the two alternatives, i.e. neither clutch size nor female size is related to social mating pattern. We also tested and rejected three alternative explanations for reduced egg size in polyandrous taxa. First, we found no evidence that polyandrous females have evolved smaller egg sizes in response to selection to match smaller size of males, which provide the parental care in these species. Second, reduction in egg size was not related to longer breeding seasons (and hence more opportunity for re-nesting). Third, reduced egg sizes were also not related to rates of clutch predation (another potential correlate of multiple clutch production). Our results are thus consistent with the hypothesis that selection for reducing laying costs explains small egg size in socially polyandrous shorebirds.     

Signaling of Reproductive Status in Captive Female Golden-Headed Lion Tamarins (Leontopithecus chrysomelas)

In species in which paternal care has an important impact on the offspring's fitness, concealment of reproductive status has been proposed as a strategy employed by females to prevent males from practicing desertion and polygamy, which would then lead to monogamous or polyandrous mating systems or both. We investigated whether the female's reproductive status is being concealed in golden-headed lion tamarins, which exhibit extensive paternal care and a mainly monogamous/polyandrous mating system. We used a combination of behavioral observations and endocrine data to determine female reproductive status and to examine changes in sociosexual behaviors over the ovarian cycle and between conceptive and nonconceptive cycles. Females clearly signaled their reproductive status by way of proceptive sexual presenting. Males showed increased frequencies of anogenital sniffing and mounting during the fertile period, indicating that they detected changes in olfactory and behavioral cues emitted by females, and they adjusted their mounting behavior accordingly. Males and females also remained in closer proximity before and during the fertile period, which suggests the existence of mate guarding. We discuss a possible function of behavioral advertisement of reproductive status in shaping the mating system in Leontopithecus chrysomelas.