Double brooding and polygyny in Sedge Warblers Acrocephalus schoenobaenus breeding in north-west England

Sedge Warbler breeding was studied at a site in Greater Manchester, UK, during 1991 and 1993. Birds were individually colour-ringed and, in the 2 years of the study, 18 pairs and 38 nesting attempts were observed. In both years, the majority of pairs (> 90%) attempted a second brood after successfully raising their first. Overall nest survival was 44% and the median reproductive output of individual males was 6.5 fledglings in each year (range 0–10 and 0–15 in 1991 and 1993, respectively). After fledging, one or both parents continued to feed the young for up to 15 days (median 11, range 0–15 days). One male was apparently bigamous in 1991 and in 1993, raising four broods successfully in 1993 and 25 fledglings over the 2 years. These observations suggest that double brooding is the normal strategy for Sedge Warblers in this population and that polygyny may be common, although easily overlooked.     

Cooperation and conflict among dunnocks, Prunella modularis, in a variable mating system

The mating system of a population of 90 breeding dunnocks (or hedge sparrow, Prunella modularis) included monogamy, polygyny, polyandry and polygynandry. Monogamous males guarded females during their fertile period to prevent neighbouring males from copulating. The most intense guarding occurred where two (unrelated) males shared a territory. Here, the alpha male tried to prevent the beta male from copulating with the female. Beta males were seen to copulate in only half the cases. They were more likely to succeed when the alpha male found it difficult to guard the female closely because her range was large, the vegetation was dense or there were other females breeding synchronously on the same territory. Close guarding and chasing by males reduced the female's feeding rate and was correlated with unhatched eggs in the nest. Females attempted to escape the alpha male's attentions and actively encouraged the beta male to mate. Beta males only helped to feed the young if they copulated with the female. Nestlings fed by two males and a female got more food and weighed more than those fed by just one male and a female. Indirect evidence suggested that when beta males failed to copulate, they destroyed eggs or young chicks. Females laid larger clutches when two males mated with them as opposed to one, thus adapting their clutch size to the amount of parental care they expected. The results of natural removal experiments and matched comparisons of the same female in different mating systems support these conclusions. For females, selection favours cooperative polyandry, whereas for males if favours polygyny; the variable mating system may reflect the different outcomes of this sexual conflict.     

Female dark‐eyed juncos Junco hyemalis thurberi produce male‐like song in a territorial context during the early breeding season

Reports of female song, once considered a rarity, have recently increased across a variety of avian taxa. Females of many species can be induced to produce male‐like song with exogenous testosterone, but observations of female song in free‐living birds remain limited by incomplete sampling of females. Here, we report three independent observations of female dark‐eyed juncos Junco hyemalis producing male‐like song early in the breeding season (i.e. post‐territory establishment, pre‐nesting) in a recently established non‐migratory, urban population. To elicit song, we presented 17 free‐living junco pairs with a live, caged female conspecific. Three unique females responded to our trials by diving at the intruding female, chasing their (male) mate, fanning their tail feathers, and singing a trilled song similar in structure to male long‐range (broadcast) song. We compared male and female songs quantitatively and found that the two sexes were statistically similar in many spectral and temporal characteristics, but female songs had significantly lower minimum and peak frequencies than males. This result is particularly surprising, as males in this urban population are known to sing at a significantly higher minimum frequency than males in a nearby montane population. Both the seasonal and social context in which these songs were observed suggest a potential function for female song in mate guarding and polygyny prevention, but more data are needed to test this hypothesis. Whether female song is common in all dark‐eyed juncos during the early breeding season or if it is restricted to this particular urban and non‐migratory population remains an important question for future research.     

Age-specific patterns of infection with haemosporidians and trypanosomes in a warbler: implications for sexual selection

Although the selective loss of individuals susceptible to disease can favor the evolution of female preference for older males, the interrelationship between age, infection, longevity, and mating success remains poorly characterized in natural populations. In a longitudinal study of 61 male common yellowthroats (Geothlypis trichas), we found that the probability of infection with hematozoa increased as males aged from 1 to 5 years. Despite a significant, negative association between infection and longevity that partially masked age-effects, the odds that a male was infected with Trypanosoma, Plasmodium, or Leucocytozoon increased 71–212% per year. Nearly 75% of males in their first breeding season were either uninfected or infected with only a single parasite, while 50% of older males were infected with at least two parasites and 16% were infected with all three. No males escaped infection after their second breeding season. Older males were also more likely to sire extra-pair young (EPY) and, as a consequence, infection with multiple parasites was associated with a fourfold increase in the odds of producing EPY. Unlike younger males, 80% of the oldest males had a history of either surviving chronic infection or recovering. Combined with previous work showing higher diversity at the major histocompatibility complex among older males, our results suggest that the song and plumage traits that signal male age in common yellowthroats also, perforce, signal resistance to parasites. By preferring older males, females may obtain good genes for disease resistance even in the absence of any effect of infection on male ornamentation.     

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.     

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.     

Sexual aggression by intruders in hooded crow <Emphasis Type="Italic">Corvus cornix</Emphasis>

The hooded crow Corvus cornix is a west Palaearctic, solitary nesting, monogamous corvid. In the breeding season, populations are characterized by a social organization wherein breeding pairs are territorial and non-breeding individuals, called floaters, live in flocks. During a study of the breeding ecology of the hooded crow, conducted in a protected flooded area, we monitored nests with video cameras. We recorded two separate incidents when intruders attacked a female at the nest. We believe that she remained in the nest in order to prevent the strangers cannibalizing the nestlings by mantling over the brood. The spatio-temporal occurrence of these attacks suggests that the observed behaviour is intraspecific sexual aggression wherein non-breeding males mounted an immobilized female.     

Intraspecific competition and the maintenance of monogamy in tree swallows

Intraspecific competition for access to breeding resources maylimit male mating success typically monogamous birds. We examinedthe potential for intraspecific competition to limit polygynyin tree swallows at Beaverhill Lake, Alberta, Canada. In thispopulation, polygynous males raisedmore fledglings than monogamousmales, and there was little or no cost to females from nestingpolygynously. Under these conditions one might expect polygynyto be more common than that observed(8% of males). We foundthat females were most aggressive toward conspecific intrudersearly in the breeding season. This aggression was associatedwith (1) females settling farther apart than expected underrandom settlement, (2) later settlement by secondary than bymonogamous females, and (3) no relationship between female settlementdate and male territory size instead of the negative correlationexpected if females settled randomly without competition. Earlyin the season, males also settled farther apart than expectedif they had settled randomly, and among males with two or morenest boxes on their territory, males with widely separated nestboxes were more likely to be polygynous. Monogamy is probablythe most common pairing association in this population becauseintraspecific competition for nest sites prevents most malesfrom gaining a territory with nest sites far enough apart topermit two females to breed without one female excluding theother. Females appeared to be defending an area surroundingtheir nest box to limit nest usurpation or intraspecific broodparasitism, rather than to limit any loss of male parental carefrom polygyny.     

Double brooding and offspring desertion in the barn owl Tyto alba

Many bird species produce two annual broods during a single breeding season. However, not all individuals reproduce twice in the same year suggesting that double brooding is condition‐dependent. In contrast to most raptors and owls, the barn owl Tyto alba produces two annual clutches in most worldwide distributed populations. Nevertheless, the determinants of double brooding are still poorly studied. We performed such a study in a Swiss barn owl population monitored between 1990 and 2014. The annual frequency of double brooding varied from 0 to 14% for males and 0 to 59% for females. The likelihood of double brooding was higher when individuals initiated their first clutch early rather than late in the season and when males had few rather than many offspring at the first nest. Despite the reproductive benefits of double brooding (single‐ and double‐brooded individuals produced 3.97 ± 0.11 and 7.07 ± 0.24 fledglings, respectively), double brooding appears to be traded off against offspring quality because at the first nest double‐brooded males produced poorer quality offspring than single‐brooded males. This might explain why females desert their first mate to produce a second brood with another male without jeopardizing reproductive success at the first nest. Furthermore, the reproductive cycle being very long in the barn owl (120 d from start of laying to offspring independence), selection may have favoured behaviours that accelerate the initiation of a second annual brood. Accordingly, half of the double‐brooded females abandoned their young offspring to look for a new partner in order to initiate the second breeding attempt, 9.48 d earlier than when producing the second brood with the same partner. We conclude that male and female barn owls adopt different reproductive strategies. Females have more opportunities to reproduce twice in a single season than males because mothers are not strictly required during the entire rearing period in contrast to fathers. A high proportion of male floaters may also encourage females to desert their first brood to re‐nest with a new male who is free of parental care duties.     

Within‐season variation in sexual selection in a fish with dynamic sex roles

The strength of sexual selection may vary between species, among populations and within populations over time. While there is growing evidence that sexual selection may vary between years, less is known about variation in sexual selection within a season. Here, we investigate within‐season variation in sexual selection in male two‐spotted gobies (Gobiusculus flavescens). This marine fish experiences a seasonal change in the operational sex ratio from male‐ to female‐biased, resulting in a dramatic decrease in male mating competition over the breeding season. We therefore expected stronger sexual selection on males early in the season. We sampled nests and nest‐holding males early and late in the breeding season and used microsatellite markers to determine male mating and reproductive success. We first analysed sexual selection associated with the acquisition of nests by comparing nest‐holding males to population samples. Among nest‐holders, we calculated the potential strength of sexual selection and selection on phenotypic traits. We found remarkable within‐season variation in sexual selection. Selection on male body size related to nest acquisition changed from positive to negative over the season. The opportunity for sexual selection among nest‐holders was significantly greater early in the season rather than late in the season, partly due to more unmated males. Overall, our study documents a within‐season change in sexual selection that corresponds with a predictable change in the operational sex ratio. We suggest that many species may experience within‐season changes in sexual selection and that such dynamics are important for understanding how sexual selection operates in the wild.     

Manipulations of male parental investment in polygynous pied flycatchers, Ficedula hypoleuca

We designed three experiments to identify important cues asto how bigamous male pied flycatchers (Ficedula hypoleuca) apportionnestling feeding between their broods. Normally, males givepriority to their primary brood, that is, the brood of theirfirst-acquired mate. In the first experiment, we reversed thehatching order of primary and secondary broods by substitutingeggs. Males responded by reallocating their efforts in favorof secondary broods. Males thus favored the brood that hatchedfirst, irrespective of female mating order. In the second experiment,carried out on the same males when the younger brood was 4 or5 days old, we transferred the older brood to the nest of theyounger, and vice versa; the males changed their investmentpattern accordingly, still giving priority to the older brood.In the third experiment, primary and secondary broods were madeto hatch on the same day. In these cases, males divided theirnestling feeding efforts fairly equally between the broods.The results reveal a remarkable flexibility of male investmentdecisions, which is discussed in light of parental investmenttheory. The fact that the degree of male assistance to secondarymates is variable and that it is to a large extent predictablefrom the nest initiation asynchrony of the two females has importantimplications for our understanding of the polyterritorial matingsystem of this species.     

Mate Choice and Mating Pattern in a Stream Goby of the Genus Rhinogobius

The mate choice and mating pattern of a benthic goby Rhinogobius sp. CB (cross band type) were investigated in the Kamo River, Shikoku, Japan. During the breeding season, gravid females assumed a nuptial color and either males or females initiated a courtship display. Males preferentially courted a female of similar size to lead her to his nest, whereas females courted more frequently when they encountered a large male. Eggs in any one nest were always at the same developmental stage. Sampling data of nesting males and females indicated that, in more than half the nests, males gathered more than one female before spawning. In some nests with eggs, two or three females had spent ovaries, indicating that the eggs were laid by multiple females within a short span of time. However, a comparison between the total number of eggs which females would spawn in one nest and the number of eggs actually deposited suggested that eggs were contributed by one female in most nests. This low level of polygyny in spite of multiple female availability is attributed to a limited available spawning area of the nest.     

Food availability and the male's role in parental care in double-brooded Treecreepers Certhia familiaris

The aim of this work was to examine differences in paternal and maternal care in a double-brooded, monogamous species, the Treecreeper Certhia familiaris, in relation to food availability. As a measure of parental care, we recorded the hourly feeding activity of parents when the nestlings from their first and second breeding attempts were 7 and 12 days old. Feeding frequency of the first brood increased with the age of the nestlings and also with the brood size when 12 days old. While the feeding activities of the females were similar with respect to the first and second broods, the males were less active and failed to provide any food to their nestlings in 15 cases out of 28 second broods. In spite of this, the fledglings from the second broods were heavier than those in the first. Such a pattern of male behaviour was possible without being a disadvantage to the chicks because the food supply increased during the breeding season and the female could provide food for the young alone. Thus paternal care was particularly important in times of poor food supply, i.e. during the first brood, where the extent of these males' activity in feeding the 7-day-old nestlings was positively correlated with the average mass of the nestlings. Our results support the idea that the male of monogamous, altricial bird species often makes important contributions to raising the young, especially during periods when it is difficult for the female to do so alone. Males show flexibility in their pattern of parental care, and male Treecreepers change their contribution to the first and second broods within the same season.     

Male ornamentation, timing of breeding, and cost of polygyny in the collared flycatcher

Highly ornamented males are often thought to be better ableto provide females with resources, parental assistance, or goodgenes. Individual variation in such male abilities may overridethe costs of polygyny and therefore largely explain within-populationvariation in mating patterns. We investigated the influenceof variation in male ornamentation and the environment on thecosts of polygyny for female collared flycatchers (Ficedulaalbicollis), using data from a long-term study involving 2733breeding attempts over 19 years. We show that females sufferreduced reproductive success when mated polygynously but thatthe costs of polygyny depend on an interaction between maleornamentation and timing of breeding. Among early breeders,polygynously mated females experience higher reproductive successwhen mated to less ornamented males, but among late breeders,females mated polygynously to highly ornamented males were moresuccessful. We suggest that a high effort spent on obtainingextrapair matings early in the season renders highly ornamentedmales less able to assist two females in caring for the young.Thus, a male's ability to simultaneously gain from extrapairmatings and polygyny may be limited through direct effects onfemale reproductive success. Given such limitation, extrapairmatings may be expected to be less frequent in species withbiparental care and a high level of social polygyny.     

Male-Biased Sexual Size Dimorphism,Resource Defense Polygyny,and Multiple Paternity in the Emei Moustache Toad (Leptobrachium boringii)

We tested the hypotheses that the Emei moustache toad (Leptobrachium boringii) exhibits resource defense polygyny and that combat led to the evolution of male-biased sexual size dimorphism. Between February and March of 2011 and 2012, 26 female and 55 male L. boringii from Mount Emei UNESCO World Heritage Site, Sichuan, China, were observed throughout the breeding season. Prior to the breeding season, males grow 10–16 keratinized maxillary nuptial spines, which fall off once the season has ended. Throughout this time, males construct and defend aquatic nests where they produce advertisement calls to attract females. In a natural setting, we documented 14 cases involving a total of 22 males where males used their moustaches for aggressive interaction, and nest takeover was observed on seven occasions. Males were also observed to possess injuries resulting from combat. Genetic analysis using microsatellite DNA markers revealed several cases of multiple paternity, both within nest and within clutch. This observation indicated that some alternative male reproductive strategy, such as satellite behaviour, is occurring, which may have led to the multiple paternity. Larger males were observed to mate more frequently, and in multiple nests, suggesting that females are selecting for larger males, or that larger males are more capable of defending high quality territories.     

Effect of female group size on harem male roosting behavior of the Indian short-nosed fruit bat <Emphasis Type="Italic">Cynopterus sphinx</Emphasis>

Mate guarding has been known to incur costs and cause constraints for harem males in many polygynous species. However, the effect of female group size on the harem male’s time budget in bats has received very limited attention. The Indian short-nosed fruit bat, Cynopterus sphinx, exhibits resource defense polygyny, in which tent roosting males construct tents and defend multiple female bats. We studied the effect of female group size on three aspects of harem male behavior: social grooming by reciprocal licking, tent maintenance, and tent guarding in the mast tree Polyalthia longifolia. In the process of reciprocal licking, all the bats in the harem were drenched in saliva before emergence, and this activity was positively and significantly correlated with female group size. Once females departed for foraging, harem males remained in their respective tents at night-time between intermittent foraging bouts and engaged in tent maintenance and tent guarding. Time invested by harem male bats in tent maintenance and tent guarding were positively and significantly correlated with female group size. Harem males extended their presence in tent by utilizing tents as feeding roosts. Female group size also influenced the emergence time of harem male bats, where males with largest group emerged later than did the smallest group. Likewise, harem male with the smallest group had more time available for foraging than the male with the largest group. Findings of this study suggest that having a larger harem may indeed be costly for the males by reducing their foraging time.     

SOCIAL ORGANIZATION AND NEST-BUILDING IN THE FOREST WEAVER BIRDS OF THE GENUS MALIMBUS (PLOCEINAE)

A comparative study was made of social organization during breeding among the genus Malimbus. In M. nitens, the male chooses the nest site, builds the nest alone, guards the nest during incubation, and feeds the young; the female incubates, broods alone and with the male feeds the young. In M. malimbicus, the male chooses the nest site, builds the nest with the female and guards the nest; the female builds the nest with the male, but incubates alone. In M. racheliae and M. cassini, the nest is built by one female and a multi-male party of two or three. One male drives off the other males when the nest is completed. One male and one female incubate alternately. The female seems to be the leader of the building group, and works like a male. In M. coronatus, the nest is built by a mixed party of males and females (3–6 birds), all working together without any overt leadership. Only one male and one female however, incubate, brood and feed the young. In their morphology and behaviour, Malimbus spp. are close to the weaver birds of the genus Ploceus. M. nitens seems the least evolved species while M. cassini and M. coronatus are behaviourally the most evolved. In the last species, which has a very elaborate nest, the pair of breeding birds is helped by one to four other birds. These helpers are birds in full adult plumage, and are probably capable of breeding and may do so at another period in the long breeding season of at least six months.     

Mate-sharing costs in polygynous Northern Lapwings Vanellus vanellus

In this study bigamous female Northern Lapwings Vanellus vanellus received significantly less incubation relief from their males than monogamous females. On average, monogamous males spent 34.3% of their time incubating and bigamous males 29.9%. Bigamous males divided their effort between their nests, incubating on average 9.4% on primary nests and 20.5% on secondary nests. Bigamous females compensated for the lack of male relief. Primary females incubated for 71.8% of their time, secondary females for 64.2%, while monogamous females spent 52.7% of their time incubating. As a result, there was no significant difference in total nest attentiveness among nests of different status. Primary and secondary females received equivalent incubation relief from the male. Bigamous males increased their contribution to incubation significantly as the season progressed. A bigamous male's distribution of incubation relief between his females was unrelated to female body mass, or to the degree of asynchrony between primary and secondary females in arrival and laying. Incubation time was significantly, negatively, correlated with total nest attentiveness. Monogamous females spent most time, secondary females spent an intermediate time, and primary females spent the least time on maintenance behaviour (foraging, comfort behaviour, inactivity). No significant differences were found in hatching success among females of different mating status. However, the ratio of unhatched to hatched eggs (i.e. the eggs that remained in the nest at the time of hatching) differed significantly: secondary females hatched a smaller proportion of their eggs than monogamous and primary females.     

THE BIRDS OF BLYTHSWOOD AND SOME NOTES ON BIRDS OF THE DISTRICT

Wintle, C. C. &; Taylor, P. B. 1993. Sequential polyandry, behaviour and moult in captive Striped Crakes Aenigmatolimnas marginalis. Ostrich 64:115-122.

Captive Striped Crakes showed sequential polyandry, the female laying for a second male when the clutch of her first mate was about to hatch. Where aviary space permitted each male set up a breeding territory and each female defended a larger area encompassing the territories of one or two males. Non-territorial subordinate males and females did not breed. The female initiated breeding by attracting the male and soliciting copulation, and the male incubated the eggs and cared for the young. Incubation took 17–18 days, the chicks left the nest at 4–5 days of age and were fully grown and capable of flight at 46–53 days. Breeding occurred from September to March and males normally reared two broods per season. Territoriality was evident only during the breeding season. Juvenile plumage was a duller version of the sexually dimorphic adult plumage; post-juvenile moult bean at 13–15 weeks and was complete at 21 weeks. Remex moult was simultaneous and a complete moult regular1 occurred twice a year in adults, in December and April (males) and September and March/April (females).     

Monogamy when there is potential for polygyny: tests of multiple hypotheses in a group-living fish

Monogamy within social groups where there exists a high potentialfor polygyny poses a challenge to our understanding of matingsystem evolution. Specifically, the traditional explanationthat monogamy evolves due to wide female dispersion, affordingmales little opportunity to defend multiple females, cannotapply. Instead, monogamy in groups potentially arises becausefemales compete for breeding resources such as breeding sites,food, and paternal care. We conducted manipulative experimentsto determine whether females compete over limiting resourceswithin groups of the obligate coral-dwelling goby, Paragobiodonxanthosomus (Gobiidae). Breeding females behaved aggressivelytoward individuals of their own sex and evicted subordinatefemales that were large and mature from the group. Experimentalremoval of nest sites caused breeding partners to breed in alternativenest sites, demonstrating that nest site limitation was notthe cause of female competition. Supplemental feeding resultedin an increase in the fecundity of breeding females but no maturationof subordinate females, demonstrating that food-limited femalefecundity was a likely cause of female competition. Finally,supplemental feeding of breeding pairs demonstrated that thedifference in eggs hatched by fed versus unfed males was lessthan the difference in eggs laid by fed versus unfed females,suggesting that paternal care limitation might also drive femalecompetition. These results suggest that competition over foodand possibly paternal care selects for dominant, breeding femalesto suppress the maturation of subordinate females to minimizecompetition. Monogamy in association with group living is thereforelikely to have evolved because female competition prevents malesfrom utilizing the potential for polygyny.