Breeding of the bellbird on the Poor Knights Islands,New Zealand

Abstract

The breeding of the bellbird (Anthornis melanura) was studied intensively over three seasons on Aorangi Island, Poor Knights Islands. Adult males defended territories all year but ventured beyond them to exploit localised food resources and to obtain water; some adults defended the same territory for at least 5 years. Adult females shared a territory with a male only during the breeding season. At other times of the year adult females were joined by juveniles and immatures and formed feeding flocks. The breeding season extended from late September to late December. A few nests were built on the ground but most were in dense vegetation, usually near the canopy. Peak egg-laying extended from mid-October to mid-November and only one clutch of two to four eggs was laid. Nest building and incubation were completed by the female alone but both parents fed nestlings. Fledglings stayed in the vicinity of the nest for several days, and were fed by both parents. Incubation and nestling periods were about 15 and 19 days respectively. Comparisons are made with the breeding biology of bellbirds and other native passerines on mainland New Zealand, and the importance of the predator-free enviomment of the Poor Knights Islands is stressed.     

Females compensate for moult‐associated male nest desertion in Hooded Warblers

Uniparental offspring desertion occurs in a wide variety of avian taxa and usually reflects sexual conflict over parental care. In many species, desertion yields immediate reproductive benefits for deserters if they can re‐mate and breed again during the same nesting season; in such cases desertion may be selectively advantageous even if it significantly reduces the fitness of the current brood. However, in many other species, parents desert late‐season offspring when opportunities to re‐nest are absent. In these cases, any reproductive benefits of desertion are delayed, and desertion is unlikely to be advantageous unless the deserted parent can compensate for the loss of its partner and minimize costs to the current brood. We tested this parental compensation hypothesis in Hooded Warblers Setophaga citrina, a species in which males regularly desert late‐season nestlings and fledglings during moult. Females from deserted nests effectively doubled their provisioning efforts, and nestlings from deserted nests received just as much food, gained mass at the same rate, and were no more likely to die from either complete nest predation or brood reduction as young from biparental nests. The female provisioning response, however, was significantly related to nestling age; females undercompensated for male desertion when the nestlings were young, but overcompensated as nestlings approached fledging age, probably because of time constraints that brooding imposed on females with young nestlings. Overall, our results indicate that female Hooded Warblers completely compensate for male moult‐associated nest desertion, and that deserting males pay no reproductive cost for desertion, at least up to the point of fledging. Along with other studies, our findings support the general conclusion that late‐season offspring desertion is likely to evolve only when parental compensation by the deserted partner can minimize costs to the current brood.     

Male parental care promotes early fledging in an open-nester, the Willow Warbler Phylloscopus trochilus

The importance of male parental care to female reproductive success was investigated in the monogamous Willow Warbler Phylloscopus trochilus by removing the male parent at two different stages of the breeding cycle. Females that were widowed at the start of egg-laying continued breeding and managed to raise their brood on their own with no apparent reductions in numbers fledged or fledgling body-mass. The widowed females compensated for the loss of male assistance by increasing their own food provisioning rate as compared with control females. However, widows spent less time brooding the small young, and the growth rate of nestlings was reduced. In nests where the male parent was removed 7 days after the eggs hatched, the subsequent growth rate of nestlings was still affected, which suggests that male care is influential throughout the nestling period. On average, broods reared by widows fledged 2 days later than did broods of control females. An extension of the nestling period may appreciably affect reproductive success, since 68% of nests failed due to predation, mostly during the nestling period. We suggest that the main role of male parental care in the Willow Warbler is to assure a high growth rate of nestlings, which leads to early fledging and hence a reduced risk of nest predation.     

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.     

The reproductive biology of the Red Shining Parrot Prosopeia tabuensis on the island of 'Eua, Kingdom of Tonga

On Eua, Red Shining Parrots Prosopeia tabuensis breed in the cool and dry season, i.e. from May to October. Climate is the main factor timing the breeding season, which gives additional evidence for an assumed origin of the genus from temperate regions. Two or three eggs are laid in cavities of forest trees; the incubation time is about 24 days. Only females were observed brooding and feeding the young. Nestlings were fed three times a day independently of age. Single nestlings and older siblings received almost twice as much food per feeding as younger siblings. Growth constants are presented for weight, the width of the upper mandible, and wing-length. The fledging age is reached after 7 weeks but none of the observed nests fledged young due to predation by man. Breeding success was calculated at more than 50% for those nestlings that would have fledged without human interference. The reproductive biology of the Red Shining Parrot does not, apparently, show island-specific adaptations such as a reduced clutch-size or a prolonged nestling period.     

Factors affecting the cost of polygynous breeding for female Great Reed Warblers Acrocephalus arundinaceus

The male's role in feeding young and his effect on the mortality of eggs and nestlings of the Great Reed Warbler Acrocephalus arundinaceus were studied in Kahokugata, central Japan, during eight breeding seasons. Three potential costs of polygynous breeding for females were examined:(i) a reduction of male parental care per brood, (ii) an attraction of predators due to concentration of nests, and (iii) an increase of unfertilized eggs due to the reduced frequency of copulation per female. Only the first cost was found to occur:polygynous males fed their young, especially of their second broods, less frequently than monogamous ones. The lowered paternal feeding frequency, however, did not increase the mortality by starvation in second-hatched broods. This was because the warm climate emancipated females from brooding and enabled them to compensate for the deficiency of feeding by males. The paper discusses possible roles of the staggering of the time of laying among harem mates and the shift of breeding status of females caused by nesting failure, in reducing potential costs of polygynous breeding for females.     

Effect of altitude on male parental expenditure in Mountain Bluebirds ( Sialia currucoides ): are higher-altitude males more attentive fathers?

Male investment of time and energy in caring for offspring varies substantially both between and within bird species. Explaining this variation is of long-standing interest to ornithologists. One factor that may affect male care is breeding site altitude, through its effects on climate. The harsher, less predictable abiotic conditions at higher altitudes are hypothesized to favour increased male investment of time and energy in offspring care. We tested this hypothesis by comparing male parental behaviour in Mountain Bluebirds (Sialia currucoides) nesting at 1500 and 2500 m a.s.l. in the Bighorn Mountains of Wyoming, USA. We compared rates of prey delivery to nestlings at these two altitudes at two times: 1–2 days after hatching, when females spend much of their time brooding young, and 12–13 days later, when brooding has ended and nestling energy demands are peaking. High-altitude males fed nestlings 18 and 28% more often than low-altitude males early and later in the nestling stage, respectively, but only the difference in late-stage feeding rates were significant. Like males, females at the high site also fed nestlings significantly more often than females at the low site later in the nestling stage (45% difference in feeding rates). Consequently, the proportion of all feeding trips made by males at the high site (40%) did not differ significantly from that at the low site (44%). Parents at the high altitude may feed nestlings more often to compensate for their greater thermoregulatory costs. Parents may also be attempting to assist nestlings in storing fat and/or attaining a large size and effective homeothermy as quickly as possible to enhance nestling ability to survive bouts of severe weather which are common at high altitudes.     

Correlations between body size, defensive behaviour and reproductive success in male Little Blue Penguins Eudyptula minor: implications for female choice

Field studies exploring relationships among body size, defensive behaviour (including vocalizations) and reproductive success in male Little Blue Penguins Eudyptula minor were conducted to identify behavioural and physical attributes that might provide females or opponents with information on male quality. Head length provided a simple estimator of body size in both sexes. Nests defended by large males produced eggs and chicks earlier in the breeding season, and contained chicks that grew more quickly than chicks in nests defended by small males. In contrast, female body size did not affect any of the reproductive parameters we measured. The pitch of male defensive calls varied significantly with male size, so females (or opponents) could potentially gauge male size by assessing the calls. Large males were also more likely to respond vocally to simulated nest intrusions. As a result, by 'keeping company' with prospective partners early in the breeding season, females can obtain information about males that would allow them to select better partners and improve their own reproductive potential.     

Intraspecific brood-parasitism and dispersal in fledgling Sparrowhawks Accipiter nisus

The progeny of early-nesting Sparrowhawks Accipiter nisus have a higher survival rate than those of late nesters. To ascertain the reasons for greater survival in early-season Sparrowhawk fledglings, I studied post-fledging dispersal behaviour in this species by direct observations and radio-tracking during 3 years in Rockingham Forest, Northamptonshire, U.K.
Post-fledging brood-parasitism was found among early-dispersed young of both sexes.
Early-dispersed young spent up to 6 days being fed by the parents of other fledged, but still dependent, broods, as far as 6 km from their own nests.
Three broods were provided with supplementary food for 4 weeks, starting 1 week before expected dispersal. These young dispersed when significantly older than young from control broods. In both groups, males dispersed, on average, 3–4 days earlier than females. The ultimate dispersal age of young in control broods was negatively correlated with their rate of mass gain during the nestling period. Unlike the young of the control broods, fledglings in broods with augmented food were usually silent.
These findings offer an explanation for why Sparrowhawk young that disperse early in the season survive better than those which disperse late.     

Active browsing by mouthbrooding females of Tropheus duboisi and Tropheus moorii (Cichlidae) to feed the young and/or themselves

Synopsis Fishes usually do not eat while brooding offspring in their mouths. In two epilithic algal eaters Tropheus duboisi and T. moorii in Lake Tanganyika, however, mouthbrooding females exhibited feeding actions. In T. duboisi, the feeding rate of mouthbrooding females was 80 percent of that of males and non brooding females irrespective of the developmental state of their offspring. In T. moori, females brooding early embryos rarely fed but their feeding rate increased with development of offspring. An examination of specimens revealed that such females took food for nourishment of themselves and the young in the former species but for nourishment of only the young in the latter.     

The role of male territories in the mating system of the Yellow-shouldered Widowbird Euplectes macrourus

The territorial system and breeding biology of the Yellow-shouldered Widowbird Euplectes macrourus (Ploceidae) was investigated in western Kenya. Yellow-shouldered Widowbirds had a resource-defence polygynous mating system: males defended large (mean = 0.95 ha) territories and built the coarse framing for the nests in tall grass. Males had up to five females nesting per territory. Females provided nearly all parental care except for a territorial male seen feeding a fledgling: the first observation of paternal care in the wild for this genus. There was considerable variation in territory size, but the cause of this variation remains unknown: territory size was not related to potential indicators of territory quality, such as grass height and abundance, did not relate to male morphology (mass, size and ornament size) or territorial behaviour (boundary displays and singing) and did not affect female preferences. Although resources (territories and nests) were defended by the males, observations that males frequently fed outside their territories and formed communal roosts during the breeding season suggest that this species represents a transitional stage between typical resource-defence polygyny and lek breeding.     

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.     

Successive nest building and polygyny of Fan-tailed Warblers Cisticola juncidis

The polygynous mating system of the Fan-tailed Warbler Cisticola juqcidis was investigated between 1978 and 1981. The male warbler builds many nests unaided; however, he has no more than one active vacant nest for courting at any time. Nest building lasted from April until August. After completing the outer fabric of a nest, the male advertised it and led a female to the nest site by a unique invitation flight. On average a male built 6.5 nests during one breeding season and three of them were accepted by females. The most successful male completed 18 nests, and mated with 11 females. Out of a total of 111 males which established a territory, 30 had no mate, 14 were monogamous, and the rest were polygynous. About 50 to 70% males were polygynous over the four years. The sex ratio varied from 1.41:1 to 2.17:1 (females: male) in the breeding population. It was partly caused by the presence of 'floating males'. After the completion of the outer fabric of the nest, the male warbler did not take any further role in nesting and caring for young.
The polygynous mating system of the Fan-tailed Warbler is characterized by successive nest building. Its extreme development results from the long breeding period and the male having no role in parental care.     

Mate choice by marsh wrens: the influence of male and territory quality

If females choose breeding situations to maximize their fitness, then those features of the male and/or territory that are important in mate selection by females should affect female fitness, be assessable prior to mating, and vary among males. The purpose of this study was to identify characteristics of male quality and territory quality that fit these criteria for marsh wrens Cistothorus palustris. The male's contribution to nest defence appeared not to affect female success or choice. However, females that received male assistance with feeding young produced more and heavier fledglings than females without assistance. Males that fed young did not attract more mates. Few males in this population fed young, and no relationship was found between feeding effort and physical or behavioural features of the male. Results of multivariate analyses suggest that, within sites, the measures of territory quality used here cannot explain the variance in the number of young fledged per female or male harem size. The examination of arbitrary territories showed that male pairing success, female settlement order and predation patterns were not significantly correlated between years. The results suggest that territory quality does not influence female success or choice. These results are discussed in the light of earlier attempts to relate features of males and territories to mate selection by female passerines.     

Effects of maternal quality and mating status on female reproductive success in the polygynous spotless starling

Female mating status may affect reproductive success in avian polygynous mating systems through a combination of differences in female parental quality and status-dependent male assistance in parental care. Traditionally the literature has emphasized male assistance, neglecting evidence for consistency in female parental quality independent of mating status or repeatability in status. We studied the effects of male assistance on breeding success and its association with female mating status in a population of the polygynous spotless starling, Sturnus unicolor, during 3 years. Nestling provisioning by males improved the fledging success of late (mostly second) but not early (mostly first) broods. Reproductive success of females was affected mainly by female maternal quality: (1) primary and secondary females did not differ in output for early broods and in seasonal output despite a greater male assistance at primary nests; (2) monogamous females were more successful despite receiving no more help than other females; and (3) primary and secondary females had different clutch sizes before male assistance in parental care could operate. Female mating status was not significantly repeatable within seasons or between years. Females who changed status between years were as successful when rearing broods as secondary females as they were as primary or monogamous females. Breeding success (proportion of eggs resulting in fledglings) and clutch size were significantly repeatable between years for late but not for early clutches. Performance showed no strong association with female age or condition. Female breeding success seems to be weakly affected by male assistance in this sedentary, colonial species, and more dependent on inherent differences between females. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Breeding adaptations of Franklin's gull (larus pipixcan) to a marsh habitat

The behaviour and ecology of Franklin's gull were studied at Agassiz National Wildlife Refuge in northwestern Minnesota to determine the adaptations of the species for nesting in marshes. Two factors seemed to be important in colony site selection: cattail dispersion pattern and cattail density. Franklin's gulls prefer to nest in cattail areas closest to open water. The number of nests per unit area decreased as cattail density increased. Nest site selection is dependent on aggression and visibility. Visibility from nest level is the result of cattail placement and height. The distance between nests was directly correlated with visibility. Aggression by gulls on nests was lowered experimentally by decreasing visibility and raised by increasing visibility. Nest platforms were constructed of cattail material, and were attached to cattail stems. Nest material was added to the nests throughout the incubation and brooding period. Material was usually added following nest relief. The egg laying period was from 6 to 28 May. There was more synchrony of egg laying in sub-areas of the colony than in the colony as a whole. Successive eggs in clutches were laid at 24- to 48-hr intervals. The distance between nests decreased during the season as pairs filled in areas that were not defended. Territorial pairs defended an area up to 10 m from their stations prior to egg laying, but defended only the area within 3 m of their nests during incubation. Both members of pairs incubated the eggs and cared for the young. The incubation period was 24 days. The primary predators on adults and young were marsh hawk, great horned owl and mink. Franklin's gulls do not eat eggs or young of gulls. Adults fed on earthworms, insects and grain. Most marked adults fed within 16 km of the colony. Chicks were fed primarily on earthworms. The hatching period was from 30 May to 21 June. Chicks of all ages tested on a visual cliff apparatus were able to perceive the drop. Chicks tested on a 30-degree incline apparatus walked up it when 6 days old and younger, and walked down at 12 days of age and older. Brood mobility was less than in ground nesting species of gulls. In an undisturbed colony the chicks remained on the nest platforms until they were 25 to 30 days old although they were capable of swimming shortly after hatching. Individual recognition between parents and chicks appeared later in this species than in ground-nesting gulls. Adults accepted alien chicks (experimentally exchanged) that were younger than about 14 days old until their own chicks were over that age. Adults accepted larger and older broods than their own, as well as broods of mixed ages. Chicks began to react differently to strange adults at about 16 days of age. The breeding chronology of Franklin's gull is compressed when compared to that of other gulls. Possible selection pressures affecting this synchrony are discussed. The behaviour of the marsh-nesting Franklin's gull is compared with that of typical ground-and cliffnesting gulls; the possibility that the ancestral gull may have been a marsh nester is discussed.     

Effects of mate removal on the behaviour and reproductive success of Reed Warblers Acrocephalus scirpaceus

The Reed Warbler Acrocephalus scirpaceus is a largely monogamous insectivorous passerine in which males and females provide equal care by day to eggs and chicks. Polygyny occurs occasionally, with males leaving one female unaided. When females were temporarily removed from three recently-completed clutches their males deserted and resumed the high song levels typical of unmated males. Males may desert either because they are physically incapable of incubation or because the energy expenditure needed for a male to return to an equivalent stage in the breeding cycle is much lower than for a female to do so. Widowed females ( n = 7 ), however, continued the breeding attempt alone, with similar incubation levels but higher provisioning rates than those of control females. In three out of four mid-season broods raised by lone females all fertile eggs were reared to healthy fledglings (in the fourth brood the female died), while only one of four late-season nests produced any fledglings (which were underweight). Late-season control nests were as successful as earlier ones. Loss of male help led to starvation of chicks, but caused no adverse effects during incubation. This explains the small changes in widows' sitting levels during incubation, but much greater effects after hatching. Females may need male help to rear late broods (but not early broods) as days are shorter and food is scarcer. Males may normally help at nests, even those in the mid-season, because in stressful spells (even for a few days) such help is vital for successful breeding but in good periods it costs the male little.     

Effects of Supplementary Feeding on the Breeding Ecology of the Buff-Throated Partridge in a Tibetan Sacred Site,China

Our goal was to document effects of year-round supplemental feeding on breeding ecology of the Buff-throated Partridge, Tetraophasis szechenyii, within a Tibetan sacred site. We evaluated effects of supplemental feeding used as religious/cultural practices which could potentially aid conservation of endangered phasianids. We compared fed breeding groups to neighboring nonfed groups. Fed groups initiated first clutches significantly earlier than nonfed groups. Earlier laying groups within fed and nonfed groups showed significantly lower hatching rates than later groups; however, fed groups showed significantly higher hatching rates than nonfed groups laying in the same period. Earlier laying increased opportunities to renest. All six fed groups with clutch failures renested compared to only one of five nonfed groups with clutch failures. Fed female breeders showed significantly greater investment in their young with larger clutches and larger eggs, which likely increased survivability of early hatchlings. We observed no predation on birds at feeding sites and recorded only four cases of predation on incubating females, which showed no detectable difference between fed and nonfed groups. Ground-nesting birds typically face high risks of predation. Ten of the 48 groups nested in trees, which occurs in few phasianid species. Tree nests showed significantly higher hatching rates compared to ground nests; however, we found no significant difference in tree nesting between fed and nonfed groups. This partridge is one of four gallinaceous species with cooperative breeding. Breeding groups with helpers had significantly greater reproductive success than single pairs, and fed female breeders with helpers laid bigger eggs than single pairs. Comparing annual reproductive output per group, fed groups not only produced significantly more independent young (≥150 days post-hatching), their young hatched significantly earlier, which likely have greater reproductive value over later hatched young of nonfed groups. Supplemental feeding year-round is likely what enabled the successes of the fed partridges.     

Breeding and management of the common trumpeter (Psophia crepitans)

The breeding biology and management of three wild-caught adult Common trumpeters (one male and two females) was documented at the Woodland Park Zoological Gardens, Seattle, from April, 1984 to August, 1986. A total of 27 eggs were laid, and eight young survived to fledging. Both sexes exhibited crane-like dances during courtship, but the male appeared to perform these behaviors more frequently and with greater intensity than the females. Courtship feeding and allo-preening also occurred. Nests consisted of simple scrapes on the ground, but the birds had no opportunity to rest in trees or other elevated sites. Clutches contained two or three eggs, and the incubation period was aproximately 28 days. Although the breeding pair was generally aggressive toward the second female, all three adults participated in incubation and in caring for the young. Parental behavior consisted of brooding, allopreening, and feeding. The male preened and fed one chick significantly more often than either of the females. Trumpeter chicks were highly precocial, but grew relatively slowly, reaching 50% of adult weight by 45 to 50 days of age. Trumpeters are difficult to maintain and breed in captivity and appear to be susceptible to mycotic diseases, such as aspergillosis. Changes in the social composition of captive groups may result in improved breeding.     

Provisioning behaviour at the nest in single-parent versus biparental nests and male versus female parents in the common redstart (<Emphasis Type="Italic">Phoenicurus phoenicurus</Emphasis>)

We analysed video-sequences of undisturbed parental provisioning behaviour on 12 nests of common redstart (Phoenicurus phoenicurus). In 4 of the 12 nests, chicks were fed by a single parent only. We compared provisioning rate of chicks, time spent on the nest and food allocation rules between nests with uniparental and biparental care and between male and female parents in biparental nests. In nests with a single parent, the frequency of feeding visits per parent was higher than in biparental nests. As a result, the rate of food provisioning of chicks was similar in uniparental and biparental nests. The food allocation rules did not differ between uniparental and biparental nests. In biparental nests, male and female provisioning behaviour was similar though with two exceptions: males had a strong preference for feeding chicks in front positions in the nest and females spent a longer time on the nest after feeding. We conclude that single common redstart parents are able to compensate fully for the absence of the other parent through increased provisioning efforts, and that in biparental nests, males and females contribute equally to the provisioning of the young.