The annual cycle and breeding behaviour of the Mauritius Fody Foudia rubra

Safford, R.J. 1997. The annual cycle and breeding behaviour of the Mauritius Fody Foudia rubra. Ostrich 68 (24): 58–67.

The Mauritius Fody Foudia rubra, a highly endangered species of weaver (Ploceidae) endemic to Mauritius, was studied from 1989–1993. The plumage, various display postures (typical of the family), and seven adult and two juvenile vocalisations are described. The diet consisted of insects, nectar and fruit. Males were in breeding plumage, and breeding took place, between late August and early April, although severe weather in February 1992 terminated breeding activity early. The species appeared to be monogamous and was suspected to maintain a long-term pair bond. Pairs occupied exclusive territories of not less than 0.9 ha. Both sexes built the nest from the outset, but only the female lined it. Between breeding attempts, some pairs habitually started but then abandoned one or more nests. Clutch sizes of two to four were recorded, three being the norm. Incubation and brooding were carried out by the female; the male joined in feeding the young after daytime brooding ceased. Juvenile dependency appeared to last around two weeks after which the young were ousted from the territory. The potential productivity of up to three broods per pair per year was not achieved by any pair studied because of poor nesting success. A complete moult followed breeding. Territorial defence continued throughout the year, and no evidence for seasonal movements was seen. Two behavioural features seem unexpected: 1) male nest invitation and nest advertisement behaviour appeared to be absent; 2) females participated from the outset of nest-building (including prospecting). These could be explained by the existence of a long-term pair bond, which needs to be confirmed but would not be surprising in a species that remains on territory all year. Hypotheses that could be tested to find a mechanism responsible for the unusual features of the fody are suggested. In comparison with other fodies studied, the breeding behaviour most resembled that of the Seychelles Fody F. sechellarum (also a monogamous, non-graminivorous omnivore that breeds in solitary pairs in evergreen forest). The annual cycle was similar to that of other native passerines and most other fody taxa.     

First record of the fledging period of Temminck's Courser,Cursorius temminckii

The Seychelles Fody, Foudia sechellarum, is a ploceid weaver occurring naturally on three islands in the Seychelles group in the Indian Ocean. The population on Cousine Island was studied between 30 June and 25 August 1997. The size of the population on the island was estimated at 458–614 individuals and densities varied in different habitat types. As Seychelles Fodies in non-breeding plumage are difficult to sex, we provide sexing criteria based on wing length. Breeding pairs form small, probably temporary, territories that are defended by both partners against other fodies, including the introduced Madagascar Fody, Foudia madagascariensis. The Seychelles Fody often breeds semi-colonially and we observed up to five nests close together. The birds are socially monogamous and both sexes share in nest building, nest defense and provisioning of the young. Only females incubate. Many nests were deserted before eggs were laid, including some that had been accepted by the female. Additional males and females were sometimes seen helping provision the young, but this cooperative breeding behaviour appeared to be uncommon. Non-breeding individuals congregate in large flocks, sometimes joined by breeding birds.     

The annual cycle of the Toc-toc Foudia sechellarum on Cousin Island, Seychelles

The Seychelles Fody or Toc-toc shows a fairly well-defined annual cycle. A full pre-nuptial moult, lasting 90–100 days, occurs from February to May at the end of the wet season. During this period males enter eclipse plumage and there is little breeding. In most years there is a peak of breeding during the dry southeast monsoon from May to September. The subsequent period of post-fledging parental care is lengthy, up to four months. I suggest that this cycle, which minimizes variation in adult energy requirements through the year, may be particularly advantageous to a generalist feeder like the Toc-toc.
Annual adult mortality is not higher than 17–21%, a figure which agrees reasonably with observed juvenile recruitment of 15–16%. The Cousin population of this globally scarce species is estimated at 1300.     

MOULT OF BUDGERIGARS MELOPSZTTACUS UNDULATUS

In captive Budgerigars Melopsitticus undulatus moult of primaries started in the middle of the tract and moved progressively inwards and outwards, the inner feathers being replaced faster than the outer ones. Full replacement of primaries took six to eight months and a new cycle of moult usually started before completion of the old cycle. Moult of secondaries followed no clear pattern and occurred less frequently than moult of primaries. Moult of rectrices started with the middle pair and moved progressively outwards on both sides. Complete moult of rectrices took about six months and a new cycle often started before completion of the old. Moult of the head and body occurred intermittently throughout the year. Birds fledged in juvenal plumage, they passed into first basic plumage with a partial moult (head and body feathers) and into definitive basic plumage with a moult of all contour feathers.
In the field in inland mid-eastern Australia, there were some birds replacing feathers and some with complete plumage in most months of the year. Birds with complete plumage may have been between moults or within a moult and between replacement of feathers. The proportion of birds in moult did not increase in intensity after breeding, or cease during breeding or before movements. Some birds of both sexes with gonads in a reproductive condition were replacing feathers. Rirds that were replacing feathers had similar lipid deposits to birds that had a complete plumage.     

A twelve-month field study of the West African thrush Turdus pelios (Passeriformes: Muscicapidae). Part 2: annual cycles

In Africa, birds inhabiting forested regions are less seasonal in their activities than those from open areas. In order to study annual cycles in forest regions of South western Nigeria, West African Thrushes (Turdus pelios) were mist-netted and banded during the last two weeks of each month. The nest is a cup-shaped structure built out of grasses, herbs, weeds, roots and earth laid out in a clockwise manner. Only the nesting tree and feeding sites were defended during the breeding period. The clutch size was 2.69 +/- 0.20 eggs with a mean incubation period of 14.11 +/- 0.26 days. The mean nestling period was 15 +/- 1.00 days. The nestlings were fed on a variety of plant and animal matter, of which grass seeds and insects were predominant. Moult was found to be protracted with a population moult period of 194 days and a much shorter individual moult period. Moult and breeding periods were spread out: moult period dovetailed into the breeding period. The birds were found to gain weight during the period but they attained their maximum weight in August after the moult period. The lowest weight was recorded in February, during the peak of the dry season, when food availability was lower.     

Delayed plumage maturation increases overwinter survival in North Island robins

Many bird species show delayed plumage maturation (DPM), retaining sub-adult plumage until after their first breeding season. Most explanations assume that DPM increases fitness over the breeding season. However, unless birds undergo a full moult before breeding, DPM could also be an adaptation to increase survival over the previous winter. The winter adaptation hypothesis has never been tested owing to the difficulty of measuring overwinter survival. We experimentally tested this hypothesis in North Island robins (Petroica longipes) using a closed island population where we could accurately estimate survival. The experiment involved dyeing 41 juveniles to mimic adult males, and comparing their survival with 41 control juveniles treated with the same peroxide base minus the pigment. The population was monitored with a series of resighting surveys, and mark-recapture analysis used to estimate overwinter survival. Survival probability was estimated to be 10% for dyed birds versus 61% for control birds in 2001, and 29% for dyed birds versus 40% for control birds in the winter of 2002, supporting the winter adaptation hypothesis for DPM. Access to suitable habitat is the key factor limiting juvenile survival in this population, and the locations where dyed juveniles were sighted suggest that they were often excluded from suitable areas.     

Why renew fresh feathers? Advantages and conditions for the evolution of complete post‐juvenile moult

Juveniles of several passerine species renew all of their fresh juvenile feathers immediately after fledging (complete post‐juvenile moult), in contrast to the majority, which perform a partial post‐juvenile moult. To understand the adaptive roles of this phenomenon we compared the quality of juvenile plumage in species that perform a complete post‐juvenile moult with that of species which perform a partial post‐juvenile moult; we similarly compared juveniles and adults in each of these groups. The quality of feathers was measured by mass of primaries, colour, and length. In species which perform a complete post‐juvenile moult the plumage quality of second‐year individuals, in their first breeding season, is similar to the plumage quality of adults, unlike those species that perform a partial post‐juvenile moult. In species which perform complete post‐juvenile moult, the quality of the feathers grown in the nest is lower than the quality of adult post‐breeding feathers. In contrast, in species which perform partial post‐juvenile moult the quality of the feathers grown in the nest is similar to that of adult post‐breeding feathers. We found that a complete post‐juvenile moult strategy is much more common 1) in residents and short‐distance migrants than in long‐distance migrants, 2) in southern latitudes, 3) in species with medium body mass and 4) in omnivores and granivores. Our results indicate two adaptive roles of the complete post‐juvenile moult strategy: 1) achieving high quality plumage in the first year which may increase individual survival probability and fitness and 2) allocating fewer resources to nestling plumage and more to nestling development, which enables the nestlings to leave the nest earlier, thus reducing the probability of encountering nest predators. We suggest that the complete post‐juvenile moult, immediately after fledging, is an optimal strategy in favourable habitats and under low time constraints, as in some tropical ecosystems.     

FORAGING, SEASONALITY AND NESTING OF SEYCHELLES SUNBIRDS NECTARINIA DUSSUMIERI

Foraging and breeding behaviour of Seychelles Sunbirds Nectarinia dussumieri was studied for a total of four months on six islands in the Seychelles. Birds obtained nectar from indigenous and introduced plants directly (inserting the bill into the corolla-tube), perched or hovering, or indirectly (piercing the corolla). Direct methods were used in plants with corolla-tubes less than 20 mm (the length of birds' bills) and indirect methods in most longer flowers. Flowers with longer corolla-tubes appeared to be preferred. In feeding groups, birds fed more rapidly if not involved in aggressive interactions, which may reflect defence of feeding areas. Insects were taken from the foliage of indigenous and introduced trees, using a variety of techniques. Flycatching was most prevalent in small-leaved species, and upright probing in large-leaved species. In Calophyllum inophyllum, which held very high densities of insects, hovering was used only in saplings, while other methods were used in both mature trees and saplings. The percentage of nests that were active declined from September to December-January, suggesting a peak of breeding during this period, the first part of the northwest monsoon season. Examination of live birds and museum skins showed that wing-moult occurs chiefly from January-May, but males on Praslin also moult in August-September. Nests varied in conspicuousness, being best hidden on Mahé and Silhouette, the only islands with populations of Seychelles Kestrels Falco araea. Males contributed to the earliest stages of nest building, but all later building, as well as all incubation and most feeding of young was done by females. Both sexes defended the nest area against other birds, especially those attempting to steal nest material. Males advertized by song from prominent perches near nests; females also sang occasionally. Males are larger in most dimensions than females, but no inter-island differences were found. The species may have evolved duller plumage and larger size than related species, but this remains uncertain without further evidence on its origins. The clutch of one egg only represents a reduction from the normal clutch size of local mainland congeners.     

Carry‐over effects and compensation: late arrival on non‐breeding grounds affects wing moult but not plumage or schedules of departing bar‐tailed godwits Limosa lapponica baueri

In the annual cycle of migratory birds, temporal and energetic constraints can lead to carry‐over effects, in which performance in one life history stage affects later stages. Bar‐tailed godwits Limosa lapponica baueri, which achieve remarkably high pre‐migratory fuel loads, undertake the longest non‐stop migratory flights yet recorded, and breed during brief high‐latitude summers, may be particularly vulnerable to persistent effects of disruptions to their rigidly‐timed annual routines. Using three years of non‐breeding data in New Zealand, we asked how arrival timing after a non‐stop flight from Alaska (>11 000 km) affected an individual godwit's performance in subsequent flight feather moult, contour feather moults, and migratory departure. Late arrival led to later wing moult, but godwits partially compensated for delayed moult initiation by increasing moult rate and decreasing the total duration of moult. Delays in arrival and wing moult up to 34–37 d had no apparent effect on an individual's migratory departure or extent of breeding plumage at departure, both of which were extraordinarily consistent between years. Thus, ‘errors’ in timing early in the non‐breeding season were essentially corrected in New Zealand prior to spring migration. Variation in migration timing also had no apparent effect on an individual's likelihood of returning the following season. The bar‐tailed godwits’ rigid maintenance of plumage and spring migration schedules, coupled with high annual survival, imply a surprising degree of flexibility to address unforeseen circumstances in the annual cycle.     

Costs and benefits of variable breeding plumage in the red-backed fairy-wren

The red-backed fairy-wren is a socially monogamous passerine bird which exhibits two distinct types of breeding male, bright males that breed in bright red and black plumage and dull males that breed in dull brown plumage. Most males spend their first potential breeding season in dull plumage and subsequent breeding seasons in bright plumage, but a relatively small proportion of males develop bright plumage in their first breeding season. This study quantifies morphology, behavior, and reproductive success of dull and bright males to assess the adaptive costs and benefits of bright plumage while controlling for age. Older bright males (two years of age or older) attempted to increase their reproductive success via copulations with extrapair females, whereas younger (one-year old) bright males and dull males did not. Thus, older bright males spent less time on their own territories, intruded on neighboring groups with fertile females more frequently, gave more courtship displays, and had larger sperm storage organs than did younger bright males and dull males. Microsatellite analyses of paternity indicate that the red-backed fairy-wren has extremely high levels of sexual promiscuity, and that older bright males had higher within-brood paternity than dull males or younger bright males. Regardless of age, bright males were more attractive to females in controlled mate choice trials than were dull males, and both age classes of bright males obtained higher quality mates earlier in the breeding season than did dull males, when nesting success was higher. In conclusion, although it appears that bright plumage increases access to higher quality mates, age also plays a central role in determining a male's overall reproductive success because of the high levels of sexual promiscuity exhibited by the red-backed fairy-wren.     

Summer food of the golden plover Pluvialis apricaria at Hardangervidda, southern Norway

Golden plover stomachs were examined through the breeding season al Hardangervidda, southern Norway. The main foods were Diptera larvae (in particular Tipula excisa ), adult Coleoptera (notably Otiorrhynchus dubius and Carabidae), and in May and August berries of Empetrum hermaphroditum. Also some larvae of Coleoptera and Lepidoptera were taken. Adult Coleoptera were considerably more common in the diet than expected from abundance data, while Araneae were unimportant as food in spite of a high abundance. The males take the major parental duties after hatching, allowing the females a more undisturbed feeding, supposed to be strongly advantageous for them due to close proximity of egg-laying and the start of the postnuptial moult. This is reflected in the diet: in July males showed a larger tropic diversity, and ate more adult insects and less insect larvae than the females.     

Fat reserves of migrating passerines at arrival on the breeding grounds in Swedish Lapland

Mist-net capture data taken during 6 years (1988–1990 and 1992–1994) of field work were used to describe the arrival sequences and fat loads of nine species of migratory passerines which breed in a near-Arctic environment in Swedish Lapland. Long-distance migrants arrived with significantly larger mean fat reserves than did medium- and short-distance migrants. Long-distance migrants carried fat loads at arrival which corresponded to potential remaining flight distances between 242 and 500 km. When females and males arrived on the breeding grounds simultaneously, females carried significantly larger energy reserves than did males in seven out of nine species. In contrast, when the sexes showed a significant difference in median arrival date (two out of nine species), there was no difference in mean fat load carried into the breeding area. A relationship was found between the migratory habits and foraging ecology of each species and the amount of fat reserves at arrival, suggesting that species-specific migratory distances and feeding habits determine the amount of fat that is needed during the transition period between migration and onset of breeding. The short growing season in the study area restricts the time available for breeding and moult, and large energy reserves at arrival may speed up the breeding schedule to counteract possible time constraints. Overloading at the last stopover site during spring migration may be an adaptation allowing birds to cope with a restricted time frame for breeding and moult at high latitudes.     

Moult in adult Fiery-necked Nightjars Caprimulgus pectoralis ringed on Ranelia Farm,Cashel, Zimbabwe

Nothing has been published on the moult of the Fiery-necked Nightjar Caprimulgus pectoralis in Zimbabwe. However, most of the birds handled on Ranelia Farm, Cashel, during a study of nightjar breeding biology over four seasons, were examined for moult. Fiery-necked Nightjars were examined on over 70 occasions. Their annual moult occurs between late October and early March, commencing with the primaries, which moult descendantly. The secondaries, which moult ascendantly, follow after P5 has been shed, and so do the rectrices, which moult centrifugally, but R5 precedes R4. Body moult, which starts about the time that R1 is shed, progresses from the head across the neck to the rest of the dorsal plumage, and then over the throat and flanks to the ventral surface. The rictal bristles moult descendantly in time with the primaries. Several birds, some with primary moult scores as high as 18, had commenced moult while still tending young from the first brood, or incubating the eggs of a second, or replacement, clutch. The moult season overlaps the breeding season by about two months.     

BIOLOGY OF THE BANK CORMORANT,PART 2: MORPHOMETRICS,PLUMAGE, BARE PARTS AND MOULT

Cooper J. 1985. Biology of the Bank Cormorant, Part 2: Morphometrics, plumage, bare parts and moult. Ostrich 56: 79–85.

The Bank cormorant Phalacrocorax neglectus is a medium-sized, heavily built but relatively short-tailed cormorant. Males average 2 107 g, females 1794 g. There are significant differences, but overlaps, in mass, wing length, bill, tarsus and tail length between the sexes of breeding adults. The species is all black; adults have a white rump patch and white filoplumes on the head and neck in prenuptial plumage. The white rump patch is lost during breeding. Bare parts are black except for the iris (the only useful aging character) which is dark brown in juveniles, green in subadults and horizontally divided orange-brown above, green below, in adults. Partial albinism is common but leucism is not. Adults can moult while breeding but it seems likely that most moult occurs outside the breeding season. Adults and a single subadult examined exhibited Staffelmauser or stepwise moult of the primaries.     

Sex‐dependent response of primary moult to simulated time constraints in the rock sparrow Petronia petronia

There is growing evidence that moult speed affects plumage quality. In many bird species, males and females differ in terms of breeding effort, survival expectation and the relationship between fitness and plumage quality. Consequently, differences in moult strategies between the sexes can be expected. The aim of this study was to assess whether, under simulated time constraints and with no parental investment in the previous breeding season, males and females differed in: a) timing and duration of primary moult, b) growth rates of individual primary feathers, and c) number of concurrently growing feathers. We investigated the effect of time constraints generated by a treatment consisting of two decreasing photoperiods (slow changing photoperiod, SCP=2 min day^?1 and fast changing photoperiod, FCP=8 min day^?1) on the primary post‐nuptial moult of captive rock sparrows Petronia petronia. Females started to moult on average 14 and 15 days later than males in both experimental groups. Primary moult duration was 10 (FCP) and 24 (SCP) days longer in males than in females, and, within sex, 34 (females) and 48 (males) days longer in SCP birds than in FCP ones. Females renewed a larger number of primaries simultaneously (5.7% in FCP and 12.8% in SCP) and had a higher total daily feather mass grown (9.9% in FCP and 22.4% in SCP), even though daily growth rates of individual primaries did not differ between sexes. As a result, males and females completed their primary moult at the same time within treatment. The observed differences in timing, duration and energy allocation for primary moult between the sexes probably have a genetic basis, as birds did not engage in reproduction during the preceding breeding season.     

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).     

4. MIGRATION OF AFRICAN BIRDS

Dean, W. R. J. 1978. Plumage, reproductive condition and moult in non-breeding Wattled Starlings. Ostrich 49:97-101. A random sample of Wattled Starlings Creatophora cinerea taken from a large flock (about 5000 birds) that was exhibiting pre-breeding displays was found to have a male:female ratio of 63:50. Less than 15% of the total sample were in breeding plumage, in spite of testis and ovarian follicle development that suggested that the whole flock was capable of breeding. The weight of dehydrated flight muscles gave a reliable indication of general body condition, but could not be correlated with reproductive condition, The presence of helminths (Diplotriaena tricuspis) does not appear to affect the birds adversely. Of the 113 birds in the sample, 13.2% were in active primary moult, 53,9% were in new plumage, and 32.9% were either old or indeterminable. Of 108 stomachs examined. 38.8% (42) contained fruit, 41,6% (45) contained insects and fruit, 17,5% (19) contained insects and 49% (1) contained a lizard.     

Differential moult patterns in relation to antipredator behaviour during incubation in four tundra plovers

Golden plovers and Grey Plovers Pluvialis spp. all have very distinct breeding plumage rich in contrast, with a conspicuous black belly and breast bordered by a bright white fringe. Eurasian Golden Plovers are known partly to replace their breeding plumage with striped yellow feathers during incubation, different from both breeding and non-breeding plumages. In this study a similar partial breeding moult was observed in Pacific Golden Plovers and American Golden Plovers caught on the nest or collected during incubation, although the feathers did not differ clearly from those of non-breeders. This moult starts during incubation and precedes the post-breeding moult into non-breeding plumage. Because the lighter feathers reduce the contrast between the black belly and the white flanks, we suggest that during incubation the plumage characteristic that plays an important role in mate choice is no longer important; at this stage it is better for the bird to be inconspicuous. Additional information on museum skins of golden plovers and of Grey Plovers indicated that only the three golden plovers undergo this partial moult, but that Grey Plovers in general retain full breeding plumage throughout incubation. The three golden plovers also resemble each other in their generally very passive nest defence strategies. In contrast, the larger Grey Plovers actively chase and attack aerial and ground predators. Thus, a reduced conspicuousness of the body plumage during incubation is likely to benefit the golden plovers more than the Grey Plover. We suggest that nest defence behaviour, plumage characteristics and perhaps size have co-evolved as a response to different selection pressures in golden plovers and Grey Plover, but alternative hypotheses are also discussed.     

THE BREEDING SEASON OF BIRDS IN THE LOWLAND RAINFOREST AND IN THE MONTANE FOREST OF WEST CAMEROON

An analysis is made of over a thousand precise bird breeding records for a forest area in West Cameroon. In the lowland forest there is no universal breeding season although most families have a preferred nesting season. On the other hand. in the montane forest breeding is seasonal and takes place in the drier months. These conclusions are supported by field observations and by the examination of over 5000 birds for gonad state, plumage wear and moult.
The remarkable difference in seasonality between the lowland and montane avifaunas is discussed with reference to climate, particularly to the mist which envelops the montane forest in the rainy season.     

A sexual conflict in collared flycatchers,Ficedula albicollis: early male moult reduces female fitness

A sexual conflict over levels of parental care occurs in most animals with biparental care, and studies of sexual differences in levels of parental care have usually focused on its intra-annual fitness consequences. We investigated inter-annual fitness consequences of a sexual difference in timing of feather replacement (moult) in collared flycatchers (Ficedula albicollis). In this study, males overlapped reproduction and moult more often than females, they also initiated their moult at an earlier stage of breeding than females. Females mated to males with a moult-breeding overlap had significantly lowered survival chances than females mated with males initiating moult after breeding. Furthermore, females mated with moulting males risked a lowered future fecundity in terms of a delayed start to breeding in the following season. However, early moulting males achieved a similar reproductive success as males initiating moult after breeding. Likewise, male survival probability to the following breeding season did not differ between early and late moulting individuals, nor was there any evidence that males gained or lost in future mating advantages by moulting early. These results show not only that a sexual conflict over timing of moult may operate, but also that it can impose severe fitness consequences, in terms of reduced future fecundity and survival probability, upon the ''losing'' sex.