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    Parental behaviour and mating system of the fairy martin, Hirundo Ariel
    Magrath, Michael J. L. (University of Melbourne, 1997)
    The birds are unique among the major animal taxa in that biparental care is common, occurring in about 90% of the 9000 or so extant species. Among these biparental species there is dramatic variation, both across and within taxa, in the extent to which males and females contribute to incubation and nestling care. However, despite considerable interest, the causes of this variation generally remain poorly understood. The aim of this thesis was to examine a range of factors that may contribute to variation in the participation of males and females within a population of the socially monogamous fairy martin, Hirundo ariel. Fairy martins are endemic to Australia and nest colonially in aggregations of enclosed mud nests. The sexes are morphologically similar, although only the female acquires a brood patch during the breeding season. In the Yarra Valley, where this study was conducted, colonies ranged in size from 8 to 29 nests. Over the three years from 1993 to 1996, birds arrived in the study area in September and usually commenced laying in October, although nesting activities were generally asynchronous both between and within colonies. Most colonies contained active nests until February. An average of at least 16% of adults and 5% of fledglings returned to the study area in the following year. Returning adults generally nested at the same colony site as the previous year (68%), while most first year birds nested at sites other than their natal colony (75%). Adult males were more likely to return than adult females. Similarly, fledgling males were more likely to return than fledgling females (assuming an equal sex ratio at fledging). Both sexes participated in building the mud nest, incubating the clutch and feeding the nestlings. Clutch size ranged from two to five (mean = 3.5) and declined during the course of the season. The clutch was attended for an average of 88% of the time during daylight hours, although attendance generally declined during the course of the day. Females were more attentive of the clutch than males (56%), resulting primarily from longer incubation bouts, but also shorter recess periods. Females almost invariably attended the clutch overnight, and were usually accompanied by the male. Total clutch attendance did not vary with clutch size, however, male attendance increased with clutch size while female attendance decreased. Total attendance increased with clutch age resulting from an increase in female but not male attentiveness. Both sexes were more attentive of the clutch on cooler days, especially in the early morning and late afternoon. The duration of the incubation period varied from 12 to 18 days (mean = 13.7), and was shorter if the clutch was attended for a greater proportion of the day. A mean of 1.8 chicks fledged per completed clutch, while 60% of these clutches produced at least one chick. The period from hatching to fledging varied from 17 to 32 days (mean = 22.1), and was longer for larger broods. Fledging success was highest during the middle of the breeding season (December). Adverse weather conditions, resulting in clutch and brood abandonment, were the most common cause of nest failure, and on several occasions also resulted in adult mortality. Pairs produced from zero to eight fledglings per season (mean = 2.8), with those that commenced nesting earlier in the season having higher annual productivity. Parents visited larger broods more frequently, although the visit rate per chick declined with brood size. Visit rate generally increased with brood age, reaching a plateau between days 8 and 16, before declining until the chicks fledged. The relative contribution of the sexes did not vary with brood size or brood age. Brood visit rate generally increased with ambient temperature and decreased during periods of rainfall. Male participation in incubation (both absolute and relative to his mate) declined with an increase in the proportion of fertile females in the colony. Males were most 'responsive' to the availability of fertile females in the early morning, when copulations are likely to occur most frequently. Furthermore, the decline in male contribution with the proportion of fertile females was greater among males with smaller than average clutches. Male contribution to brood visits also tended to decline with an increase in the proportion of fertile females over the most demanding days of the nestling period. These patterns, each demonstrated for the first time, provide strong support for the presence of a reproductive trade-off for males between parental and extra-pair copulation effort. This trade-off may be widespread among the 90% of bird species where males contribute to parental care. Microsatellite parentage analysis revealed that 14% (29/207) of young were not sired by the putative father, while 2% (4/207) appeared to be the result of conspecific brood parasitism. The frequency of extra-pair fertilizations was not influenced by nesting synchrony, but tended to increase with colony size. The genetic fathers of extra-pair young had larger tarsi, greater previous breeding experience, and were more attentive to their own clutch than both the male that they cuckolded and the other males in the colony. Moreover, males with previous breeding experience had greater paternity of their own brood than those without. These results are consistent with female extra-pair mate choice for 'good genes'. Male incubation attendance, both across and matched for bird, was lower for clutches with reduced paternity. Furthermore, males with reduced paternity took longer in returning to their clutch after a disturbance than males with complete paternity. Together, these results suggested that males may adjust their level of parental investment in relation to their confidence of paternity. However, these results were confounded by the relationship between male contribution to incubation and the availability of fertile females in the colony. Mean nestling mass was lower in broods with reduced paternity however there was no evidence that this resulted from a reduced rate of feeding by cuckolded males. Among the Hirundinidae, and birds in general, species with male incubation are usually sexually monomorphic and have relatively low frequencies of extra-pair paternity. This study indicates that the biparental fairy martin conforms to this pattern. The evolution/ maintenance of male incubation in fairy martins may, in part, relate to the small body size and high nesting density of this species.