Abstract

Factors contributing to breeding success of puffins were studied on Great Island, Newfoundland, in 1968 and 1969. Puffin burrow density was negatively correlated with distance from the cliff edge and positively correlated with angle of slope. These correlations are biologically significant in that close to the cliff edge, where the angle of slope was steep, breeding success was significantly higher than on adjacent level habitat. In spring both habitats were occupied simultaneously, and nest—site tenacity was equally strong in them. During settlement the frequency of fighting was higher and the peak was reached earlier on slope habitat. Males were heavier on slope than on level habitat just after peak egg laying, although wing lengths were similar. Females were similar on the two habitats. Measurements of eggs from both habitat were the same. Egg—laying dates were also similar, but annual variation was greater on level than on slope habitat, as was variation within a single year. Hatching success was higher on slope habitat, mainly because the incidence of egg disappearance was lower during incubation; also the frequency of infertile eggs was greater on level habitat. Fledging success was higher on slope habitat, and higher on both habitats in 1969. Frequency of chick deaths in the nest and disappearance before fledging was higher on level habitat in both years. Fledging success was higher for early—hatched chicks in both habitats. Total breeding success was higher on slope habitat, and higher in both habitats in 1969. Fledging condition of chicks varied according to the habitat and time period in which they were raised. On the average, body weight at fledging was greater and less variable for birds on slope habitat; early—hatched chicks were heaviest in both habitats. Wing length of young at fledging did not differ. Early—hatched chicks on slope habitat fledged quickest (also quicker than early—hatched chicks on level habitat); late—hatched birds on slope fledged slower than late—hatched birds on level habitat. Age at fledging was greater in 1968 than in 1969 on both habitats. Meal size delivered to chicks by parents was the same on the two habitats, but frequency of feeding was greater on slope than level habitat. Breeding pairs on slope habitat were unable to raise two chicks (artificial twins) to fledging; similarly, one adult could not rear a single chick. Adults feeding chicks on level habitat were attacked and robbed more frequently by gulls than birds on slope habitat, probably because escape (take—off) from an attack was quicker on slope than on level ground. Experiments performed to determine causes of the differential egg and chick loss before fledging in the two habitats showed that during incubation the proportion of eggs displaced to the burrow entrance by incubating birds when leaving the burrow in a hurry was the same, but the rate of panic flights was higher on level habitat. Also, chicks when starved spent more time at the burrow entrance than when fed regularly. It is concluded that the difference in breeding success on the two habitats was due to a higher exposure of eggs and chicks to gull predation on level habitat. The primary cause for this differential exposure was that adults on slope habitat were less vulnerable to gull disturbance during incubation and gull robbery when feeding their chicks. Thus breeding failures resulted from the interactions of food shortage and gull interference. This conclusion was tested by comparing breeding performance of birds at Great Island with birds at two islands where gull interference was absent. Egg and chick survival was greater under "gull—free" conditions. In addition, body weights at fledging were higher and less variable at the colonies without gull interference than at Great Island. The ways in which natural selection acts upon puffins at the breeding colony are considered; a model relating puffin nest distribution and habitat features is presented, and predictions which might be tested are outlined.

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