--African
Penguins (Spheniscus demersus), which hrecd more successfully in burrows than on the
surface (Frost ctal. 1976a), arc subject to heat stress, which can cause them to desert their
nests (Randall 1983, Williams and Cooper 1984). As burrow-nesting reduces the effects of
ambient temperature extremes experienced by surface-nesting birds (Frost et al. 1976b), the
incidence of surface-nesting would be expected to be higher in the austral summer than in
winter when ambient temperatures arc higher and insolation most intense. Also, as burrows
in sandy soil are prone to collapse and flood during heavy rains (Frost et al. 1976b, pets.
ohs.), a higher incidence of surface-nesting would be expected in sandy areas. l tested these
two hypotheses during visits to Dassen Island (33ø25'S, 18ø05'E), on the west coast of South
Africa, where peak hrccding occurs in the austral winter (June) and summer (November-
December) (Cooper 1980).
From 1983 to 1985, two 200-m 2 burrow-nesting sites were checked for occupied nests.
132 THE WILSON BULLETIN ß Vol. 100, No. 1, March 1988
Site 1 was in an area of sandy soil; Site 2 was in an area of firmly packed gravel. The
numbers of surface nests and burrows in use at each site were counted during five visits in
winter (June-August) and summer (February).
African Penguins nested on the surface at sandy Site I more frequently in winter (19
surface nests, 111 burrows) than in summer (3 surface nests, 60 burrows) (X 2 = 4.08, df =
1, P < 0.05). Season had no effect on the number of surface nests at Site 2 (X 2 = 0.025,
df = 1, P < 0.01) (winter: 3 surface nests, 97 burrows; summer 3 surface nests, 85 burrows),
which is not susceptible to burrow-flooding and collapse. In winter a significantly greater
proportion of penguins nested on the surface at Site 1 (the sandy site), which was susceptible
to burrow-flooding and collapse than at Site 2 (X 2 = 8.81, P ( 0.01, df = 1). In summer
there was no significant difference in the proportion of surface nesters between the two sites
(X 2 = 0.176, P > 0.05, df = 1).
The results indicate that, for African Penguins, ambient temperature and insolation affect
the proportion of the population that occupies surface nests. Scolaro (1984) found that
Magellanic Penguins (S. magellanicus) show a tendency to nest in soils of high clay content
compared to soils of low clay content. The results of this study confirm the tendency of
penguins preferentially to burrow in suitable substrata and suggest that the quality of the
burrowing substratum is important for spheniscid penguins.
Acknowledgments.--I thank the Sea Fisheries Research Institute for transport to and
accommodation on Dassen Island, and D.C. Duffy and S. Hunter for commenting on the
manuscript. This work forms part of the Benguela Ecology Programme, sponsored by the
South African National Committee for Oceanographic Research.
LITERATURE CITED
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conservation. Pp. 227-231 in Proc. 4th Pan-African Ornithol. Congr., Mahe 1976.
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FROST, P. G. H., W. R. SIEGFRIED, AND J. COOPER. 1976a. Conservation of the Jackass
Penguin (Spheniscus demersus) (L.). Biol. Conserv. 9:79-99.
--,ADA.E. BuRGER. 1976b. Behavioural adaptations ofthe Jackass Penguin
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RANIALL, R.M. 1983. Biology of the Jackass Penguin (Spheniscus demersus) (L.) at St.
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