Evolutionary and ecological aspects of some Antarctic and sub-Antarctic penguin distributions

Penguins probably originated in the core of Gondwanaland when South America, Africa, and Antarctica were just beginning to separate. As the continents drifted apart, the division filled with what became the southern ocean. One of the remaining land masses moved south and was caught at the pole by the Earth's rotation. It became incrusted with ice and is now known as East Antarctica. Linking it to South America was a series of submerged mountain ranges that formed a necklace of islands. The northern portion of the necklace, called the Scotia Arc, is now the "fertile crescent" of the Southern Ocean. The greatest numbers and biomass of penguins are found here as well as that of krill, the primary prey species of most penguins, and many other marine predators. Today penguins are found throughout the sub-Antarctic islands and around the entire Antarctic continent. Using satellite transmitters and time-depth recorders, while taking advantage of the parental dedication of breeding birds, numerous investigators have described foraging habits of several species of penguins. The information obtained is labor intensive and costly so that studies are restricted to certain species, areas and seasons. Here I review the patterns evident among six of the most abundant and completely studied of the penguins. The variation in behavior is considerable from those species that seldom dive deeper than 20 m in search of prey to those that will dive to depths >500 m to catch mesopelagic fish and squid. Foraging trips from breeding colonies vary among species and with the season. Often the birds travel no more than 30 km and at other times the trips may exceed 600 km. Sub-Antarctic species often reach more productive waters near or within the Antarctic Polar Front zone, where the mixing of Antarctic and sub-Antarctic waters provide rich resources for their prey. Antarctic species usually remain close to shore, along the continental slope, or near the sea ice edge. Less is known about penguins during the pelagic phase between breeding cycles. What we do know is surprising in regard to their dispersal, which ranges from hundreds to thousands of kilometers from the breeding colonies