Abstract
Seabirds have become adapted for foraging in an oceanic environment that can be highly dynamic. Oceanographic processes determine the spatial distribution of seabird prey, while seasonality often has a temporal influence on prey availability. In penguins, these factors are reflected in the different species' foraging strategies. Penguins can broadly be categorized as inshore foragers that live in subtropical to temperate regions and profit from a stable food supply throughout the year close to their breeding sites, and offshore foragers that breed in a pelagic environment at higher latitudes where oceanographic processes and seasonality create much more dynamic, temporally limited prey situations. In this light, offshore foragers can be expected to be much more flexible in their foraging behaviour so as to quickly respond to changes in a dynamic marine environment, while inshore foragers are more likely to exhibit predictable foraging patterns. I examined the foraging ecology of two New Zealand penguin species - the offshore foraging Snares penguin Eudyptes robustus and the inshore foraging Yellow-eyed penguin Megadyptes antipodes and how their foraging strategies reflect an adaptation to the marine environment they exploit.
Diet composition of breeding Snares penguins (incubation and early chick-guard) was determined using the water-offloading method. Before the chicks hatched, the penguins generally brought little food back from their long foraging trips. During chick-guard, the stomach contents comprised mainly of crustaceans (~55%), fish (~24%) and cephalopods (~21 %). However, the presence at times of many fish otoliths and squid beaks suggests that the latter two prey classes may play an even more important role in the adults' diet than the simple percentages based on mass suggest. The penguins' nesting routines were strongly synchronised between the years and correlated with the onset of the spring planktonic bloom. Using GPS data loggers and dive recorders I found that during the incubation phase, male penguins that performed long (ea. 2 week) foraging trips exhibited a strong affinity to forage in the Subtropical Front some 200 km east of the Snares. At that stage (late mid-October) the front featured elevated chlorophyll a concentrations, a pattern that can be observed every year. Thus, it seems that the front represents a reliable and predictable source of food for the male penguins. After the males returned, the female penguins also performed long foraging trips (<1 week) but never reached the front, primarily because they had to time their return to the hatching of their chicks. After the chicks had hatched, the female Snares penguins were the sole providers of food. At this stage, the penguins performed short foraging trips (1-3 days) and foraged halfway between the Snares and Stewart Island (ea. 70-90 km north of the Snares), where nutrient-rich coastal waters flow eastwards to form the Southland Current. The penguins concentrated their diving effort in these waters, underlining the importance of the warm coastal waters as a food source for breeding Snares penguins. However, diving behaviour between 2003 and 2004 differed with penguins searching for prey at greater depths in the latter year. This underlines the Snares penguins' behavioural flexibility in response to a changing marine environment.
The Yellow-eyed penguins as typical inshore foragers showed very consistent foraging patterns at all stages. GPS logger deployments on penguins at Oamaru revealed that the birds foraged almost exclusively at the seafloor and targeted specific areas that featured reefs or epibenthic communities. As a result, the penguins' at-sea movements appeared conservative and at times almost stereotypic. Nevertheless, a comparison of Yellow-eyed penguins breeding on the adjacent Codfish and Stewart islands revealed a degree of plasticity in the species' foraging behaviour. Birds from Codfish Island extended their foraging ranges considerably and switched from primarily bottom to mid-water foraging during the post-guard stage of breeding. It seems likely that this switch is a result of enhanced feeding conditions (e.g. increased prey abundance/quality) in an area further away from the island, but the time required to get there renders this strategy not viable when chicks are small and need to be guarded and fed on a daily basis. As such, the change of behaviour represents a traditional pattern rather than a dynamic response to a sudden change in the marine environment. In comparison, penguins from Stewart Island showed consistent foraging patterns during all
stages of breeding. Given the high levels of chick starvation on Stewart Island, the lack of plasticity in foraging behaviour is surprising and might indicate that Yellow-eyed penguins find it difficult to react quickly to a sub-optimal food situation.
Overall, it seems that Yellow-eyed penguins show a specialisation for a consistent benthic environment and, thus, lack the behavioural flexibility apparent in Snares penguins, which find their food in a changing pelagic marine environment.