New fossil penguins (Aves, Sphenisciformes) from the Oligocene of New Zealand reveal the skeletal plan of stem penguins

Abstract

ABSTRACT Three skeletons collected from the late Oligocene Kokoamu Greensand of New Zealand are among the most complete Paleogene penguins known. These specimens, described here as Kairuku waitaki, gen. et sp. nov., and Kairuku grebneffi, sp. nov., reveal new details of key elements of the stem penguin skeleton associated with underwater flight, including the sternum, flipper, and pygostyle. Relative proportions of the trunk, flippers, and hind limbs can now be determined from a single individual for the first time, offering insight into the body plan of stem penguins and improved constraints on size estimates for ‘giant’ taxa. Kairuku is characterized by an elongate, narrow sternum, a short and flared coracoid, an elongate narrow flipper, and a robust hind limb. The pygostyle of Kairuku lacks the derived triangular cross-section seen in extant penguins, suggesting that the rectrices attached in a more typical avian pattern and the tail may have lacked the propping function utilized by living penguins. New materials described here, along with re-study of previously described specimens, resolve several long-standing phylogenetic, biogeographic, and taxonomic issues stemming from the inadequate comparative material of several of the first-named fossil penguin species. An array of partial associated skeletons from the Eocene–Oligocene of New Zealand historically referred to Palaeeudyptes antarcticus or Palaeeudyptes sp. are recognized as at least five distinct species: Palaeeudyptes antarcticus, Palaeeudyptes marplesi, Kairuku waitaki, Kairuku grebneffi, and an unnamed Burnside Formation species.