The phylogeny of the living and fossil Sphenisciformes (penguins)

  title={The phylogeny of the living and fossil Sphenisciformes (penguins)},
  author={Daniel T. Ksepka and Sara Bertelli and Norberto P. Giannini},
We present the first phylogenetic analysis of the Sphenisciformes that extensively samples fossil taxa. Combined analysis of 181 morphological characters and sequence fragments from mitochondrial and nuclear genes (12S, 16S, COI, cytochrome b, RAG‐1) yields a largely resolved tree. Two species of the New Zealand Waimanu form a trichotomy with all other penguins in our result. The much discussed giant penguins Anthropornis and Pachydyptes are placed in two clades near the base of the tree… 
A new Miocene penguin from Patagonia and its phylogenetic relationships
We describe a new medium−sized penguin, Madrynornis mirandus gen. et sp. nov., from the early late Miocene Puerto Madryn Formation, Chubut Province, Argentina. Although it is evident that extant and
Chatham Island Paleocene fossils provide insight into the palaeobiology, evolution, and diversity of early penguins (Aves, Sphenisciformes)
Numerous skeletal remains recovered in situ from the late early to middle Paleocene Takatika Grit of Chatham Island, New Zealand, are among the oldest known fossils attributed to the penguin clade
Phylogenetic Characters in the Humerus and Tarsometatarsus of Penguins
The present review aims to improve the scope and coverage of the phylogenetic matrices currently in use, as well as explore some aspects of the relationships among Paleogene penguins, using two key skeletal elements, the humerus and tarsometatarsus, using a corrected subset of morphological characters.
A review of Australian fossil penguins (Aves: Sphenisciformes)
Five named species are based on type specimens of Eocene, Miocene—Pliocene and Holocene age collected from South Australia, Victoria and Tasmania, and the phylogenetic affinities of these taxa remain unresolved.
Phylogenetic affinities and taxonomy of the Oligocene Diomedeoididae, and the basal divergences amongst extant procellariiform birds
A comprehensive collection of diomedeoidid fossils from the Rupelian stratotype in Belgium are described, which allows the recognition of previously unknown features of phylogenetic significance and contradicting recent proposals that Oceanitinae (southern storm-petrels) are the earliest diverging crown group Procellariiformes.
Patterning and Microstructure of Penguin Plumage APPROVED BY SUPERVISING COMMITTEE:
The synapomorphies recovered from the reassessed dataset differed relative to those previously recovered for extant penguin clades, and most of the differences involved characters that were rescored or rewritten due to inconsistencies between the original scoring and images or drawings of live birds.
A history of shifting fortunes for African penguins
Recovery of Miocene penguin remains is in accordance with earlier predictions of multiple pre-Pliocene colonizations of Africa and supports a higher level of ecological diversity amongst African penguins in the past.
Taxonomic revision and phylogenetic analysis of the flightless Mancallinae (Aves, Pan-Alcidae)
P phylogenetic results support placement of Mancallinae as the sister taxon to all other Alcidae, indicating that flightlessness evolved at least twice in the alcid lineage.
Phylogenetic Analysis of Pelecaniformes (Aves) Based on Osteological Data: Implications for Waterbird Phylogeny and Fossil Calibration Studies
Relationships of extant pelecaniforms inferred from morphology are more congruent with molecular phylogenies than previously assumed, though notable conflicts remain.
Bone histology in extant and fossil penguins (Aves: Sphenisciformes)
New observations on the long bone microstructure of penguins are presented, based on histological sections from two extant taxa and eight fossil specimens belonging to stem lineages, indicating that the modification of flipper boneMicrostructure continued long after the initial loss of flight in penguins.


Redescription and Phylogenetic Position of the Early Miocene Penguin Paraptenodytes Antarcticus from Patagonia
The placement of Paraptenodytes outside the crown clade of extant penguins reveals the order in which many spheniscid synapomorphies were acquired and lends support to the hypothesis that modern penguins had Subantarctic ancestors.
A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology and mitochondrial sequences
Molecular and morphological results indicated that the most significant problem in the phylogeny of extant penguins is rooting the ingroup, and the mutual interaction of molecular and Morphological data decreases the ambiguity regarding the placement of the root, and provides a resolved, relatively well‐supported phylogeny.
The deep divergences of neornithine birds: a phylogenetic analysis of morphological characters
A broad array of morphological characters (including both cranial and postcranial characters) are analyzed for an ingroup densely sampling Neornithes, with crown clade outgroups used to polarize these characters.
Fossil penguin (Aves: Sphenisciformes) cranial material from the Eocene of Seymour Island (Antarctica)
Our knowledge of the cranial morphology of early penguins remains poor, particularly for Paleogene taxa. This paper describes a partial penguin skull and additional isolated cranial elements from the
Description of the Earliest Fossil Penguin from South America and First Paleogene Vertebrate Locality of Tierra Del Fuego, Argentina
The discovery of the first vertebrate from the Paleogene of Tierra del Fuego (Isla Grande), Argentina, in southernmost South America is reported, with the earliest known penguin (pansphenisciform) from South America being found.
Taxonomic revision of Eocene Antarctic penguins based on tarsometatarsal morphology
Gradual cooling of climate, changes of environment and trophic relationships, that lasted several millions years, were most probably responsible for the intense speciation and taxonomic diversification of the Middle-Late Eocene La Meseta penguins.
Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution.
A test for events around the Late Cretaceous is reported by describing the earliest penguin fossils, analyzing complete mitochondrial genomes from an albatross, a petrel, and a loon, and describing the gradual decline of pterosaurs at the same time modern birds radiate.
A phylogeny of extant penguins (18 forms) was estimated on the basis of 70 integumentary and breeding characters, which recovered monophyly of Sphenisciformes and all the traditional genera.
Eocene penguins of Seymour Island, Antarctica: Taxonomy
It is suggested that ten species grouped in six genera are a minimal reliable estimate of the Eocene Antarctic penguin diver− sity, which may have co−existed in the Antarctic Peninsula region during the Late Eocene epoch.