The basal penguin (Aves: Sphenisciformes) Perudyptes devriesi and a phylogenetic evaluation of the penguin fossil record

Bulletin of the American Museum of Natural History

Volumn , Issue 337, 2010, Pages 1-77

  Ksepka, Daniel T ^a   Clarke, Julia A ^b  

^a NORTH CAROLINA STATE UNIVERSITY   (United States)

^b UNIVERSITY OF TEXAS AT AUSTIN   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

AVES; EUDYPTES; EUDYPTULA; MEGADYPTES; SPHENISCIDAE; SPHENISCIFORMES; SPHENISCUS;

EID: 77953503093     PISSN: 00030090     EISSN: None     Source Type: Book Series    
DOI: 10.1206/653.1     Document Type: Article

References (178)

1. Acosta Hospitaleche, C. 2004. Los pinguinos (Aves, Sphenisciformes) fosiles de Argentina. Sistematica, biogeografia y evolution. Ph.D. dissertation, Universidad Nacional de La Plata.
2. Acosta Hospitaleche, C. 2005. Systematic revision of Arthrodytes Ameghino, 1905 (Aves, Spheniscidae) and its assignment to the Paraptenodytinae. Neues Jahrbuch für Geologie und Palaöntologie - Abhandlungen 7: 404-414.
3. Acosta Hospitaleche, C. 2007. Revísion sistemá-tica de Palaeospheniscus biloculata (Simpson) nov. comb. (Aves, Spheniscidae) de la Formacio ́n Gaiman (Mioceno Temprano), Chubut, Argentina. Ameghiniana 44: 417-426.
4. Acosta Hospitaleche, C., and J. Canto. 2005. Primer registro de cráneos asignados a Palaeospheniscus (Aves, Spheniscidae) procedentes de la Formación Bahía Inglesa (Mioceno Medío-Tardio), Chile. Revista Chilena de Historia Natural 78: 219-231.
5. Acosta Hospitaleche, C., L. Castro, C. Tambussi, and R.A. Scasso. 2008. Palaeospheniscus patagonicus (Aves, Sphenisciformes): new discoveries from the early Miocene of Argentina. Journal of Paleontology 82: 565-575.
6. Acosta Hospitaleche, C., M. Chávez, and O. Fritis. 2006. Pingüinos fó siles (Pygoscelis calderensis nov. sp.) en la Formación Bahía Inglesa (Mioceno Medio-Plioceno), Chile. Revista Geolo ́ gica de Chile 33: 327-338.
7. Acosta Hospitaleche, C., and M. Reguero. In press. First articulated skeleton of Palaeeudyptes gunnari from the late Eocene of Isla Marambio (Seymour Island), Antarctica Antarctic Science.
8. Acosta Hospitaleche, C., and M. Stucchi. 2005. Nuevos restos terciarios de Spheniscidae (Aves, Sphenisciformes) procedentes de la costa del Perú . Revista Española de Paleontología 20: 1-5.
9. Acosta Hospitaleche, C., C. Tambussi, and M. Cozzuol. 2004. Eretiscus tonnii (Simpson) (Aves, Sphenisciformes): materiales adicionales, status taxonómico y distribución geográ fica. Revista del Museo Argentino de Ciencias Naturales 6: 233-237.
10. Acosta Hospitaleche, C., C. Tambussi, M. Donato, and M. Cozzuol. 2007. A new Miocene penguin from Patagonia and its phylogenetic relationships. Acta Palaeontologica Polonica 52: 299-314.
11. Ameghino, F. 1905. Enumeración de los impennes fosiles de Patagonia y de la Isla Seymour. Anales del Museum Nacional de Buenos Aires (36) 6: 97-167.
12. Ando, T. 2007. New Zealand fossil penguins: origin, pattern, and process. Ph.D. dissertation, University of Otago.
13. Baker, A.J., S.L. Pereira, O.P. Haddrath, and K.-A. Edge. 2006. Multiple gene evidence for expansion of extant penguins out of Antarctica due to global cooling. Proceedings of the Royal Society Series B 217: 11-17.
14. Banks, J., A. van Buren, Y. Cherel, and J.B. Whitfield. 2006. Genetic evidence for three species of rockhopper penguins, Eudyptes chrysocome. Polar Biology 30: 61-67.
15. Baumel, J.J., and L.M. Witmer. 1993. Osteologia. In J.J. Baumel, A.S. King, J.E. Breazile, H.E. Evans, and J.C. Vanden Berge (editors), Handbook of avian anatomy: nomina anatomica avium: 45-132. Cambridge: Nuttall Ornithology Club.
16. Bertelli, S., and N.P. Giannini. 2005. A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology and mitochondrial sequences. Cladistics 21: 209-239.
17. Bertelli, S., N.P. Giannini, and D.T. Ksepka. 2006. Redescription and phylogenetic position of the early Miocene penguin Paraptenodytes antarcticus from Patagonia. American Museum Novitates 3525: 1-36.
18. Boessenkool, S., et al. 2009. Relict or colonizer? Extinction and range expansion of penguins in southern New Zealand. Proceedings of the Royal Society Series B 276: 815-821.
19. Bourdon, E., B. Bouya, and M. Iarochene. 2005. Earliest African neornithine bird: a new species of Prophaethontidae (Aves) from the Paleocene of Morocco. Journal of Vertebrate Paleontology 25: 157-170.
20. Chubb, A.L. 2004. New nuclear evidence for the oldest divergences among neognath birds: phylogenetic utility of ZENK (I). Molecular Phylogenetics and Evolution 30: 140-151.
21. Clarke, J.A., et al. 2007. Paleogene equatorial penguins challenge the proposed relationship between biogeography, diversity, and Cenozoic climate change. Proceedings of the National Academy of Sciences 104: 11545-11550.
22. Clarke, J.A., E.B. Olivero, and P. Puerta. 2003. Description of the earliest fossil penguin from South America and first Paleogene vertebrate locality of Tierra del Fuego, Argentina. American Museum Novitates 3423: 1-18.
23. Cooper, A., and D. Penny. 1997. Mass survival of birds across the Cretaceous-Tertiary boundary: molecular evidence. Science 275: 1109-1113.
24. Cooper, R.A. 2004. The New Zealand geological timescale. Lower Hutt, New Zealand: Institute of Geological and Nuclear Sciences Monograph 22.
25. Cracraft, J. 1985. Monophyly and phylogenetic relationships of the Pelecaniformes: a numerical cladistic analysis. Auk 102: 834-853.
26. Cracraft, J. 1988. The major clades of birds. In M.J. Benton (editor), The phylogeny and classification of the tetrapods. Vol.1. Amphibians, reptiles, birds: 339-361. Oxford: Systematics Association.
27. Cracraft, J., et al. 2004. Phylogenetic relationships among modern birds (Neornithes): towards an avian tree of life. In J. Cracraft and M.J. Donoghue (editors), Assembling the tree of life: 468-489. New York: Oxford Press.
28. Davis, P.G., and D.E.G. Briggs. 1998. The impact of decay and disarticulation on the preservation of fossil birds. Palaios 13: 3-13.
29. del Valle, R.A., D. Montalti, and M. Inbar. 2002. Mid-Holocene macrofossil-bearing raised marine beaches at Potter Peninsula, King George Island, South Shetland Islands. Antarctic Science 14: 263-269.
30. DeVries, T.J. 1998. Oligocene deposition and Cenozoic sequence boundaries in the Pisco Basin (Peru). Journal of South American Earth Sciences 11: 217-231.
31. DeVries, T.J. 2007. Cenozoic Turritellidae (Gastropoda) from southern Peru. Journal of Paleontology 81: 331-351.
32. DeVries, T.J., Y. Narváez, A. Sanfilippo, N. Malumian, and P. Tapia. 2006. New microfossil evidence for a late Eocene age of the Otuma Formation (southern Peru). In Extended Abstracts, 13 Congreso Peruana de Geología: 615-618. Lima: Sociedad Geoló gica del Perú .
33. Dingle, R.V., W.G. Siesser, and A.R. Newton. 1983. Mesozoic and Tertiary geology of Southern Africa. Rotterdam: Balkema.
34. Dunbar, R.B., R.C. Marty, and P.A. Baker. 1990. Cenozoic marine sedimentation in the Sechura and Pisco basins, Peru. Palaeogeography Palaeoclimatology Palaeoecology 77: 235-261.
35. Dutton, A.L., K.C. Lohmann, and W.J. Zinmeister. 2002. Stable isotope and minor element proxies for Eocene climate of Seymour Island, Antarctica. Paleoceanography 17: 1016-1029.
36. Elliot, D.H., and T.A. Trautman. 1982. Lower Tertiary strata on Seymour Island, Antarctic Peninsula. In C. Craddock (editor), Antarctic geoscience: 287-297. Madison: University of Wisconsin Press.
37. Emslie, S.D., L. Coats, and K. Licht. 2007. A 45,000 yr record of Adélie penguins and climate change in the Ross Sea, Antarctica. Geology 35: 61-64.
38. Emslie, S.D., and C. Guerra Correa. 2003. A new species of penguin (Spheniscidae: Spheniscus) and other birds from the late Pliocene of Chile. Proceedings of the Biological Society of Washington 116: 308-316.
39. Emslie, S.D., and W.P. Patterson. 2007. Abrupt recent shift in dC and dN values in Adélie penguin eggshell in Antarctica. Proceedings of the National Academy of Sciences 104: 11666-11669.
40. Emslie, S.D., and E.J. Woehler. 2005. A 9000-year record of Adélie penguin occupation and diet in the Windmill Islands, East Antarctica. Antarctic Science 17: 57-66.
41. Ericson, P.G.P., et al. 2006. Diversification of Neoaves: integration of molecular sequence data and fossils. Biology Letters 4: 543-547.
42. Esperante, R., L. Brand, K.E. Nick, O. Poma, and M. Urbina. 2008. Exceptional occurrence of fossil baleen in shallow marine sediments of the Neogene Pisco Formation, Southern Peru. Palaeogeography Palaeoclimatology Palaeoecology 257: 344-360.
43. Fain, M.G., and P. Houde. 2004. Parallel radiations in the primary clades of birds. Evolution 58: 2558-2573.
44. Feldmann, R.M., and I.W. Keyes. 1992. Systematic and stratigraphic review with catalogue and locality index of the Mesozoic and Cenozoic decapod Crustacea of New Zealand. New Zealand Geological Survey Record 45: 1-73.
45. Feldmann, R.M., C.E. Schweitzer, and D. Mc-Lauchlan. 2006. Additions to the records for decapod Crustacea from Motunau and Glenafric Beaches, North Canterbury, New Zealand. New Zealand Journal of Geology and Geophysics 49: 417-427.
46. Flynn, J.J., and C.C. Swisher, III. 1995. Cenozoic South American Land Mammal Ages: correlation to global geochronologies. In W.A. Berggren, D.V. Kent, M.-P. Aubry, and J. Hardenbol (editors), Geochronology, time scales and global stratigraphic correlation (SEPM Special Publication 54): 317-333. Tulsa, OK: Society for Sedimentary Geology.
47. Fordyce, R.E. 1991. A new look at the fossil vertebrate record of New Zealand. In P. Vickers-Rich, J.M. Monaghan, R.F. Baird, and T.H. Rich (editors), Vertebrate palaeontology of Australasia: 1191-1316. Melbourne: Pioneer Design Studio and Monash University.
48. Fordyce, R.E. 2003. Cetacean evolution and Eocene-Oligocene oceans revisited. In D.R. Prothero, L.C. Ivany, and E.A. Nesbitt (edi-tors), From greenhouse to icehouse: the marine Eocene-Oligocene transition: 154-170. New York: Columbia University Press.
49. Fordyce, R.E., and C.M. Jones. 1990. Penguin history and new fossil material from New Zealand. In L.S. Davis and J.T. Darby (editors), Penguin biology: 419-446. San Diego, CA: Academic Press.
50. Gadzdzicki, A., A. Tatur, U. Hara, and R.A. del Valle. 2004. The Weddell Sea formation: post Late-Pliocene terrestrial glacial deposits on Seymour Island, Antarctic Peninsula. Polish Polar Research 25: 189-204.
51. Gauthier, J. 1986. Saurischian monophyly and the origin of birds. Memoirs of the California Academy of Sciences 8: 185-197.
52. Giannini, N.P., and S. Bertelli. 2004. Phylogeny of extant penguins based on integumentary and breeding characters. Auk 121: 422-434.
53. Gill, E.D. 1959. Provenance of a fossil penguin from western Victoria. Proceedings of the Royal Society of Victoria 71: 121-123.
54. Göhlich, U.B. 2007. The oldest fossil record of the extant penguin genus Spheniscus - a new species from the Miocene of Peru. Acta Paleontologica Polonica 52:285-298.
55. Goloboff, P.A., J.S. Farris, and K.C. Nixon. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24: 1-13.
56. Grant-Mackie, J.A., and G.G. Simpson. 1973. Tertiary penguins from the North Island of New Zealand. Journal of the Royal Society of New Zealand 3: 441-452.
57. Graur, D., and D. Martin. 2004. Reading the entrails of chickens: molecular timescales of evolution and the illusion of precision. Trends in Genetics 20: 80-86.
58. Groth, J.G., and G.F. Barrowclough. 1999. Basal divergences in birds and the phylogenetic utility of the nuclear RAG-1 gene. Molecular Phylogenetics and Evolution 12: 115-123.
59. Hackett, S.J., et al. 2008. A phylogenomic study of birds reveals their evolutionary history. Science 320: 1763-1768.
60. Harrison, G.L., et al. 2004. Four new avian mitochondrial genomes help get to basic evolutionary questions in the Late Cretaceous. Molecular Biology and Evolution 21: 974-983.
61. Hartley, A.J., G. Chong, J. Houston, and A.E. Mather. 2005. 150 million years of climactic stability: evidence from the Atacama Desert, northern Chile. Journal of the Geological Society (London) Special Paper 162: 421-424.
62. Hebert, P.D.N., M.Y. Stoeckle, T.S. Zemlak, and C.M. Francis. 2004. Identification of birds through DNA barcodes. PLoS Biology 2: e312.
63. Huxley, T.H. 1859. On a fossil bird and a cetacean from New Zealand. Quarterly Journal of the Geological Society of London 15: 670-677.
64. Jadwiszczak, P. 2001. Body size of Eocene Antarctic penguins. Polish Polar Research 22: 147-158.
65. Jadwiszczak, P. 2003. The early evolution of Antarctic penguins In A.H.L. Huiskes, et al. 2003. Antarctic biology in a global context: 148-151. Leiden: Bachhuys.
66. Jadwiszczak, P. 2006a. Eocene penguins of Seymour Island, Antarctica: taxonomy. Polish Polar Research 27: 3-62.
67. Jadwiszczak, P. 2006b. Eocene penguins of Seymour Island, Antarctica: the earliest record, taxonomic problems and some evolutionary considerations. Polish Polar Research 27: 287-302.
68. Jadwiszczak, P. 2008. An intriguing penguin bone from the late Eocene of Seymour Island, Antarctic Peninsula. Antarctic Science 20: 589-590.
69. Jenkins, R.J.F. 1974. A new giant penguin from the Eocene of Australia. Palaeontology 17: 291-310.
70. Jenkins, R.J.F. 1985. Anthropornis nordenskjoeldi Wiman, 1905: Nordenskjoeld's giant penguin. In P.V. Rich and G.F. van Tets (editors), Kadimakara, extinct vertebrates of Australia: 183-187. Victoria: Pioneer Design Studio.
71. Jesinkey, C., R.D. Forsythe, C. Mpodozis, and J. Davidson. 1987. Concordant late Paleozoic paleomagnetizations from the Atacama Desert: implications for tectonic models of the Chilean Andes. Earth and Planetary Science Letters 85: 461-472.
72. Jouventin, P., P.M. Nolan, J. Ornborg, and F.S. Dobson. 2005. Ultraviolet beak spots in King and Emperor penguins. Condor 107: 144-150.
73. Kandefer, H.M. 1994. Róznorodnosc fauny pingwino ́w kopalnych antarktycznej Wyspy Seymour w oparciu o analize humeri z kolekcji Instytutu Biologii Filii Uniwersytetu Warszawskiego w Bialymstoku. Dissertation, Warszawski University.
74. Keller, G., et al. 1997. The Cretaceous/Tertiary boundary event in Ecuador: reduced biotic effects due to eastern boundary current setting. Marine Micropaleontology 31: 97-133.
75. Kerr, K.C.R., et al. 2007. Comprehensive DNA barcode coverage of North American birds. Molecular Ecology Notes 7: 535-543.
76. Kooyman, G.L. 2002. Evolutionary and ecological aspects of some Antarctic and sub-Antarctic penguin distributions. Oecologia 130: 485-495.
77. Ksepka, D.T. 2007. Phylogeny, histology and functional morphology of fossil penguins (Aves: Sphenisciformes). Ph.D. dissertation, Columbia University.
78. Ksepka, D.T., and S. Bertelli. 2006. Fossil penguin (Aves: Sphenisciformes) cranial material from the Eocene of Seymour Island (Antarctica). Historical Biology 18: 389-395.
79. Ksepka, D.T., S. Bertelli, and N.P. Giannini. 2006. The phylogeny of the living and fossil Sphenisciformes (penguins). Cladistics 22: 412-441.
80. Ksepka, D.T., J.A. Clarke, T.J. DeVries, and M. Urbina. 2008. Osteology of Icadyptes salasi, a giant penguin from the Eocene of Peru. Journal of Anatomy 213: 131-147.
81. Lambert, D.M., et al. 2002. Rates of evolution in ancient DNA from Adélie penguins. Science 295: 2270-2273.
82. Lambert, O., G. Bianucci, and C. de Muizon. A new stem-sperm whale (Cetacea, Odontoceti, Physeteroidea) from the Latest Miocene of Peru. Comptes Rendus Palevol 7: 361-369.
83. Livezey, B.C. 1989. Morphometric patterns in recent and fossil penguins (Aves, Sphenisciformes). Journal of the Zoological Society of London 219: 269-307.
84. Livezey, B.C., and R.L. Zusi. 2006. Higher-order phylogenetics of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. I. Methods and characters. Bulletin of the Carnegie Museum of Natural History 37: 1-544.
85. Livezey, B.C., and R.L. Zusi. 2007. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linnean Society 149: 1-95.
86. MacPhee, R.D.E. 2005. 'First' appearances in the Cenozoic land-mammal record of the Greater Antilles: significance and comparison with South American and Antarctic records. Journal of Biogeography 32: 551-564.
87. Maddison, W.P., and D.R. Maddison. 1992. MacClade: analysis of phylogeny and character evolution, Version 4.08. Sunderland, MA: Sinauer Associates.
88. Manegold, A. 2006. Two additional synapomorphies of grebes Podicipedidae and flamingos Phoenicopteridae. Acta Ornithologica 41: 79-82.
89. Marenssi, S.A., S.N. Santillana, and C.A. Rinaldi. 1998. Stratigraphy of the La Meseta Formation (Eocene), Marambio (Seymour) Island, Antarctica. In S. Casadío (editor), Paleógeno de América del Sur y de la Península Antarctica. Asociacón Paleontoló gica Argentia Publicación Especial 5: 137-146.
90. Marples, B.J. 1952. Early Tertiary penguins of New Zealand. New Zealand Geological Survey Paleontological Bulletin 20: 1-66.
91. Marples, B.J. 1953. Fossil penguins from the mid-Tertiary of Seymour Island. Falkland Islands Dependencies Survey Scientific Reports 5: 1-15.
92. Marples, B.J. 1960. A fossil penguin from the Late Tertiary of North Canterbury. Records of the Canterbury Museum 7: 185-195.
93. Marples, B.J. 1962. Observations on the history of penguins. In G.W. Leeper (editor), The evolution of living organisms: 408-416. London: Cambridge University Press.
94. Marples, B.J., and C.A. Fleming. 1963. A fossil penguin bone from Kawhia, New Zealand. New Zealand Journal of Geology and Geophysics 6: 189-192.
95. Mayr, G. 2004. Morphological evidence for sister group relationship between flamingos (Aves: Phoenicopteridae) and grebes (Podicipedididae). Zoological Journal of the Linnean Society 140: 157-169.
96. Mayr, G. 2005. Tertiary plotopterids (Aves, Plotopteridae) and a novel hypothesis on the phylogenetic relationships of penguins (Spheniscidae). Journal of Zoological Systematics and Evolutionary Research 43: 61-71.
97. Mayr, G. 2007. The contribution of fossils to the reconstruction of the higher-level phylogeny of birds. Species, Phylogeny and Evolution 1: 58-64.
98. Mayr, G., and J. Clarke. 2003. The deep divergences of neornithine birds: a phylogenetic analysis of morphological characters. Cladistics 19: 527-553.
99. McEvey, A.R., and W.J.M. Vestjens. 1973. Fossil penguin bones from Macquarie Island, Southern Ocean. Proceedings of the Royal Society of Victoria 86: 151-174.
100. McKitrick, M.C. 1991. Phylogenetic analysis of avian hindlimb musculature. Miscellaneous Publications University of Michigan Museum of Zoology 179: 1-85.
101. Muizon, C. de. 1981. Les vertébrés fossiles de la formation Pisco (Pérou). Première partie: deux nouveaux Monachinae (Phocidae: Mammalia) du Pliocène inférieur de Sud-Sacaco. Travaux de l'Institut Français d'É tudes Andines 22: 1-160.
102. Muizon, C. de. 1984. Les vertébrés fossiles de la formation Pisco, Pérou. Deuxième partie: les Odontocètes (Cetacea, Mammalia) du Pliocène inférieur de Sud-Sacaco. Travaux de l'Institut Français d'É tudes Andines 27: 1-888.
103. Muizon, C. de. 1988. Les vertébrés fossiles de la formation Pisco, Pérou. Troisième partie: les Odontocètes (Cetacea, Mammalia) du Miocène. Travaux de l'Institut Français d'É tudes Andines 42: 1-244.
104. Muizon, C. de. 1993. Walrus-like feeding adaptation in a new cetacean from the Pliocene of Peru. Nature 365: 745-748.
105. Muizon, C. de., and T.J. Devries. 1985. Geology and paleontology of late Cenozoic marine deposits in the Sacaco area (Peru). International Journal of Earth Sciences 74: 547-563.
106. Muizon, C. de., G. McDonald, R. Salas, and M. Urbina. 2003. A new early species of the aquatic sloth Thalassocnus (Mammalia, Xenarthra) from the late Miocene of Peru. Journal of Vertebrate Paleontology 23: 886-894.
107. Muizon, C. de., G. McDonald, R. Salas, and M. Urbina. 2004. The evolution of feeding adaptations of the aquatic sloth Thalassocnus. Journal of Vertebrate Paleontology 24: 401-413.
108. Mü ller, J., and R.R. Reisz. 2005. Four wellconstrained calibration points from the vertebrate fossil record for molecular clock estimates. BioEssays 27: 1069-1075.
109. Myrcha, A., et al. 2002. Taxonomic revision of Eocene Antarctic penguins based on tarsometatarsal morphology. Polish Polar Research 23: 5-46.
110. Myrcha, A., A. Tatur, and R. del Valle. 1990. A new species of fossil penguin from Seymour Island, West Antarctica. Alcheringa 14: 195-205.
111. Norell, M.A. 1992. The effect of phylogeny on temporal diversity and evolutionary tempo. In M.J. Novacek and Q.D. Wheeler (editors), Extinction and phylogeny: 89-118. New York: Columbia University Press.
112. Nunn, G.B., J. Cooper, P. Jouventin, C.J.R. Robertson, and C.G. Robertson. 1996. Evolutionary relationships among extant albatrosses (Procellariiformes: Diomedeidae) established from complete cytochrome b gene sequences. Auk 113: 784-801.
113. Nunn, G.B., and S.E. Stanley. 1998. Body size effects and rates of cytochrome b evolution in tube-nosed seabirds. Molecular Biology and Evolution 15: 1360-1371.
114. O'Hara, R.J. 1989. Systematics and the study of natural history, with an estimate of the phylogeny of penguins (Aves: Spheniscidae). Ph.D. dissertation, Harvard University.
115. Olson, S.L. 1983. Fossil seabirds and changing marine environments in the Late Tertiary of South Africa. South African Journal of Science 79: 399-402.
116. Olson, S.L. 1985. The fossil record of birds. In D.S. Framer, J.R. King, and K.C. Parkes (editors), Avian biology: 79-238. New York: Academic Press.
117. Olson, S.L., and Y. Hasegawa. 1979. Fossil counterparts of giant penguins from the North Pacific. Science 330: 51-57.
118. Olson, S.L., and Y. Hasegawa. 1996. A new genus and two new species of gigantic Plotopteridae from Japan (Aves: Plotopteridae). Journal of Vertebrate Paleontology 16: 742-751.
119. Palazzesi, L., V. Barreda, and R.A. Scasso. 2006. Early Miocene spore and pollen record of the Gaiman Formation (northeastern Patagonia, Argentina): correlations and paleoenvironment implications. In Abstracts 4th Congreso Latinoamericano de Sedimentología and 11th Reunion Argentina de Sedimentología: 11. San Carlos de Bariloche.
120. Parras, A., et al. 2008. Correlation of marine beds based on Sr- and Ar-date determinations and faunal affinities across the Paleogene/Neogene boundary in southern Patagonia, Argentina. Journal of South American Earth Sciences 26: 204-216.
121. Paterson, A.M., G.P. Wallis, and R.D. Gray. 1995. Penguins, petrels, and parsimony: does cladistic analysis of behavior reflect seabird phylogeny? Evolution 49: 974-989.
122. Pycraft, W.P. 1907. On some points in the anatomy of the emperor and adelie penguins. British National Antarctic Expedition (1901- 1904) Report 2: 1-21.
123. Reisz, R.R., and J. Mü ller. 2004. Molecular timescales and the fossil record: a paleontological perspective. Trends in Genetics 20: 569-597.
124. Rinaldi, C.A., A. Massabie, J. Morelli, H.L. Rosenman, and R. del Valle. 1978. Geologia de la isla Vicecomodoro Marambio. Contribuciones del Instituto Antá rtico Argentino 217: 5-43.
125. Ritchie, P.A., C.D. Millar, G.C. Gibb, C. Baroni, and D.M. Lambert. 2004. Ancient DNA enables timing of the Pleistocene origin and Holocene expansion of two Adélie penguin lineages in Antarctica. Molecular Biology and Evolution 21: 240-248.
126. Rivera, R. 1957. Muluscos fó siles de la Formación Parcas, departmento de Ica. Boletín de la Sociedad Geoló gica del Perú 32: 165-220.
127. Scarlett, R.J. 1983. Tereingaornis moisleyi - a new Pliocene penguin. New Zealand Journal of Geology and Geophysics 26: 419-428.
128. Scasso, R.A., J.M. McArthur, C.J. del Río, S. Martínez, and M.F. Thirlwall. 2001. 87Sr/86Sr Late Miocene age of fossil molluscs in the 'Entrerriense' of the Valdés Peninsula (Chubut, Argentina). Journal of South American Earth Sciences 14: 319-329.
129. Schreiweis, D.O. 1982. A comparative study of the appendicular musculature of penguins (Aves: Sphenisciformes). Smithsonian Contributions to Zoology 341: 1-46.
130. Shaul, S., and D. Graur. 2002. Playing chicken (Gallus gallus): methodological inconsistencies of molecular divergence date estimates due to secondary calibrations. Gene 300: 59-61
131. Sibley, C.G., and J.E. Ahlquist. 1990. Phylogeny and classification of birds: a study in molecular evolution. New Haven, CT: Yale University Press.
132. Simpson, G.G. 1946. Fossil penguins. Bulletin of the American Museum of Natural History 87(1): 1-99.
133. Simpson, G.G. 1957. Australian fossil penguins, with remarks on penguin evolution and distribution. Records of the South Australian Museum 13: 51-70.
134. Simpson, G.G. 1959. A new fossil penguin from Australia. Proceedings of the Royal Society of Victoria 71: 113-119.
135. Simpson, G.G. 1965. New record of a fossil penguin in Australia. Proceedings of the Royal Society of Victoria 79: 91-93.
136. Simpson, G.G. 1970. Miocene penguins from Victoria, Australia, and Chubut, Argentina. Memoirs of the National Museum of Victoria 31: 17-24.
137. Simpson, G.G. 1971a. Review of fossil penguins from Seymour Island. Proceedings of the Royal Society of London B Biological Sciences 178: 357-387.
138. Simpson, G.G. 1971b. A review of the pre-Pliocene penguins of New Zealand. Bulletin of the American Museum of Natural History 144(5): 321-378.
139. Simpson, G.G. 1971c. Fossil penguin from the Late Cenozoic of South Africa. Science 171: 1144-1145.
140. Simpson, G.G. 1972a. Conspectus of Patagonian fossil penguins. American Museum Novitates 2488: 1-37.
141. Simpson, G.G. 1972b. Pliocene penguins from North Canterbury. Records of the Canterbury Museum 9: 159-182.
142. Simpson, G.G. 1973. Tertiary penguins (Sphenisciformes, Spheniscidae) from Ysterplaats, Cape Town, South Africa. South African Journal of Science 69: 342-344.
143. Simpson, G.G. 1975. Notes on variation in penguins and on fossil penguins from the Pliocene of Langebaanweg, Cape Province, South Africa. Annals of the South African Museum 69: 59-72.
144. Simpson, G.G. 1976. Penguins: past and present, here and there. New Haven, CT: Yale University Press.
145. Simpson, G.G. 1979a. Tertiary penguins from the Duinefontein site, Cape Province, South Africa. Annals of the South African Museum 79: 1-17.
146. Simpson, G.G. 1979b. A new genus of Late Tertiary penguin from Langebaanweg, South Africa. Annals of the South African Museum 78: 1-9.
147. Simpson, G.G. 1981. Notes on some fossil penguins, including a new genus from Patagonia. Ameghiniana 18: 266-272.
148. Slack, K.E., A. Janke, D. Penny, and U. Amason. 2003. Two new avian mitochondrial genomes (penguin and goose) and a summary of bird and reptile mitochondrial features. Gene 302: 43-52.
149. Slack, K.E., et al. 2006. Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution. Molecular Biology and Evolution 23: 1144-1155.
150. Somoza, R., and A. Tomlinson. 2002. Paleomagnetism in the Precordillera of northern Chile (22u309S): implications for the history of tectonic rotations in the Central Andes. Earth and Planetary Science Letters 194: 369-381.
151. Stanley, S.E., and R.G. Harrison. 1999. Cytochrome b evolution in birds and mammals: an evaluation of the avian constraint hypothesis. Molecular Biology and Evolution 16: 1575-1585.
152. Storer, R.W. 1971. Adaptive radiation of birds. In D.S. Farmer and J.R. King (editors), Avian biology, Vol.1: 149-188. New York: Academic Press.
153. Stucchi, M. 2002. Una nueva especie de Spheniscus (Aves: Spheniscidae) de la Formación Pisco, Perú . Boletín de la Sociedad Geoló gica del Perú 94: 17-24.
154. Stucchi, M. 2003. Los Piqueros (Aves: Sulidae) de la Formación Pisco. Boletín de la Sociedad Geoló gica del Perú 95: 75-91.
155. Stucchi, M. 2007. Los pingüinos de la Formación Pisco (Neógeno), Perú . In E. Díaz-Martínez and I. Rábano (editors), 4th European Meeting on the Palaeontology and Stratigraphy of Latin America Cuadernos del Museo Geominero 8: 367-373. Madrid: Instituto Geoló gico y Minero de España.
156. Stucchi, M., and S.D. Emslie. 2005. A new condor (Ciconiiformes, Vulturidae) from the late Miocene/ early Pliocene Pisco formation, Peru. Condor 107: 107-113.
157. Stucchi, M., and M. Urbina. 2004. Ramphastosula (Aves, Sulidae): a new genus from the Pisco formation, Peru. Journal of Vertebrate Paleontology 24: 974-978.
158. Stucchi, M., M. Urbina, and A. Giraldo. 2003. Una nueva specie de Spheniscidae del Mioceno Tardío de la Formacíon Pisco, Perú . Bulletin de l'Institut Français d'É tudes Andines 32: 361-375.
159. Swofford, D.L. 2003. PAUP*. Phylogenetic analysis using parsimony (* and other methods). Version 4. Sunderland, MA: Sinauer Associates. Tambussi, C.P., C.I. Acosta Hospitaleche, M.A.
160. Reguero, and S.A. Marenssi. 2006. Late Eocene penguins from West Antarctica: systematics and biostratigraphy. Geological Society of London Special Publications 258: 145-161.
161. Tambussi, C.P., M.A. Reguero, S.A. Marenssi, and S.N. Santillana. 2005. Crossvalia unienwillia, a new Spheniscidae (Sphenisciformes, Aves) from the late Paleocene of Antarctica. Geobios 38: 667-675.
162. Thompson, J.D., T.G. Gibson, F. Plewniak, F. Jeanmougin, and D.G. Higgins. 1997. The ClustalX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research 25: 4876-4882.
163. Uhen, M.D., and N.D. Pyenson. 2007. Diversity estimates, biases, and historographic effects: resolving cetacean diversity in the Tertiary. Palaeontologia Electronica 10 (2): 11A: 22 pp. (available at http://www.palaeo-electronica.org/ 2007-2/00123/index.html).
164. van Tets, G.F., and S. O'Conner. 1983. The Hunter Island penguin, an extinct new genus and species from a Tasmanian midden. Records of the Queen Victoria Museum 81: 1-12.
165. van Tuinen, M., and S.B. Hedges. 2001. Calibration of avian molecular clocks. Molecular Biology and Evolution 18: 206-213.
166. van Tuinen, M., C.G. Sibley, and S.B. Hedges. 2000. The early history of modern birds inferred from DNA sequences of nuclear and mitochondrial genes. Molecular Biology and Evolution 17: 451-457.
167. Walsh, S.A., and M.E. Suárez. 2006. New penguin remains from the Pliocene of Northern Chile. Historical Biology 18: 115-126.
168. Warheit, K.I., and D.R. Lindberg. 1988. Interactions between seabirds and marine mammals through time: interference competition at breeding sites. In J. Burger (editor), Seabirds and Other marine vertebrates. competition, predation and other interactions: 292-328. New York: Columbia Univerity Press.
169. Watanabe, M., et al. 2006. New candidate species most closely related to penguins. Gene 378: 65-73.
170. Watson, M. 1883. Report on the anatomy of the Spheniscidae collected during the voyage of H.M.S. Challenger. In J. Murray (editor), Report of the scientific results of the voyage of H.M.S. Challenger during the years 1873-1876, Zoology. Vol.7: 1-244. Edinburgh: Neill and Company.
171. Wiens, J.J. 1998. Incomplete taxa, incomplete characters, and phylogenetic accuracy: is there a missing data problem? Journal of Vertebrate Paleontology 23: 297-310.
172. Wilkinson, M. 1995. Coping with abundant missing data entries in phylogenetic inference using parsimony. Systematic Biology 44: 501-514.
173. Williams, T.D. 1995. The penguins. Oxford: Oxford University Press.
174. Wiman, C. 1905a. Vorfläufige Mitteilung über die alttertiären Vertebraten der Seymourinsel. Bulletin of the Geological Institute of Uppsala 6: 247-253.
175. Wiman, C. 1905b. Uber die alttertiären Vertebraten der Seymourinsel. Wissenschaftliche Ergebnisse der Schwedischen Südpolar-Expedition 1901-1903 3: 1-37.
176. Worthy, T.H. 1997. The identification of fossil Eudyptes and Megadyptes bones at Marfels Beach, Marlborough, South Island. New Zealand Natural Science 23: 71-85.
177. Worthy, T.H., and J.A. Grant-Mackie. 2003. Late-Pleistocene avifaunas from Cape Wanbrow, Otago, South Island, New Zealand. Journal of the Royal Society of New Zealand 33: 427-485.
178. Zusi, R.L. 1975. An interpretation of skull structure in penguins. In B. Stonehouse (editor), The biology of penguins: 55-84. Baltimore, MD: University Park Press.