Peer-Reviewed Papers
XX. Ksepka, D.T. and A.J.D. Tennyson. 2024. Oldest fossil record of Sulidae from New Zealand. Journal of Ornithology 165: 277-280
XX. Ksepka, D.T., A.J.D. Tennyson, M.D. Richards, and R.E. Fordyce. 2023. Stem albatrosses wandered far: a new species of Plotornis (Aves, Pan-Diomedeidae) from the earliest Miocene of New Zealand. Journal of the Royal Society of New Zealand, 1-17
XX. Ksepka, D.T., D.J. Field, T.A. Heath, W. Pett, D.B. Thomas, S. Giovanardi, and A.J.D. Tennyson. 2023 Largest-known fossil penguin provides insight into the early evolution of sphenisciform body size and flipper anatomy. Journal of Paleontology 97: 434-453
Thomas, D.B., A.J.D. Tennyson, F.G. Marx and D.T. Ksepka, D.T. 2023. Pliocene fossils support a New Zealand origin for the smallest extant penguins. Journal of Paleontology 97: 711-721
64. Ksepka, D.T., C.M. Early, K.M. Dzikiewicz, and A.M. Balanoff. 2022. Osteology and neuroanatomy of a phasianid (Aves: Galliformes) from the Miocene of Nebraska. Journal of Paleontology. [PDF]
63. Cole, T. L., C. Zhou, M. Fang, H. Pan, D. T. Ksepka, S. R. Fiddaman, C. A. Emerling, D. B. Thomas, X. Bi, Q. Fang, M. R. Ellegaard, S. Feng, A. L. Smith, T. A. Heath, A. J. D. Tennyson, P. G. Borboroglu, J. R. Wood, P. W. Hadden, S. Grosser, C.-A. Bost, Y. Cherel, T. Mattern, T. Hart, M.-H. S. Sinding, L. D. Shepherd, R. A. Phillips, P. Quillfeldt, J. F. Masello, J. L. Bouzat, P. G. Ryan, D. R. Thompson, U. Ellenberg, P. Dann, G. Miller, P. Dee Boersma, R. Zhao, M. T. P. Gilbert, H. Yang, D.-X. Zhang, and G. Zhang. 2022. Genomic insights into the secondary aquatic transition of penguins. Nature Communications 13:3912. [PDF]
62. Smith, N.A., K.L Koeller, J.A. Clarke, D.T. Ksepka, J.S. Mitchell, A. Nabavizadeh, R.C Ridgley, and L.M. Witmer. 2021. Convergent evolution in dippers (Aves, Cinclidae): the only wing-propelled diving songbirds. The Anatomical Record, 10.1002/ar.24820.
61. Giovanardi, S., D.T. Ksepka, and D.B. Thomas. 2021. A giant Oligocene fossil penguin from the North Island of New Zealand. Journal of Vertebrate Paleontology: e1953047.
60. Smaers, J.B., R.S. Rothman, D.R. Hudson, A.M. Balanoff, B. Beatty, D.K. Dechmann, D. de Vries, J.C. Dunn, J.G. Fleagle, C.C. Gilbert, A. Goswami, A.N. Iwaniuk, W.L. Jungers, M. Kerney, D.T. Ksepka, P.R. Manger, C.S. Mongle, F.J. Rohlf, N.A. Smith, C. Soligo, V. Weisbecker, and K. Safi. 2021. The evolution of mammalian brain size. Science Advances 7: 2101.
59. Yuan, M., D.Q. Li, D.T. Ksepka, and H.Y. Yi. 2021. A juvenile skull of the longirostrine choristodere (Diapsida: Choristodera), Mengshanosaurus minimus gen. et sp. nov., with comments on neochoristodere ontogeny. Vertebrata PalAsiatica 59: 213-228.
58. Thomas, D.B., A.J.D. Tennyson, R.P. Scofield, T.A. Heath, W. Pett, and D.T. Ksepka. 2020. Ancient crested penguin constrains timing of recruitment into seabird hotspot Proceedings of the Royal Society B 287: 20201497.
57. Ksepka, D.T., A.M.Balanoff, N.A. Smith, G.S.Bever. B.A.S. Bhullar, E. Bourdon, E.L. Braun, J.G. Burleigh, J.A.Clarke, M.W. Colbert, J.R. Corfield, F.J.Degrange, V.L.De Pietri, C.M. Early, D.J.Field, P.M. Gignac, M.E. L. Gold, R.T. Kimball, S. Kawabe, L. Lefebvre, J. Marugán-Lobón, C.S. Mongle, A. Morhardt, M.A. Norell, R.C.Ridgely, R.S. Rothman, R.P. Scofield, C.P. Tambussi, C.R.Torres, M. van Tuinen, S.A. Walsh, A. Watanabe, L.M. Witmer, A.K. Wright, L.E. Zanno, E.D.Jarvis, and J.B.Smaers. 2020. Tempo and pattern of avian brain size evolution. Current Biology 30: 2026–2036.
56. Field, D.J., J. Benito, A. Chen, J.W.M. Jagt, and D.T. Ksepka. 2020. Late Cretaceous neornithine from Europe illuminates the origins of crown birds. Nature 579: 397-401
55. Thomas, D.B., D.T. Ksepka, E.J. Holvast, A.J.D. Tennyson, P. Scofield. 2019. Re-evaluating New Zealand’s endemic Pliocene penguin genus. New Zealand Journal of Geology and Geophysics.
54. Pan, H., T. L. Cole, X. Bi, M. Fang, C. Zhou, Z. Yang, D.T. Ksepka, T. Hart, J. L. Bouzat, L. S. Argilla, M. F. Bertelsen, P. D. Boersma, C.-A. Bost, Y. Cherel, P. Dann, S. R. Fiddaman, P. Howard, K. Labuschagne, T. Mattern, G. Miller, P. Parker, R. A. Phillips, P. Quillfeldt, P. G. Ryan, H. Taylor, D. R. Thompson, M. J. Young, M. R. Ellegaard, M. T. P. Gilbert, M.-H. S. Sinding, G. Pacheco, L. D. Shepherd, A. J. D. Tennyson, S. Grosser, E. Kay, L. J. Nupen, U. Ellenberg, D. M. Houston, A. H. Reeve, K. Johnson, J. F. Masello, T. Stracke, B. McKinlay, P. G. Borboroglu, D.-X. Zhang, and G. Zhang. 2019. High-coverage genomes to elucidate the evolution of penguins. Gigascience 8.
52. Kimball, R.T., C.H. Oliveros, N. Wang, N.D. White, F.K. Barker, D.J. Field, D.T. Ksepka, R.T. Chesser, R.G. Moyle, M.J. Braun and R.T. Brumfield. 2019. A Phylogenomic supertree of birds. Diversity 11: 109. [PDF]
51. Musser, G., D.T. Ksepka and D.J. Field. 2019. New material of Paleocene-Eocene Pellornis (Aves: Gruiformes) clarifies the pattern and timing of the extant gruiform radiation. Diversity 11: 102. [PDF]
50. Oliveros, C.H., D.J. Field, D.T. Ksepka, F.K. Barker, A. Aleixo, M.J. Andersen, P. Alström, B.W. Benz, E.L. Braun, M.J. Braun, G.A. Bravo, R.T. Brumfield, R.T. Chesser, S. Claramunt, J. Cracraft, A.M. Cuervo, E.P. Derryberry, T.C. Glenn, M.G. Harvey, P.A. Hosner, L. Joseph, R. Kimball, A.L. Mack, C.M. Miskelly, A.T. Peterson, M.B. Robbins, F.H. Sheldon, L.F. Silveira, B.T. Smith, N.D. White, R.G. Moyle, B.C. Faircloth. 2019. Earth history and the passerine superradiation. Proceedings of the National Academy of Sciences 116: 7916-7925. [PDF]
49. Ksepka, D.T., L. Grande and G. Mayr. 2019. Oldest finch-beaked birds reveal parallel ecological radiations in the earliest evolution of passerines, Current Biology 29: 657-663.
48. Cole T. L., D.T. Ksepka, K.J. Mitchell, A.J.D. Tennyson, D.B. Thomas, H. Pan, G. Zhang, N.J. Rawlence, J.R. Wood, P. Bove, J.L. Bouzat, A. Cooper, S. Fiddaman, T. Hart, G. Miller, P.G. Ryan, L.D. Shepherd, J.M. Wilmshurst, J.M. Waters. 2019. Mitogenomes uncover extinct penguin taxa and reveal island formation as a key driver of speciation. Molecular Biology and Evolution 36: 784-797.
47. Degrange, F.D., D.T. Ksepka, and C.P. Tambussi. 2018. Redescription of the oldest crown clade penguin: cranial osteology, jaw myology, neuroanatomy, and phylogenetic affinities of Madrynornis mirandus. Journal of Vertebrate Paleontology 38: e1445636.
46 Ksepka, D.T., T.A. Stidham, and T.E. Williamson. 2017. Early Paleocene landbird supports rapid phylogenetic and morphological diversification of crown birds after the K–Pg mass extinction. Proceedings of the National Academy of Sciences 114: 8047-8052. [PDF]
45. Gavryushkina, A., T.A. Heath, D.T. Ksepka, T. Stadler, D. Welch, and A.J. Drummond. 2017. Bayesian total evidence dating reveals the recent crown radiation of penguins. Systematic Biology 66: 57-73. [PDF]
44. Thomas, D.B and D.T. Ksepka. 2016. The Glen Murray fossil penguin from the North Island of New Zealand extends the geographic range of Kairuku. Journal of the Royal Society of New Zealand 46: 200-213.
43. Ksepka, D.T., S. Werning, M. Sclafani, and Z.M. Boles. 2015. Bone histology in extant and fossil penguins (Aves: Sphenisciformes). Journal of Anatomy 227: 611-630.
42. Balanoff, A.M., G.S. Bever, M.W. Colbert, J.A. Clarke, D.J. Field, P.M. Gignac, D.T. Ksepka, R. Ridgely, N.A. Smith, C.R. Torres, S. Walsh, and L. Witmer. In press. Best practices for digitally constructing endocranial casts: examples from birds and their dinosaurian relatives. Journal of Anatomy.
41. Tambussi, C.P., F.J Degrange, and D.T. Ksepka. 2015. Endocranial anatomy of Antarctic Eocene stem penguins: implications for sensory system evolution in Sphenisciformes (Aves). Journal of Vertebrate Paleontology: e981635.
40. Ksepka, D.T. and M.J. Phillips. 2015. Avian diversification patterns across the K-Pg boundary: influence of calibrations, datasets, and model misspecification. Annals of the Missouri Botanical Garden 100: 300-328.
39. Ksepka, D.T., J.F. Parham, J.F. Allman, M.J. Benton, M.T. Carrano, K.A. Cranston, P.C. J. Donoghue, J.J. Head, E.J. Hermsen, R.B. Irmis, W.G. Joyce, M. Kohli, K.S. Lamm, D. Leehr, J.S.L. Patané, P.D. Polly, M.J. Phillips, N.A. Smith, N.D. Smith, M. van Tuinen, J.L. Ware, R.C.M. Warnock. 2015. The Fossil Calibration Database, a new resource for divergence dating. Systematic Biology 64: 853-859.
38. Smith, N.D. and Ksepka, D.T. 2015. Five well-supported fossil calibrations within the "Waterbird" assemblage (Tetrapoda, Aves). Palaeontologia Electronica 18.1.7FC.
37. Ksepka, D.T. and J.A. Clarke. 2015. Phylogenetically vetted and stratigraphically constrained fossil calibrations within Aves. Palaeontologia Electronica 18.1.3FC.
36. Ksepka, D.T. 2014. Flight performance of the largest volant bird. Proceedings of the National Academy of Sciences 111: 10624-10629. [PDF]
35. Ksepka, D.T., J.L. Ware and K.S. Lamm. 2014. Flying rocks and flying clocks: disparity in fossil and molecular dates for birds. Proceedings of the Royal Society B 281: 20140677. [PDF]
34. Thomas, D.B. and D.T. Ksepka. 2013. A history of shifting fortunes for African penguins. Zoological Journal of the Linnean Society 168: 207-219. [PDF]
33. Ksepka, D.T., J.A Clarke, S.J. Nesbitt, F. Kulpe and L. Grande. 2013. Fossil evidence of wing shape in a stem relative of swifts and hummingbirds (Aves, Pan-Apodiformes). Proceedings of the Royal Society B 280: 20130580. [PDF]
32. Ksepka, D.T., A.M. Balanoff, M.A. Bell and M.D. Houseman. 2013. Fossil grebes from the Truckee Formation (Miocene) of Nevada and a new phylogenetic analysis of Podicipediformes (Aves). Palaeontology 56: 1149–1169. [PDF]
31. *Cadena, E.A., D.T. Ksepka and M.A. Norell. 2013. New material of Mongolemys elegans Khosatzky and Mlynarski, 1971 (Testudines: Lindholmemydidae), from the Late Cretaceous of Mongolia with comments on bone histology and phylogeny. American Museum Novitates 3766: 1-27. [PDF]
30. Ksepka, D.T. A.M. Balanoff, S. Walsh, *A. Revan and *A. Ho. 2012. Evolution of the brain and sensory organs in Sphenisciformes: new data from the stem penguin Paraptenodytes antarcticus. Zoological Journal of the Linnean Society 166: 206-219. [PDF]
29. Ksepka, D.T., R.E. Fordyce, T. Ando and C.M. Jones. 2012. New fossil penguins (Aves: Sphenisciformes) from the Oligocene of New Zealand reveal the skeletal plan of stem penguins. Journal of Vertebrate Paleontology 32: 235-254. (Cover Article) [PDF]
28. Ksepka, D.T. and C.A. Boyd. 2012. Quantifying historical trends in the completeness of the fossil record and the contributing factors: an example using Aves. Paleobiology 38: 826-839. [PDF]
27. Ksepka, D.T. and D.B. Thomas. 2012. Multiple Cenozoic invasions of Africa by penguins (Aves, Sphenisciformes). Proceedings of the Royal Society B Biological Sciences 279: 1027-1032. [PDF]
26. Ksepka, D.T. and J.A. Clarke. 2012. A new stem parrot from the Green River Formation and the complex evolution of the grasping foot in Pan-Psittaciformes. Journal of Vertebrate Paleontology 32: 395-406. [PDF]
25. *Cadena, E.A., D.T. Ksepka, C.A. Jaramilo and J.I. Bloch. 2012. New pelomedusoid turtles (Testudines, Panpleurodira) from the late Paleocene Cerrejón Formation of Colombia and implications for phylogeny and body size evolution. Journal of Systematic Palaeontology. 10: 313-331. [PDF]
24. Parham, J.F., P.C.J. Donoghue, C.J. Bell, T.D. Calway, J.J. Head, P.A. Holroyd, J.G. Inoue, R.B. Irmis, W.G. Joyce, D.T. Ksepka, J.S.L. Patané, N.D. Smith, J.E. Tarver, M. Van Tuinen, Z. Yang, K.D. Angielczyk, J. Greenwood, C.A. Hipsley, L. Jacobs, P.J. Makovicky, J. Müller, K.T. Smith, J.M. Theodor, R.C.M. Warnock, and M.J. Benton. 2012. Best practices for justifying fossil calibrations. Systematic Biology 61: 346-359. [PDF]
23. Ksepka, D.T., J.A Clarke, and L. Grande. 2011. Stem parrots (Aves, Halcyornithidae) from the Green River Formation and a combined phylogeny of Pan-Psittaciformes. Journal of Paleontology 85: 835-854.(Cover Article)
22. Nesbitt, S.J., D.T. Ksepka and J.A. Clarke. 2011. Podargiform affinities of the enigmatic Fluvioviridavis platyrhamphus and the early diversification of Strisores (‘‘Caprimulgiformes’’ + Apodiformes). PLoS One: e26350.
21. Thomas, D.B., D.T. Ksepka, and R.E. Fordyce. 2011. Penguin heat retention structures evolved in a Greenhouse Earth. Biology Letters 7: 461-464.
20. Ksepka, D.T. and J.A. Clarke. 2010. New fossil mousebird (Aves: Coliiformes) with feather preservation provides insight into the ecological diversity of an Eocene North American avifauna. Zoological Journal of the Linnean Society 160: 685-706.
19. Clarke, J.A., D.T. Ksepka, R. Salas-Gismondi, A.J. Altamirano, M.D. Shawkey, L. D'Alba, J. Vinther, T.J. DeVries and P. Baby. 2010. Fossil evidence for evolution of the shape and color of penguin feathers. Science 330: 954-957. (Cover Article)
18. Ksepka, D.T. and M.A. Norell. 2010. The illusory evidence for Asian Brachiosauridae: new specimens of Erketu ellisoni and a phylogenetic reappraisal of basal Titanosauriformes. American Museum Novitates 3700: 1-27.
17. Ksepka, D.T. and J.A. Clarke. 2010. The basal penguin (Aves: Sphenisciformes) Perudyptes devriesi and a phylogenetic evaluation of the penguin fossil record. Bulletin of the American Museum of Natural History 337: 1-77.
16. Ksepka, D.T. and J.A. Clarke. 2010. Primobucco mcgrewi (Aves: Coracii) from the Eocene Green River Formation: new anatomical data from the earliest definitive record of stem rollers. Journal of Vertebrate Paleontology 30: 215-225.
15. Hou, L.-H., P.-P. Li, D.T. Ksepka, K.-Q. Gao and M.A. Norell. 2010. Implications of flexible-shelled eggs in a Cretaceous choristoderan reptile. Proc. Royal Society B 277: 1235-1239.
14. Ksepka, D.T. and J.A. Clarke. 2009. Affinities of Palaeospiza bella and the phylogeny and biogeography of mousebirds (Coliiformes). Auk 126: 245-259. (Cover Article)
13. Ksepka, D.T. 2009. Broken gears in the avian molecular clock: new phylogenetic analyses support stem galliform status for Gallinuloides wyomingensis and rallid affinities for Amitabha urbsinterdictensis. Cladistics 25: 173-197.
12. Clarke, J.A., D.T. Ksepka, N.A. Smith and M.A. Norell. 2009. Combined phylogenetic analysis of a new North American fossil species confirms widespread Eocene distribution for stem rollers (Aves, Coracii). Zoological Journal of the Linnean Society 157: 586-611.
11. Ksepka, D.T., J.A. Clarke, T. DeVries and M. Urbina. 2008. Osteology of Icadyptes salasi, a giant penguin from the Eocene of Peru. Journal of Anatomy 213: 131-147.
10. Ksepka, D.T. and J. Cracraft. 2008. An avian tarsometatarsus from near the K-T boundary of New Zealand. Journal of Vertebrate Paleontology 28: 1224-1227.
9. Gao, K-Q. and D.T. Ksepka. 2008. Osteology and taxonomic revision of Hyphalosaurus (Diapsida: Choristodera) from the Lower Cretaceous of Liaoning, China. Journal of Anatomy 212: 747-760.
8. Clarke, J.A., D.T. Ksepka, M. Stucchi, M. Urbina, N. Giannini, S. Bertelli, Y. Naraez and C. Boyd. 2007. Paleogene equatorial penguins challenge the proposed relationship between biogeography, diversity, and Cenozoic climate change. PNAS 104: 11545-11550.
7. Gao, K-Q., D. Ksepka, H. Lianhai, D. Ye and H. Dongyu. 2007. Cranial morphology of an Early Cretaceous monjurosuchid (Reptilia: Diapsida) from Liaoning Province of China and evolution of the choristoderan palate. Historical Biology 19: 215-224.
6. Ksepka, D.T., S. Bertelli and N. Giannini. 2006. The phylogeny of the living and fossil Sphenisciformes (penguins). Cladistics 22: 412-441.
5. 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.
4. Bertelli, S., N.P. Giannini and D.T. Ksepka. 2006. Redescription and phylogenetic position of the early Miocene penguin Paraptenodytes antarcticus from Patagonia. Am. Mus. Novitates 3525: 1-36.
3. Ksepka, D.T. and M.A. Norell. 2006. Erketu ellisoni, a long-necked sauropod from Bor Guve (Dornogov Aimag, Mongolia). American Museum Novitates 3508: 1-16.
2. Ksepka, D.T., K. Gao and M.A. Norell. 2005. A new choristodere from the Cretaceous of Mongolia. American Museum Novitates 3468: 1-22.
1. Ksepka, D.T. and M.A. Norell. 2004. Ornithomimosaur cranial material from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates 3448: 1-4.
Popular Articles
Editorials, Reports, and Review Articles
6. Ksepka, D.T. 2022. Developmental Biology: A Dinosaur in a Quail Egg. Current Biology 32: R964-R967.
5. Ksepka, D.T. 2020. Feathered Dinosaurs. Current Biology 30: R1347-R1353.
4. Smith, N.A., A.M. Balanoff, and D.T. Ksepka. 2016. Symposium on ‘Evolving approaches for studying the anatomy of the avian brain’: Introduction Journal of Anatomy 229: 171-172.
3. Polly, P.D., D.T. Ksepka and J.F. Parham. 2015. Announcing the Fossil Calibration Series and Database. Palaeontologia Electronica 18: 1-5.
2. Ksepka, D.T. 2014. Evolution: A rapid flight towards birds. Current Biology 24: R1052-R1055. (Dispatch Article)
1. Ksepka, D.T., M.J. Benton, M.T. Carrano, M.A. Gandolfo, J.J. Head, E. J. Hermsen, W.G. Joyce, K.S. Lamm, J.S.L. Patané, M.J. Phillips, P.D. Polly, M. van Tuinen, J.L. Ware, R.C.M. Warnock, and J.F. Parham. 2011. Synthesizing and databasing fossil calibrations: divergence dating and beyond. Biology Letters 7: 801-803. (Meeting Report)
Invited Book Chapters
2. Ksepka, D.T. 2022. Evolution of Birds. In C.G. Scanes and S. Dridi (eds), Sturkie’s Avian Physiology (7th Edition): 83-107. Academic Press: London.
1. Ksepka, D.T. and T. Ando. 2011. Penguins past, present and future: trends in the evolution of the Sphenisciformes. In G. Dyke and G. Kaiser (eds), Living Dinosaurs: The Evolutionary History of Modern Birds: 155-186. John Wiley and Sons: Chichester.