This order is restricted to the Spheniscidae (penguins).

Species-level timetree of extant Spheniscidae based on Cole et al. (2019, 2022), Fernandes et al. (2019), Pan et al. (2019), and Vianna et al. (2020), with the distribution of each species being indicated by the colour-code used throughout this website (see Distribution code). Divergence-time estimates are based on Vianna et al. (2020). Their divergence-time estimates are close to those of Fernandes (2019), but significantly younger than those of Cole et al. (2019), which are based on mitogenomes.  


Baker AJ, Pereira SL, Haddrath OP, and Edge KA (2006), Multiple gene evidence for expansion of extant penguins out of Antarctica due to global cooling, Proc. Biol. Sci. 273(1582), 11-17. (pdf)

Banks JC, Mitchell AD, Waas JR, and Paterson AM (2002), An unexpected pattern of molecular divergence within the blue Penguin (Eudyptula minor) complex, Notornis 49(1), 29-38. (pdf)

Banks J, van Buren A, Cherel Y, and Whitfield JB (2006), Genetic evidence for three species of penguins, Eudyptes chrysocome, Polar Biol. 30(1), 61-67. (abstract)

Bertelli S, and Giannini NP (2005), A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology and mitochondrial sequences, Cladistics 21(3), 209-239. (pdf)


Cole TL, Rawlence NJ, Dussex N, Ellenberg U, Houston DM, Mattern T, Miskelly CM, Morrison KW, Scofield RP, Tennyson AJD, Thompson DR, Wood JR, and Waters JM,  (2019), Ancient DNA of crested penguins: testing for temporal genetic shifts in the world´s most diverse penguin clade. Mol. Phylogenet. Evol. 131, 72-79. (abstract)



Cole TL, Ksepka DT, Mitchell KJ, Tennyson AJD, Thomas DB, Pan H, Zhang G, Rawlence NJ, Wood JR, Boer P, Bouzat JL, CooperA, Fiddaman SR, Hart T, Miller G, Ryan PG, Shepherd LD, Wilmshurst JM, and Waters JM (2019), Mitogenomes uncover extinct penguin taxa and reveal island formation as a key driver of speciation. Mol. Biol. Evol. 36, 784-797. (pdf)


Cole TL, Zhou C, Fang M, Pan H, Ksepka DT, Fiddaman SR, Emerling CA, Thomas DB, Bi X, Fang Q, Ellegaard MR,Feng S, Smith AL, Heath TA, Tennyson AJD, Borboroglu PG, Wood JR, Hadden PW, Grosser S, Bost CA, Cherel Y, Mattern T, Hart T, Sinding MHS, Shepherd LD, Phillips RA, Quillfeldt P, Masello JF, Bouzat JL, Ryan PG, Thompson DR, Ellenberg U, Dann P, Miller G, Boersma PD, Zhao R, Gilbert MTP, Yang H, Zhang DX, and Zhang G (2022), Genomic insights into the secondary aquatic transition of penguins, Nature Commun. 13, e:3912. (pdf)

Frugone  MJ, Lowther A, Noll D, Ramos B, Pistorius P, Dantas GPM, Petry MV, Bonadonna F, Steinfurth A, Polanowski A, and Rey AR (2018), Contrasting phylogeographic pattern among Eudyptes penguins around the Southern Ocean, Sci. Rep. 8(1), e17481. (pdf)

Gavryushkina A, Heath TA, Ksepka DT, Stadler T, Welch D, and Drummond AJ (2013), Bayesian total-evidence dating reveals the recent crown radiation of penguins, Syst. Biol. 66(16), 57-73. (pdf)


Ksepka DT, Bertelli S, and Giannini NP (2006), The phylogeny of the living and fossil Sphenisciformes (penguins), Cladistics 22(5), 412-441. (pdf)


Pan H, Cole TL, Bi X, Fang M, Zhou C, Yang Z, Ksepka DT, Hart T, Bouzat JL, Argilla LS, Bertelsen MF, Boersman PD, Bost CA, Cherel Y, Dann P, Fiddaman SR, Howard P, Labuschagne K, Mattern T, Miller G, Parker P, Phillips RA, Quillfeldt P, Ryan PG, Taylor H, Thompson DR, Young MJ, Ellegaard MR, Gilbert MTP, Sinding MHS, Pacheco G, Shepherd LD, Tennyson AJD, Grosser S, Kay E, Nupen LJ, Ellenberg U, Houston DM, Hart A, Reeve, Johnson K, Masello JF, Stracke T, McKinlay B, García Borboroglu P, Zhang DX, and Zhang G (2019), High-coverage genomes to elucidate the evolution of penguins. GigaScience 8, 1-17.



Pelegrín JS, and Hospitaleche CA (2022), Evolutionary and biogeographical history of penguins (Sphenisciformes): review of the dispersal patterns and adaptations in a geologic and paleoecological context, Diversity 14, e:255. (free pdf)

Ramos B, González-Acuña D, Loyola DE, Johnson WE, Parker PG, Massaro M, Dantas GP, Miranda MD, and Vianna JA (2018), Landscape genomics: natural selection drives the evolution of mitogenome in penguins, BMC Genomics 19(1), e53. (pdf)


Subramanian S, Beans-Picón G, Swaminathan SK, Millar CD, and Lambert DM (2013), Evidence for a recent origin of penguins, Biol. Lett. 9, e20130748. (pdf)


Thomas DB, Tennyson AJD, Scofield RP, Heath TA, Pett W, and Ksepka DT (2020), Ancient crested penguin constrains timing of recruitment into seabird hotspot, Proc. R. Soc. B 287, e:20201497. (pdf)

Vianna JA, Fernandes FAN, Frugone MJ, Figueiró HV, Pertierra LR, Noll D, Bi K, Wang-Claypool CY, Lowther A, Parker P, Le Bohec C, Bonadonna F, Wienecke B, Pistorius P, Steinfurth A, Burridge CP, Dantas GPM, Poulin E, Simison WB, Henderson J, Eizirik E, Nery MF, and Bowie RCK (2020), Genome-wide analyses reveal drivers of penguin diversification, PNAS 117(36), 22303-10. (pdf)