Cited by 38Evolution and Paleontology StudiesPaleontology and Evolutionary BiologyAvian ecology and behavior

Multiple cenozoic invasions of Africa by penguins (Aves, Sphenisciformes)

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Ksepka, Daniel T., Thomas, Daniel B.·2012·Proceedings of the Royal Society B: Biological Sciences

Abstract

Africa hosts a single breeding species of penguin today, yet the fossil record indicates that a diverse array of now-extinct taxa once inhabited southern African coastlines. Here, we show that the African penguin fauna had a complex history involving multiple dispersals and extinctions. Phylogenetic analyses and biogeographic reconstructions incorporating new fossil material indicate that, contrary to previous hypotheses, the four Early Pliocene African penguin species do not represent an endemic radiation or direct ancestors of the living Spheniscus demersus (blackfooted penguin). A minimum of three dispersals to Africa, probably assisted by the eastward-flowing Antarctic Circumpolar and South Atlantic currents, occurred during the Late Cenozoic. As regional sea-level fall eliminated islands and reduced offshore breeding areas during the Pliocene, all but one penguin lineage ended in extinction, resulting in today's depleted fauna.

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  • The genus Spheniscus comprises two more extant species besides Humboldt and African penguins: the Magellanic penguin and the Galapagos penguin.

    “The genus Spheniscus comprises 2 more extant species, other than Humboldt and African penguins: the Magellanic penguin (S. magellanicus) and the Galapagos penguin (S. mendiculus) (Ksepka and Thomas, 2012).”

    From Molecular Genetics Unveiled Unknown Family Relationships and Hybrids in an Ex-Situ Colony of African Penguins (Spheniscus demersus) (2018) · cites this paper as Ksepka and Thomas, 2012

    The Results (page unavailable) state that Spheniscus has four extant species and identify S. magellanicus as the sister species of the African penguin, S. demersus. Our copy does not identify the other two extant species as the Humboldt and Galapagos penguins, so it does not fully support the cited species list. (LLM, conf=0.84)

    Experimental — accuracy not yet measured
  • Based on molecular and morphological data, Galapagos and Humboldt penguins are considered sister species, as are Magellanic and African penguins.

    “Based on molecular and morphological dataset, Galapagos and Humboldt penguins are considered sister species, as are Magellanic and African penguins (Ksepka and Thomas 2012).”

    From Molecular Genetics Unveiled Unknown Family Relationships and Hybrids in an Ex-Situ Colony of African Penguins (Spheniscus demersus) (2018) · cites this paper as Ksepka and Thomas 2012

    The Methods describe an analysis combining morphological and molecular data, and the Results explicitly call African penguins (S. demersus) and Magellanic penguins (S. magellanicus) sister species. The passages in our copy do not establish the claimed sister relationship between Galápagos and Humboldt penguins. (LLM, conf=0.86)

    Experimental — accuracy not yet measured

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  1. [1]Bost C. A., et al. 1997. Foraging habitat and food intake of satellite-tracked king penguins during the austral summer at Crozet Archipelago. Mar. Ecol. Prog. Ser. 150, 21–33 10.3354/meps150021 (doi:10.3354/meps150021) [DOI] [Google Scholar] · DOI
  2. [2]Kooyman G. L. 2002. Evolutionary and ecological aspects of some Antarctic and sub-Antarctic penguin distributions. Oecologia 130, 485–495 10.1007/s00442-001-0836-x (doi:10.1007/s00442-001-0836-x) [DOI] [PubMed] [Google Scholar] · DOI
  3. [3]Clarke J. A., Ksepka D. T., Stucchi M., Urbina M., Giannini N., Bertelli S., Narváez Y., Boyd C. A. 2007. Paleogene equatorial penguins challenge the proposed relationship between biogeography, diversity, and Cenozoic climate change. Proc. Natl Acad. Sci. USA 104, 11 545–11 550 10.1073/pnas.0611099104 (doi:10.1073/pnas.0611099104) [DOI] [PMC free article] [PubMed] [Google Scholar] · DOI
  4. [4]Fordyce R. E., Jones C. M. 1990. Penguin history and new fossil material from New Zealand. In Penguin biology (eds Davis L. S., Darby J. T.), pp. 419–446 San Diego, CA: Academic Press [Google Scholar]
  5. [5]Jadwiszczak J. 2010. Penguin response to the Eocene climate and ecosystem change in the northern Antarctic Peninsula region. Polar Sci. 4, 229–235 10.1016/j.polar.2010.03.001 (doi:10.1016/j.polar.2010.03.001) [DOI] [Google Scholar] · DOI
  6. [6]Ksepka D. T., Bertelli S., Giannini N. P. 2006. The phylogeny of the living and fossil Sphenisciformes (penguins). Cladistics 22, 412–441 10.1111/j.1096-0031.2006.00116.x (doi:10.1111/j.1096-0031.2006.00116.x) [DOI] [Google Scholar] · DOI
  7. [7]Bertelli S., Giannini N. P. 2005. A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology and mitochondrial sequences. Cladistics 21, 209–239 10.1111/j.1096-0031.2005.00065.x (doi:10.1111/j.1096-0031.2005.00065.x) [DOI] [Google Scholar] · DOI
  8. [8]Olson S. L. 1985. An Early Pliocene marine avifauna from Duinefontein, Cape Province, South Africa. Ann. S. Afr. Mus. 95, 147–164 [Google Scholar]
  9. [9]Baker A. J., Pereira S. L., Haddrath O. P., Edge A. 2006. Multiple gene evidence for expansion of extant penguins out of Antarctica due to global cooling. Proc. R. Soc. B 217, 11–17 10.1098/rspb.2005.3260 (doi:10.1098/rspb.2005.3260) [DOI] [PMC free article] [PubMed] [Google Scholar] · DOI
  10. [10]Simpson G. G. 1946. Fossil penguins. Bull. Am. Mus. Nat. Hist. 87, 7–99 [Google Scholar]
  11. [11]Jadwiszczak J. 2009. Penguins past: the current state of knowledge. Pol. Polar Res. 30, 3–28 [Google Scholar]
  12. [12]Ksepka D. T., Clarke J. A. 2010. The basal penguin (Aves: Sphenisciformes) Perudyptes devriesi and a phylogenetic evaluation of the penguin fossil record. Bull. Am. Mus. Nat. Hist. 337, 1–77 10.1206/653.1 (doi:10.1206/653.1) [DOI] [Google Scholar] · DOI
  13. [13]Slack K. E., Jones C. M., Ando T., Harrison G. L., Fordyce R. E., Arnason U., Penny D. 2006. Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution. Mol. Biol. Evol. 23, 1144–1155 10.1093/molbev/msj124 (doi:10.1093/molbev/msj124) [DOI] [PubMed] [Google Scholar] · DOI · Paper page
  14. [14]Simpson G. G. 1971. Fossil penguin from the Late Cenozoic of South Africa. Science 171, 1144–1145 10.1126/science.171.3976.1144 (doi:10.1126/science.171.3976.1144) [DOI] [PubMed] [Google Scholar] · DOI
  15. [15]Simpson G. G. 1973. Tertiary penguins (Sphenisciformes, Spheniscidae) from Ysterplaats, Cape Town, South Africa. S. Afr. J. Sci. 69, 342–344 [Google Scholar]
  16. [16]Simpson G. G. 1979. Tertiary penguins from the Duinefontein site, Cape Province, South Africa. Ann. S. Afr. Mus. 79, 1–17 [Google Scholar]
  17. [17]Simpson G. G. 1979. A new genus of Late Tertiary penguin from Langebaanweg, South Africa. Ann. S. Afr. Mus. 78, 1–9 [Google Scholar]
  18. [18]Hendey Q. B. 1973. Fossil occurrences at Langebaanweg, Cape Province. Nature 244, 13–14 10.1038/244013a0 (doi:10.1038/244013a0) [DOI] [Google Scholar] · DOI
  19. [19]Göhlich U. B. 2007. The oldest fossil record of the extant penguin genus Spheniscus–a new species from the Miocene of Peru. Acta Palaeontol. Pol. 52, 285–298 [Google Scholar]
  20. [20]Simpson G. G. 1975. Notes on variation in penguins and on fossil penguins from the Pliocene of Langebaanweg, Cape Province, South Africa. Ann. S. Afr. Mus. 69, 59–72 [Google Scholar]
  21. [21]Olson S. L. 1983. Fossil seabirds and changing marine environments in the Late Tertiary of South Africa. S. Afr. J. Sci. 79, 399–402 [Google Scholar]
  22. [23]Roberts D. L. 2006. Varswater Formation including the Langeeheid Clayey Sand, Konings Vlei Gravel, Langeberg Quartz Sand and Muishond Fontein Phosphatic Sand Members. In Catalogue of South African lithostratigraphic units (ed. Johnson M. R.), pp. 97–931 Pretoria, South Africa: Council for Geoscience [Google Scholar]
  23. [24]Roberts D. L., et al. 2011. Regional and global context of the Late Cenozoic Langebaanweg (LBW) palaeontological site: west coast of South Africa. Earth Sci. Rev. 106, 191–214 10.1016/j.earscirev.2011.02.002 (doi:10.1016/j.earscirev.2011.02.002) [DOI] [Google Scholar] · DOI
  24. [25]Swofford D. L. 2003. PAUP*. Phylogenetic analysis using parsimony (*and other methods). Sunderland, MA: Sinauer Associates [Google Scholar]
  25. [26]Maddison W. P., Maddison D. R. 2005. MacClade. Version 4.08. Sunderland, MA: Sinauer Associates [Google Scholar]
  26. [27]Yu Y., Harris A. J., He X. J. 2011. RASP (reconstruct ancestral state in phylogenies). See http://mnh.scu.edu.cn/soft/blog/RASP [DOI] [PMC free article] [PubMed]
  27. [28]Lamm K. S., Redelings B. D. 2009. Reconstructing ancestral ranges in historical biogeography: properties and prospects. J. System. Evol. 47, 369–382 10.1111/j.1759-6831.2009.00042.x (doi:10.1111/j.1759-6831.2009.00042.x) [DOI] [Google Scholar] · DOI
  28. [29]Gallagher K., Brown R., Osmaston M., Ebinger C., Bishop P. 1999. Denudation and uplift at passive margins: the record on the Atlantic margin of southern Africa. Phil Trans. R. Soc. Lond. A 357, 835–859 10.1098/rsta.1999.0354 (doi:10.1098/rsta.1999.0354) [DOI] [Google Scholar] · DOI
  29. [30]Roberts D. L., Botha G. A., Maud R. R., Pether J. 2007. Coastal Cenozoic deposits. In The geology of South Africa (eds Johnson M. R., Anhaeusser C. R., Thomas R. J.), pp. 605–628 Pretoria, South Africa: Council for Geoscience [Google Scholar]
  30. [31]Cruz-Uribe K., Klein R. G., Avery G., Avery M., Halkett D., Hart T., Milo R. G., Sampson C. G., Volman T. P. 2003. Excavation of buried Late Acheulean (Mid-Quaternary) land surfaces at Duinefontein 2, Western Cape Province, South Africa. J. Archaeol. Sci. 30, 229–575 10.1016/S0305-4403(02)00202-9 (doi:10.1016/S0305-4403(02)00202-9) [DOI] [Google Scholar] · DOI
  31. [32]Klein R. G., Avery G., Cruz-Uribe K., Halkett D., Hart T., Milo R. G., Volman T. P. 1999. Duinefontein 2: an Acheulean site in the Western Cape Province of South Africa. J. Hum. Evol. 37, 153–190 10.1006/jhev.1999.0307 (doi:10.1006/jhev.1999.0307) [DOI] [PubMed] [Google Scholar] · DOI
  32. [33]Lawver L. A., Gahagan L. M. 2003. Evolution of Cenozoic seaways in the circum-Antarctic region. Palaeogeogr. Palaeoclimatol. 198, 11–37 10.1016/S0031-0182(03)00392-4 (doi:10.1016/S0031-0182(03)00392-4) [DOI] [Google Scholar] · DOI
  33. [34]Diester-Haass L., Meyers P. A., Vidal L. 2002. The Late Miocene onset of high productivity in the Benguela Current upwelling system as part of a global pattern. Mar. Geol. 180, 87–103 10.1016/S0025-3227(01)00207-9 (doi:10.1016/S0025-3227(01)00207-9) [DOI] [Google Scholar] · DOI
  34. [35]Martínez S., del Río C. J. 2002. Late Miocene molluscs from the southwestern Atlantic Ocean (Argentina and Uruguay): a palaeobiogeographic analysis. Palaeogeogr. Palaeoclimatol. 188, 167–187 10.1016/S0031-0182(02)00551-5 (doi:10.1016/S0031-0182(02)00551-5) [DOI] [Google Scholar] · DOI
  35. [36]Williams T. D. 1995. The penguins. Oxford, UK: Oxford University Press [Google Scholar]
  36. [37]Crawford R. J. M., Underhill L. G., Upfold L., Dyer B. M. 2007. An altered carrying capacity of the Benguela upwelling ecosystem for African penguins (Spheniscus demersus). ICES J. Mar. Sci. 64, 570–576 10.1093/icesjms/fsm009 (doi:10.1093/icesjms/fsm009) [DOI] [Google Scholar] · DOI
  37. [38]Stramma L., England M. 1999. On the water masses and mean circulation of the South Atlantic Ocean. J. Geophys. Res. 104, 20 863–20 883 10.1029/1999JC900139 (doi:10.1029/1999JC900139) [DOI] [Google Scholar] · DOI
  38. [39]Robert C., Diester-Haass L., Paturel J. 2005. Clay mineral assemblages, siliciclastic input and paleoproductivity at ODP Site 1085 off Southwest Africa: a late Miocene–early Pliocene history of Orange river discharges and Benguela current activity, and their relation to global sea level change. Mar. Geol. 216, 221–238 10.1016/j.margeo.2005.02.024 (doi:10.1016/j.margeo.2005.02.024) [DOI] [Google Scholar] · DOI
  39. [40]Miller K. G., et al. 2005. The Phanerozoic record of global sea-level change. Science 310, 1293–1298 10.1126/science.1116412 (doi:10.1126/science.1116412) [DOI] [PubMed] [Google Scholar] · DOI · Paper page
  40. [41]Zachos J., Pagani M., Sloan L., Thomas E., Billups K. 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292, 686. 10.1126/science.1059412 (doi:10.1126/science.1059412) [DOI] [PubMed] [Google Scholar] · DOI · Paper page

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Authors
Ksepka, D. T., & Thomas, D. B.
Affiliations
  • North Carolina Museum of Natural Sciences (US)
  • University of Cape Town (ZA)
Corresponding author
Ksepka, Daniel T.
Venue
Proceedings of the Royal Society B: Biological Sciences
Published
2011-09-07
Volume
279
Issue
1730
Pages
1027-1032
ISSN
  • 0080-4649 (ISSN-L)
  • 0962-8452
  • 1471-2954
OpenAlex
W2146233142
Cited by
38
References
0
Citations by year
2024: 3 · 2023: 3 · 2022: 4 · 2020: 1 · 2019: 4 · 2018: 2 · 2017: 1 · 2016: 2 · 2015: 1 · 2014: 6 · 2013: 5 · 2012: 6
FWCI
3.78
Type
journal article
Language
en
Open access
Locations
  • Proceedings of the Royal Society B Biological Sciences (publishedVersion)
  • Europe PMC (PubMed Central) (publishedVersion)
  • PubMed Central (publishedVersion)
  • PubMed (publishedVersion)
Topics
  • Physical Sciences › Earth and Planetary Sciences › Paleontology › Evolution and Paleontology Studies
  • Physical Sciences › Earth and Planetary Sciences › Paleontology › Paleontology and Evolutionary Biology
  • Physical Sciences › Environmental Science › Ecology › Avian ecology and behavior
Keywords
penguins, Sphenisciformes, African penguin, Spheniscus demersus, fossil penguins, biogeography, dispersal, extinction, Cenozoic, Pliocene, sea-level change, ocean currents, phylogenetics
MeSH terms
Africa, Animals, Female, Fossils, Male, Phylogeny, Population Dynamics, Water Movements, Animal Migration, Spheniscidae, Extinction, Biological, Phylogeography

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