Journal of Mammalian Evolution

, Volume 22, Issue 4, pp 535–560 | Cite as

The Earliest Gulp-Feeding Mysticete (Cetacea: Mysticeti) from the Oligocene of New Zealand

  • Cheng-Hsiu TsaiEmail author
  • R. Ewan Fordyce
Original Paper


Horopeta umarere is a new genus and species of extinct baleen whales from the Kokoamu Greensand (early Chattian, Oligocene, in the range 25–27 Ma), Hakataramea Valley, New Zealand. The geological age makes Horopeta umarere one of the earliest named baleen whales. Phylogenetic analysis indicates that Horopeta umarere may be the earliest crown Mysticeti (the sister taxon to Cetotheriidae), or the sister species to the crown Mysticeti; it is clearly not a species of Eomysticetidae. Estimated skull and body length of Horopeta umarere are 1.5-1.6 m and 6.5-7.5 m, respectively. Horopeta umarere shows some features that are linked to gulp feeding as seen in living humpback and rorquals: laterally bowed and robust mandible, D-shaped to teardrop-shaped mandible in cross-section, and posterolaterally deflected triangular coronoid process of the mandible. The sternum of Horopeta umarere is elongate, rod-shaped, and dorsoventrally stout with bilateral anterior and posterior rugose protrusions, indicating the presence of at least two pairs of ribs or costal cartilages. The structure of the skull and mandible are consistent with the use of gulp feeding, but the sternal morphology and rib attachments suggest an early evolutionary stage in gulp feeding employment, where more complex rib attachment may restrict the volume of water and food taken in one gulp compared to living humpback and rorquals. Thus, the morphology of Horopeta umarere has implications for the emergence of gulp feeding in baleen whale evolution as well as the emergence of the crown Mysticeti.


Baleen whale Rorqual Skull Fossil Evolution 



We thank Foveran Station and Haughs’ quarry operators for access and permission to collect OU21982; the late A. Grebneff and C. M. Jones for helping REF collect OU21982; A. Grebneff for initial preparation; S. E. White for assisting CHT in finalizing the preparation; J. Mead, C. Potter, J. Ososky, N. Pyenson, D. Bohaska (National Museum of Natural History, Smithsonian Institution, USA), J. Dines, D. Janiger (Natural History Museum of Los Angeles County, USA), A. Stewart, T. Schultz (National Museum of New Zealand, NZ), E. Fitzgerald, K. Roberts (Museum Victoria, Australia), T. Yamada, N. Kohno, Y. Tajima (National Museum of Nature and Science, Japan), C. Kemper, D. Stemmer, N. Pledge, M.-A. Binnie (South Australian Museum, Australia), E. Westwig (American Museum of Natural History, USA), P. Holroyd (University of California Museum of Paleontology, USA), A. Sanders (The Charleston Museum, USA) for access to collections and allowing photography during CHT’s and/or REF’s visits. We thank O. Lambert and J. Geisler for constructive reviews and comments; the editor, J. R. Wible, for helpful advice; G. Aguirre-Fernandez, F. G. Marx, R. Boessenecker, Y. Tanaka, and J. Corrie for discussion; S. E. White for discussion of Maori. CHT thanks J. Mead (Washington DC), R. Boessenecker (California), E. Fitzgerald, K. Roberts (Melbourne), F. Marx, I. Aguirre (Tsukuba), and A. Fukumoto (Yokohama) for accommodations during various research trips. This study is part of CHT’s PhD thesis and funded by a University of Otago Doctoral Scholarship. Grants 3542–87 and 3657–87 from the National Geographic Society to REF supported field and laboratory work on OU21982.

Ethical Statement

This work has not been submitted to more than one journal for consideration simultaneously, and this work has not been published previously elsewhere.


  1. Arvy L, Pilleri G (1977) The sternum in Cetacea. Investigations on Cetacea 8:123–148Google Scholar
  2. Benham WB (1937) Fossil Cetacea of New Zealand II.-On Lophocephalus, a new genus of zeuglodont Cetacea. Trans Roy Soc of New Zealand 67:1–7Google Scholar
  3. Bisconti M (2014) Anatomy of a new cetotheriid genus and species from the Miocene of Herentals, Belgium, and the phylogenetic and palaeobiogeographical relationships of Cetotheriidae s.s. (Mammalia, Cetacea, Mysticeti). J Syst Palaeontol 1–19. doi: 10.1080/14772019.2014.890136
  4. Bouetel V, Muizon C de (2006) The anatomy and relationships of Piscobalaena nana (Cetacea, Mysticeti), a Cetotheriidae s.s. from the early Pliocene of Peru. Geodiversitas 28:319–395Google Scholar
  5. Brodie PF (1993) Noise generated by the jaw actions of feeding fin whales. Can J Zool 71:2546–2550CrossRefGoogle Scholar
  6. de Buffrenil V, de Ricqles A, Ray CE, Dorning DP (1990) Bone histology of the ribs of the archaeocetes (Mammalia: Cetacea). J Vertebr Paleontol 10:455–466CrossRefGoogle Scholar
  7. Deméré TA, McGowen MR, Berta A, Gatesy J (2008) Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales. Syst Biol 57:15–37CrossRefPubMedGoogle Scholar
  8. Ekdale EG, Berta A, Deméré TA (2011) The comparative osteology of the petrotympanic complex (ear region) of extant baleen whales (Cetacea: Mysticeti). PLoS ONE 6:e21311. doi: 10.1371/journal.pone.0021311 PubMedCentralCrossRefPubMedGoogle Scholar
  9. El Adli JJ, Deméré TA, Boessenecker RW (2014) Herpetocetus morrowi (Cetacea: Mysticeti), a new species of diminutive baleen whale from the upper Pliocene (Piacenzian) of California, USA, with observations on the evolution and relationships of the Cetotheriidae. Zool J Linn Soc 170:400–466CrossRefGoogle Scholar
  10. Fitzgerald EMG (2006) A bizarre new toothed mysticete (Cetacea) from Australia and the early evolution of baleen whales. Proc Roy Soc B: Biol Sci 273:2955–2963CrossRefGoogle Scholar
  11. Fitzgerald EMG (2010) The morphology and systematics of Mammalodon colliveri (Cetacea: Mysticeti), a toothed mysticete from the Oligocene of Australia. Zool J Linn Soc 158:367–476CrossRefGoogle Scholar
  12. Fordyce RE (2002) Oligocene origins of skim-feeding right whales: a small archaic balaenid from New Zealand. In: Abstract of Papers of 62nd Annual Meeting Society of Vertebrate Paleontology, Oklahoma, USA, p 54Google Scholar
  13. Fordyce RE, Marx FG (2013) The pygmy right whale Caperea marginata: the last of the cetotheres. Proc Roy Soc B: Biol Sciences 280:20122645. doi:  10.1098/rspb.2012.2645
  14. Fraser FC, Purves PE (1960) Hearing in cetaceans. Evolution of the accessory air sacs and the structure and function of the outer and middle ear in recent cetaceans. Bull Brit Mus (Nat Hist) Zool 7:1–140Google Scholar
  15. Geisler JH, Sanders AE (2003) Morphological evidence for the phylogeny of Cetacea. J Mammal Evol 10:23–129CrossRefGoogle Scholar
  16. Goldbogen JA, Calambokidis J, Friedlaender AS, Francis J, DeRuiter SL, Stimpert AK, Falcone E, Southall BL (2013) Underwater acrobatics by the world's largest predator: 360° rolling manoeuvres by lunge-feeding blue whales. Biol Lett 9:20120986. doi:  10.1098/rsbl.2012.0986 PubMedCentralCrossRefPubMedGoogle Scholar
  17. Goloboff PA, Farris JS, Nixon KC (2008) TNT, a free program for phylogenetic analysis. Cladistics 24:774–786CrossRefGoogle Scholar
  18. Hoelzel AR, Dorsey EM, Stern J (1989) The foraging specializations of individual minke whales. Anim Behav 38:786–794CrossRefGoogle Scholar
  19. Ingebrigtsen A (1929) Whales caught in the North Atlantic and other areas. Rapports et Proces-Verbaux des Reunions - Conseil Permanent International Pour l'Exploration de la Mer 56:1–26Google Scholar
  20. Kasuya T (1973) Systematic consideration of recent toothed whales based on the morphology of tympano-periotic bone. Scientific Reports of the Whales Research Institute Tokyo 25:1–103Google Scholar
  21. Kellogg R (1923) Description of two squalodonts recently discovered in the Calvert Cliffs, Maryland; and notes on the shark-toothed cetaceans. Proc US Natl Mus 62:1–69CrossRefGoogle Scholar
  22. Kellogg R (1965) A new whalebone whale from the Miocene Calvert formation. US Natl Mus Bull 247:1–45Google Scholar
  23. Kimura T (2002) Feeding strategy of an early Miocene cetothere from the Toyama and Akeyo Formations, central Japan. Paleontol Res 6:179–189Google Scholar
  24. Kimura T, Hasegawa Y (2010) A new baleen whale (Mysticeti: Cetotheriidae) from the earliest late Miocene of Japan and a reconsideration of the phylogeny of cetotheres. J Vertebr Paleontol 30:577–591CrossRefGoogle Scholar
  25. Lambertsen R, Ulrich N, Straley J (1995) Frontomandibular stay of Balaenopteridae: a mechanism for momentum recapture during feeding. J Mammal 76:877–899CrossRefGoogle Scholar
  26. Luo Z, Gingerich PD (1999) Terrestrial Mesonychia to aquatic Cetacea: transformation of the basicranium and evolution of hearing in whales. Univ Michigan Pap Paleontol 31:1–98Google Scholar
  27. Marx FG (2011) The more the merrier? A large cladistic analysis of mysticetes, and comments on the transition from teeth to baleen. J Mammal Evol 18:77–100CrossRefGoogle Scholar
  28. Mead JG, Fordyce RE (2009) The therian skull: a lexicon with emphasis on the odontocetes. Smithsonian Contrib Zool 627:1–248CrossRefGoogle Scholar
  29. Mitchell ED (1989) A new cetacean from the late Eocene La Meseta Formation Seymour Island, Antarctic Peninsula. Can J Fish Aquatic Sci 46:2219–2235CrossRefGoogle Scholar
  30. Moran MM, Bajpai S, George JC, Suydam R, Usip S, Thewissen JGM (2014) Intervertebral and epiphyseal fusion in the postnatal ontogeny of cetaceans and terrestrial mammals. J Mammal Evol doi: 10.1007/s10914-014-9256-7 Google Scholar
  31. Nemoto T (1957) Foods of baleen whales in the northern Pacific. Scientific Reports of the Whales Research Institute Tokyo 12:33–89Google Scholar
  32. Noble ER, Smoker WR (1996) The forgotten condyle: the appearance, morphology, and classification of occipital condyle fractures. Am J Neuroradiol 17:507–513PubMedGoogle Scholar
  33. Oishi M, Hasegawa Y (1995) Diversity of Pliocene mysticetes from eastern Japan. The Island Arc 3:436–452CrossRefGoogle Scholar
  34. Potvin J, Goldbogen JA, Shadwick RE (2012) Metabolic expenditures of lunge feeding rorquals across scale: Implications for the evolution of filter feeding and the limits to maximum body size. PLoS ONE 7:e44854. doi: 10.1371/journal.pone.0044854 PubMedCentralCrossRefPubMedGoogle Scholar
  35. Pyenson ND, Goldbogen JA, Shadwick RE (2013) Mandible allometry in extant and fossil Balaenopteridae (Cetacea: Mammalia): the largest vertebrate skeletal element and its role in rorqual lunge feeding. Biol J Linn Soc 108:586–599CrossRefGoogle Scholar
  36. Pyenson ND, Goldbogen JA, Vogl AW, Szathmary G, Drake RL, Shadwick RE (2012) Discovery of a sensory organ that coordinates lunge feeding in rorqual whales. Nature 485:498–501CrossRefPubMedGoogle Scholar
  37. Pyenson ND, Sponberg SN (2011) Reconstructing body size in extinct crown Cetacea (Neoceti) using allometry, phylogenetic methods and tests from the fossil record. J Mammal Evol 18:269–288CrossRefGoogle Scholar
  38. Raine J, Beu A, Boyes A, Campbell H, Cooper R, Crundwell M (2012) New Zealand Geological Timescale v. 2012/1 Lower Hutt, GNS Science 1 (downloaded this New Zealand Geological Timescale v.2012/1 at:
  39. Sanders AE, Geisler JH (2015) A new basal odontocete from the upper Rupelian of South Carolina, U.S.A, with contributions to the systematics of Xenorophus and Mirocetus (Mammalia, Cetacea). J Vertebr Paleontol. doi: 10.1080/02724634.2014.890107 Google Scholar
  40. Sasaki T, Nikaido M, Hamilton H, Goto M, Kato H, Kanda N, Pastene LA, Cao Y, Fordyce RE, Hasegawa M (2005) Mitochondrial phylogenetics and evolution of mysticete whales. Syst Biol 54:77–90CrossRefPubMedGoogle Scholar
  41. Schulte HVW (1916) Anatomy of a foetus of Balaenoptera borealis. Mem Am Mus Nat Hist 1:394–502Google Scholar
  42. Steeman ME (2007) Cladistic analysis and a revised classification of fossil and recent mysticetes. Zool J Linn Soc 150:875–894CrossRefGoogle Scholar
  43. Steeman ME, Hebsgaard MB, Fordyce RE, Ho SYW, Rabosky DL, Nielsen R, Rahbek C, Glenner H, Sørensen MV, Willerslev E (2009) Radiation of extant cetaceans driven by restructuring of the oceans. Syst Biol 58:573–585PubMedCentralCrossRefPubMedGoogle Scholar
  44. True FW (1904) The whalebone whales of the western North Atlantic compared with those occurring in European waters with some observations on the species of the North Pacific. Smithsonian Contrib Knowledge 33:1–332Google Scholar
  45. Tsai CH, Fordyce RE (2014a) Juvenile morphology in baleen whale phylogeny. Naturwissenschaften 101:765–769CrossRefPubMedGoogle Scholar
  46. Tsai CH, Fordyce RE (2014b) Disparate heterochronic processes in baleen whale evolution. Evol Biol 41:299–307CrossRefGoogle Scholar
  47. Tsai CH, Fordyce RE (2015) Ancestor–descendant relationships in evolution: origin of the extant pygmy right whale, Caperea marginata. Biol Lett 11 doi: 10.1098/rsbl.2014.0875
  48. Tsai CH, Fordyce RE, Chang C-H, Lin L-K (2014) Quaternary fossil gray whales from Taiwan. Paleontol Res 18:82–93CrossRefGoogle Scholar
  49. Walsh BM, Berta A (2011) Occipital ossification of balaenopteroid mysticetes. Anat Record 294:391–398CrossRefGoogle Scholar
  50. Werth A (2000) Feeding in marine mammals. In: Schwenk K (ed) Feeding: Form, Function, and Evolution in Tetrapod Vertebrates Academic Press, San Diego pp 487–526Google Scholar

Copyright information

© Springer Science+Business Media New York 2015

Authors and Affiliations

  1. 1.Department of GeologyUniversity of OtagoDunedinNew Zealand

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