page_title: Evolution of cetaceans
section_title: Skeletal evolution
hierarchical_section_title: Evolution of cetaceans / Skeletal evolution
caption_attribution_description: English: Skeleton of a Baleen whale circling the hind limbs and pelvic structure.
context_page_description: The evolution of cetaceans is thought to have begun in the Indian subcontinent, from even-toed ungulates 50 million years ago, over a period of at least 15 million years. Cetaceans are fully aquatic marine mammals belonging to the order Artiodactyla, and branched off from other artiodactyls around 50 mya. Cetaceans are thought to have evolved during the Eocene or earlier, sharing a closest common ancestor with hippopotamuses. Being mammals, they surface to breathe air; they have 5 finger bones in their fins; they nurse their young; and, despite their fully aquatic life style, they retain many skeletal features from their terrestrial ancestors. Research conducted in the late 1970s in Pakistan revealed several stages in the transition of cetaceans from land to sea.
  The two modern parvorders of cetaceans – Mysticeti and Odontoceti – are thought to have separated from each other around 28-33 million years ago in a second cetacean radiation, the first occurring with the archaeocetes. The adaptation of animal echolocation in toothed whales distinguishes them from fully aquatic archaeocetes and early baleen whales.
context_section_description: Modern cetaceans have internal, rudimentary hind limbs, such as reduced femurs, fibulas, and tibias, and a pelvic girdle. Indohyus has a thickened ectotympanic internal lip of the ear bone. This feature compares directly to that of modern cetaceans. Another similar feature was the composition of the teeth, which contained mostly calcium phosphate which is needed for eating and drinking by aquatic animals, though, unlike modern day toothed whales, they had a heterodont (more than one tooth morphology) dentition as opposed to a homodont (one tooth morphology present) dentition. Although they somewhat resembled a wolf, the fossils of pakicetids showed the eye sockets were much closer to the top of their head than that of other terrestrial mammals, but similar to the structure of the eyes in cetaceans. Their transition from land to water led to reshaping of the skull and food processing equipment because the eating habits were changing. The change in position of the eyes and limb bones is associated with the pakicetids becoming waders. The ambulocetids also began to develop long snouts, which is seen in current cetaceans. Their limbs (and hypothesized movement) were very similar to otters.
  Limblessness in cetaceans does not represent a regression of fully formed limbs nor the absence of limb bud initiation, but rather arrest of limb bud development. Limb buds develop normally in cetacean embryos. Limb buds progress to the condensation phase of early skeletogenesis, where nerves grow into the limb bud and the apical ectodermal ridge (AER), a structure that ensures proper limb development, appears functional. Occasionally, the genes that code for longer extremities cause a modern whale to develop miniature legs (atavism).
  Pakicetus had a pelvic bone most similar to that of terrestrial mammals. In later species, such as Basilosaurus, the pelvic bone, no longer attached to the vertebrae and the ilium, was reduced. Certain genes are believed to be responsible for the changes that occurred to the cetacean pelvic structure, such as BMP7, PBX1, PBX2, PRRX1, and PRRX2. The pelvic girdle in modern cetaceans were once thought to be vestigial structures that served no purpose at all. The pelvic girdle in male cetaceans is different in size compared to females, and the size is thought to be a result of sexual dimorphism. The pelvic bones of modern male cetaceans are more massive, longer, and larger than those of females. Due to the sexual dimorphism displayed, they were most likely involved in supporting male genitalia that remain hidden behind abdominal walls until sexual reproduction occurs.
  Early archaeocetes such as Pakicetus had the nasal openings at the end of the snout, but in later species such as Rodhocetus, the openings had begun to drift toward the top of the skull. This is known as nasal drift. The nostrils of modern cetaceans have become modified into blowholes that allow them to break to the surface, inhale, and submerge with convenience. The ears began to move inward as well, and, in the case of Basilosaurus, the middle ears began to receive vibrations from the lower jaw. Today's modern toothed whales use their melon organ, a pad of fat, for echolocation.
caption: The skeleton of a bowhead whale with the hind limb and pelvic bone structure circled in red. This bone structure stays internal during the entire life of the species.
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