Archaeoceti
Archaeoceti Temporal range:
| |
---|---|
Cynthiacetus and Ambulocetus skeletons | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Artiodactyla |
Infraorder: | Cetacea |
Informal group: | †Archaeoceti Flower, 1883 |
Families and clades | |
See text |
Archaeoceti ("ancient whales"), or Zeuglodontes in older literature, is a
All archaeocetes from the
The archaeocetes are paraphyletic in relation to their extant modern descendants, the
Description
Pakicetidae
First identified as cetaceans by
Dozens of fossils are known, but only of skulls, teeth, and jaw fragments; no complete skeletons have been found. The dentition varied; the smallest species had teeth like modern fish eaters, and the largest were more like modern hyenas. The pakicetids may have been predators or carrion feeders. Neither the skull nor the dentition of pakicetids resembles those of modern whales, but the
Ambulocetidae
The next diverging family of whales, the Ambulocetidae, were large, already fully aquatic,[8] and crocodile-like with large feet and a strong tail. Sediments indicate that they lived in coastal areas and their compact bones suggest that they were ambush rather than fast-pursuit predators. Also known exclusively from Pakistan and India, the ambulocetids include the oldest known whale, Himalayacetus, which is believed to be 53.5 million years old, some 4 million years older than the rest of its family.[7]
Of the less than 10 fossils that have been described, one, Ambulocetus natans, is nearly complete and the main source of information concerning early cetacean evolution. The size of a male sea lion, it had a large head with a long snout and robust, strongly worn teeth. The lower jaw shows that Ambulocetus had an unusual soft tissue connecting the back of the jaw to the middle ear — a small equivalent to the large sound-receiving fat pad in modern odontocetes. Its eyes were placed dorsally on the head, but were facing laterally. The musculature of the head, neck, and back was strong and the fluke-less tail was long. The hind limbs were short, but equipped with long feet. The fore limbs were also short and equipped with five short hooves. Ambulocetus probably swam with its hind feet like a modern otter, and was incapable of supporting its own weight on land. It probably was an ambush hunter like modern crocodiles.[2]
Remingtonocetidae
The Remingtonocetidae had short limbs, and a strong and powerful tail with flattened vertebrae. Their long snout, tiny eyes, and ear morphology suggest their vision was poor and that hearing was their dominant sense. They, too, have only been found in Pakistan and India, and sediments suggest that they lived in turbid waters in coastal areas. Though they were probably able to live on land, they apparently used their tails to swim.[7]
Dozens of fossils have been described, but most are only skulls and lower jaws with few dental and postcranial remains. Remingtonocetids probably varied in size with the smallest species matching Pakicetus and the largest Ambulocetus. Remingtonocetids had longer snouts than other archaeocetes, except that the cranial morphology also varied considerably, probably reflecting different diets. The eyes were small, but the ears were large and set far apart — probably reflecting an increased emphasis on underwater hearing. The fragmentary remains of remingtonocetid postcrania suggest that they had a long neck and large hind limbs that were probably able to support the body weight on land.[2]
The remaining families and later crown cetaceans form a clade united by six
Protocetidae
The Protocetidae, known from both Africa and America, were a diversified family with hind limbs and a strong tail, indicating that they were strong swimmers that colonized shallow and warm oceans, such as reefs. They greatly affected cetacean evolution 47 to 41 million years ago, because they spread across Earth's oceans.[7] They had long snouts, large eyes, and a nasal opening located farther up the head than in earlier archaeocetes — suggesting they could breathe with the head held horizontally, similar to modern cetaceans — a first step towards a blowhole. Their dentition varied, but started to evolve towards the nonmasticating teeth of modern cetaceans, and they were probably active hunters. Their ability to move on land seems to have been variable: in Rodhocetus and Peregocetus possess a sacroiliac joint, indicating they could move on land.[10] In other genera (Georgiacetus and Aegicetus), the pelvis was not connected to the vertebral column, suggesting the hind limbs could not have supported the body weight.[2] Some genera (Rodhocetus) had large hind feet forming large paddles, while Aegicetus seems to have relied more on its tail to propel itself through the water.[11]
Basilosauridae
Taxonomy
The Archaeoceti include five well-established families:[13] The status of the Kekenodontidae is still disputed, and the family is placed in either the Archaeoceti, Mysticeti,[14] or even Delphinoidea.[15]
- Cetartiodactyla
- Protocetidae (Stromer 1908)
- Georgiacetinae (Gingerich et al. 2005)
- Makaracetinae (Gingerich et al. 2005)
- Protocetinae (Gingerich et al. 2005)
- Aegyptocetus (Bianucci & Gingerich 2011)
- Artiocetus (Gingerich et al. 2001)
- Dhedacetus
- Gaviacetus (Gingerich, Arif & Clyde 1995)
- Indocetus (Sahni & Mishra 1975)
- Maiacetus (Gingerich et al. 2009)
- Peregocetus
- Protocetus (Fraas 1904)
- Qaisracetus (Gingerich et al. 2001)
- Rodhocetus (Gingerich et al. 1994)
- Takracetus (Gingerich, Arif & Clyde 1995)
- Togocetus (Gingerich & Cappetta 2014)
- Basilosauridae
- Basilosaurinae
- Dorudontinae
- Ancalecetus (Gingerich & Uhen 1996)
- Basilotritus (Goldin & Zvonok 2013)
- Chrysocetus (Uhen & Gingerich 2001)
- Cynthiacetus (Uhen 2005)
- Dorudon (Gibbes 1845)
- Masracetus (Gingerich 2007)
- Ocucajea (Uhen et al. 2011)
- Pontogeneus
- Saghacetus (Gingerich 1992)
- Stromerius (Gingerich 2007)
- Supayacetus (Uhen et al. 2011)
- Tutcetus (Antar et al. 2023)
- Zygorhiza (True 1908)
- Pachycetinae
- Kekenodontidae
- Protocetidae (Stromer 1908)
Phylogeny
Notes
- ^ "Archaeoceti". Paleobiology Database. Retrieved 20 September 2018.
- ^ a b c d e Thewissen 2002, pp. 36–8
- ^ Fordyce 2002, p. 216
- ^ Geisler, Sanders & Luo 2005, Biogeography, pp. 50–2
- ^ Steeman et al. 2009, p. 573
- ^ Fordyce 2008, p. 758
- ^ a b c d e Bajpai, Thewissen & Sahni 2009, p. 675
- PMID 27396988.
- ^ Geisler, Sanders & Luo 2005, Discussion, p. 45
- ^ Olivier L, Bianucci G, Salas-Gismondi R, Di Celma C, Steurbaut E, Urbina M & de Muizon C (2019). "An amphibious whale from the Middle Eocene of Peru reveals early South Pacific dispersal of quadrupedal cetaceans". Current Biology 29(8): p. 1352–1359.e3.
- ^ Gingerich P.D., Antar M.S.M. & Zalmout I.S. (2019). "Aegicetus gehennae, a new late Eocene protocetid (Cetacea, Archaeoceti) from Wadi Al Hitan, Egypt, and the transition to tail-powered swimming in whales". PLOS ONE 15(3): e0230596
- ^ Thewissen et al. 2001, p. 277
- ^ Rose 2006, p. 273
- ^ Clementz et al. 2014, Fig. 1
- ^ Gingerich 2005, Table 15.1
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External links
- Media related to Archaeoceti at Wikimedia Commons