Basiloterus

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Basiloterus
Temporal range: Bartonian
~40.4–40  Ma
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Basiloterus husseini.png
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Artiodactyla
Infraorder: Cetacea
Family: Basilosauridae
Genus: Basiloterus
Gingerich et al., 1997
Species:
B. husseini
Binomial name
Basiloterus husseini
Gingerich et al., 1997

Basiloterus is an extinct genus of late-Eocene archaeocete whale from the Drazinda Formation in southwestern Punjab, Pakistan and possibly also the Barton Group (originally Barton Beds) of England. Known from two isolated lumbar vertebrae, the elongated nature of these elements has been taken as possible evidence that Basiloterus was a close relative of the better-known Basilosaurus . This was also the reasoning behind its name, which roughly translates to "another king". However, publications since then not only lead to some major changes of the internal relationships within Basilosauridae but have also called into question how diagnostic elongated vertebrae are for members of this group, as other early whales have developed similar anatomy independently. Though the identity of Basiloterus as a basilosaurid is generally maintained, its exact position within more recent interpretations of the family is unclear.

Contents

History and naming

The fossil remains of Basiloterus were discovered in 1996 in the green shales that compose the middle parts of the Drazinda Formation in Pakistan. The genus was described in 1997 by Gingerich et al. alongside the species Basilosaurus drazindai on the basis of two lumbar vertebrae thought to represent a single individual. Gingerich and colleagues tentatively refer a vertebral centrum from the Barton Group of England to this genus as well on the basis of its age, size and general morphology. [1]

The scientific name derives from the Ancient Greek "basileus" and "heteros" or "oteros", in combination translating to "other king" or "another king". [2] The name was explained as being chosen as a companion name to Basilosaurus, whose name translates to "king reptile". The species name of Basiloterus husseini honors Dr. S. Taseer Hussain, a researcher who warked at the Howard University in Washington DC and sponsored the first expedition to the Drazinda Formation. [1]

Description

Basiloterus differs from species of Basilosaurus in that the centra of the lumbar vertebrae were less elongated relative to their height compared to those of Basilosaurus. The crosssections of the vertebrae are more circular than in Pachycetus, but the front and back surfaces are still not perfectly round. Instead, the anterior epiphyseal surface, the part of the centrum that faces the vertebra before it, is described as mildly saddle-shaped, whereas the surface that faces backwards is more oval and flattened. The actual epiphysis are however missing in both specimens. The preserved parts of the vertebrae measure 18.5 cm (7.3 in) in length, accounting for the missing epiphysis this means would suggest that the vertebrae measured up to 20 cm (7.9 in) long if complete. The vertebrae narrow towards the middle from all sides, giving them a dumbbell-shape as described by Gol'din and Zvonok, which sets them apart from those of Basilotritus. [2] The lower surface of the vertebrae is concave and does not feature a keel, instead displaying three vascular foramina that are located within the concavity. [1]

The transverse processes, which project from the sides of the centrum, are described as being oriented anteroventrally. This downturned state is thought to allow for an increased arc of movement of the iliocostalis muscle. [1] The transverse processes are less elongated than those of Basilotritus. [2]

The laminae at the base of the neural spine is thin and leads into the spinous process while also enclosing the moderately large neural canal in junction with the pedicles. The pedicles connect to the laminae at an angle and are described as being situated closer to the front of the vertebra, something considered to be typical for basilosaurids by Gingerich. The metapophysis, which are robust pyramidal structures that project from the pedicles, are described as being oriented much more vertically than those of Basilosaurus. They extend beyond the front-most edge of the centrum, but do not flare out much. The spinous process extends to a height of 17.5 cm (6.9 in) above the vertebral centrum, all the while being directed at a 10° angle forward. The base of the neural spine is 12 cm (4.7 in) long from the front to the back but is only 1.3 cm (0.51 in) thick. The anterior to posterior length of the neural spine is another feature that differentiates Basiloterus from Basilotritus, as this element is less elongated. [2] The tip of the spinous process does not expand, suggesting only weak ligaments and muscle attachments. [1]

According to Gingerich and colleagues, Basiloterus was smaller than Basilosaurus, and Gol'din & Zvonok state that the vertebrae are of similar size to those of Basilotritus. [2] The size of the vertebrae has also been compared to Pachycetus robustus, with Gingerich determining that those of Basiloterus were around 25% larger. [1]

Phylogeny

Gingerich and colleagues considered Basiloterus to be a close relative of Basilosaurus based on the size and shape of the lumbar vertebrae as well as the fact that the two genera were contemporary. [1] Later, Mark D. Uhen argued that this first identification may not be as certain as initially believed. Although Uhen agrees that elongated vertebrae are a distinguishing feature of basilosaurines within the family Basilosauridae, he also points out that other groups evolved this trait independently. According to Uhen, this means that Basiloterus and Basilosaurus drazindai could also be protocetids similar to Eocetus , which also developed elongated vertebrae. [3] While few papers examined or even acknowledge Basiloterus following the type description, those that do feature it continue to regard it as a basilosaurid. [3] [4] [5] [6]

The phylogenetic tree provided by Uhen in 2009's Encyclopedia of Marine Mammals suggests a paraphyletic Basilosauridae leading up to modern cetaceans, [3] although some more recent studies have proposed monophyletic solutions for Basilosauridae as well. [4] Despite his reservations towards the identity of Basiloterus as a basilosaurine, the tree still depicts it as the immediate sister taxon to Basilosaurus as proposed by Gingerich and colleagues. [3]

Pakicetidae

Ambulocetidae

Remingtonocetidae

Protocetidae

Basilosauridae
Basilosaurinae

Cynthiacetus

Basiloterus

Basilosaurus

Dorudontinae

Dorudon

Ancalecetus

Zygorhiza

Saghacetus

Chrysocetus

Mysticeti

Odontoceti

The elongation of the vertebrae as a convergent trait is again noted by Gol'din and Zvonok, who propose that it had been independently gained by Basiloterus, Basilotritus and Basilosaurus. They argue that this independent acquisition of elongated lumbars renders it an unreliable character to separate the subfamilies Basilosaurinae and Dorudontinae, further supported by the close relationship they recover between Basilosaurus and Dorudon . In their results, Dorudon is the closest relative of Basilosaurus in place of Basiloterus, which is not included in their results. [2] Similar results also appear in later papers such as the description of the small-bodied basilosaurid Tutcetus, which also argues against the concept of a distinct Basilosaurinae and Dorudontinae and does not discuss Basiloterus. [7] The description of Cynthiacetus peruvianus also fails to recover two distinct subfamilies and simply refers to Basiloterus as a basilosaurid of uncertain relation to the other taxa within the family. [4]

Paleobiology

Although not found in the same area, Basiloterus likely coexisted with Basilosaurus drazindai as they both come from the same green shales of the Drazinda Formation, [1] which also yielded the remains of the giant softshell turtle Drazinderetes . The Drazinda Formation consists of shallow marine sediments. [8]

See also

Related Research Articles

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Rodhocetus is an extinct genus of protocetid early whale known from the Lutetian of Pakistan. The best-known protocetid, Rodhocetus is known from two partial skeletons that taken together give a complete image of an Eocene whale that had short limbs with long hands and feet that were probably webbed and a sacrum that was immobile with four partially fused sacral vertebrae. It is one of several extinct whale genera that possess land mammal characteristics, thus demonstrating the evolutionary transition from land to sea.

<span class="mw-page-title-main">Pakicetidae</span> Family of mammals

Pakicetidae is an extinct family of Archaeoceti that lived during the Early Eocene in Pakistan.

<i>Ambulocetus</i> Genus of extinct mammals of the order Cetacea

Ambulocetus is a genus of early amphibious cetacean from the Kuldana Formation in Pakistan, roughly 48 or 47 million years ago during the Early Eocene (Lutetian). It contains one species, Ambulocetus natans, known solely from a near-complete skeleton. Ambulocetus is among the best-studied of Eocene cetaceans, and serves as an instrumental find in the study of cetacean evolution and their transition from land to sea, as it was the first cetacean discovered to preserve a suite of adaptations consistent with an amphibious lifestyle. Ambulocetus is classified in the group Archaeoceti—the ancient forerunners of modern cetaceans whose members span the transition from land to sea—and in the family Ambulocetidae, which includes Himalayacetus and Gandakasia.

<i>Basilosaurus</i> Prehistoric cetacean genus from the Late Eocene epoch

Basilosaurus is a genus of large, predatory, prehistoric archaeocete whale from the late Eocene, approximately 41.3 to 33.9 million years ago (mya). First described in 1834, it was the first archaeocete and prehistoric whale known to science. Fossils attributed to the type species B. cetoides were discovered in the United States. They were originally thought to be of a giant reptile, hence the suffix "-saurus", Ancient Greek for "lizard". The animal was later found to be an early marine mammal, which prompted attempts at renaming the creature, which failed as the rules of zoological nomenclature dictate using the original name given. Fossils were later found of the second species, B. isis, in 1904 in Egypt, Western Sahara, Morocco, Jordan, Tunisia, and Pakistan. Fossils have also been unearthed in the southeastern United States and Peru.

<span class="mw-page-title-main">Basilosauridae</span> Family of mammals

Basilosauridae is a family of extinct cetaceans. They lived during the middle to the early late Eocene and are known from all continents, including Antarctica. They were probably the first fully aquatic cetaceans. The group is noted to be a paraphyletic assemblage of stem group whales from which the monophyletic Neoceti are derived.

<i>Dorudon</i> Genus of ancient whales

Dorudon ("spear-tooth") is a genus of extinct basilosaurid ancient whales that lived alongside Basilosaurus 40.4 to 33.9 million years ago in the Eocene. It was a small whale, with D. atrox measuring 5 metres (16 ft) long and weighing 1–2.2 metric tons. Dorudon lived in warm seas around the world and fed on small fish and mollusks. Fossils have been found along the former shorelines of the Tethys Sea in present-day Egypt and Pakistan, as well as in the United States, New Zealand and Western Sahara.

<span class="mw-page-title-main">Archaeoceti</span> Paraphyletic group of primitive cetaceans from Early Eocene to Late Oligocene

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<i>Zygorhiza</i> Genus of mammals

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<span class="mw-page-title-main">Basilosaurinae</span> Subfamily of mammals

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References

  1. 1 2 3 4 5 6 7 8 Gingerich, Philip D.; Arif, M.; Bhatti, M. Akram; Anwar, M.; Sanders, William J. (1997). "Basilosaurus drazindai and Basiloterus hussaini, New Archaeoceti (Mammalia, Cetacea) from the Middle Eocene Drazinda Formation, with a Revised Interpretation of Ages of Whale-Bearing Strata in the Kirthar Group of the Sulaiman Range, Punjab (Pakistan)" (PDF). Contributions from the Museum of Paleontology, University of Michigan. 30 (2): 55–81.
  2. 1 2 3 4 5 6 Gol’din, P.; Zvonok, E. (2013). "Basilotritus uheni, a new cetacean (Cetacea, Basilosauridae) from the late middle Eocene of eastern Europe". Journal of Paleontology. 87 (2): 254–268. Bibcode:2013JPal...87..254G. doi:10.1666/12-080r.1.
  3. 1 2 3 4 Uhen, M. D. (2009). "Basilosaurids". Encyclopedia of Marine Mammals. pp. 91–94. doi:10.1016/b978-0-12-373553-9.00026-2. ISBN   978-0-12-373553-9.
  4. 1 2 3 Martínez-Cáceres, M.; Lambert, O.; De Muizon, C. (2017). "The anatomy and phylogenetic affinities ofCynthiacetus peruvianus, a large Dorudon-like basilosaurid (Cetacea, Mammalia) from the late Eocene of Peru" (PDF). Geodiversitas. 39 (1): 7–163. doi:10.5252/g2017n1a1. S2CID   90142285.
  5. Uhen, M.D.; Taylor, D. (2020). "A basilosaurid archaeocete (Cetacea, Pelagiceti) from the Late Eocene of Oregon, USA". PeerJ. 8: e9809. doi: 10.7717/peerj.9809 . PMC   7534682 . PMID   33062412.
  6. Gingerich, P.D. (2003). "Land-to-sea transition in early whales: evolution of Eocene Archaeoceti (Cetacea) in relation to skeletal proportions and locomotion of living semiaquatic mammals" (PDF). Paleobiology. 29 (3): 429–454. doi:10.1666/0094-8373(2003)029<0429:LTIEWE>2.0.CO;2. S2CID   86600469.
  7. Antar, M.S.; Glaohar, A.S.; El-Desouky, H.; Seiffert, E.R.; El-Sayed, S.; Claxton, A.G.; Sallam, H.M. (2023). "A diminutive new basilosaurid whale reveals the trajectory of the cetacean life histories during the Eocene". Commun Biol. 6 (707): 707. doi: 10.1038/s42003-023-04986-w . PMC   10415296 . PMID   37563270.
  8. Head, J.J.; Raza, S.M.; Gingerich, P.D. (1999). "DRAZINDERETES TETHYENSIS, A NEW LARGE TRIONYCHID (REPTILIA: TESTUDINES) FROM THE MARINE EOCENE DRAZINDA FORMATION OF THE SULAIMAN RANGE, PUNJAB (PAKISTAN)". Contributions from the Museum of Paleontology University of Michigan. 30 (7): 199–214.