Basilosaurus | |
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B. cetoides skeleton, National Museum of Natural History, Washington DC | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Artiodactyla |
Infraorder: | Cetacea |
Family: | † Basilosauridae |
Subfamily: | † Basilosaurinae |
Genus: | † Basilosaurus Harlan 1834 |
Species | |
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Synonyms | |
Basilosaurus (meaning "king lizard") 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. [2] 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, prompting 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. [3] Fossils have also been unearthed in the southeastern United States and Peru. [4] [5] [6]
Basilosaurus is thought to have been common in the Tethys Ocean. [7] [8] It was one of the largest animals of the Paleogene. It was the top predator of its environment, preying on sharks, large fish and other marine mammals, namely the dolphin-like Dorudon , which seems to have been their predominant food source. Based on the localities where its fossils are discovered, Basilosaurus would have preferred to live in the shallows, specifically in the middle to outer neritic zones of the inland sea. [4]
Basilosaurus was at one point a wastebasket taxon before the genus slowly started getting reevaluated, with many species of different Eocene cetacean being assigned to the genus in the past. However, most are invalid or have been reclassified under a new or different genus, leaving only 2 confirmed species. Basilosaurus may have been one of the first fully aquatic cetaceans, [2] sometimes referred to as the Pelagiceti. [9] Basilosaurus, unlike modern cetaceans, had various types of teeth–such as canines and molars–in its mouth (heterodonty), and it probably was able to chew its food, in contrast to modern cetaceans which swallow their food whole. [10] [11] Despite there being whales of a similar size and lifestyle to Basilosaurus, Basilosaurus was still the apex predator of the Eocene seas.
The two species of Basilosaurus are B. cetoides, whose remains were discovered in the United States, and B. isis, which was discovered in Egypt. B. cetoides is the type species for the genus. [12] [13] The holotype of B. cetoides was found in Ouachita Parish, Louisiana. [14] [15] Vertebrae were sent to the American Philosophical Society by a Judge Henry Bry of Ouachita Parish, Louisiana and Judge John Creagh of Clarke County, Alabama. Both fossils ended up in the hands of the anatomist Richard Harlan, who requested more examples from Creagh. [16] [17] The first bones were unearthed when rain caused a hillside full of sea shells to slide. The bones were lying in a curved line "measuring upwards of four hundred feet in length, with intervals which were vacant." Many of these bones were used as andirons and destroyed; Bry saved the bones he could find, but was convinced more bones were still to be found on the location. Bry speculated that the bones must have belonged to a "sea monster" and supplied "a piece having the appearance of a tooth" to help determine which kind. [18]
Harlan identified the tooth as a wedge-shaped shell and instead focused on "a vertebra of enormous dimensions" which he assumed belonged to the order "Enalio-Sauri of Conybeare", "found only in the sub-cretaceous series." [19] He noted that some parts of the vertebra were similar to those of Plesiosaurus and skull was similar to Mosasaurus , but that they were completely different in proportions. Comparing his vertebra to those of large dinosaurs such as Megalosaurus and Iguanodon , Harlan concluded that his specimen was considerably larger—he estimated the animal to have been no less than 80–100 ft (24–30 m) long—and therefore suggested the name Basilosaurus, meaning "king lizard". [20]
Harlan brought his assembled specimens (including fragments of jaw and teeth, humerus, and rib fragments) to the UK where he presented them to anatomist Richard Owen. Owen concluded that the molar teeth were two-rooted, a dental morphology unknown in fishes and reptiles, and more complex and varied than in any known reptile, and therefore that the specimen must be a mammal. Owen correctly associated the teeth with cetaceans, but he thought it was an herbivorous animal, similar to sirenians. [21] Consequently, Owen proposed renaming the find Zeuglodon cetoides ("whale-like yoke teeth" in reference to the double-rooted teeth) and Harlan agreed. [22]
Wādī al-Ḥītān (Arabic : وادي الحيتان, lit. 'Valley of the Whales') is an Egyptian sandstone formation where many early-whale skeletons were discovered. [23] German botanist Georg August Schweinfurth discovered the first archaeocete whale in Egypt (Zeuglodon osiris, now Saghacetus osiris) in 1879. He visited the Qasr el Sagha Formation in 1884 and 1886 and missed the now famous Wadi El Hitan by a few kilometers. German paleontologist Wilhelm Barnim Dames described the material, including the type specimen of Z. osiris, a well-preserved dentary. [24]
Hugh Beadnell, head of the Geological Survey of Egypt 1896–1906, [24] named and described Zeuglodon isis in Andrews 1904 based on a partial mandible and several vertebrae from Wadi El Hitan in Egypt. [25] Andrews 1906 [26] described a skull and some vertebrae of a smaller archaeocete and named it Prozeuglodon atrox, now known today as Dorudon atrox. Kellogg 1936 discovered deciduous teeth in this skull and it was then believed to be a juvenile [Pro]zeuglodon isis for decades before more complete fossils of mature Dorudon were discovered. [27] [24] [28]
In the 1980s, Elwyn L. Simons and Philip D. Gingerich started to excavate at Qasr el-Sagha and Wadi El Hitan with the hope of finding material that could match archaeocete fossils from Pakistan. Since then, over 500 archaeocete skeletons have been found at these two locations, of which most are B. isis or D. atrox, several of the latter carrying bite marks assumed to be from the former. [24] A 1990 paper described additional fossils including foot bones and speculated that the reduced hind limbs were used as copulatory guides. [29] One thing that was noted, was that whale fossils were so common, that when a mason company looked at their newest table counter, they realized that they had created a cross section of a 40 million year old basilosaurid fossil. This find was another thing that caught the eye of Gingerich. [30]
In 2015, a complete skeleton, the first-ever such find for Basilosaurus, was uncovered in Wadi El Hitan, preserved with the remains of its prey, including a Dorudon and several species of fish. [31] The whale's skeleton also shows signs of scavenging or predation by large sharks such as the otodontid Carcharocles sokolovi.
Many dubious species have been assigned to Basilosaurus in the past which have since been invalidated or were too incomplete to determine anything.
A nomen dubium is a scientific name that is of unknown or doubtful application. There are a few documented cases of this being applied to Basilosaurus in the past.
Basilosaurus is one of the largest animals known to exist between the K–Pg extinction event 66 million years ago (mya) and around 15 million years ago when modern cetaceans began to reach enormous sizes. [51] [52] B. cetoides measured 17–20 metres (56–66 ft) long and weighed more than 5.8 metric tons (6.4 short tons), while B. isis measured 15–18 metres (49–59 ft) long and weighed nearly 6.5 metric tons (7.2 short tons). [31] [53] Basilosaurus is distinguished from other genera of basilosaurids by its larger body size and its more elongated posterior thoracic, lumbar, and anterior caudal vertebrae. Basilosaurus does not have the vertically oriented metapophyses seen in its closest relative the basilosaurid known as Basiloterus . Basilosaurus is considered to be the largest of archeocete whales. [39] [54]
The dental formula for B. isis is 3.1.4.23.1.4.3. The upper and lower molars and second to fourth premolars are double-rooted and high-crowned. [55]
The head of Basilosaurus did not have room for a melon like modern toothed whales, and the brain was smaller in comparison, as well. They are not believed to have had the echolocation capabilities nor the social dynamics of extant cetaceans.
A 2011 study concluded that the skull of Basilosaurus is asymmetrical like in modern toothed whales, and not, as previously assumed, symmetrical like in baleen whales and artiodactyls (which are closely related to cetaceans). In modern toothed whales, this asymmetry is associated with high-frequency sound production and echolocation, neither of which is thought to have been present in Basilosaurus. This probably evolved to detect sound underwater, with a fatty sound-receiving pad in the mandible. [56]
In the skull, the inner and middle ear are enclosed by a dense tympanic bulla. [57] The synapomorphic cetacean air sinus system is partially present in basilosaurids, including the pterygoid, peribullary, maxillary, and frontal sinuses. [58] The periotic bone, which surrounds the inner ear, is partially isolated. The mandibular canal is large and laterally flanked by a thin bony wall, the pan bone or acoustic fenestra. These features enabled basilosaurs to hear directionally in water. [57]
The ear of basilosaurids is more derived than those in earlier archaeocetes, such as remingtonocetids and protocetids, in the acoustic isolation provided by the air-filled sinuses inserted between the ear and the skull. The basilosaurid ear did, however, have a large external auditory meatus, strongly reduced in modern cetaceans, but, though this was probably functional, it can have been of little use under water. [59]
A 16-meter (52 ft) individual of B. isis had 35-centimeter-long (14 in) hind limbs with fused tarsals and only three digits. The limited size of the limb and the absence of an articulation with the sacral vertebrae make a locomotory function unlikely. [60] Analysis has shown that the reduced limbs could rapidly adduct between only two positions. [29] Possible uses for the structure have been given, such as clasper-like body functions (compare to the function of pelvic spurs, the last vestiges of limbs in certain modern snakes). These limbs would have been used to guide the animals' long bodies during mating. [61]
A complete Basilosaurus skeleton was found in 2015, and several attempts have been made to reconstruct the vertebral column from partial skeletons. Kellogg 1936 estimated a total of 58 vertebrae, based on two partial and nonoverlapping skeletons of B. cetoides from Alabama. More complete fossils uncovered in Egypt in the 1990s allowed a more accurate estimation: the vertebral column of B. isis has been reconstructed from three overlapping skeletons to a total of 70 vertebrae with a vertebral formula interpreted as seven cervical, 18 thoracic, 20 lumbar and sacral, and 25 caudal vertebrae. The vertebral formula of B. cetoides can be assumed to be the same. [12]
Basilosaurus has an anguilliform (eel-like) body shape because of the elongation of the centra of the thoracic through anterior caudal vertebrae. In life, these vertebrae were filled with marrow, and because of the enlarged size, this made them buoyant. Basilosaurus probably swam predominantly in two dimensions at the sea surface, in contrast to the smaller Dorudon , which was likely a diving, three-dimensional swimmer. [62] The skeletal anatomy of the tail suggests that a small fluke was probably present, which would have aided only vertical motion. [63]
Similarly sized thoracic, lumbar, sacral, and caudal vertebrae imply that it moved in an anguilliform fashion, but predominantly in the vertical plane. Paleontologist Philip D. Gingerich theorized that Basilosaurus may also have moved in a very odd, horizontal anguilliform fashion to some degree, something completely unknown in modern cetaceans. The vertebrae appear to have been hollow, and likely also fluid-filled. This would imply that Basilosaurus typically functioned in only two dimensions at the ocean surface, compared with the three-dimensional habits of most other cetaceans. Judging from the relatively weak axial musculature and the thick bones in the limbs, Basilosaurus is not believed to have been capable of sustained swimming or deep diving, or terrestrial locomotion. [64] Basilosaurus did still have an elbow joint in its flipper like a seal.
The cheek teeth of Basilosaurus retain a complex morphology and functional occlusion. Heavy wear on the teeth reveals that food was first chewed then swallowed. [57] Scientists were able to estimate the bite force of Basilosaurus isis by analyzing the scarred skull bones of another species of prehistoric whale, Dorudon, and concluded that it could exert a maximum bite force of at least 16,400 newtons (3,700 lbf) and could possibly exceed 20,000 newtons (4,500 lbf), [65] roughly equivalent to the range between modern alligators [66] and crocodiles. [67]
Analyses of the stomach contents of B. cetoides has shown that this species fed exclusively on fish and large sharks, while bite marks on the skulls of juvenile Dorudon have been matched with the dentition of B. isis, suggesting a dietary difference between the two species, similar to that found in different populations of modern killer whales. [55] It was probably an active predator rather than a scavenger. [68] The discovery of juvenile Dorudon at Wadi Al Hitan bearing distinctive bite marks on their skulls indicates that B. isis would have aimed for the skulls of its victims to kill its prey, and then subsequently torn its meals apart, based on the disarticulated remains of the Dorudon skeletons. The finding further cements theories that B. isis was an apex predator that may have hunted newborn and juvenile Dorudon at Wadi Al Hitan when mothers of the latter came to give birth. [31] The stomach contents of an elderly male B. isis not only includes Dorudon but the fish Pycnodus mokattamensis. [31]
Basilosaurus would have been the top predator of its environment. [69] It lived in the warm tropical environment of the Eocene in areas abundant with sea grasses, such as Thalassodendron , Thalassia (also known as turtle grass) and Halodule . [70] [71] [72] It would have coexisted with the dolphin-like Dorudon, [73] the whales Cynthiacetus and Basiloterus , [74] the primitive sirenian Protosiren , the early elephant Moeritherium , [75] [76] [77] the sea turtle Puppigerus [78] and many sharks, such as Galeocerdo alabamensis , [79] Physogaleus , Otodus , Squatina prima , Striatolamia , Carcharocles sokolovi and Isurus praecursor . [80]
Basilosaurus fossil record seems to end at about 35–33.9 mya. [81] Basilosaurus extinction coincides with the Eocene–Oligocene extinction event which happened 33.9 mya, [82] which also resulted in the extinction of almost all other archaeocetes. [83] The event has been attributed to volcanic activity, meteor impacts, or a sudden change in climate (such as the environment getting cooler), the latter of which might have caused changes in the ocean by disrupting oceanic circulation, thus limiting the numbers of prey for predators like Basilosaurus to feed on. [84] [85] [86] Basilosaurus went extinct leaving no descendants, along with the rest of the Archaeocetes. After their extinction, the new currents and deep ocean upwelling created a new environment that favored the evolutionary diversification of modern cetaceans (Neocetes) such as early toothed and baleen whales, from more advanced Archaeocetes that evolved the traits associated with Neocetes.
Below is the phylogenetic analysis on the placement of Basilosaurus. Two subfamilies exist in Basilosauridae: Basilosaurinae which includes Basilosaurus, and Dorudontinae. These groups have been declared invalid in the past. [87] [88] Dorudon remains were once thought to represent juvenile Basilosaurus. [89]
The species B. cetoides is the state fossil of Alabama [84] and Mississippi. [90] [91] During the early 19th century, B. cetoides fossils were so common (and sufficiently large) that they were regularly used as furniture in the American South. [92]
Basilosaurus is featured in the BBC's Walking with... series in Walking with Beasts and Sea Monsters . [93] [94] [95]
In the novel Moby-Dick by Herman Melville, Ishmael cites the Basilosaurus during his studies as a possible whale fossil. [96]
Pakicetidae is an extinct family of Archaeoceti that lived during the Early Eocene in Pakistan. Unlike modern cetaceans, they had well developed limbs and were capable of walking.
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.
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.
Archaeoceti, or Zeuglodontes in older literature, is a paraphyletic group of primitive cetaceans that lived from the Early Eocene to the late Oligocene. Representing the earliest cetacean radiation, they include the initial amphibious stages in cetacean evolution, thus are the ancestors of both modern cetacean suborders, Mysticeti and Odontoceti. This initial diversification occurred in the shallow waters that separated India and Asia 53 to 45 mya, resulting in some 30 species adapted to a fully oceanic life. Echolocation and filter-feeding evolved during a second radiation 36 to 35 mya.
Pycnodus is an extinct genus of ray-finned fish from the Eocene period. It is wastebasket taxon, although many fossils from Jurassic or Cretaceous are assigned to this genus, only Eocene species, P. apodus is valid. As its name suggests, it is the type genus of Pycnodontiformes.
Zygorhiza ("Yoke-Root") is an extinct genus of basilosaurid early whale known from the Late Eocene of Louisiana, Alabama, and Mississippi, United States, and the Bartonian to the late Eocene of New Zealand . Specimens reported from Europe are considered Dorudontinae incertae sedis.
Pachycetus is an extinct genus of pachycetine basilosaurid from Middle Eocene of the eastern United States and Europe. The best known remains generally suggest that Pachycetus lived during the Bartonian, however, fossil finds have also been recovered from sediments of less certain age that could suggest that it may have also lived during the Late Lutetian and Early Priabonian. Pachycetus is primarily known from vertebrae and ribs and is characterized by its highly osteosclerotic and pachyostotic skeleton. This means the bones not only featured thickened rings of cortical bone surrounding the internal cancellous bone, but the cortical bone was furthermore much denser than in other basilosaurids. Two species of Pachycetus are recognized: Pachycetus paulsonii from Europe and Pachycetus wardii from the United States. A third species might be represented by "Zeuglodon" wanklyni.
Georgiacetus is an extinct genus of ancient whale known from the Eocene period of the United States. Fossils are known from Georgia, Alabama, and Mississippi and protocetid fossils from the right time frame, but not yet confirmed as Georgiacetus, have been found in Texas and South Carolina.
Cynthiacetus is an extinct genus of basilosaurid early whale that lived during the Late Eocene Specimens have been found in the southeastern United States and Peru.
Remingtonocetidae is a diverse family of early aquatic mammals of the order Cetacea. The family is named after paleocetologist Remington Kellogg.
Basiloterus is an extinct genus of late-Eocene archaeocete whale from the Drazinda Formation in southwestern Punjab, Pakistan and possibly also the Barton Group 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.
Ancalecetus is an extinct genus of early whale known from the Late Eocene Birket Qarun Formation in Wadi Al-Hitan, Egypt. The species is named after anthropologist and primate researcher Elwyn L. Simons who discovered the type specimen in 1985.
Chrysocetus is a genus of extinct early whale known from Late Eocene-aged fossils of the eastern United States and western Africa. It is the second smallest basilosaurid after Saghacetus.
The Qasr el Sagha Formation is a geological formation located in Egypt. The formation is part of the Wadi El Hitan World Heritage Site. The Qasr el Sagha Formation overlies the Birket Qarun Formation and is overlain by the Gebel Qatrani Formation. The sandstones and shales of the formation were deposited in a deltaic to shallow marine environment. It dates to the Late Eocene.
Pontogeneus is a genus of extinct cetacean known from fossils recovered from the Late Eocene sediments of the southeastern United States.
Saghacetus is an extinct genus of basilosaurid early whale, fossils of which have been found in the Upper Eocene Qasr el Sagha Formation, Egypt.
Supayacetus is an extinct genus of basilosaurid cetacean from the Middle Eocene Paracas Formation of Peru. It has been noted for its relatively small size and basal morphology, with the sternum bearing close resemblance to those of protocetids. Due to this, it has been traditionally placed as one of the basalmost basilosaurids, except for a 2023 study that places it within the family Pachycetinae close to Neoceti. Supayacetus is monotypic, meaning the genus includes only a single species: S. muizoni.
Masracetus is an extinct genus of basilosaurid ancient whale known from the Late Eocene of Egypt.
Eocetus is an extinct protocetid early whale known from the early-late Eocene Giushi Formation in Gebel Mokattam, outside Cairo, Egypt. Fossil remains have also been discovered in the Aridal Formation of the Sahara Desert in southwestern Morocco.
Pachycetinae is an extinct subfamily of basilosaurid cetaceans that lived during the middle Eocene. The best-dated remains stem from Bartonian strata, but some finds suggest that they could have first appeared during the Lutetian and may have survived until the Priabonian. Fossils of pachycetines are chiefly known from the southern United States, Ukraine, Morocco and Germany, among others. They differ from other basilosaurids in having pachyostotic and osteosclerotic vertebrae and ribs, making them denser and heavier by comparison. Based on this it has been suggested that these whales lived in shallow waters and that these thickened bones act as a buoyancy control as seen in sirenians. Analysis of the teeth suggests that pachycetines had a varying diet, with the robust teeth of the larger Pachycetus indicating that it possibly fed on sharks, whereas the more gracile teeth of Antaecetus suggest a diet of smaller prey items. The clade currently only includes two genera, Antaecetus and Pachycetus, but a 2023 study suggests that the Peruvian Supayacetus may at least be a close relative.
But by far the most wonderful of all cetacean relics was the almost complete vast skeleton of an extinct monster, found in the year 1842, on the plantation of Judge Creagh, in Alabama. The awe-stricken credulous slaves in the vicinity took it for the bones of one of the fallen angels. The Alabama doctors declared it a huge reptile, and bestowed upon it the name of Basilosaurus. But some specimen bones of it being taken across the sea to Owen, the English Anatomist, it turned out that this alleged reptile was a whale, though of a departed species. A significant illustration of the fact, again and again repeated in this book, that the skeleton of the whale furnishes but little clue to the shape of his fully invested body. So Owen rechristened the monster Zeuglodon; and in his paper read before the London Geological Society, pronounced it, in substance, one of the most extraordinary creatures which the mutations of the globe have blotted out of existence.
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