Evolution of cetaceans

The evolution of cetaceans is thought to have begun in the Indian subcontinent from even-toed ungulates 50 million years ago (mya) and to have proceeded 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 (56-34 mya), the second epoch of the present-extending Cenozoic Era. Molecular and morphological analyses suggest Cetacea share a relatively recent closest common ancestor with hippopotami and that they are sister groups. Being mammals, they surface to breathe air; they have 5 finger bones (even-toed) in their fins; they nurse their young; and, despite their fully aquatic life style, they retain many skeletal features from their terrestrial ancestors.Thewissen, J. G. M., L. N. Cooper, J. C. George, and S. Bajpai. 2009. From land to water: the origin of whales, dolphins, and porpoises. Evolution: Education and Outreach 2:272–288. 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 (baleen whales) and Odontoceti (toothed whales) – are thought to have separated from each other around 28–33 mya 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. The presence of baleen in baleen whales occurred gradually, with earlier varieties having very little baleen, and their size is linked to baleen dependence (and subsequent increase in filter feeding).

Early evolution

The aquatic lifestyle of cetaceans first began in the Indian subcontinent from even-toed ungulates 50 million years ago, over a period of at least 15 million years, but a jawbone discovered in Antarctica may reduce this to 5 million years. Archaeoceti is an extinct parvorder of Cetacea containing ancient whales. The traditional hypothesis of cetacean evolution, first proposed by Van Valen in 1966, was that whales were related to the mesonychids, an extinct order of carnivorous ungulates (hoofed animals) that resembled wolves with hooves and were a sister group of the artiodactyls (even-toed ungulates). This hypothesis was proposed due to similarities between the unusual triangular teeth of the mesonychids and those of early whales. However, molecular phylogeny data indicates that whales are very closely related to the artiodactyls, with hippopotamuses as their closest living relative. Because of this observation, cetaceans and hippopotamuses are placed in the same suborder, Whippomorpha. Cetartiodactyla (formed from the words Cetacea and Artiodactyla) is a proposed name for an order that includes both cetaceans and artiodactyls. However, the earliest anthracotheres, the ancestors of hippos, do not appear in the fossil record until the Middle Eocene, millions of years after '' Pakicetus'', whereas the first known whale ancestor appeared during the Early Eocene; this difference in timing implies that the two groups diverged well before the Eocene. Molecular analysis identifies artiodactyls as being very closely related to cetaceans, so mesonychids are probably an offshoot from Artiodactyla, and cetaceans did not derive directly from mesonychids, but the two groups may share a common ancestor.

The molecular data are supported by the discovery of ''Pakicetus'', the earliest archaeocete. The skeletons of ''Pakicetus'' show that whales did not derive directly from mesonychids. Instead, they are artiodactyls that began to take to the water soon after artiodactyls split from mesonychids. Archaeocetes retained aspects of their mesonychid ancestry (such as the triangular teeth) which modern artiodactyls, and modern whales, have lost. The earliest ancestors of all hoofed mammals were probably at least partly carnivorous or scavengers, and today's artiodactyls and perissodactyls became herbivores later in their evolution. Whales, however, retained their carnivorous diet because prey was more available and they needed higher caloric content in order to live as marine endotherms (warm-blooded). Mesonychids also became specialized carnivores, but this was likely a disadvantage because large prey was uncommon. This may be why they were out-competed by better-adapted animals like the hyaenodontids and later Carnivora.

''Indohyus''

''Indohyus'' was a small chevrotain-like animal that lived about 48 million years ago in what is now Kashmir. It belongs to the artiodactyl family Raoellidae, which is believed to be the closest sister group of Cetacea. ''Indohyus'' is identified as an artiodactyl because it has two trochlea hinges, a trait unique to artiodactyls. Approximately the size of a raccoon or domestic cat, this omnivorous creature shared some traits of modern whales, most notably the involucrum, a bone growth pattern which is the diagnostic characteristic of any cetacean; this is not found in any other species. It also showed signs of adaptations to aquatic life, including dense limb bones that reduce buoyancy so that they could stay underwater, which are similar to the adaptations found in modern aquatic mammals such as the hippopotamus. This suggests a similar survival strategy to the African mousedeer or water chevrotain which, when threatened by a bird of prey, dives into water and hides beneath the surface for up to four minutes.

The first fossils of the Indohyus were unearthed by Indian geologist A. Ranga Rao. He discovered a few teeth and a jawbone amongst rocks that he had collected. After his death, his widow Leelavathi Rao donated the rocks to professor Hans Theweissan. His technician accidentally broke open a couple of the donated rocks and discovered additional Indohyus fossils.

Pakicetidae

The pakicetids were digitigrade hoofed mammals that are thought to be the earliest known cetaceans, with ''Indohyus'' being the closest sister group. They lived in the early Eocene, around 50 million years ago. Their fossils were first discovered in North Pakistan in 1979, located at a river not far from the shores of the former Tethys Sea. After the initial discovery, more fossils were found, mainly in the early Eocene fluvial deposits in northern Pakistan and northwestern India. Based on this discovery, pakicetids most likely lived in an arid environment with ephemeral streams and moderately developed floodplains millions of years ago. By using stable oxygen isotopes analysis, they were shown to drink fresh water, implying that they lived around freshwater bodies. Their diet probably included land animals that approached water for drinking or some freshwater aquatic organisms that lived in the river. The elongated cervical vertebrae and the four, fused sacral vertebrae are consistent with artiodactyls, making ''Pakicetus'' one of the earliest fossils to be recovered from the period following the Cetacea/Artiodactyla divergence event.

Pakicetids are classified as cetaceans mainly due to the structure of the auditory bulla (ear bone), which is formed only from the ectotympanic bone. The shape of the ear region in pakicetids is highly unusual and the skull is cetacean-like, although a blowhole is still absent at this stage. The jawbone of pakicetids also lacks the enlarged space (mandibular foramen) that is filled with fat or oil, which is used in receiving underwater sound in modern cetaceans. They have dorsal orbits (eye sockets facing up), which are similar to crocodiles. This eye placement helps submerged predators observe potential prey above the water. According to a 2009 study, the teeth of pakicetids also resemble the teeth of fossil whales, being less like a dog's incisors, and having serrated triangular teeth, which is another link to more modern cetaceans.
It was initially thought that the ears of pakicetids were adapted for underwater hearing, but, as would be expected from the anatomy of the rest of this creature, the ears of pakicetids are specialized for hearing on land. However, pakicetids were able to listen underwater by using enhanced bone conduction, rather than depending on the tympanic membrane like other land mammals. This method of hearing did not give directional hearing underwater.

Pakicetids have long thin legs, with relatively short hands and feet which suggest that they were poor swimmers. To compensate for that, their bones are unusually thick (osteosclerotic), which is probably an adaptation to make the animal heavier to counteract the buoyancy of the water. According to a 2001 morphological analysis by Thewissen et al., pakicetids display no aquatic skeletal adaptation; instead they display adaptations for running and jumping. Hence pakicetids were most likely aquatic waders.

Ambulocetidae

'' Ambulocetus'', which lived about 49 million years ago, was discovered in Pakistan in 1994. They were vaguely crocodile-like mammals, possessing large brevirostrine jaws. In the Eocene, ambulocetids inhabited the bays and estuaries of the Tethys Sea in northern Pakistan. The fossils of ambulocetids are always found in near-shore shallow marine deposits associated with abundant marine plant fossils and littoral mollusks. Although they are found only in marine deposits, their oxygen isotope values indicate that they consumed a range of water with different degrees of salinity, with some specimens having no evidence of sea water consumption and others that did not ingest fresh water at the time when their teeth were fossilized. It is clear that ambulocetids tolerated a wide range of salt concentrations. Hence, ambulocetids represent a transition phase of cetacean ancestors between fresh water and marine habitat.

The mandibular foramen in ambulocetids had increased in size, which indicates that a fat pad was likely to be housed in the lower jaw. In modern toothed whales, this fat pad in the mandibular foramen extends posteriorly to the middle ear. This allows sounds to be received in the lower jaw, and then transmitted through the fat pad to the middle ear. Similar to pakicetids, the orbits of ambulocetids are on the top of the skull, but they face more laterally than in pakicetids.

Ambulocetids had relatively long limbs with particularly strong hind legs, and they retained a tail with no sign of a fluke. The hindlimb structure of Ambulocetids shows that their ability to engage in terrestrial locomotion was significantly limited compared to that of contemporary terrestrial mammals, and likely did not come to land at all. The skeletal structures of the knee and ankle indicates that the motion of the hindlimbs was restricted into one plane. This suggests that, on land, propulsion of the hindlimbs was powered by the extension of dorsal muscles. They probably swam by pelvic paddling (a way of swimming which mainly utilizes their hind limbs to generate propulsion in water) and caudal undulation (a way of swimming which uses the undulations of the vertebral column to generate force for movements), as otters, seals and modern cetaceans do. This is an intermediate stage in the evolution of cetacean locomotion, as modern cetaceans swim by caudal oscillation (a way of swimming similar to caudal undulation, but is more energy efficient).

Recent studies suggest that ambulocetids were fully aquatic like modern cetaceans, possessing a similar thoracic morphology and being unable to support their weight on land. This suggests that complete abandonment of the land evolved much earlier among cetaceans than previously thought.

Remingtonocetidae

Remingtonocetids lived in the Middle-Eocene in South Asia, about 49 to 43 million years ago. Compared to family Pakicetidae and Ambulocetidae, Remingtonocetidae was a diverse family found in north and central Pakistan and western India. Remingtonocetids were also found in shallow marine deposits, but they were obviously more aquatic than ambulocetidae. This is demonstrated by the recovery of their fossils from a variety of coastal marine environments, including near-shore and lagoonal deposits. According to stable oxygen isotopes analysis, most remingtonocetids did not ingest fresh water, and had hence lost their dependency on fresh water relatively soon after their origin.

The orbits of remingtonocetids faced laterally and were small. This suggests that vision was not an important sense for them. The nasal opening, which eventually becomes the blowhole in modern cetaceans, was located near the tip of the snout. The position of the nasal opening had remained unchanged since pakicetids. One of the notable features in remingtonocetids is that the semicircular canals, which are important for balancing in land mammals, had decreased in size. This reduction in size had closely accompanied the cetacean radiation into marine environments. According to a 2002 study done by Spoor et al., this modification of the semicircular canal system may represent a crucial ‘point of no return’ event in early cetacean evolution, which excluded a prolonged semi-aquatic phase.

Compared to ambulocetids, remingtonocetids had relatively short limbs. Based on their skeletal remains, remingtonocetids were probably amphibious cetaceans that were well adapted to swimming, and likely to swim by caudal undulation only.

Protocetidae

The protocetids form a diverse and heterogeneous group known from Asia, Europe, Africa, and North America. They lived in the Eocene, approximately 48 to 35 million years ago. The fossil remains of protocetids were uncovered from coastal and lagoonal facies in South Asia; unlike previous cetacean families, their fossils uncovered from Africa and North America also include open marine forms. They were probably amphibious, but more aquatic compared to remingtonocetids. Protocetids were the first cetaceans to leave the Indian subcontinent and disperse to all shallow subtropical oceans of the world.
There were many genera among the family Protocetidae. There were different degrees of aquatic adaptations in this group, with some able to support their weight on land, and others not. The discovery of the southeastern Pacific '' Peregocetus'' indicates they crossed the Atlantic and achieved circumstances-equatorial distribution by 40 mya. Their amphibious nature is supported by the discovery of a pregnant ''Maiacetus'', in which the fossilised fetus was positioned for a head-first delivery, suggesting that ''Maiacetus'' gave birth on land. If they gave birth in the water, the fetus would be positioned for a tail-first delivery to avoid drowning during birth.

Unlike remingtonocetids and ambulocetids, protocetids have large orbits which are oriented laterally. Increasingly lateral-facing eyes might be used to observe underwater prey, and are similar to the eyes of modern cetaceans. Furthermore, the nasal openings were large and were halfway up the snout. The great variety of teeth suggests diverse feeding modes in protocetids.
In both remingtonocetids and protocetids, the size of the mandibular foramen had increased. The large mandibular foramen indicates that the mandibular fat pad was present. However the air-filled sinuses that are present in modern cetaceans, which function to isolate the ear acoustically to enable better underwater hearing, were still not present. The external auditory meatus (ear canal), which is absent in modern cetaceans, was also present. Hence, the method of sound transmission that were present in them combines aspects of pakicetids and modern odontocetes (toothed whales). At this intermediate stage of hearing development, the transmission of airborne sound was poor due to the modifications of the ear for underwater hearing while directional underwater hearing was also poor compared to modern cetaceans.

Some protocetids had short, wide fore- and hindlimbs that were likely to have been used in swimming, but the limbs gave a slow and cumbersome locomotion on land. It is possible that some protocetids had flukes. However, it is clear that they were adapted even further to an aquatic life-style. In ''Rodhocetus'', for example, the sacrum (a bone that, in land-mammals, is a fusion of five vertebrae that connects the pelvis with the rest of the vertebral column) was divided into loose vertebrae. However, the pelvis was still connected to one of the sacral vertebrae. The ungulate ancestry of these archaeocetes is still underlined by characteristics like the presence of hooves at the ends of the toes in ''Rodhocetus''.

The foot structure of ''Rodhocetus'' shows that protocetids were predominantly aquatic. A 2001 study done by Gingerich et al. hypothesized that ''Rodhocetus'' locomoted in the oceanic environment similarly to how ambulocetids pelvic paddling, which was supplemented by caudal undulation. Terrestrial locomotion of Rodhocetus was very limited due to their hindlimb structure. It is thought that they moved in a way similar to how eared seals move on land, by rotating their hind flippers forward and underneath their body.

Basilosauridae

Basilosaurids and dorudontines lived together in the late Eocene around 41 to 33.9 million years ago, and are the oldest known obligate aquatic cetaceans. They were fully recognizable whales which lived entirely in the ocean. This is supported by their fossils usually found in deposits indicative of fully marine environments, lacking any freshwater influx. They were probably distributed throughout the tropical and subtropical seas of the world. Basilosaurids are commonly found in association with dorudontines, and were closely related to one another. The fossilised stomach contents in one basilosaurid indicates that it ate fish.

Although they look very much like modern cetaceans, basilosaurids lacked the ' melon organ' that allows toothed whales to use echolocation. They had small brains; this suggests they were solitary and did not have the complex social structures of some modern cetaceans. The mandibular foramen of basilosaurids covered the entire depth of the lower jaw as in modern cetaceans. Their orbits faced laterally, and the nasal opening had moved even higher up the snout, closer to the position of the blowhole in modern cetaceans. Furthermore, their ear structures were functionally modern, with the insertion of air-filled sinuses between ear and skull. Unlike modern cetaceans, basilosaurids retained a large external auditory meatus.

Both basilosaurids have skeletons that are immediately recognizable as cetaceans. A basilosaurid was as big as the larger modern whales, with genera like ''Basilosaurus'' reaching lengths of up to long; dorudontines were smaller, with genera like ''Dorudon'' reaching about long. The large size of basilosaurids is due to the extreme elongation of their lumbar vertebrae. They had a tail fluke, but their body proportions suggest that they swam by caudal undulation and that the fluke was not used for propulsion. In contrast, dorudontines had a shorter but powerful vertebral column. They too had a fluke and, unlike basilosaurids, they probably swam similarly to modern cetaceans, by using caudal oscillations. The forelimbs of basilosaurids were probably flipper-shaped, and the external hind limbs were tiny and were certainly not involved in locomotion. Their fingers, however, retained the mobile joints of their ambulocetid relatives. The two tiny but well-formed hind legs of basilosaurids were probably used as claspers when mating. The pelvic bones associated with these hind limbs were not connected to the vertebral column as they were in protocetids. Essentially, any sacral vertebrae can no longer be clearly distinguished from the other vertebrae.

Both basilosaurids and dorudontines are relatively closely related to modern cetaceans, which belong to parvorders Odontoceti and Mysticeti. However, according to a 1994 study done by Fordyce and Barnes, the large size and elongated vertebral body of basilosaurids preclude them from being ancestral to extant forms. As for dorudontines, there are some species within the family that do not have elongated vertebral bodies, which might be the immediate ancestors of Odontoceti and Mysticeti. The other basilosaurids became extinct.

Evolution of modern cetaceans

Baleen whales

All modern baleen whales or mysticetes are filter-feeders which have baleen in place of teeth, though the exact means by which baleen is used differs among species (gulp-feeding within balaenopterids, skim-feeding within balaenids, and bottom plowing within eschrichtiids). The first members of both groups appeared during the middle Miocene. Filter feeding is very beneficial as it allows baleen whales to efficiently gain huge energy resources, which makes the large body size in modern varieties possible. The development of filter feeding may have been a result of worldwide environmental change and physical changes in the oceans. A large-scale change in ocean current and temperature could have contributed to the radiation of modern mysticetes. The earlier varieties of baleen whales, or "archaeomysticetes", such as ''Janjucetus'' and ''Mammalodon'' had very little baleen and relied mainly on their teeth.

There is also evidence of a genetic component of the evolution of toothless whales. Multiple mutations have been identified in genes related to the production of enamel in modern baleen whales. These are primarily insertion/deletion mutations that result in premature stop codons. It is hypothesized that these mutations occurred in cetaceans already possessing preliminary baleen structures, leading to the pseudogenization of a "genetic toolkit" for enamel production. Recent research has also indicated that the development of baleen and the loss of enamel-capped teeth both occurred once, and both occurred on the mysticete stem branch.

Generally it is speculated the four modern mysticete families have separate origins among the cetotheres. Modern baleen whales, Balaenopteridae (rorquals and humpback whale, ''Megaptera novaengliae''), Balaenidae (right whales), Eschrichtiidae (gray whale, ''Eschrictius robustus''), and Neobalaenidae (pygmy right whale, ''Caperea marginata'') all have derived characteristics presently unknown in any cetothere and vice versa (such as a sagittal crest). Mysticetes are also known for their gigantism, as baleen whales are among the largest organisms to ever have lived; they reach lengths greater than 20 m and weigh more than 100,000 kg. This gigantism is directly related to their feeding mechanism - mysticete size has been found to be dependent on the amount of baleen a mysticete can use to filter its prey. Additionally, size is a positively selected trait that gives mysticetes a boost in fitness. Mysticete populations will therefore slowly become even more gigantic as whales with larger amounts of baleen are selected.

Toothed whales

The adaptation of echolocation occurred when toothed whales (Odontoceti) split apart from baleen whales, and distinguishes modern toothed whales from fully aquatic archaeocetes. This happened around 34 million years ago in a second cetacean radiation. Modern toothed whales do not rely on their sense of sight, but rather on their sonar to hunt prey. Echolocation also allowed toothed whales to dive deeper in search of food, with light no longer necessary for navigation, which opened up new food sources. Toothed whales echolocate by creating a series of clicks emitted at various frequencies. Sound pulses are emitted, reflected off objects, and retrieved through the lower jaw. Skulls of ''Squalodon'' show evidence for the first hypothesized appearance of echolocation. ''Squalodon'' lived from the early to middle Oligocene to the middle Miocene, around 33–14 million years ago. ''Squalodon'' featured several commonalities with modern toothed whales: the cranium was well compressed (to make room for the melon, a part of the nose), the rostrum telescoped outward into a beak, a characteristic of the modern toothed whales that gave ''Squalodon'' an appearance similar to them. However, it is thought unlikely that squalodontids are direct ancestors of modern toothed whales.

The first oceanic dolphins such as kentriodonts, evolved in the late Oligocene and diversified greatly during the mid-Miocene. The first fossil cetaceans near shallow seas (where porpoises inhabit) were found around the North Pacific; species like ''Semirostrum'' were found along California (in what were then estuaries). These animals spread to the European coasts and Southern Hemisphere only much later, during the Pliocene

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