From Land to Water: the Origin of Whales, Dolphins, and Porpoises
© The Author(s) 2009
Received: 5 September 2008
Accepted: 26 March 2009
Published: 16 April 2009
Cetaceans (whales, dolphins, and porpoises) are an order of mammals that originated about 50 million years ago in the Eocene epoch. Even though all modern cetaceans are obligate aquatic mammals, early cetaceans were amphibious, and their ancestors were terrestrial artiodactyls, similar to small deer. The transition from land to water is documented by a series of intermediate fossils, many of which are known from India and Pakistan. We review raoellid artiodactyls, as well as the earliest families of cetaceans: pakicetids, ambulocetids, remingtonocetids, protocetids, and basilosaurids. We focus on the evolution of cetacean organ systems, as these document the transition from land to water in detail.
Whales, dolphins, and porpoises together constitute the Cetacea (English: cetaceans). All modern Cetacea live in water and cannot survive out of the water. In spite of this, cetaceans are mammals. Like other mammals and unlike other vertebrates, they nurse their young; they have three ear bones that are involved in sound transmission (hammer, anvil, and stirrup), and their lower jaws consist of a single bone (the dentary).
Cetacea includes one of the largest species of animal ever, the blue whale (27 m in length, 136,000 kg) but also has some very small modern representatives, e.g., the vaquita (1.4 m in length, 42 kg). In spite of the variation in body size, all modern Cetacea are relatively similar in shape: they have a horizontal tail fluke used in swimming; their forelimbs are flippers; there are no external hind limbs; their neck is short, and their body is streamlined.
Cetaceans are unrelated to other marine mammals, the sirenians (manatees and dugongs) and the pinnipeds (seals, sea lions, walruses). Sirenians are most closely related to elephants, and pinnipeds are related to land carnivores (e.g., dogs and bears). In some regards, all cetaceans, sirenians, and pinnipeds are similar; they are all adapted to life in water. For instance, they all have streamlined bodies, short limbs, and fin-shaped hands and feet. In other regards, these three groups are dissimilar. For instance, cetaceans and sirenians lack (nearly all) body hair, whereas pinnipeds have dense fur. On the other hand, whereas the main propulsive organ of cetaceans and sirenians is the tail, sea lions swim with their forelimbs, and seals with their hind limbs.
Even in Darwin's time, it was known that cetaceans had land ancestors, but fossils that recorded the transition from land to water were not known: all fossil whales bore great similarity to modern whales. This changed in the early 1990s, when paleontologists unearthed the first of a series of fossil cetaceans, mostly in India and Pakistan, documenting the transition from land to water in detail in the Eocene Period (which lasted from approximately 54 to 34 million years ago). Now, cetacean origin is one of the best known examples of macroevolution documented in the fossil record.
Phylogeny: What Cetaceans Are Related To
Almost as soon as scientists realized that cetaceans had land ancestors, they tried to identify what the closest relatives of cetaceans were. Cetaceans are so different from land mammals that it was difficult to find significant similarities in the anatomy between cetaceans and land mammals. Molecular biology came to the rescue, identifying genetic similarities between cetaceans and artiodactyls (English: even-toed ungulates) that were not present in other mammals. Modern representatives of artiodactyls include pigs, hippos, camels, deer, sheep, cattle, and giraffe, and, of these, hippos are thought to be the closest living relatives of cetaceans (Nikaido et al. 1999; Gatesy and O'Leary 2001).
However, the oldest whale fossils known are approximately 50 million years old, and it is unlikely that the closest relatives of whales are still living. Therefore, it was up to paleontologists to find the artiodactyl that is most closely related to whales among the extinct diversity of even-toed ungulates. This happened in 2007, when skeletons for raoellids were found in the Himalayas that were shown to be the closest relatives to whales (Thewissen et al. 2007).
Here, we will present an overview of the most important players in the origin of cetaceans. We will discuss them, starting with raoellids and continuing with archaeocetes, the archaic whales that lived in the Eocene, approximately between 55 and 37 million years ago. We will discuss these following the order of the cladogram. Cetacean evolution continued after that with the two suborders of whales that have modern representatives, Odontoceti (toothed whales, which includes porpoises and dolphins) and Mysticeti (baleen whales), but their evolution is not discussed here. There are several recent reviews of the evolution of odontocetes and mysticetes (Fordyce and Muizon 2001; Bianucci and Landini 2007).
Raoellidae: the Closest Relatives of Cetaceans
Raoellidae is one of the families of artiodactyls. It contains a small group of species, most of which are only known from teeth and jaws (Thewissen et al. 2001, 2007). Skulls and skeletons are known for a single raoellid: Indohyus (Thewissen et al. 2007). Raoellids are only known from Pakistan and western India and are restricted to the lower and middle Eocene, approximately between 55 and 45 million years ago.
Further evidence of the aquatic habitat for Indohyus comes from the chemical composition of its teeth. Teeth consist mostly of calcium phosphate. Oxygen in the molecules that make up the teeth comes from the drinking water and food that the animal ingests. Two isotopes, forms of elements that are chemically identical but have heavier atoms because of excess neutrons in the nucleus, are common in nature: Oxygen-16 and Oxygen-18 (where the number reflects the mass of the atom). Oxygen-16 is by far the more common isotope (over 99% in nature), but the ratio between Oxygen-16 and Oxygen-18 varies in different environments, and animals living in water have a different ratio compared to animals living on land (Roe et al. 1998; Clementz et al. 2006). A stable isotope study of the teeth of Indohyus also suggested that it lived in water (Thewissen et al. 2007).
These results suggest that Indohyus was aquatic and thus that cetaceans originated from aquatic ancestors. It may seem odd that a 47-million-year-old artiodactyl that looks like a tiny deer is aquatic, but this behavior is reminiscent of one species of modern artiodactyl. The African mouse deer (Hyemoschus aquaticus) lives on the forest floor of central Africa, feeding mostly on fruits and flowers. It always stays near water, and when in danger from a predator, Hyemoschus jumps in the water and scurries to safety fully submerged. A remarkable video of this behavior is posted on www.youtube.com and is called Eagle versus Water Chevrotain (chevrotain is the French name for African mouse deer).
Hyemoschus is not osteosclerotic and spends relatively little time in the water. Given its morphology, it appears that Indohyus is more aquatic than Hyemoschus and may have spent much of its life in water. It is possible that it fed on water plants, but it is also possible that it came on land to feed on land plants, in a way similar to modern hippos.
With aquatic origins for cetaceans now being known to occur within the artiodactyls, the search is on for the discovery of the terrestrial relatives of raoellids. It is possible that these relatives are also closely related to hippopotamids, which would make molecular and morphological phylogenies consistent.
Pakicetidae: the First Cetaceans
Externally, pakicetids look nothing like a modern cetacean. They are more similar to a wolf with a long nose and tail (Thewissen et al. 2001; www.neoucom.edu/DEPTS/ANAT/Thewissen/whale_origins/whales/Pakicetid.html). However, the details of the pakicetid skeleton tell a different story; this was not an ordinary land predator. The skulls show that the orbits (the sockets of the eyes) of these cetaceans were located close together on top of the skull, as is common in aquatic animals that live in water but look at emerged objects. Just like Indohyus, limb bones of pakicetids are osteosclerotic (Madar 2007), also suggestive of aquatic habitat, an interpretation consistent with stable isotope evidence (Roe et al. 1998; Clementz et al. 2006).
Summarizing, pakicetids inherited the aquatic lifestyle from their raoellid ancestors. The position of the eyes, osteosclerosis of the limb bones, sedimentological data, and stable isotope data are consistent, and all suggest that pakicetids were waders in shallow freshwater.
Ambulocetidae: the First Marine Cetaceans
The skull of Ambulocetus has a long snout, as evidenced by the long lower jaw (much of the upper jaw is not preserved). In pakicetids, the eyes faced upward, whereas in Ambulocetus, they face toward the sides, although they are still located high on the skull (Nummela et al. 2006). This eye position occurs in aquatic mammals such as hippopotamus.
Ambulocetus fossils have only been found in rocks that were formed in a shallow sea, possibly in a coastal swamp or forest. Stable isotope data indicate that Ambulocetus lived in environments that were partly freshwater, possibly implying that they were near a river mouth (Roe et al. 1998).
Remingtonocetidae: Long-Snouted Cetaceans
The oldest representatives of the Remingtonocetidae are found at the same fossil localities as Ambulocetus, but the greatest diversity of remingtonocetids is known from younger rocks, between 48 and 41 million years ago in India and Pakistan (Gingerich et al. 1997). In all, there are four or five genera of remingtonocetids, characterized by a long snout, which makes up nearly two thirds of the length of the skull.
Dentally, remingtonocetids are specialized (Thewissen and Bajpai 2001a); their molars have lost the crushing basins of pakicetids and ambulocetids. This suggests that the diet of remingtonocetids is different from that of earlier cetaceans.
The morphology of the sense organs suggests that hearing was important for Remingtonocetus but that vision was not. This is consistent with the environmental evidence from the rocks that the fossils are found in. Indian Remingtonocetus probably lived in a muddy bay protected from the ocean by islands or peninsulas. Rivers may have brought sediment into this bay, and the water may not have been transparent.
The postcranial skeleton of remingtonocetids (Bajpai and Thewissen 2000) shows that these whales had short legs but a very long powerful tail. Consistent with Fish's hypothesis regarding the evolution of cetacean locomotion, these cetaceans may have used their tail as the main propulsive organ in the water and only used their limbs for steering, and they were probably fast swimmers, although the semicircular canals indicate that there was limited ability for locomotion on land. Modern giant South American river otters (Pteronura brasiliensis) have a long tail that is flat dorsoventrally and that is swept up and down during swimming. This type of locomotion may be a good model for swimming in Remingtonocetus. Therefore, externally, remingtonocetids may have resembled enormous otters with long snouts (www.neoucom.edu/DEPTS/ANAT/Thewissen/whale_origins/whales/Remi.html).
Protocetids: the Cetaceans that Conquered the Oceans
The earliest cetaceans, pakicetids, ambulocetids, and remingtonocetids are only known from India and Pakistan. With the origin of protocetids, cetaceans spread across the globe. Protocetids are known from low latitudes of Asia, Africa, Europe, and North America, and it is likely that they had a worldwide distribution in the middle Eocene between 49 and 40 million years ago (Gingerich et al. 1997; Williams 1998; Geisler et al. 2005).
Locomotor abilities in water may also differ between protocetids. While early reports on protocetid skeletons proposed that a fluke was present (Gingerich et al. 1994), it is now generally accepted that protocetids lacked a fluke (Gingerich et al. 2001b; Buchholtz 1998). Swimming may have been a combination of paddling with the hind limbs and dorsoventral undulations of the tail.
Little is known about the diet and feeding morphology of protocetid cetaceans, but, there too, variation appears to be common. Protocetids such as Babiacetus have heavy jaws (Fig. 23) with large teeth, suggestive of a diet that includes hard elements (such as bones of large fish or other vertebrates). For other protocetids, a diet of smaller fish has been suggested (O'Leary and Uhen 1999).
Basilosauridae: the First Fully Aquatic Cetaceans
In the late middle Eocene, around 41 million years ago, a new kind of cetacean emerged, the first one that resembles modern cetaceans: Basilosauridae (Uhen 1998). Basilosaurids have a nasal opening that has shifted back far toward the eyes to form a blowhole and have flippers for forelimbs, a fluke at the end of the tail, and tiny hind limbs, too tiny to support the body weight on land. In all these features, basilosaurids are more similar to modern cetaceans than to protocetids, and it is likely that they did not leave the oceans and were the first obligate cetaceans (Kellogg 1936; Uhen 2004).
There are approximately seven genera of basilosaurid cetaceans, but basically they can be divided into two body types. The first occurs in the genus Basilosaurus which had a snake-like body with a maximum length of approximately 17 m long. Basilosaurus may have swum by sinuous movements of its entire body (Buchholtz 1998). The second body type among basilosaurids is shorter, as short as 4 m. These basilosaurids, called dorudontines (Uhen 1998), had dolphin-shaped bodies and swam by up-and-down motions of their tail fluke. Basilosaurids are known from all the New World and the Old World and probably lived in all seas between 41 and 35 million years ago. The great length of the vertebral column of basilosaurids can be attributed to the increase in the number of lumbar vertebrae in the taxon but also by the increase in length of each individual vertebra.
Similar to earlier archaeocetes and unlike most later cetaceans, basilosaurids retained a heterodont dentition, with clear morphological differences between incisors, canines, premolars, and molars (Uhen 2004). This is unlike modern (odontocete) cetaceans in which the teeth along the tooth row are all very similar (a condition called homodonty). Unlike earlier archaeocetes, which all had 11 teeth per half jaw (44 teeth in all), basilosaurids had lost one tooth in each upper jaw, bringing their total number to 42. Their molars differed greatly from those of protocetids and ambulocetids, there not being a central depression surrounded by three cusps in the upper molars (O'Leary and Uhen 1999). As such, these teeth are not suitable for crushing food.
In the forelimb, basilosaurids resemble modern cetaceans, in that their elbow joint is not separately mobile and their hand webbed with individual digits not recognizable (Uhen 2004). Basilosaurids are like most mammals in that there are only three phalanges per finger, whereas in modern cetaceans this number is commonly increased.
Odontoceti and Mysticeti: Modern Cetaceans
Around 34 million years ago, the first representatives of the modern groups of whales, odontocetes and mysticetes are found. It is now generally assumed that odontocetes and mysticetes (together called Neoceti) arose from a common Eocene cetacean ancestor and are thus monophyletic. The most important innovation of the odontocete body plan is the acquisition of echolocation: These animals produce sounds that are reflected from objects that surround them, and these reflections enable them to image their surroundings. Mysticetes acquired a novel feeding mechanism: they filter feed for bulk prey (e.g., krill), using strainers in their mouth, the baleen plates. Although echolocation and filter feeding are important evolutionary themes of odontocetes and mysticetes, respectively, both of these suborders are diverse, feeding on different prey and using different hunting techniques.
Odontocetes and mysticetes conquered nearly all of the oceans: they include coastal and off-shore forms, arctic and tropical waters, shallow water, deep sea, and riverine forms. Good introductions to the evolutionary history of odontocetes and mysticetes have been published (Fordyce and Muizon 2001; Bianucci and Landini 2007).
In spite of our advances in understanding of the pattern of cetacean origins, it remains unclear which process caused this pattern: Why did cetaceans enter the oceans? The availability of rich new food sources has been proposed as a reason for the cetacean entry into the water, but this is unlikely, given that cetacean ancestors already lived in very shallow freshwater. The new find of aquatic behaviors in raoellids suggests that these animals used the water as a refuge against danger. Raoellid teeth are very different from those of early cetaceans, suggesting that a dietary shift took place after the habitat change and may have been critical in the early diversification of cetaceans but not in their entry into the water. On the other hand, it is not clear what raoellids ate, and neither raoellid nor early cetacean dentitions have good modern analogs. It has been suggested that early cetaceans ate fish (O'Leary and Uhen 1999).
The rich fossil record that has emerged can now be used to enrich other subfields of evolutionary science, including developmental biology, comparative anatomy, and molecular systematics. We hope that a detailed understanding of evolutionary patterns will allow us to determine the processes that drove cetacean evolution.
We thank the Geological Survey of Pakistan for collaborating in collecting and studying Pakistani fossils and for logistic support, and Dr. S. Taseer Hussain for his leadership of the Howard University-Geological Survey of Pakistan project. We thank Ajay Thakore and the Gujarat Mining Development Corporation for assistance with fieldwork in Gujarat, and Mr. Bhatti of Bhuj for help with logistics. We thank the Alaska Eskimo Whaling Commission and the Barrow Whaling Captains Association for access to specimens and contributing to their scientific study. The Bowhead whale specimens were collected under NMFS marine mammal collection permit 814-1899. We also thank the Department of Wildlife, North Slope Borough, and the Barrow Arctic Science Consortium for logistic support and assistance in the acquisition of specimens. This work was supported by grants from the Indian Department of Science and Technology (to Sunil Bajpai) and the US National Science Foundation (to J. G. M. Thewissen).
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