The Evolution of Cetaceans: Evolutionary Phylogeny, Fossil Records and Evolutionary Novelties1

Jayson Garcia Jacob Labios Abigail Larios

A term paper submitted in partial fulfillment of the requirements in Evolutionary Biology under Dr. William SM. Gruezo, 1st sem., 2011-2012.

I. Main Group of Cetaceans The word cetacean comes from the Latin cetus which means a large sea animal, and the Greek word ketos, meaning sea monster (Kennedy, 2011). Order Cetaceais comprised of the most uncharacteristic species falling under the Class Mammalia. They are fully adapted to their highly specialized way of life through alterations in the basic structures common to other mammals. Early in their history, they diverged from their primitive eutherian ancestors and were isolated. This isolation led to the rapid evolutionary changes within the species belonging to this group when they shift from terrestrial to aquatic environment (Colbert, 1980). Cetaceans include the whales, dolphins and porpoisesin about 87 species, 14 families and two sub-orders. The two sub-orders are Suborder Odontoceti or the toothed whales and Suborder Mysticeti or the baleen whales. Odontocetes have teeth for feeding and a single blowhole. There are more species falling under this taxon as compared to the other subgroup. They are also smaller than baleen whales and with an asymmetrical skull. Mysticetes, on the other hand, have baleen plates instead of teeth for filter feeding and two blowholes for respiration. In contrast to toothed whales, their skulls are symmetrical. II. Unifying Characteristics/Features Being part of Class Mammalia, cetaceans are warm blooded. They have no hair surrounding their body. These features lead to the development of a heavy fat layer underneath the skin called blubber for insulation by keeping the whales body warm.The whales went back to the sea to take the place of the extinct ichthyosaurs and to adapt to a life that is similar to fishes. Their body is torpedo shaped without a distinct neck. They have streamlined body that enabled them to move efficiently in their new habitat. They also developed a tail that resembles that of the fishes. This tail propelled them through the open oceans. Their tail leads to a horizontal end fin (flukes) that moves up and down, unlike the

fishes vertical fin which moves sideways. The legs became paddles that enabled them to balance their relatively long and large bodies (Colbert, 1980). Whales also acquired a unique mode of respiration. They have blowhole/s (one for toothed whales and two for baleen whales) through which they exchange gases. Whales go up to the surface of the water where they exhale and inhale using this blowhole and can stay underwater for a long time and dive to great depths due to their flexible lungs and rib cages. Whales have very poor sense of smell compared to other mammals but are very sensitive to vibrations in the water. They use these vibrations or echoes for echolocation and communication (Kennedy, 2011). Echolocation allows cetaceans to review their environment by emitting sounds and listening to echoes (Milinkovitch and Lambert, 2006). Through this mechanism, they are able to detect preys, predators and obstacles. They even use this to communicate with each other by sending in vibrations of different frequencies (Colbert, 1980). They have highly developed brains and some experiments showed that they are intelligent mammals (Colbert, 1980). When they reproduce, they give birth to live young whales which are large and welldeveloped. As soon as they are born, they are being pushed upwards by their mothers to the surface of the water to get their first dosage of air (Colbert 1980). The baby whales then swim beside their mother for nourishment. The mammary glands of the mothers are contained in a pocket where the young whales are being nursed without transporting a lot of sea water. III. Evolutionary Changes Evolutionary Changes: Microevolution When a blue whale opens its mouth, tons of water surge in. The whale then forces the water back out with its tongue, in such a way that it flows through the baleen combs in the front of its mouth. These baleen plates can filter up to half a million calories worth of plankton, krill and small fish out of the water. The earliest ancestors of the blue whale fed

themselves differently. They still had teeth, and no baleen. Some modern whales still have teeth (the Odontoceti literally toothed whales), but they form a separate group from the baleen whales (Mysticeti) who have replaced all their teeth with baleen. This transition to toothlessness is documented by multiple fossil whales. Each of these fossils provides a snapshot of what must have been gradual change. Several early baleen whales such as Janjucetus and Mammalodon still had fully developed enamel-covered teeth. Eomysticetus had already exchanged its teeth for baleen plates but it is Aetiocetus that captures this evolutionary change in its entirety. This was a whale that still had teeth, but that also carried baleen, as small modifications in its skull reveal.Whale embryos also contain hints that their distant ancestors once bore teeth. They still grow tooth buds that disappear before the young whale is born. Aside from the evidence from fossils and whale embryos, the loss of enamel-capped teeth also left traces in the genomes of modern whales. All reptiles and mammals have genes that produce proteins that mineralize the enamel of teeth. Since baleen whales have no teeth as adults, they have no need for these proteins. Over time such unnecessary genes tend to acquire mutations that impair the protein. This is exactly what happened in baleen whales. In all species of baleen whale, up to three tooth genes turned into pseudogenes (remnants of genes that can no longer produce a functional protein, but are still recognizable as former genes).

But there is something strange about how these mutations are distributed: every species of whale has a different set of mutations. Humpback whales have a mutated enamelin gene, for example. Blue whales carry a different mutation in enamelin. And sei whales have a mutation in a different tooth gene altogether.

Fig.1. The family tree of whales, including extinct relatives. Baleen whales (top) and some pygmy sperm whales (bottom) have mutations in their tooth genes. Every orange symbol denotes a mutation; different letters represent different genes.

Such an uneven distribution of mutations can mean a couple of things. One explanation could be that all the baleen whales lost their enamel independently from each other, due to different

mutations in each lineage. Another possibility is that a hitherto unknown mutation that can be found in all baleen whales is responsible for the loss of enamel. Enamel-covered teeth would have been only lost once by the common ancestor of baleen whales. The fossil evidence supports this scenario: the distribution of toothed baleen whales is not nearly as patchy as the distribution of tooth gene mutations. The MMP20 protein is involved in processing tooth proteins such as enamelin and ameloblastin. A mutation in MMP20 could affect multiple enamel proteins downstream. Moreover, humans and mice that have a defective MMP20 gene develop bad and brittle enamel (amelogenesis imperfecta).In all four whales, a stretch of DNA (a SINE) had inserted itself right inside MMP20, splitting the gene in two. When they extended their search to other species, they found that whale after whale had the same DNA insertion inside MMP20. It is this insertion that rung the death knell for the whales teeth. (Meredith, 2011)

Evolutionary Changes: Macroevolution

The basic theme of whale evolution is the development of large animals from much smaller ancestors--and nowhere is this more evident than in the case of multi-ton sperm and gray whales, whose ultimate forebears were small, dog-sized prehistoric mammals that prowled the riverbeds of central Asia 50 million years ago. Perhaps more intriguingly, whales are also a case study in the gradual evolution of mammals from fully terrestrial to fully marine lifestyles, with corresponding adaptations (elongated bodies, webbed feet, blowholes, etc.) at various key intervals along the way.

All members of the order cetacea are believed to have evolved from terrestrial hoofed mammals like cows, camels and sheep some 45 million years. Throughout their evolution, cetaceans have become perfectly suited to an aquatic environment, and are virtually incapable

of leaving it. Cetaceans illustrate an example of adaptive radiation among mammals. Adaptive radiation allows mammals, as a group, to effectively inhabit the land, the sea, and the air through the development of special adaptations needed to survive in each of these environments. Members of the order cetacea have undergone a number of changes or adaptations needed to fare well in their watery home: their bodies have become streamlined for efficient movement through the water; their forelimbs have been modified into flippers which aid them in steering; their hind limbs have disappeared almost completely; their tail has become broadened horizontally and consists of two large flukes which propel them powerfully through the water by moving up and down, rather than side to side like a fish; in place of hair they have developed a thick layer of fat called blubber under their skin that insulates them from the cold and provides buoyancy ; and the position of their nostrils has shifted to the top of their head creating a blowhole that allows them to effectively come to the surface for air. A whale's blowhole generally reaches the surface before the rest of its body. In addition, a number of other changes have taken place to help whales adapt to life in the sea. Many of these changes are related to the position and abilities of their sensory organs, as life in the water is not the same as life on the land. Sound and light travel differently in water than they do in the air. As a result, whales have developed unique ways of hearing and seeing. Hearing in particular is highly developed in whales, so much so that they depend on it in the same way that we depend on the combination of our eyes, ears and nose to understand the world around. Many of the whale's sensory and reproductive organs have been internalized to reduce drag while swimming. For example, whales do not have external ears, but rely on an internal system of air sinuses and bones to detect sounds. Changes in their reproductive and parental behaviors have also taken place, enabling whales to provide optimum care for their young in the cold, large ocean. Along with these differences, cetaceans do, however, possess many of the same physiological systems - circulatory,

digestive, respiratory, and nervous as the land mammals from which they evolved. For instance, many species possess multi-chambered stomachs even though there is no obvious advantage to having such an arrangement(EarthTrust, 2010) Whales also share brain cells with humans according to a new study published online November 27, 2006 in The Anatomical Record, the official journal of the American Association of Anatomists. The research suggests that "certain cetaceans and hominids may have evolved side by side." Examining brains from various cetaceans, the group of marine mammals that includes whales and dolphins, Patrick R. Hof and Estel Van der Gucht of the Department of Neuroscience at Mount Sinai School of Medicine in New York, NY found substantial variability between the cell structures of the cortex in humpback whales compared to toothed whales like dolphins. The authors say that "these differences may indicate differences in brain function and behavior in aquatic species that are not yet understood," according to a news release from John Wiley & Sons, Inc. The release continues, one feature that stood out in the humpback whale brain was the modular organization of certain cells into "islands" in the cerebral cortex that is also seen in the fin whale and other types of mammals. The authors speculate that this structural feature may have evolved in order to promote fast and efficient communication between neurons. The other notable feature was the presence of spindle cells in the humpback cortex in areas comparable to hominids and in other areas of the whale brain as well. Although the function of spindle neurons is not well understood, they are thought to be involved in cognitive processes and are affected by Alzheimer's disease and other debilitating brain disorders such as autism and schizophrenia. Spindle neurons were also found in the same location in toothed whales with the largest brains, which suggests that they may be related to brain size. The authors note that spindle neurons probably first appeared in the common ancestor of hominids about 15 million years ago, since they are observed in

great apes and humans, but not in lesser apes and other primates; in cetaceans they evolved earlier, possibly as early as 30 million years ago. It is possible that they were present in the ancestors of all cetaceans, but were retained only in those with the largest brains during their evolution. It may also be that they evolved several times independently in the two cetacean suborders; part of this process may have taken place at the same time as they appeared in the ancestor of great apes, which would be a rare case of parallel evolution. "In spite of the relative scarcity of information on many cetacean species, it is important to note, in this context, that sperm whales, killer whales, and certainly humpback whales, exhibit complex social patterns that included intricate communication skills, coalition-formation, cooperation, cultural transmission and tool usage," the authors of the The Anatomical Recordstate. "It is thus likely that some of these abilities are related to comparable histologic complexity in brain organization in cetaceans and in hominids." "Cetacean and primate brains may be considered as evolutionary alternatives in neurobiological complexity and as such, it would be compelling to investigate how many convergent cognitive and behavioral features result from largely dissimilar neocortical organization between the two orders," they conclude. (Hof, et.al. 2006) Cetaceans always excrete urine with an osmolality markedly higher than that of plasma. Although the mechanisms by which cetaceans concentrate urine have not been elucidated, data support a role for medullary urea accumulation in this process, as is the case for terrestrial mammals. Therefore, we hypothesized that facilitated urea transporters are present in the kidney of cetaceans. It was conclude that renal urea transporters are highly conserved among mammals inhabiting terrestrial and pelagic environments. A urea-based concentrating mechanism, presumably evolved to meet the demands of an arid terrestrial environment, may have contributed a fortuitous preadaptation that enabled the ancestors of cetaceans to reinvade the sea. (Janech, 2002)

IV. Fossil Records Fossil cetaceans are classified on the evidence of skeletons (Fig.2.). No other significant body parts preserve, and fossils have not yet produced biomolecules useful in molecular taxonomy. Skulls are by far the most versatile and thus important elements in classification, but teeth and, rarely, other bones (vertebrae, limb elements) have been used at times. A. Modes of Fossil Preservation Fossil cetaceans occur in sedimentary rocks. Originally, remains accumulated in mud, silt, sand, or gravel which, as flesh decayed, was buried and turned to rock through compaction and/or deposition of cementing minerals. Sedimentary rocks are recognized as discrete formations (genetically unified bodies of strata), and are named formally, e.g. the Calvert Formation, Maryland. Marine mammals come from strata including sandstone, mudstone, limestone, greensand, and phosphorite, most of which are marine rocks now exposed on land. Rare fossils have been recovered from the sea floor. Because broadly similar rock types may form at different times and places, sedimentary rocks must be dated to establish their time relationships. (Perrin, et.al, 2008) B. Center of Fossil Distribution The oldest known cetaceans have been recovered from coastal and fluvial deposits in the Indo-Pakistan region of South Asia, dating to approximately 55 Ma, by Philip Gingerich. Later in 1995, Hans Thewissen found Ambolectus. Whales with legs are now known from Pakistan, India, Egypt and USA. Collectively known as Archaeoceti, these early cetaceans had four limbs, heterodont dentitions, and shared basicranial, facial, vertebral, and forelimb characters with modern cetaceans. Archaeocete fossils from the Eocene are also represented in higher temperate latitudes

including Europe, North America, New Zealand, and Antarctica. Later diverging protocetids and basilosaurids were also among archaeocetes for their dispersal throughout the Tethys seaway, producing significant fossil localities throughout present-day Jordan, Egypt, and the southeastern United States. (Estes, 2006) C. Range in Geologic Time Scale The relative timescale has named intervals in an agreed international sequence: Eocene, Oligocene, Miocene, Pliocene, and Pleistocene. These epochs are usually subdivided into early, middle and late. Cetacean diversity in the early Oligocene shows a marked reduction in archaeotes, concurrent with the first appearances of echolating and baleen-bearing cetaceans (odontocetes and mysticetes). Marine rock outcrop from the early Oligocene is generally poor; however, middle to late Oligocene localities in New Zealand, Japan and North America indicate that early odontocetes and mysticetes were quite unlike modern forms. The largest diversification of cetaceans appears to have occurred during the Miocene (23.5-5 Ma) and especially the middle (14.3-11.1 Ma) and late Miocene (11.1-5.1 Ma). (Figure 5). Researchers have long recognized this spike in diversity, although this taxonomic peak may be the result of multiple historiographic, geologic, or taphonomic biases. By the middle Miocene, most representatives of extant families appear, including the mysticete families (Balaenopteridae and Balaenidae) as well as the odontocete families (Delphinidae, Physeteridae, Monodontidae, Phocoenidae, Ziphidae). (Perrin, et.al, 2008)

Fig.5. Changes in the Generic Diversity of Cetaceans

D. Evolutionary Changes Whales and sirenians are the only marine mammals to live their entire lives in water. A thick layer of blubber, rather than hair or fur, insulates them. The hind limbs have been lost and they use the horizontal tail flukes for propulsion. Steering and maintenance of stability when moving is accomplished by a pair of paddle-shaped foreflippers. Estimates of the divergence time for the mysticete-odontocete split differ depending on data and method. According to the fossil record, mysticetes and odontocetes diverged fromacommon archaeocete ancestor about 35 Ma. On the basis of mitochondrial genomic analyses, Arnason et.al, postulated a 35-Ma age for the split between odontocetes and baleen whales in agreement with the fossil record.

Modern whales differ from archaeocetes because they possess a number of derived characters not seen in archaeocetes. Arguably one of the most obvious features is the relationship of the bones in the skull to one another in response to the migration of the nasal openings (blowholes) to the top of the skull. Termed telescoping, the modern whale

skull has premaxillary and maxillary bones that have migrated far posteriorly and presently form most of the skull roof resulting in a long rostrum (beak) and dorsal nasal openings. The occipital bone forms the back of the skull and the nasal, frontal, and parietal bones are sandwiched between the other bones. Another derived feature of modern whales is a fixed elbow joint. The laterally flattened forelimbs are usually short and rigid with an immobile elbow. Archaeocetes have flexible elbow joints, capable of rotation. (Berta, 2006)

V. Adaptations and Evolutionary Novelties Through the course of cetacean evolution, there are major problems that this group has to resolve before they were fully adapted to the aquatic environment. First, they should evolve mechanisms for movement by modifying the parts they acquired from their terrestrial, tetrapod ancestor. Next, they should have an efficient mode of respiration and lastly, an effective mode of reproduction.Cetaceans are considered as the fastest swimmers since they have adapted stream-lined bodies for reducing drag (Reidenberg, 2007). Other changes that allowed them to be efficient swimmers are the loss of their hind limbs and enlargement of their tails. Cetaceans use their large tails called flukes to propel. According to Fish et al, the shape of those flukes was useful in decreasing drag as cetaceans travel quickly underwater. Their shape were also said to be appropriate for decreasing the likelihood of the exterior part

of their flukes to separate from flow (Reidenberg, 2007). According to Gingerich et al. (1994), it is clear that flukes evolved on middle Eocene as seen from fossils of different cetacean families (Fish, 1998). However the progression of the flukes modification is still not definite since direct proof showing their origin are not available (Fish, 1998). Their forelimbs were also modified into flippers. According to Reidenberg (2007), the shape of their flippers also determines their function. He argued that slender and elongated flippers help them to swim quickly, while broad flippers assist them in turning slowly. Whales retained their lungs but they have modified their skull for adaptations to breathing. One evolutionary novelty in this group is the appearance of a blowhole which they use to respire. They changed the configuration of their cranium to shift their external nostrils to the top and back of the skull (Colbert, 1980). According to Colbert (1980), these nostrils can be closed by valves that enabled them to take in a large volume of oxygen so they can stay at depth for a longer period of time before they resurface again to breath for air. They also adapted features for diving. Diving poses difficulty to cetaceans since pressure increases with depth so during a deep dive, their lung volumes collapse (Boyd, 1975; Ridgway and Howard, 1979 as cited by Reidenberg ,2007). Kanatous et al (2002) argued that the shrinking lungs are being hold since the thorax is squeezed due to a collapsible rib (Reidenberg, 2007). Cetacean eyes also adapted features for vision not only underwater but also above water since they still spend time breathing above water level (Mass and Supin, 2007). Mass and Supin (2007) said that these modifications are noticeable in iris and pupil of cetaceans and that sight of cetaceans can be influenced by drastic change in light brightness as they move deeper underwater. As stated by Colbert(1980), embryonic development of cetaceans does not create any problems because they have adapted some modifications for their young to survive in water. They lay their young alive and were fed via mammary glands.

Each of the adaptive features of cetaceans has been very useful in helping the cetaceans to live efficiently in the new environment where they have chosen to exist. They have acquired those adaptive features as they act in response to aquatic life. It will then be not unusual to see other aquatic mammals to possess some of those adaptive features. Their flippers were also evident in other mammals that live primarily in water. Fish et al (1988) said that seals use alternating horizontal sweeps of their hind flippers (Reidenberg, 2007). Flippers of fur seals and sea lions also help them propel by beating their fore flippers (English, 1977; Feldkamp, 1987 as cited by Reidenberg (2007). The hind limbs of sirenians (manatees and dugongs) were also lost so they move forward with the help of their flukes just like the cetaceans (Reidenberg, 2007).Adaptive features are believed to be a beneficial product of natural selection and may have started as a small modification of a specific structure. Those small changes of a certain structure may also come together and lead to formation of novel features (Campbell et al, 1999). According to Campbell et al, exaptation may explain these evolutionary novelties. They said that novel features form step and they have specific functions at a certain stage. Another thing that may also explain the occurrence of evolutionary novelties is the control of certain genes in the development of an organism (Campbell et al, 1999). VI. The Organisms Future Cetaceans are of great importance, not only to humans, but also to the environment. They play a vital role in maintaining the balance within the aquaticecosystem. But due to several predisposing factors, a number of cetacean species are starting to decline. Continuous growth in the volume of carbon dioxide emission through the years leads to worsening problem regarding global warming. Global warming will then result to an increase in global atmospheric temperature every year and thus warming of the waters. Amplification of water temperature would greatly affect all the thriving marine species, thereby disrupting the flow

of energy in the system. This would result to the death of whales in the aquatic environment.The migration pattern of some species of whales will also be altered which would lead to the disturbance in the reproduction of these species. Hotter waters also have lower capacity of holding oxygen. Insufficient oxygen leads to the development of dead zones or areas where no life could thrive (Than, 2009). Growth of organisms like phytoplankton will be affected. Phytoplankton are part of the main diet of baleen whales. Carnivorous toothed whales will also have less food available. The worsening problem on the pollution of the oceans also leads to the death of whales. Chemicals from factories and oil spills from shipping vessels greatly affect the condition of the aquatic environment. Not only whales but other marine life is being poisoned. Despite of the conservation efforts done by different concerned groups whaling is still on spread. Baleen whales are being hunted for whale oil that is being used in production of soaps and grease. Whale meat is being eaten as delicacy in some countries. Bones are being gathered for building materials and fertilizer (Colbert, 1980). Of the 11 genera of Mysticetes, 9 are endangered due to massive hunting. Odontocetes like dolphins and porpoises are being sought after and are used in amusement parks to attract tourists. Most of the cetacean genera are endangered or nearly facing extinction. There are many other factors that affect the proliferation of whales in their natural habitat. Conservation efforts on cetaceans are being pushed by concerned groups like the International Whaling Commission (IWC) and Whale and Dolphin Conservation Society (WDCS) (Michaels, 2007).Cetaceans were also included in the protection list covered by the Marine Mammal Protection Act(MMPA) and Endangered Species Act (ESA). Whales which are endangered are being protected and all attempts to increase their number are being undertaken.

Despite all these conservation efforts, not all aspects related to the preservation of cetaceans are being considered (e.g. global warming and water pollution). Extrapolating from the scenarios, it can be concluded that a million years from now, most genera of cetaceans, if not all, may face extinction due to a drastic change in the condition of the aquatic environment. Increasing effect of global warming can overwhelm marine life leading to mass extinction of the life in the waters.

Literature Cited BBC UK. 2003. Whale Evolution The http://www.bbc.co.uk/dna/h2g2/A1283186 Fossil Evidence. Retrieved from

Bejder, L., B.K. Hall. 2002. Limbs in Whales and Limblessness in other Vertebrates: Mechanisms of Evolutionary and Developmental Transformation and Loss. Evolution & Development 4:6, 445-458 Berta, A., J.L. Sumich, K.M. Kovacs. 2006. Marine Mammals: Evolutionary Biology. USA: Academic Press Campbell N., J. Reece, L. Mitchell. (1999). Biology.5th ed. CA: Addison Wesley Longman, Inc. pp. 458-460 and p. 1126. Colbert, E.H. (1980). Evolution of the Vertebrates: A history of the backboned animals through time. 3rd ed. USA: Wiley-Interscience Publication. pp. 325-332. EarthTrust.org. 2010. Retrieved from http://earthtrust.org/wlcurric/whales.html Estes, J.A. 2006. Whales, Whaling and Ocean Ecosystems. USA: University of California Press. Fish F. (1998). Biochemical Perspective of the Origin of Cetacean Flukes. September 10, 2011 from http://darwin.wcupa.edu/~biology/fish/pubs/pdf/1998FlukeMech.pdf Hoelzel, A.R. 1998. Genetic Structure of Cetacean Populations in Sympatry, Parapatry, and Mixed Assemblages: Implications for Conservation Policy. The American Genetic Association. 89:451-458 Hof, P.R., E.V. Gucht. 2006. The Structure of the Cerebral Cortex of the Humpback Whale, Megapteranovaeangliae (Cetacea, Mysticeti, Balaenopteridae).The Anatomical Record. 290:1-31 Janech, M.G., R. Chen, J. Klein, M.W. Nowak, W. McFee, R.V. Paul, W.R. Fitzgibbon, D.W. Ploth. 2002. Molecular and Functional Characterization of a Urea Transporter from the Kidney of a Short-Finned Pilot Whale. Am J Physiol Renal Physiol: 2003,284 (5): F996-F1005 Kennedy, J. 2011. Cetaceans- Whales, Dolphins and Porpoises. Retrieved from http://marinelife.about.com/od/marinelifeprofiles/p/cetaceans.htm Luskin, C. 2010. Of Whale and Feather Evolution: Natures Two Macroevolutionary Lumps of Coal. Retrieved from http://www.evolutionnews.org/2010/08/of_whale_and_feather_evolution037221.html Mass, A., A. Supin. (2007). Adaptive Feature of Aquatic Mammals Eye. The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology, 290(6), 701-715 doi: 10.1002/ar.20529

Meredith, R.W., J. Gatesy, J. Cheng, M.S. Springer. 2011. Pseudogenization of the tooth gene enamelysin (MMP20) in the Common Ancestor of Extant Baleen Whales. Proceedings. Biological Sciences / The Royal Society. 278 (1708), 993-1002 Michaels, P.A. (2007). Commercial Whaling and the International Whaling Commission. Retrieved from http://greennature.com/article16.html Milinkovitch, Michel C. and Olivier Lambert. 2006. Cetacea. Whales, dolphins, and porpoises. Version 07 August 2006 (under construction). http://tolweb.org/Cetacea/15977/2006.08.07in The Tree of Life Web Project, http://tolweb.org/ Perrin, W.F., B.G. Wursig, J.G.M. Thewissen. 2008. Encyclopedia of Marine Mammals. USA: Academic Press. Reidenberg, J. (2007). Anatomical Adaptations of Aquatic Mammals. The Anatomical Record: Advances in Integrative Anatomy and Evolutionary Biology, 290(6), 507-513 doi:10.1002/ar.20541 Strauss, B. Prehistoric Whales The Story of Whale Evolution. About.com Guide. Retrieved from http://dinosaurs.about.com/od/otherprehistoriclife/a/ancientwhales.htm Thewissen, J.G.M. 1998. The Emergence of Whales: Evolutionary Patterns in the Origin of Cetacea. USA: Springer Than, K. (2009). Global Warming to Create Permanent Ocean Dead Zones? Retrieved from http://news.nationalgeographic.com/news/2009/01/090128-ocean-deadzones.html University of California Museum of Paleontology. (2008). The Evolution of Whales [Photograph]. Retrieved September 10, 2011, from: http://evolution.berkeley.edu/evolibrary/article/evograms_03 Wikipedia: The Free Encyclopedia. Evolution of Cetaceans. http://en.wikipedia.org/wiki/Evolution_of_cetaceans Retrieved from

Fig.2. Cetacean Fossils

Fig.3. The cladogram of cetaceans.

Fig.4. Cetacean Morphological and Trophic Evolution