The Evolution of Cetaceans: Evolutionary phylogeny, fossil records and evolutionary novelties I.

Main Groups of Cetaceans Cetaceans: From Latin cetus = a large sea animal and Greek ketos = sea monster Uncharacteristic of the Class Mammalia due to modifications of the basic mammalian structure adaptation to transition from terrestrial to aquatic habitat Isolated from primitive eutherian ancestors leads to rapid evolutionary changes Include whales, dolphins and porpoises in about 87 species, 14 families and 2 sub-orders

Kingdom Animalia Class Mammalia Order Cetacea Suborder Odontoceti Suborder Mysticeti Suborder Odontoceti toothed whales with teeth for feeding and a single blowhole for respiration asymmetrical skull include more species and are smaller than the other subgroup Suborder Mysticeti baleen whales with baleen plates for filter feeding and to blowholes for respiration symmetrical skull

II. Unifying Characteristics/Features warm-blooded no hairs in the body uses blubber, a heavy fat layer underneath the skin, for insulating or warming of the body torpedo shaped body with no distinct neck streamlined body and horizontal end fin (flukes) for efficient swimming paddles for balancing their bodies blowholes for exchange of gases poor sense of smell good sense of hearing for sensing vibrations underwater for echolocation o Echolocation allows cetaceans to review their environment by emitting sounds and listening to echoes. It enables them to detect

preys, predators and obstacles and also for communicating with other whales highly developed brains give birth to live and well-developed young and nourish their offspring through their mammary glands

III. Evolutionary Changes Microevolution: Transition to toothlessness baleen plates of mysticetes evolved gradually through time early baleen whales such as Janjucetus and Mammalodon still had fully developed enamel-covered teeth Eomysticetus had already exchanged its teeth for baleen plates Aetiocetus has both teeth and baleen as evident in its skull whale embryos also show that their distant ancestors once bore teeth. They still grow tooth buds that disappear before the young whale is born. through time, proteins needed to encode enamel on teeth of whales are impaired via mutation all species of baleen whales have a different set of at least three tooth genes that mutated into pseudogenes (remnants of genes that can no longer produce a functional protein, but are still recognizable as former genes) Example: Humpback whales, blue whales, and sei whales have different mutations in the tooth gene enamelin Possible explanations: baleen whales lost enamel independent from each other or their common ancestor lost enamel once Evidence: fossil records show a modification in the MMP20 gene (gene involved in processing tooth proteins such as enamelin and ameloblastin) which lead to the permanent lost of teeth of baleen whales o a stretch of DNA (a SINE) had inserted itself right inside MMP20, splitting the gene in two

Macroevolution and Evolutionary Novelties gradual evolution from fully terrestrial to fully marine lifestyle with corresponding adaptations (elongated bodies, webbed feet, blowholes, etc.) at various key intervals along the way order cetacea are believed to have evolved from smaller terrestrial hoofed mammals like cows, camels and sheep some 45 million years they are a good example of adaptive radiation in 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 Their body became streamlined; forelimbs modified into flippers; limbs lost; tail broadened horizontally all for efficient motility in water

blubbers, in place of hair, for buoyancy and insulation nostrils shifted to the top of the head to create blowholes for effective respiration when they reach the surface of the water their sensory organs have been modified to adapt to the differences in the life on land and the life on sea hearing is highly developed sensory and reproductive organs have been internalized to reduce drag while swimming (e.g. 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 Since renal urea transporters are highly conserved among mammals inhabiting terrestrial and pelagic environments, urea-based concentrating mechanism, presumably evolved to meet the demands of an arid terrestrial environment, may have contributed a unexpected preadaptation that enabled the ancestors of cetaceans to reinvade the sea

Parallel Evolution: whale and Hominid brain whales share brain cells with humans (American Association of Anatomists, 2006) suggesting that certain cetaceans and hominids may have evolved side by side there is substantial variability between the cell structures of the cortex in humpback whales compared to toothed whales like dolphins modular organization of certain cells into "islands" in the cerebral cortex of humpback whales promotes fast and efficient communication between neurons spindle cells in the humpback cortex and in other areas of the whale brain is comparable to that of the spindle cells of hominids. Spindle cells 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 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

IV. Fossil Records fossil cetaceans are classified based on skeletons (other fossil parts have not yet produced biomolecules useful in molecular taxonomy) teeth and, rarely, other bones (vertebrae, limb elements) have been used at times

A. Modes of fossil preservation fossil cetaceans occur in sedimentary rocks 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

B. Center of Fossil Distribution oldest known cetaceans from coastal and fluvial deposits in the IndoPakistan region of South Asia, dating to approximately 55 Ma, by Philip Gingerich Hans Thewissen found Ambolectus later in 1995 Archaeoceti Early cetacean fossils with four limbs, heterodont dentitions, and shared basicranial, facial, vertebral, and forelimb characters with modern cetaceans from Pakistan, India, Egypt and USA later separated into protocetids and basilosaurids and then dispersed Jordan, Egypt, and the southeastern United States

C. Range in Geologic Time Scale Eocene, Oligocene, Miocene, Pliocene, and Pleistocene the early Oligocene reduction in archaeotes and first appearances of echolating and baleen-bearing cetaceans (odontocetes and mysticetes) Miocene diversification of cetaceans middle Miocene most representatives of extant families appear

VI. The Organisms Future increasing volume of CO2 emissions through the years Global Warming increase in temp of the waters o alterations in the migration pattern of some species of whales o lower capacity of holding O2 dead zones o decrease in the population of phytoplankton main diet of baleen whales water pollution 11 of 9 genera of mysticetes are endangered odontocetes are being hunted for tourism (e.g. amusement parks) insufficient conservation efforts (e.g. IWC) due to unstoppable problem in whaling After a million of years EXTINCTION!