J. G. M. Thewissen

Whales originated from aquatic artiodactyls in the Eocene epoch of India

Although the first ten million years of whale evolution are documented by a remarkable series of fossil skeletons, the link to the ancestor of cetaceans has been missing. It was known that whales are related to even-toed ungulates (artiodactyls), but until now no artiodactyls were morphologically close to early whales. Here we show that the Eocene south Asian raoellid artiodactyls are the sister group to whales. The raoellid Indohyus is similar to whales, and unlike other artiodactyls, in the structure of its ears and premolars, in the density of its limb bones and in the stable-oxygen-isotope composition of its teeth. We also show that a major dietary change occurred during the transition from artiodactyls to whales and that raoellids were aquatic waders. This indicates that aquatic life in this lineage occurred before the origin of the order Cetacea.

Vestibular evidence for the evolution of aquatic behaviour in early cetaceans

Early cetaceans evolved from terrestrial quadrupeds to obligate swimmers, a change that is traditionally studied by functional analysis of the postcranial skeleton. Here we assess the evolution of cetacean locomotor behaviour from an independent perspective by looking at the semicircular canal system, one of the main sense organs involved in neural control of locomotion. Extant cetaceans are found to be unique in that their canal arc size, corrected for body mass, is approximately three times smaller than in other mammals. This reduces the sensitivity of the canal system, most plausibly to match the fast body rotations that characterize cetacean behaviour. Eocene fossils show that the new sensory regime, incompatible with terrestrial competence, developed quickly and early in cetacean evolution, as soon as the taxa are associated with marine environments. Dedicated agile swimming of cetaceans thus appeared to have originated as a rapid and fundamental shift in locomotion rather than as the gradual transition suggested by postcranial evidence. We hypothesize that the unparalleled modification of the semicircular canal system represented a key ‘point of no return’ event in early cetacean evolution, leading to full independence from life on land.

Morphoregulation of teeth: modulating the number, size, shape and differentiation by tuning Bmp activity

Summary During development and evolution, the morphology of ectodermal organs can be modulated so that an organism can adapt to different environments. We have proposed that morphoregulation can be achieved by simply tilting the balance of molecular activity. We test the principles by analyzing the effects of partial downregulation of Bmp signaling in oral and dental epithelia of the keratin 14‐Noggin transgenic mouse. We observed a wide spectrum of tooth phenotypes. The dental formula changed from 1.0.0.3/1.0.0.3 to 1.0.0.2(1)/1.0.0.0. All mandibular and M3 maxillary molars were selectively lost because of the developmental block at the early bud stage. First and second maxillary molars were reduced in size, exhibited altered crown patterns, and failed to form multiple roots. In these mice, incisors were not transformed into molars. Histogenesis and differentiation of ameloblasts and odontoblasts in molars and incisors were abnormal. Lack of enamel caused misocclusion of incisors, leading to deformation and enlargement in size. Therefore, subtle differences in the level, distribution, and timing of signaling molecules can have major morphoregulatory consequences. Modulation of Bmp signaling exemplifies morphoregulation hypothesis: simple alteration of key signaling pathways can be used to transform a prototypical conical‐shaped tooth into one with complex morphology. The involvement of related pathways and the implication of morphoregulation in tooth evolution are discussed.

Fossil Evidence for the Origin of Aquatic Locomotion in Archaeocete Whales

Recent members of the order Cetacea (whales, dolphins, and porpoises) move in the water by vertical tail beats and cannot locomote on land. Their hindlimbs are not visible externally and the bones are reduced to one or a few splints that commonly lack joints. However, cetaceans originated from four-legged land mammals that used their limbs for locomotion and were probably apt runners. Because there are no relatively complete limbs for archaic archaeocete cetaceans, it is not known how the transition in locomotory organs from land to water occurred. Recovery of a skeleton of an early fossil cetacean from the Kuldana Formation, Pakistan, documents transitional modes of locomotion, and allows hypotheses concerning swimming in early cetaceans to be tested. The fossil indicates that archaic whales swam by undulating their vertebral column, thus forcing their feet up and down in a way similar to modern otters. Their movements on land probably resembled those of sea lions to some degree, and involved protraction and retraction of the abducted limbs.

Phylogenetic aspects of Cetacean origins: A morphological perspective

The evolutionary steps leading up to the origin of cetaceans involved pervasive changes in the masticatory apparatus, the ear, and limb morphology. These changes bear heavily on the phylogenetic relationships of Cetacea, and are investigated here on the basis of two of its earliest members:Pakicetus andAmbulocetus. A phylogenetic analysis of cetaceans, five groups of mesonychians, and five other groups of ungulates indicates thatPakicetus is the sister group to all other cetaceans, and that Cete (mesonychians and Cetacea) is a monophyletic group.

Locomotor evolution in the earliest cetaceans; functional model, modern analogues, and paleontological evidence

We discuss a model for the origin of cetacean swimming that is based on hydrodynamic and kinematic data of modern mammalian swimmers. The model suggests that modern otters (Mustelidae: Lutrinae) display several of the locomotor modes that early cetaceans used at different stages in the transition from land to water. We use mustelids and other amphibious mammals to analyze the morphology of the Eocene cetacean Ambulocetus natans , and we conclude that Ambulocetus may have locomoted by a combination of pelvic paddling and dorsoventral undulations of the tail, and that its locomotor mode in water resembled that of the modern otter Lutra most closely. We also suggest that cetacean locomotion may have resembled that of the freshwater otter Pteronura at a stage beyond Ambulocetus.

Early Eocene warming events and the timing of terrestrial faunal exchange between India and Asia

The timing of initiation of continent-continent collision between Asia and India is controversial, but this major tectonic event is generally thought to have occurred in the Early Eocene, ca. 50 Ma. New and independent data from strontium isotopes, stable carbon isotopes, microfossil biostratigraphy, and mammal fossils from an Early Eocene marginal marine sequence (Cambay Shale) at the Vastan Lignite Mine of western India indicate that terrestrial faunal exchanges, and therefore continental collision, between Asia and the Indian subcontinent took place before 53.7 Ma. This age coincides with the second Eocene Thermal Maximum (ETM2), a short-lived warming episode that followed the Paleocene-Eocene Thermal Maximum (PETM) ca. 55.5 Ma. Our data also document, for the first time, a clear record of the ETM2 in terrestrial organic material from a low-latitude site, which is represented by a 3‰−4‰ carbon isotope excursion (CIE) in lignite and dispersed organic carbon δ13C values. The magnitude of the CIE at this location closely matches that observed in marine cores from the Arctic Ocean, which supports an interpretation that this hyperthermal event, though of lower magnitude, was similar in character to that of the PETM, being a global phenomenon that affected both terrestrial and marine ecosystems.

The role of phenacodontids in the origin of the modern orders of ungulate mammals

ABSTRACT We investigate the role of the ancestral ungulates Phenacodontidae in the evolution of Perissodactyla, Hyracoidea, Proboscidea, Sirenia, and Desmostylia on the basis of parsimony analysis of dental, cranial, and postcranial data. The monophyletic order Phenacodonta (Phenacodontidae and Meniscotheriidae) is the sistergroup to the mirorder Pantomesaxonia (consisting of the five mammalian orders mentioned), and these two clades are united as superorder Paenungulata. The relationships of taxa within Pantomesaxonia remain unresolved. Important characters of the ancestor of Pantomesaxonia include partially molarized premolars and a cursorial limb skeleton.

From Land to Water: the Origin of Whales, Dolphins, and Porpoises

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.

Eocene evolution of whale hearing

The origin of whales (order Cetacea) is one of the best-documented examples of macroevolutionary change in vertebrates. As the earliest whales became obligately marine, all of their organ systems adapted to the new environment. The fossil record indicates that this evolutionary transition took less than 15 million years, and that different organ systems followed different evolutionary trajectories. Here we document the evolutionary changes that took place in the sound transmission mechanism of the outer and middle ear in early whales. Sound transmission mechanisms change early on in whale evolution and pass through a stage (in pakicetids) in which hearing in both air and water is unsophisticated. This intermediate stage is soon abandoned and is replaced (in remingtonocetids and protocetids) by a sound transmission mechanism similar to that in modern toothed whales. The mechanism of these fossil whales lacks sophistication, and still retains some of the key elements that land mammals use to hear airborne sound.