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Most authors have identified two rapid increases in relative brain size (encephalization quotient, EQ) in cetacean evolution: first at the origin of the modern suborders (odontocetes and mysticetes) around the Eocene-Oligocene transition, and a second at the origin of the delphinoid odontocetes during the middle Miocene. We explore how methods used to estimate brain and body mass alter this perceived timing and rate of cetacean EQ evolution. We provide new data on modern mammals (mysticetes, odontocetes, and terrestrial artiodactyls) and show that brain mass and endocranial volume scale allometrically, and that endocranial volume is not a direct proxy for brain mass. We demonstrate that inconsistencies in the methods used to estimate body size across the Eocene-Oligocene boundary have caused a spurious pattern in earlier relative brain size studies. Instead, we employ a single method, using occipital condyle width as a skeletal proxy for body mass using a new dataset of extant cetaceans, to clarify this pattern. We suggest that cetacean relative brain size is most accurately portrayed using EQs based on the scaling coefficients as observed in the closely related terrestrial artiodactyls. Finally, we include additional data for an Eocene whale, raising the sample size of Eocene archaeocetes to seven. Our analysis of fossil cetacean EQ is different from previous works which had shown that a sudden increase in EQ coincided with the origin of odontocetes at the Eocene-Oligocene boundary. Instead, our data show that brain size increased at the origin of basilosaurids, 5 million years before the Eocene-Oligocene transition, and we do not observe a significant increase in relative brain size at the origin of odontocetes., Competing Interests: The authors have declared that no competing interests exist.

Studying ontogeny in both extant and extinct species can unravel the mechanisms underlying mammal diversification and specialization. Among mammalian clades, Cetartiodactyla encompass species with a wide range of adaptations, and ontogenetic evidence could clarify longstanding debates on the origins of modern specialized families. Here, we study the evolution of dental eruption patterns in early diverging cetartiodactyls to assess the ecological and biological significance of this character and shed new light on phylogenetic issues. After investigation of the ontogenetic dental series of 63 extinct genera, our parsimony reconstructions of eruption state evolution suggest that the eruption of molars before permanent premolars represents a plesiomorphic condition within Cetartiodactyla. This result substantially differs from a previous study based on modern species only. As a result, the presence of this pattern in most ruminants might represent an ancestral condition contributing to their specialized herbivory, rather than an original adaptation. In contrast, the late eruption of molars in hippopotamoids is more likely related to biological aspects, such as increases in body mass and slower pace of life. Our study mainly shows that eruption sequences reliably characterize higher level cetartiodactyl taxa and could represent a new source of phylogenetic characters, especially to disentangle the origin of hippopotamoids and cetaceans.

Von Economo neurons (VENs) are specialized projection neurons with a characteristic spindle-shaped soma and thick basal and apical dendrites. VENs have been described in restricted cortical regions, with their most frequent appearance in layers III and V of the anterior cingulate cortex, anterior insula, and frontopolar cortex of humans, great apes, macaque monkeys, elephants, and some cetaceans. Recently, a ubiquitous distribution of VENs was reported in various cortical areas in the pygmy hippopotamus, one of the closest living relatives of cetaceans. That finding suggested that VENs might not be unique to only a few species that possess enlarged brains. In the present analysis, we assessed the phylogenetic distribution of VENs within species representative of the superordinal clade that includes cetartiodactyls and perissodactyls, as well as afrotherians. In addition, the distribution of fork cells that are often found in close proximity to VENs was also assessed. Nissl-stained sections from the frontal pole, anterior cingulate cortex, anterior insula, and occipital pole of bowhead whale, cow, sheep, deer, horse, pig, rock hyrax, and human were examined using stereologic methods to quantify VENs and fork cells within layer V of all four cortical regions. VENs and fork cells were found in each of the species examined here with species-specific differences in distributions and densities. The present results demonstrated that VENs and fork cells were not restricted to highly encephalized or socially complex species, and their repeated emergence among distantly related species seems to represent convergent evolution of specialized pyramidal neurons. The widespread phylogenetic presence of VENs and fork cells indicates that these neuron morphologies readily emerged in response to selective forces,whose variety and nature are yet to be identified.

Odontocetes and mysticetes are two extant suborders of cetaceans. It is reported that the former have no sense of olfaction, while the latter can smell in air. To explain the ecological reason why mysticetes still retain their sense of smell, two hypotheses have been proposed - the echolocation-priority hypothesis, which assumes that the acquisition of echolocation causes the reduction of the importance of olfaction, and the filter-feeder hypothesis, which assumes that olfactory ability is important for filter-feeders to locate their prey because clouds of plankton give off a peculiar odor. The olfactory marker protein (OMP) is almost exclusively expressed in vertebrate olfactory receptor neurons, and is considered to play important roles in olfactory systems. In this study, full-length open reading frames of OMP genes were identified in 6 cetacean species and we analyzed the nonsynonymous to synonymous substitution rate ratio based on the maximum likelihood method. The evolutionary changes of the selective pressures on OMP genes did fit better to the filter-feeder hypothesis than to the echolocation-priority hypothesis. In addition, no pseudogenization mutations are found in all five odontocetes OMP genes investigated in this study. It may suggest that OMP retains some function even in 'anosmic' odontocetes., (Copyright © 2011 Elsevier B.V. All rights reserved.)

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.

Modern members of the mammalian order Cetacea (whales, dolphins and porpoises) are obligate aquatic swimmers that are highly distinctive in morphology, lacking hair and hind limbs, and having flippers, flukes, and a streamlined body. Eocene fossils document much of cetaceans' land-to-water transition, but, until now, the most primitive representative for which a skeleton was known was clearly amphibious and lived in coastal environments. Here we report on the skeletons of two early Eocene pakicetid cetaceans, the fox-sized Ichthyolestes pinfoldi, and the wolf-sized Pakicetus attocki. Their skeletons also elucidate the relationships of cetaceans to other mammals. Morphological cladistic analyses have shown cetaceans to be most closely related to one or more mesonychians, a group of extinct, archaic ungulates, but molecular analyses have indicated that they are the sister group to hippopotamids. Our cladistic analysis indicates that cetaceans are more closely related to artiodactyls than to any mesonychian. Cetaceans are not the sister group to (any) mesonychians, nor to hippopotamids. Our analysis stops short of identifying any particular artiodactyl family as the cetacean sister group and supports monophyly of artiodactyls.

Recent molecular studies are inconsistent with ungulate phylogenetic trees that are based on morphological traits. These inconsistencies especially relate to the position of cetaceans and perissodactyls. Evaluation of the close phylogenetic ties between artiodactyls and cetaceans has been hampered by the absence of tarsal bones of primitive cetaceans, as artiodactyls are often diagnosed on the basis of their tarsus. We here describe newly discovered tarsal bones that are the oldest cetacean tarsals known. We present a character analysis for primitive ungulate tarsals and evaluate their impact on the ungulate phylogenetic tree. Tarsal data are consistent with some molecular studies in suggesting that the extant sister group of Cetacea is Artiodactyla or that Cetacea should be included within the latter order. Tarsal data do not support Cete (Mesonychia plus Cetacea) and are consistent with the exclusion of perissodactyls from paenungulates as suggested by some molecular studies.

The embryology of cetaceans documents features that elucidate the interaction between form and function in modern species, as well as their evolution from extinct ones. Prenatal specimens of the dolphin Stenella attenuata demonstrate critical aspects of the embryonic and fetal development of cetaceans, the pharyngeal arches, ear ossicles, nasal opening, teeth, fore limb, hind limb, and flukes. In addition, specimens of the bowhead whale (Balaena mysticetus) illustrate the development of baleen and hair. [ABSTRACT FROM AUTHOR]

Abstract: Tympanic bullae and baleen plates from bowhead whales of the Western Arctic population were examined. Growth layer groups (GLGs) in the involucrum of the tympanic bone were used to estimate age of the whales, and compared to stable isotope signatures along transects of baleen plates and the involucrum. The involucrum of the tympanic bone consists of three regions that form in utero, during nursing in the first year, and during the first decades of life, respectively. Life history events, such as annual migration, are recorded in the bowhead tympanic bulla. It is likely that bone growth in the bowhead tympanic occurs during periods of high food intake, while slow or arrested growth occurs during periods of low food intake. Comparisons between numbers of GLGs in the tympanic, number of isotopic oscillations in a baleen plate, length of the baleen plate, and total whale length show correlation coefficients as high as 0.97. The tympanic GLG method is particularly useful for estimating the age of whales up to 20 yr old. [ABSTRACT FROM AUTHOR]

In utero, baleen whales initiate the development of several dozens of teeth in upper and lower jaws. These tooth germs reach the bell stage and are sometimes mineralized, but toward the end of prenatal life they are resorbed and no trace remains after birth. Around the time that the germs disappear, the keratinous baleen plates start to form in the upper jaw, and these form the food-collecting mechanism. Baleen whale ancestors had two generations of teeth and never developed baleen, and the prenatal teeth of modern fetuses are usually interpreted as an evolutionary leftover. We investigated the development of teeth and baleen in bowhead whale fetuses using histological and immunohistochemical evidence. We found that upper and lower dentition initially follow similar developmental pathways. As development proceeds, upper and lower tooth germs diverge developmentally. Lower tooth germs differ along the length of the jaw, reminiscent of a heterodont dentition of cetacean ancestors, and lingual processes of the dental lamina represent initiation of tooth bud formation of replacement teeth. Upper tooth germs remain homodont and there is no evidence of a secondary dentition. After these germs disappear, the oral epithelium thickens to form the baleen plates, and the protein FGF-4 displays a signaling pattern reminiscent of baleen plates. In laboratory mammals, FGF-4 is not involved in the formation of hair or palatal rugae, but it is involved in tooth development. This leads us to propose that the signaling cascade that forms teeth in most mammals has been exapted to be involved in baleen plate ontogeny in mysticetes. [ABSTRACT FROM AUTHOR]

The processes of lipid deposition and utilization, via the gene leptin ( Lep), are poorly understood in taxa with varying degrees of adipose storage. This study examines how these systems may have adapted in marine aquatic environments inhabited by cetaceans. Bowhead ( Balaena mysticetus) and beluga whales ( Delphinapterus leucas) are ideal study animals-they possess large subcutaneous adipose stores (blubber) and undergo bi-annual migrations concurrent with variations in food availability. To answer long-standing questions regarding how (or if) energy and lipid utilization adapted to aquatic stressors, we quantified variations in gene transcripts critical to lipid metabolism related to season, age, and blubber depth. We predicted leptin tertiary structure conservation and assessed inter-specific variations in Lep transcript numbers between bowheads and other mammals. Our study is the first to identify seasonal and age-related variations in Lep and lipolysis in these cetaceans. While Lep transcripts and protein oscillate with season in adult bowheads reminiscent of hibernating mammals, transcript levels reach up to 10 times higher in bowheads than any other mammal. Data from immature bowheads are consistent with the hypothesis that short baleen inhibits efficient feeding. Lipolysis transcripts also indicate young Fall bowheads and those sampled during Spring months limit energy utilization. These novel data from rarely examined species expand the existing knowledge and offer unique insight into how the regulation of Lep and lipolysis has adapted to permit seasonal deposition and maintain vital blubber stores. [ABSTRACT FROM AUTHOR]

Cetaceans are obligate aquatic mammals derived from terrestrial artiodactyls. The defining characteristic of cetaceans is a thick and dense lip (pachyosteosclerotic involucrum) of an ear bone (the tympanic). This unique feature is absent in modern terrestrial artiodactyls and is suggested to be important in underwater hearing. Here, we investigate the mineralogical and biochemical properties of the involucrum, as these may hold clues to the aquatic adaptations of cetaceans. We compared bioapatites (enamel, dentine, cementum, and skeletal bone) of cetaceans with those of terrestrial artiodactyls and pachyosteosclerotic ribs of manatees (Sirenia). We investigated organic, carbonate, and mineral composition as well as crystal size and crystallinity index. In all studied variables, bioapatites of the cetacean involucrum were intermediate in composition and structure between those of tooth enamel on the one hand and those of dentine, cementum, and skeletal bone on the other. We also studied the amino acid composition of the cetacean involucrum relative to that of other skeletal bone. The central involucrum had low glycine and hydroxyproline concentrations but high concentrations of nonessential amino acids, unlike most bone samples but similar to the tympanic of hippos and the (pachyosteosclerotic) ribs of manatees. These amino acid results are evidence of rapid bone development. We hypothesize that the mineralogical and amino acid composition of cetacean bullae differs from that of other bone because of (1) functional modifications for underwater sound reception and (2) structural adaptations related to rapid ossification. [ABSTRACT FROM AUTHOR]

The family Remingtonocetidae is a basal family of Eocene cetaceans only known from near shore marine environments of India and Pakistan. We describe a new skull for Remingionocetus harudiensis which elucidates the anatomy and functional morphology of the head and provides new details on cranial cavity and nasopharyngeal region. We suggest that Remingtonocetus was an ambush predator that hunted from a perch on the ocean floor, and that hearing was its most important sense. We speculate that the greatly elongated rostrum is an adaptation for water retention because these are some of the earliest whales living in seawater. [ABSTRACT FROM AUTHOR]

The article presents a description of two new species of fossil cetaceans, the Pakicetus calcis and the Pakicetus chittas, which was collected from the Shepherd's Lake locality of the Kuldana Formation of northern Pakistan. It notes that these pakicetid dentitions are the most complete as of December 2009. Information with regards to the dentitions of the pakicetid Nalacetus ratimitus, the remingtonocetid Attockicetus and the holotype of Ambulocetus natans is offered. It notes that archaic Eocene whales (archaeocetes) are included in the fossil relatives of modern cetaceans (whales, dolphins, and porpoises).

The Eocene cetacean genera Andrewsiphius and Kutchicetus are systematically revised, their anatomy described, and their phylogenetic position analyzed. Each genus contains a single species, A. sloani and K minimus, and both are known only from the middle Eocene of the Indian, Subcontinent. Andrewsiphius and Kutchicetus differ in a number of respects, the most important dental difference being that P2, P3, p2, and p3 are double-rooted in Andrewsiphius and single-rooted in Kutchicetus. Lower molars are separated by diastemata in Kutchicetus, but not in Andrewsiphius. Postcranially, Andrewsiphius has caudal vertebrae that are far more robust than those of Kutchicetus. We propose the new clade Andrewsiphiinae for these two genera, based on their unique characters: the extremely slender jaw, fused mandibular symphysis, narrow palate and rostrum, and lower molars that have a low crown with three cusps lined up rostro-caudally. A phylogenetic analysis indicates that andrewsiphiines are either a subfamily of Remingtonocetidae or an independent branch on the Eocene cetacean lineage. Interpreting conservatively, we classify them as remingtonocetids. Andrewsiphiines have a long, robust, dorso-ventrally flattened tail and short limbs, suggesting that they swam using dorsoventral undulation of the tail. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Introduces the families of archaic cetaceans that lived in the Eocene, discusses the organ systems which underwent change, and puts the functional morphology and evolution of the locomotion and osmoregulation organ systems in a broader perspective. View that the origin of whales offers an example of evolutionary change; Opposition to the theory by creationists; Topic of adaptations of early cetaceans.

A correction to this paper has been published: https://doi.org/10.1007/s10914-021-09554-9 [ABSTRACT FROM AUTHOR]

During the land-to-water transition in the Eocene epoch, the cetacean skeleton underwent modifications to accommodate life in the seas. These changes are well-documented in the fossil record. The forelimb transformed from a weight-bearing limb with mobile joints to a flipper with an immobile carpus. We used micro-CT imaging to assess evolutionary changes in carpal size, orientation, and articulation within Eocene cetacean taxa associated with the transition from a terrestrial to amphibious niche. We compared Ambulocetus natans, a well-preserved amphibious archaeocete, with other archaeocetes, and with Eocene terrestrial artiodactyls, the sister group to Cetacea. A cylindrical carpus in terrestrial taxa evolved into a mediolaterally flattened, cambered carpus in the semi-aquatic and fully aquatic cetaceans. Specifically, the pisiform bone shifted from a ventral orientation in terrestrial taxa to a lateral orientation, in plane with the carpus, within semi-aquatic and fully aquatic taxa. Flattening of the carpus, including lateral rotation of the pisiform, likely relates to functional shifts from weight-bearing terrestrial locomotion to aquatic locomotion. This laterally projecting pisiform morphology is retained in all extant cetaceans. Our results suggest this shift, along with other modifications to the carpus, predominantly occurred during the middle Eocene and facilitated an obligatorily aquatic lifestyle in late Eocene cetaceans. [ABSTRACT FROM AUTHOR]