Lisa Noelle Cooper

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.

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.

Evolution of hyperphalangy and digit reduction in the cetacean manus

Cetaceans (whales, dolphins, and porpoises) have a soft tissue flipper that encases most of the forelimb, and elongated digits with an increased number of phalanges (hyperphalangy). In addition, some cetaceans exhibit a reduction in digit number. Although toothed cetaceans (odontocetes) are pentadactylous, most baleen whales (mysticetes) are tetradactylous and also lack a metacarpal. This study conducts a survey of cetacean metacarpal and phalangeal morphologies, traces the evolution of hyperphalangy in a phylogenetic context, optimizes characters onto previously published cetacean phylogenies, and tests various digit loss hypotheses. Dissections were performed on 16 cetacean flippers representing 10 species (8 mysticetes, 2 odontocetes). Phalangeal count data were derived from forelimb radiographs (36 odontocetes, 5 mysticetes), osteological specimens of articulated forelimbs (8 mysticetes), and were supplemented with published counts. Modal phalangeal counts were coded as ordered and unpolarized characters and optimized onto two known cetacean phylogenies. Results indicate that digital ray I is reduced in many cetaceans (except Globicephala) and all elements of digital ray I were lost in tetradactylous mysticetes. Fossil evidence indicates this ray may have been lost approximately 14 Ma. Most odontocetes also reduce the number of phalangeal elements in digit V, while mysticetes typically retain the plesiomorphic condition of three phalanges. Results from modal phalangeal counts show the greatest degree of hyperphalangy in digits II and III in odontocetes and digits III and IV in mysticetes. Fossil evidence indicates cetacean hyperphalangy evolved by at least 7–8 Ma. Digit loss and digit positioning may underlie disparate flipper shapes, with narrow, elongate flippers facilitating fast swimming and broad flippers aiding slow turns. Hyperphalangy may help distribute leading edge forces, and multiple interphalangeal joints may smooth leading edge flipper contour. Anat Rec, 290:654–672, 2007.

Anthracobunids from the middle eocene of India and pakistan are stem perissodactyls.

Anthracobunidae is an Eocene family of large mammals from south Asia that is commonly considered to be part of the radiation that gave rise to elephants (proboscideans) and sea cows (sirenians). We describe a new collection of anthracobunid fossils from Middle Eocene rocks of Indo-Pakistan that more than doubles the number of known anthracobunid fossils and challenges their putative relationships, instead implying that they are stem perissodactyls. Cranial, dental, and postcranial elements allow a revision of species and the recognition of a new anthracobunid genus. Analyses of stable isotopes and long bone geometry together suggest that most anthracobunids fed on land, but spent a considerable amount of time near water. This new evidence expands our understanding of stem perissodactyl diversity and sheds new light on perissodactyl origins.

Neuromuscular anatomy and evolution of the cetacean forelimb.

The forelimb of cetaceans (whales, dolphins, and porpoises) has been radically modified during the limb‐to‐flipper transition. Extant cetaceans have a soft tissue flipper encasing the manus and acting as a hydrofoil to generate lift. The neuromuscular anatomy that controls flipper movement, however, is poorly understood. This study documents flipper neuromuscular anatomy and tests the hypothesis that antebrachial muscle robustness is related to body size. Data were gathered during dissections of 22 flippers, representing 15 species (7 odontocetes, 15 mysticetes). Results were compared with published descriptions of both artiodactyls and secondarily aquatic vertebrates. Results indicate muscle robustness is best predicted by taxonomic distribution and is not a function of body size. All cetaceans have atrophied triceps muscles, an immobile cubital joint, and lack most connective tissue structures and manus muscles. Forelimbs retain only three muscle groups: triceps (only the scapular head is functional as the humeral heads are vestigal), and antebrachial extensors and flexors. Well‐developed flexor and extensor muscles were found in mysticetes and basal odontocetes (i.e., physeterids, kogiids, and ziphiids), whereas later diverging odontocetes (i.e., monodontids, phocoenids, and delphinids) lack or reduce these muscles. Balaenopterid mysticetes (e.g., fin and minke whales) may actively change flipper curvature, while basal odontocetes (e.g., sperm and beaked whales) probably stiffen the flipper through isometric contraction. Later diverging odontocetes lack musculature supporting digital movements and are unable to manipulate flipper curvature. Cetacean forelimbs are unique in that they have lost agility and several soft tissue structures, but retain sensory innervations. Anat Rec, 290:1121–1137, 2007.

Hydrodynamic performance of the minke whale (Balaenoptera acutorostrata) flipper.

SUMMARY Minke whales (Balaenoptera acutorostrata) are the smallest member of balaenopterid whales and little is known of their kinematics during feeding maneuvers. These whales have narrow and elongated flippers that are small relative to body size compared to related species such as right and gray whales. No experimental studies have addressed the hydrodynamic properties of minke whale flippers and their functional role during feeding maneuvers. This study integrated wind tunnel, locomotion and anatomical range of motion data to identify functional parameters of the cambered minke whale flipper. A full-sized cast of a minke whale flipper was used in wind tunnel testing of lift, drag and stall behavior at six speeds, corresponding to swimming speeds of 0.7–8.9 m s–1. Flow over the model surface stalled between 10° and 14° angle of attack (α) depending on testing speed. When the leading edge was rotated ventrally, loss in lift occurred around –18° α regardless of speed. Range of mobility in the fresh limb was approximately 40% greater than the range of positive lift-generating angles of attack predicted by wind tunnel data (+14°α ). Video footage, photographs and observations of swimming, engulfment feeding and gulping minke whales showed limb positions corresponding to low drag in wind tunnel tests, and were therefore hydrodynamically efficient. Flippers play an important role in orienting the body during feeding maneuvers as they maintain trim of the body, an action that counters drag-induced torque of the body during water and prey intake.

Postcranial morphology and locomotion of the Eocene raoellid Indohyus (Artiodactyla: Mammalia)

Raoellids are small, raccoon-sized Eocene artiodactyls, closely related to archaic cetaceans (archaeocetes) that have poor representation of postcranial elements in the fossil record. Little is known of the aquatic and terrestrial locomotor affinities of raoellids due to the paucity of their fossil record, leaving a critical gap in our understanding of the earliest portion of the artiodactyl marine invasion. To address this gap, a comparative morphological analysis of the postcranial elements was undertaken based on newly recovered elements of the raoellid Indohyus, archaeocetes and extant artiodactyls. Greater than 200 postcranial elements of Indohyus were described, and some limb element cross-sections were visualised via paleohistological thin sections and CT scans. Results show that during terrestrial locomotion, Indohyus probably had a digitigrade posture and mediolaterally stabilised limbs that functioned mostly in flexion and extension within the parasagittal plane. Quantification of midshaft cross-sectional area for some elements of Indohyus showed an osteosclerotic cortex, a skeletal characteristic associated with aquatic behaviours. Indohyus may represent a critical intermediate in the evolution of the cetacean terrestrial-to-aquatic body plan, as it bears gracile postcranial element proportions similar to a terrestrial artiodactyl but also an incipient form of osteosclerosis compared to pakicetid archaeocetes.

Developmental Biology Enriches Paleontology

ABSTRACT Paleontology provides information about the history of morphological transformations, whereas developmental biology provides information about how such transformations happen at a mechanistic level. As such, developmental evidence enriches paleontology in formulating and assessing hypotheses of homology, character definition, and character independence, as well as providing insights into patterns of heterochrony, evolvability of features, and explanations for differential rates of evolution. The focus of this article is to review a series of case studies that illustrate how our understanding of paleontology is enriched by data generated by developmental biologists. The integration of paleontological and developmental data leads to a greater understanding of evolution than either of these sciences could have reached alone. Our case studies range from fish to mammals and involve somite and vertebral formation, limb loss, hand and foot patterning, and tooth formation.

New Middle Eocene Archaeocetes (Cetacea:Mammalia) from the Kuldana Formation of Northern Pakistan

ABSTRACT Two new species of fossil cetaceans (Pakicetus calcis and Pakicetus chittas) are described from a new locality in the Kuldana Formation (Lutetian, middle Eocene) of the Kala Chitta Hills in Northern Pakistan. Additional dentitions of the pakicetid Nalacetus ratimitus, the remingtonocetid Attockicetus, and additional dental material of the holotype of Ambulocetus natans are also described. Dental morphology of Nalacetus is intermediate between pakicetids and ambulocetids, as indicated by the presence of an ambulocetid-like P4, but pakicetid-like molars. Premolars of the specimen tentatively described as Attockicetus imply that some pakicetids and remingtonocetids were coeval. Furthermore, the new pakicetid dentitions described here are the most complete to date and further our understanding of pakicetid dental diversity.

Macroevolutionary developmental biology: Embryos, fossils, and phylogenies

The field of evolutionary developmental biology is broadly focused on identifying the genetic and developmental mechanisms underlying morphological diversity. Connecting the genotype with the phenotype means that evo‐devo research often considers a wide range of evidence, from genetics and morphology to fossils. In this commentary, we provide an overview and framework for integrating fossil ontogenetic data with developmental data using phylogenetic comparative methods to test macroevolutionary hypotheses. We survey the vertebrate fossil record of preserved embryos and discuss how phylogenetic comparative methods can integrate data from developmental genetics and paleontology. Fossil embryos provide limited, yet critical, developmental data from deep time. They help constrain when developmental innovations first appeared during the history of life and also reveal the order in which related morphologies evolved. Phylogenetic comparative methods provide a powerful statistical approach that allows evo‐devo researchers to infer the presence of nonpreserved developmental traits in fossil species and to detect discordant evolutionary patterns and processes across levels of biological organization. Developmental Dynamics 244:1184–1192, 2015.