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Review

Cetaceans as Exemplars of Evolution and Evolutionary Ecology: A Glossary

Department of Biology, Hampden-Sydney College, Hampden-Sydney, VA 23943, USA
Oceans 2020, 1(2), 56-76; https://doi.org/10.3390/oceans1020006
Submission received: 1 May 2020 / Revised: 18 May 2020 / Accepted: 21 May 2020 / Published: 25 May 2020
(This article belongs to the Special Issue Marine Mammals in a Changing World)

Abstract

:
Extant cetaceans (whales, dolphins, and porpoises) and their extinct ancestors offer some of the strongest and best-known examples of macroevolutionary transition as well as microevolutionary adaptation. Unlike most reviews of cetacean evolution, which are intended to chronicle the timeline of cetacean ancestry, document the current knowledge of cetacean adaptations, or simply validate the brute fact of evolution, this review is instead intended to demonstrate how cetaceans fittingly illustrate hundreds of specific, detailed terms and concepts within evolutionary biology and evolutionary ecology. This review, arrayed in alphabetical glossary format, is not meant to offer an exhaustive listing of case studies or scholarly sources, but aims to show the breadth and depth of cetacean research studies supporting and investigating numerous evolutionary themes.

1. Introduction

Perhaps no story within the field of evolutionary biology has attracted more popular attention over the past half century as the evolution of whales. This is undoubtedly due to the steady stream of striking, significant, and substantial fossil finds, and to the general appeal of whales and dolphins to scientists and non-scientists alike. No general textbook of modern biology is complete without at least a minor section or box feature outlining the reversion of early cetaceans to the watery habitat of their pre-mammalian tetrapod ancestors, and the many consequent anatomical and ecological changes that followed this major shift [1,2,3,4,5]. This is a story well and broadly told in print, online, and in superb, instructive video documentaries. Where textbooks of the preceding century could reliably be counted on to depict evolution with the history of horses from Eohippus to Equus, cetaceans are now justifiably cited as prime exemplars of biological evolution. Given this ubiquity, Thewissen and Bajpai [6] crowned cetaceans as the current “poster child for macroevolution”.
This paper is intended neither to reiterate the utility of cetaceans in validating the brute fact of evolution—a point well made in numerous popular books [7,8,9,10], magazines [11,12,13], and websites [14,15,16,17,18], and underscored by many excellent museum exhibits and other public resources—nor to review the current knowledge of cetacean ancestry, a timeline chronicled by a growing array of scholarly and popular works [19,20,21,22,23,24,25,26,27,28,29,30,31,32,33]. Instead, the aim is more narrowly targeted: To show how cetaceans aptly demonstrate specific evolutionary topics. I outline numerous examples of ways in which cetaceans fittingly illustrate detailed terms and concepts within evolutionary biology and evolutionary ecology. These are presented in alphabetical glossary form. They can be used by teachers or scholars searching for examples, or they can simply raise awareness about cetacean research. Neither the examples nor the cited references are meant to offer an exhaustive listing of case studies and illustrations. Rather, the aim is to provide readers, specialists, and non-specialists alike, with an appreciation for the breadth and depth of cetacean research studies.
Given their relatively rapid return to the sea, and hence major change in environment, it is instructive to examine all features of Cetacea within light of the terrestrial-to-aquatic transition. For example, the multi-chambered cetacean stomach is well known. Is this a legacy of cetacean ancestry (specifically their close relation to—indeed, their classification within—Artiodactyla, many of which have stomachs with multiple compartments), or does this instead represent a functional adaptation: A mill for gastric breakdown of ingested food items in the absence of cusped teeth and mastication typical of mammals?
Like the compartmentalized stomach, many aspects of cetacean bodies and life history offer prime examples for explicating and elucidating evolution. The following list of examples runs the gamut from anatomical, behavioral, genetic, and physiological traits, all intended to demonstrate the ease and effectiveness with which Cetacea provides a deep, rich well of exemplars for teaching and studying evolution.

2. Examples of Evolutionary Terms and Concepts

Adaptation: There are dozens of excellent articles and books in both the classic and contemporary literature explaining numerous examples of cetacean adaptations. Chief among these are explanations of skull telescoping [34], the origins of echolocation and related changes to the ear [35,36,37,38], thermoregulation [39], different visual pigments in the eye [40], and different types of myoglobin and other respiratory pigments that bind oxygen [41,42].
Adaptive radiation: Adaptive radiations have been important in cetacean evolutionary history, both in the initial appearance of cetaceans from (presumed) raoellid ancestors as well as in other periods, such as the origin of Odontoceti and Mysticeti (for example, with the evolution of filter feeding) as well as smaller subgroups such as individual genera (e.g., Stenella dolphins) or families (e.g., beaked whales, Ziphiidae), or superfamilies (e.g., Delphinoidea) [31,43,44,45,46].
Aging/lifespan: The discovery of an old stone point embedded in a bowhead whale, Balaena mysticetus, launched a thriving subfield of cetacean studies focused on the remarkably long lifespan (100+ years) of mysticetes, with obvious ramifications for cetacean evolution [47,48]. Such research includes examination of the wax “glove finger” of the mysticete ear, isotopic studies of baleen, or racemization of eyeball amino acids.
Albinism: Albinism has been observed in many cetacean species including sperm, humpback, and killer whales and bottlenose dolphins [49]. This trait is normally governed by a simple genetic mutation, and demonstrates the impact of body coloration and social organization on survival.
Allen’s rule: One of several so-called bioclimatic “rules”, Allen’s rule explains that organisms living at higher latitudes (with cooler climates) tend to have smaller extremities and appendages and thus have less relative surface area over which heat can be lost to the environment. This can be seen with bowhead whale flukes and flippers.
Alloparenting: Alloparenting behavior has been documented in several cetacean species, such as sperm whales that “babysit” juveniles while parents dive deep to forage, or bottlenose dolphin “aunts” (which may or may not be genetically related to true mothers) which help to rear young animals [50,51,52].
Allopatry: Although it is difficult for some people to see how marine species can be fully isolated geographically (i.e., in disparate and non-contiguous distributions) and thus open to allopatric speciation, demographic studies of whale and dolphin populations support cetacean allopatry [53].
Altruism (intraspecific): Altruistic behavior, which benefits another individual at potential cost to the animal performing the behavior, has been documented in many cetacean species, raising important evolutionary questions about the social circumstances that underlie the possible roots of moral behavior. Dolphins in particular have attracted attention for their apparent altruism to conspecifics, suggesting they may be reciprocal altruists [54].
Altruism (interspecific): Many examples of possible or probable altruistic behavior have also been documented in which a whale or dolphin comes to the aid of individuals of another species (as in epimeletic or care-giving behavior, listed below). This includes many anecdotes from both contemporary and ancient, classical times of whales or dolphins protecting smaller, vulnerable animals, such as seals or human swimmers, from drowning or from predators such as sharks [55]. The extent to which such behaviors might be instinctive, and to which cetaceans performing such behaviors recognize that the animals they are aiding are not members of their own species, is the object of much speculation.
Anagenesis: The possibility of cetacean taxa evolving via connected “straight-line” evolution (i.e., without cladogenesis) has been discussed [56].
Analogy/analogous features: Apart from examples of convergent evolution within Cetacea (see below), there are many obvious examples of cetaceans sharing analogous features (i.e., bearing similar form or function yet without a common evolutionary basis) with other taxa. Classic examples include the general fusiform body shape and stabilizing dorsal and pectoral plus propulsive caudal fins in sharks, ichthyosaurs, and dolphins [1,2,3].
Ancestral state reconstruction: Given their striking evolutionary history and increasingly well-known fossil record, reconstruction of ancestral states of cetaceans have been involved in many projects, often focused on body size (and in particular gigantism; see below) [57] but also involving changes in feeding mode over time [58,59].
Apomorphy: Examples of novel, derived traits that distinguish taxa, such as Cetacea or Mysticeti, include many detailed studies of ear bones [35,36,37,38,60,61], as well as analyses of dental and baleen morphology [62], plus highly derived bones or other cranial features related to feeding.
Atavism: The “reappearance” of “lost” hindlimbs or rudimentary yet enlarged pelvic elements are among the most notable and distinct examples of atavisms in any living animals [63].
Bergmann’s rule: Like Allen’s rule (see above), this bioclimatic “rule” holds that body size and shape vary by latitude, in this case with larger animals being found at higher latitudes, such that they have a relatively lower surface-to-volume ratio and correspondingly less heat loss from the body to cold ambient waters. The stout bowhead whale—which is not only more rotund than whales of other families but also of closely related right whales—is a prime example of this finding.
Biogeographical distribution: There are several good examples of cetacean distribution relating to evolution, such as the antitropical distribution of corresponding northern and southern species (such as right whale dolphins) [64], as well as the riverine and estuarine distribution of closely related or convergent taxa (such as river dolphins) [65].
Biological magnification: Cetaceans offer prime examples of biomagnification due to accumulation in tissues of neurotoxins from “red tides” as well as many types of anthropogenic pollutants including methylmercury and organochlorines [66,67,68,69].
Biostratigraphy: As is typical throughout paleontology, the fossil layers (geologic strata) in which cetacean fossils are found offer numerous clues to aid in dating the fossils as well as establish paleoecological or climatological information, making biostratigraphy a common and essential element of any fossil study [70].
Bone bed formation: Although some whale fossils, including whole skeletons, may be found isolated from bones of other individuals, the bones are frequently found in mixed “bone beds” containing fossils of multiple individuals, and sometimes multiple species. This is true of some of the oldest known archaeocetes as well as more recent whales and dolphins in many rich fossil localities, such as the Sharktooth Hill bonebed from the middle Miocene of California [71].
Capital breeding: As opposed to income breeders that use exogenous energy sources to “finance” reproduction, capital breeders build up and maintain internal energy stores before reproducing, a state that has been characterized for many mysticete species [72,73].
Carbon pump: The “fertilization” of primary production in epipelagic seas by the return of carbon, nitrogen, and other nutrients to surface waters via whale feces, has been documented and much discussed in recent years [74,75], with major implications not only for global nutrient cycles but also the key role of pre- and post-whaling cetacean populations in modulating and regulating oceanic ecosystems.
Character displacement: Basic studies of resource partitioning in cetacean taxa include studies of body size, skull telescoping and migration of external nares through evolution, and head shape and dental loss in many odontocete species [76,77].
Chronospecies: The idea that a single species might evolve directly (without divergence) into a morphologically distinct form, yielding a single species (or closely related sister species) connected over time in the fossil record (see entry for anagenesis), has been discussed for mysticetes such as minke whales [78].
Circumpolar: The bowhead whale is a good example of a single cetacean species with multiple breeding stocks in circumpolar distribution (e.g., from the Arctic Ocean to North Atlantic or North Pacific), which has clear implications for speciation, as the now-extinct population of Atlantic gray whales attests [79].
Coevolution: The contemporary and linked evolution of cetaceans with ectoparasitic whale “lice” provide a clear and strong example of host–parasite coevolution [80].
Coloration patterns: There have been several analyses of pigmentation and its relation to character displacement and species recognition, as well as to confusing or startling prey (as by the bright white flippers of humpback whales or asymmetrically colored jaws of fin whales); see countershading entry below [81].
Competition: Competition plays a major role in ecology and evolution, whether intraspecific, such as sperm competition between conspecific males [82], or interspecific, such as between whales and penguins and other species for krill and other food [83].
Constraints: Evolutionary outcomes depend on raw materials but also intrinsic (genetic) and extrinsic (environmental) constraints, which may explain why suspension feeding evolved in cetaceans but not marine reptiles [84], although a plesiosaur has recently been described that possibly was a filter feeder [85].
Convergence: There are many examples of convergent evolution within Cetacea but perhaps none more striking than the tusked, walrus-convergent Odobenocetops of the Miocene and Pliocene, whose skull, dentition, and presumed lifestyle appear to have closely mirrored that of the living walrus, Odobenus [86,87]. Another fine example of convergence involves a beaked whale-like Pliocene dolphin excavated from Antarctica [88].
Cooperation: Many cooperative behaviors, ranging from cooperative foraging to defense against predators, have been documented in cetaceans [89]. Some of these involve complex and instinctive or possibly learned traits. There also appears to be cooperative fishing with humans by some dolphins.
Countershading: A common coloration pattern in aquatic as well as terrestrial and aerial species, especially large predators, leads to light coloration on the animal’s underside (so that it blends in with light from above) and dark coloration on the dorsal surface (so that an animal is hard to detect when viewed from above), and this countershading is prevalent in many cetacean species [81,90].
Culture: Several aspects of distinct cultural transmission of behaviors have been described in odontocete and mysticete species [91,92].
Death and dying behavior: There have been numerous documented instances of dolphins or other cetaceans carrying bodies of dead offspring, or of close attention paid to dead animals by various related and unrelated individuals in many cetacean species [93].
Degeneracy/“devolution”: The loss of complex enamel structure (typical of nearly all non-cetacean mammals) is likely linked to the loss of dental function (typical mammalian mastication) in odontocetes; this loss of complexity appears to be an example of “devolution” [94,95]. Because such losses represent potential decline of reversal of evolution, they are often referred to as degeneracy.
Demography: The possible role of post-menopausal females in cetacean populations has spurred demographic studies focused on population breakdowns and genetics [96].
Disjunction distribution: The majority of whales, dolphins, and porpoises have disjointed (noncontinuous) geographic distributions, also known as range fragmentation. This is significant for demographic, genetic, and evolutionary reasons as well as ecological conditions [97,98].
Disruptive selection: An example of disruptive natural selection in cetaceans, apart from the obvious cases of ecological divergence based on foraging or biogeographic distribution, may involve a natural “knockout” of a basic cytokine protein that acts as an immune signal [99].
Dwarfism: The pygmy blue whale and dwarf and pygmy sperm whales are examples of subspecies and related species, respectively, that demonstrate evolutionary changes in body size (see also gigantism).
Ecotypes: One of the best examples of evolutionary divergence and resource-based character displacement involves the diversity of ecotypes among killer whales, long thought to represent a single species (Orcinus orca) but which may represent distinct species or subspecies in addition to clear ecotypes, such as stocks that feed mainly on salmon or other bony fish, those that prey largely on large sharks, and those that feed mainly on other marine mammals, including small porpoises and seals as well as huge whales [100,101].
Embryology: Although some morphologists studied cetacean embryos and fetuses hundreds of years ago, there is now a much more systematic study of the development of cetacean species, and particularly by using modern molecular and histological methods [102,103].
Encephalization and brain evolution: The story of cetacean brain evolution has attracted much attention in recent years, especially with regard to comparisons with the evolution of large, complex brains in hominins and other primates [104]. This promises to be fertile ground for evolutionary studies.
Endangered/vulnerable species: Several cetacean species have, for various reasons (largely whaling, ship strikes, fishing gear entanglement, and habitat destruction), been and in some cases continue to be highly endangered, including the North Atlantic right whale and the vaquita (a porpoise endemic to the Sea of Cortez which is nearly extinct).
Endemic: Like many organisms, cetaceans demonstrate endemicity, being confined to particular regions. In Cetacea this most commonly occurs in riverine or coastal dolphins [105].
Epigenetics: Apart from various means used to age individual whales and dolphins, such as accumulation of layers of dental tissues or clock-based degradation of molecules [106], epigenetic explanations (i.e., beyond genes) have been proposed to explain how cetaceans may fight cancer [107].
Epimeletic behavior: Care-giving behavior may include the altruistic behaviors described above (such as saving people or other animals from drowning or protecting them from predators) or simply devoting much care and attention to unrelated individuals [108,109].
Evolution of complexity: Just as the loss of odontocete enamel relates to loss of complex structure and function, the evolution of echolocation and vocalizing structures and behaviors in various cetaceans relates to the gain of complexity [33,36,95,110,111], as does the origin and evolution of baleen.
Evolutionarily stable strategy (ESS): Unlike an evolutionarily stable state, an ESS is a behavioral strategy that is fixed or cannot be “invaded” by an alternative gene-based behavior. Various altruistic and other cultural behaviors of dolphins or other cetaceans may qualify [54].
Evolutionary turnover: According to the turnover-pulse hypothesis, major environmental changes often spur major turnover and adaptive radiation of taxa, as is presumed to have occurred during the spread of Neoceti (crown cetaceans) due to Oligocene oceanographic changes [44,112].
Evolutionary-developmental biology (“evo-devo”): The role of regulatory genes leading to morphological changes in dentition and hindlimbs has been the subject of several studies [30,113,114].
Evolvability: The capacity of cetaceans for adaptive evolution due to their molecular and morphogenetic changes after leaving behind their terrestrial ancestry has attracted attention, often involving osteological changes related to ears and hearing or other parts of the cranium [115].
Exaptation: Much speculation about exaptation (“preadaptation”) in cetaceans focuses on the hearing capabilities of the first cetaceans, which were amphibious and had water-adapted hearing that “exapted” them to evolve further into more fully aquatic habitats, and which led to diverse vocalizations (with corresponding brain and anatomical specialization) that ultimately led to echolocatory abilities [116,117].
Extinction: There have, of course, been many cetacean taxa that went extinct and are now known solely from fossil material, but there has also been a recent instance of a living species (the baiji or Yangtze River dolphin, Lipotes vexillifer) which was recently declared extinct [118].
Finite element analysis (FEA): Bite strength in living and extinct cetacean taxa has been analyzed by FEA [119].
Fission–fusion structure: Although more commonly studied in primates, this social structure (in which individuals of a species temporarily join, then go separate ways) has also been documented in various cetacean taxa [120].
Fitness: Among the many examples of evolutionary fitness in cetaceans, the role of the narwhal tusk stands out as an interesting and good example [121,122].
Food fall: There have been many studies investigating the trophic and other ecological roles of cetacean carcasses that decompose over many years on the seafloor (see entry on whalefall communities). These involve observations, experiments, and even analyses of fossil material [123,124].
Fossil dating: Fossils have been used to date divergence ages of extant lineages [125], and multiple methods have been employed to determine the age of cetacean fossils, ranging from traditional isotopic analysis and other molecular methods to geologic formation analysis and use of index fossils [126,127].
Fossil lagerstätte: A lagerstätte is a fossil locality with high diversity, often with numerous complete skeletons (such as the “valley of whales” bearing many basilosaurids and protocetids in Fayum, Egypt), and excellent quality of preservation, such as the Pisco Basin of Peru, where conditions led to preservation of baleen and even digestive tract contents [128,129].
Fossil reconstruction: Many aspects of cetacean form have been reconstructed in extinct taxa, ranging from body size and shape to the curvature and proportions of the spine and their role in locomotion [56,57,130,131].
Genetic drift—bottleneck: The effects of severe population size decrease, namely seemingly random fluctuations in allele and genotype frequencies, have been studied in populations of mysticetes and odontocetes [132,133].
Genetic drift—founder effect: Similar to bottlenecks, small groups can also have random genetic effects when new populations are founded by a very few individuals; this has been studied in various dolphin species [134,135].
Genomics: Several studies have looked at the evolution of the whole genome in particular cetacean species plus higher-level taxa [136,137,138].
Gigantism: Multiple recent studies have looked at the evolution of extreme body size in Cetacea with relation to various factors such as trophic ecology or biomechanics and morphology [139,140,141]. This includes not only the obvious mysticetes but also gigantic toothed whales [142].
Group selection: The concept of group selection (and levels of selection in general) is controversial in evolutionary biology, but studies of whale and dolphin sociality relate directly or indirectly to this topic [92,143].
Habitat loss: Sadly, there are many examples of the evolutionary effects of habitat loss affecting diverse cetaceans in prehistoric and modern times—with the latter obviously involving human impact [144].
Homology: Among the many obvious examples of homologous morphological structures in cetaceans are varied bones [145] and teeth [146]; there are of course homologous chromosomes and genes too.
Host–parasite interactions: There are many records of endo- and ectoparasites on and within different cetaceans, but also interesting evolutionary stories of “switching” of hosts inferred by DNA [147]. See entries below on parasitism and phoresis.
Human impact: Humans have played and continue to play a large role in influencing the evolution of cetaceans through such acts as whaling, driving climate change, and destroying habitats (see habitat loss) [148].
Human impact-anthropophily: A fascinating story of dolphins adapting to, and working cooperatively with, human fishing (in multiple locations around the world) reflects the roles of genes, instincts, and learning in driving cetacean social and behavioral evolution [149,150].
Hybridization: There have been widely reported instances of interspecific hybrid “wholphins” in captivity, but also numerous documented cases of hybrid dolphins and large whales (e.g., blue/fin whales) in natural habitats, spurring speculation as to speciation and genetic divergence [151].
Infectious disease evolution: Changes over time in frequency or at least documentation of various diseases in wild cetacean populations has focused not only on the diseases but also the role of human impact (from pollution and poor sanitation, etc.) in altering the evolution of these diseases and the way they infect whales and dolphins [152].
Irreducible complexity: Critics of evolution often argue that many traits are too complex to have evolved. Traits such as the large brains and echolocatory abilities have been mentioned among cetaceans. Darwin himself speculated, in the first edition of the Origin of Species, about the evolution of baleen and complex filtering form, function, and behavior from swimming bears catching aquatic insects that he presumed might someday evolve into whale-like creatures [153].
Iteroparity: Unlike semelparous organisms that reproduce just once before dying, iteroparous organisms have multiple reproductive events over their lifespan; in cetaceans the timing of this often depends on energy state and accumulation of nutrients [73].
Key innovation: There are many obvious examples of key innovations that ushered in major changes in the ecology and evolution of cetaceans, including the origins of baleen, echolocation, large brains, and structures associated with producing and receiving sound waves (such as the melon, mandibular “pan bone,” and inner ear). The axial skeleton has also been offered as an example of a key innovation important to cetacean evolution [154].
Keystone predators: Killer whales have been proposed as a classic example of a keystone predator whose action helps to regulate the population dynamics and ecology of multiple species (from fish and sea otters to seals, sea lions, and other odontocetes) in marine ecosystems [155].
Kin selection: Alloparenting and related caregiving behaviors by sperm whales and post-menopausal “grandmother” killer whales has been posited as potentially being related to kin selection [96,156].
K strategy: Whales, dolphins, and porpoises are often presented as classic examples of the K-selected life history strategy favoring large bodies with slow growth and maturity, long lifespan, great devotion of resources to few offspring, and so on.
Life history: Analysis of the ways in which various life history factors (such as body size, lifespan, age of maturity, number of offspring, and so on) relate to cetacean evolution has been conducted [157].
Living fossil: The pygmy right whale, Caperea, has been proposed as a remnant of an otherwise ancient and extinct family of early mysticetes, the cetotheriids [158], although it appears that cetotheres persisted into the Pleistocene [159]. The Ganges river dolphin, Platanista, is similarly a remnant of a formerly diversified clade (Platanistoidea).
“Lumpers vs. splitters”: These colloquial terms refer to the preferences among systematists to classify taxa into as few or as many species (or other taxonomic ranks) as possible. Depending on one’s view, there may be 75–120 different extant cetacean species, with much of the disagreement involving dolphins, beaked whales, and rorquals of the genus Balaenoptera.
Mating and social systems: The intricacies of mating systems among diverse whale and dolphin species are often complex (befitting their social complexity) and interesting, as with the intense sperm competition of right whales, and have attracted much scrutiny [160,161].
Metonym (taxonomic synonym): There have been several instances of systematists taking names of extant or extinct cetacean taxa and reusing them to apply to a new taxon [162].
Migration: The relation of long migrations undertaken by whales and dolphins for mating, feeding, and other important activities related to survival and reproduction has been studied not only for specific taxa but in general terms [163].
Mimicry: A frequently cited instance of likely mimicry in Cetacea involves the shark-like appearance (with underslung jaws bearing sharp teeth, plus pigmented false gill slits) of dwarf and pygmy sperm whales, Kogia, although there are also many mentions of possible vocal mimicry.
Modularity: The concept of modularity, in which a structural or functional system can be subdivided into sets of autonomous yet interacting elements (which are altered and interrelated via “evo-devo” changes), has received much recent attention within evolutionary biology. This includes examples within Cetacea [164].
Molecular clock: Many estimates of the origins, divergences, and lifespans of various cetacean taxa have been derived from molecular data by many researchers [136,165,166,167].
Morphological disparity/phenotypic diversity: Many cetaceans display remarkable disparity (e.g., members of the beaked whale genus Mesoplodon), which has been used to study the evolution of Cetacea [44].
Morphological vs. molecular data: The long-standing issue of agreement between anatomical (often osteological) and molecular findings in resolving phylogenetic issues also includes several thorough analyses of cetaceans [117,140,168,169].
Morphometrics: Detailed morphometric studies of nearly every conceivable aspect of cetacean form have been carried out, ranging from overall body size and shape (as relates to locomotion or thermoregulation) as well as teeth, skin, brains, and varied skull structures; several such studies relate directly to phylogenetics and evolution [170].
Mosaic evolution: The extent to which cetacean form and function represents a blend of ancestral and derived characters has been considered in multiple studies involving various organs such as the brain [171]. The gradual loss of hindlimbs in archaeocetes and transition of the forelimb into the flipper also involve mosaic evolution.
Mutation: Along with general exploration of mutations involved in the terrestrial-to-aquatic transition of early cetaceans [1,2,3], many specific studies have examined specific gene mutations and their consequences in cetaceans, most notably involving key events in cetacean evolution (such the loss of body hair) and mutations related to olfaction, gustation, vision, and other senses [172,173,174].
Mutualism: Among the many described instances of mutualism in Cetacea are cases involving whales and non-cetacean taxa (such as seabirds, where the interaction may involve cleaning of parasites from whales as well as location of food sources), as well as discussions of mutualistic interactions (for example, for feeding or defense against predators) involving different cetacean species including interactions between dolphins and large whales [175,176].
Natural vs. artificial selection: The extent to which large-scale twentieth century industrial whaling may have inadvertently altered whale behavior, size, ecology, distribution, and so on, affords an excellent opportunity to compare the effects of human versus natural influences on evolution.
Neoteny and pedomorphosis: Among the many recent investigations focused on changes in developmental timing (see “evo-devo”), and in particular the retention of juvenile features, are comparative studies that closely examined the skulls of extant and extinct whales, dolphins, and porpoises [177,178].
Neuroscience: Outside of Primates, Cetacea is one of the most actively studied groups within the burgeoning field of evolutionary neuroscience, with many projects and publications looking at absolute and relative brain size, the organization of neural networks and brain regions, and the relationship between brain and behavior, including vocalization, sensation, and sociality.
Neutral theory: Many mutations within Cetacea are presumed to have had little to no effect on fitness, yet may elucidate phylogenetics or demographics [179].
Niche separation: The partitioning of food and other resources by contemporaneous humpback and minke whales in Antarctic waters offers a prime example of niche separation and competitive exclusion in Cetacea [180].
Nomenclature: The Latin binomials of many cetaceans—such as the beluga (Delphinapterus leucas, or “white dolphin without a fin”), narwhal (Monodon monoceros, or “one tooth, one tusk”), and humpback whale (Megaptera novaeangliae, or “big-winged New Englander”)—offer good lessons in the principles and practice of naming taxa for experts and beginners alike, as do the common names of these species, along with others (killer whale vs. orca, rorqual, etc.).
Nutrient distribution/trophic connections: In addition to several recent studies that have looked at the role of whales in distributing nutrients throughout marine ecosystems, other investigations have explored trophic interactions between cetaceans and other marine predators (e.g., sharks and penguins) for food [83].
Opportunism: Whereas some cetaceans appear to be highly specialized, others, such as the bottlenose dolphin, Tursiops spp., are successful ecological opportunists, with obvious evolutionary ramifications.
Organ systems: All cetaceans offer prime examples of organs and organ systems highly modified by evolution, such as kidneys that adapted to the switch from a terrestrial to marine environment (with corresponding lack of fresh drinking water), or the lungs and diaphragm modified for greater tidal volume and more efficient pulmonary ventilation, etc. Vascular (often retial) adaptations for diving and thermoregulation are also excellent examples of fundamental evolutionary changes.
Orthogenesis: Studies of cetacean evolution and diversity have not provided evidence for the claim of progressive, directed (i.e., non-random) evolution, although the concept has been discussed [181].
Osteological correlates: The study of bony landmarks and their significance in denoting major functional changes important during cetacean evolution (such as palatal sulci relating to vasculature for baleen, or muscle scars relating to origin/insertion attachment points) has proven invaluable in cetacean paleontology and morphology.
Outgroup comparison: Numerous studies have affirmed the relationship of Cetacea within Artiodactyla (or Cetartiodactyla), with hippopotamuses as the outgroup [182]; other studies have examined outgroups within Cetacea, such as the placement of porpoises within Delphinoidea.
Pair bonding: The tucuxi (Sotalia spp.) is sometimes offered as an example of a cetacean with a pair-bonded mating system [183].
Paleoecology: Among the many studies of cetacean paleoecology are fascinating stories of the likely predation by the extinct giant shark Carcharocles megalodon on baleen whales of all sizes [184].
Parallel evolution: Comparisons between bats and odontocetes as a good example of parallel evolution of echolocation are common [185].
Paraphyly: Paraphyletic groups have been noted in cetacean systematics, especially with older classifications of river dolphins, and more recently with genetic analyses of delphinine dolphins including Stenella and Tursiops [186].
Parasitism: Cetaceans are definitive hosts for numerous ecto- and endoparasites, including Anisakis worms, which easily spread to humans who eat raw or undercooked fish.
Phenetic vs. cladistic systematics: Just as there have been comparisons of molecular and morphological findings in resolving systematic and phylogenetic debates about cetacean taxonomy, so too differing results of phenetics (systematics based on similarity of form) and cladistics have yielded different conclusions, and debate [170].
Phoresis: Cetaceans are well known to “carry” (in a sort of commensal mutualism) many species of ectoparasitic barnacles, worms, and whale lice, along with remoras [187]. This is related to parasitism (see entry above).
Play behavior: Play, considered an important element and indicator of complex social interaction and cognitive ability, has been documented within numerous wild and captive cetaceans [188].
Pleiotropy: The reproductive tracts of some cetaceans may demonstrate pleiotropic genetic interactions [189].
Plesiomorphy: Enamel patterns [190] within cetaceans have been cited as an example of ancestral (plesiomorphic) traits persisting in modern taxa; to a lesser extent, ear bones, although highly modified in Cetacea, also demonstrate some plesiomorphic features [60,61].
Polydactyly vs. polyphalangy: Whales largely retain the plesiomorphic condition of five digits in the forelimb (flipper), although there are instances of digit reduction in cetaceans [191]. Cetaceans, like many other aquatic tetrapods, also are unusual in having an abnormally large number of phalangeal bones within each digit, such that flipper osteology offers a good example of mosaic evolution.
Polygyny: Polygynous mating systems, in which one male has access to a “harem” of several reproductively receptive females, can be found in many cetacean species ranging from sperm to humpback whales [192].
Polymorphism: There are many examples on (and many published papers on) diverse morphological, behavioral, and molecular traits within Cetacea [193].
Polyphyly: The diverse dolphin genus Lagenorhynchus is sometimes offered as an example of a potentially polyphyletic taxonomic grouping within Cetacea [132,193].
Rate of evolution: The extent to which evolution occurs via incremental gradualism or sudden leaps plays a role in studies of neutral mutations, molecular clocks, and regulatory gene interactions (see entries for all items in this list), and cetaceans have provided fodder for interpretations of both slow and rapid evolutionary change, often dependent on environmental factors and the origin of key innovations [194].
Red Queen hypothesis: A spiraling evolutionary “arms race” between cetacean immune systems and pathogens, possibly including neurotoxins from paralytic shellfish poisoning, has been cited as an example of Van Valen’s famous “Red Queen hypothesis,” in which taxa must “keep moving just to stay in one place” [56,195].
Regulatory genes: Homeobox and other regulatory genes (like sonic hedgehog and Hox) have been cited as important in major evolutionary transformations throughout the history of Cetacea involving form and function, such as limb and dental loss and the origins of baleen and echolocation [137].
Relict populations: Diverse river dolphin taxa as well as some oceanic dolphins, notably the rough-toothed dolphin (Steno), have been proposed as ancestral relict populations throughout their distribution or in some locales [196].
Reproductive isolation: Reproductive isolation between cetacean populations and subpopulations have been cited as important steps toward speciation or other genetic and cultural divergence [60,64,197].
Reproductive senescence: Like some higher primates, several cetacean species have been noted as having frequent and important post-reproductive phases, including “grandmother” killer whales that potentially pass along cultural knowledge [198,199].
Resource partitioning: There have been several published examples of resource partitioning within Cetacea (e.g., of different Antarctic whale species partitioning prey or feeding habitat) as well as of cetaceans partitioning resources with other marine species including sharks and seabirds [200].
Ring species: Although there is no definitive example of a cetacean ring species, the presence of numerous subspecies, interspecific hybrids, and intergraded populations of widely dispersed species (often with circumpolar distributions, as in killer whales) makes the prospect of the ring species concept within Cetacea a distinct possibility.
Scaling (isometry vs. allometry): Dozens of publications demonstrate the roles of linear and nonlinear scaling effects in the evolution of cetacean structures ranging from bones, limbs, organs, tissues (such as skin thickness), and other features. These indicate that scaling has played a prominent role in cetacean polymorphism and phenotypic disparity [201].
Selective sweep: Studies of genetic diversity within various cetacean taxa, including some (like sperm whales) with less diversity than expected, demonstrate the likely role that strong selective sweeps play in fixing alleles within a population [202].
Sensory biology: Although studies of cetacean ears and hearing have long attracted research interest, many recent studies have demonstrated that other sensory modalities (vision, olfaction, etc.) are far more complicated and important in cetacean ecomorphology and evolution than generally appreciated [172,173,174].
Sexual conflict: The likelihood of diverging male and female reproductive strategies (and counterstrategies) in mating behavior and anatomy has led to a greater recognition of and research interest in applications of sexual conflict theory within Cetacea [203].
Sexual dimorphism: Sexual dimorphism is common in many cetaceans. In mysticetes, females are generally larger than males, but in odontocetes males are typically larger, often dramatically so (e.g., sperm and killer whales).
Sexual selection: The large tusk of narwhals and prominent mandibular teeth of ziphiids (which in many taxa erupt solely in males) are commonly cited examples of sexual selection in Cetacea. These teeth are often considered to play a role in competition for females, either via display or direct male-to-male fighting [56,204].
Speciation: Antitropical distributions have been cited as a means of enabling allopatric speciation in Cetacea, although other rapid genetic changes or morphological and behavioral disparity (leading to reproductive isolation) might in turn lead to sympatric or parapatric speciation [56,64].
Stabilizing selection: Several genes have been described as having been “purified” or fixed and stabilized at high levels within cetacean populations [42].
Stable isotope analysis: For several decades stable isotopes isolated from various tissues (bones, baleen, etc.) have been widely used to indicate many parameters including distinct populations, ecological states (e.g., trophic levels), and physiological condition (e.g., stress, reproductive condition, age, etc.) [205,206].
Subspecies: Genetically and morphologically distinct subspecies have been described for many species of dolphins and whales, including, for example, humpback and fin whales [207].
Symbiosis: Among the many symbiotic interactions involving cetaceans, familiar examples include multiple species of barnacles that depend exclusively on whales or dolphins for dispersal.
Sympatry: Many cetacean species have been described as sympatric, including river dolphins sharing isolated habitats as well as oceanic whales and dolphins with neritic or pelagic habitats [208,209].
Taphonomy: The condition and potential taphonomic alteration of many fossil cetacean materials has been well described [71,210,211,212].
Tool use: Researchers have described the use of natural sponges or similar materials by dolphins searching for prey in benthic sediments, a behavior that appears to be culturally transmitted [213].
Top-down vs. bottom-up trophic cascades: Researchers have described examples of differing trophic cascades involving whales, dolphins, and porpoises, where either the cetacean (as a large predator) heavily influences the presence and abundance of primary producers and consumers, in a so-called top-down cascade [214], as well as cases of bottom-up cascades where species lower in a trophic pyramid influence the abundance of cetaceans and other large predators [83], such that both types of trophic cascades apply to cetaceans [215,216].
Transitional fossils: Considering the remarkable evolutionary transitions that have occurred throughout cetacean history, such as wholly living, locomoting, and hearing in water instead of air, or capturing prey by filtering with baleen instead of grasping with teeth, there have been many taxa described as classic transitional fossils, indicating forms intermediate between other known forms [1,2,3,4,5,6,7,8,9,10,11,12].
Vestigial features: Cetaceans are among the best-known and frequently cited examples of organisms displaying vestigial features. These include pelvic and limb bones, and hairs and hair follicles.
Vicariance: Extant river dolphins and extinct cetotheres are among the notable taxa whose vicariance (geographic separation into discontinuously distributed groups) has been studied [217,218].
Whalefall community: As previously noted in the entry on food falls, cetacean carcasses often remain on the seafloor as an important contributor to benthic trophic webs. These dead bodies, which may take years to be fully digested and decomposed, form the basis for distinctly unique benthic “whalefall” communities [123,124].

3. Conclusions

Every group of organisms can be used to highlight specific facets of evolution and evolutionary ecology, but extant and extinct whales, dolphins, and porpoises perhaps demonstrate the breadth of evolutionary topics better than any other taxon. The items listed in this glossary are intended to demonstrate the wide range of topics studied by cetacean scientists (they are not meant to provide a complete, exhaustive listing), and this list will undoubtedly grow as new methods yield new insights and discoveries. As surely as cetaceans continue to evolve, so too the fields of study involving them continue to evolve as well.

Funding

This research received no external funding.

Acknowledgments

I thank my many research colleagues and students who have helped to teach and learn with me. I am especially grateful to Olivier Lambert and two anonymous reviewers for providing helpful suggestions including relevant topics and references.

Conflicts of Interest

The author declares no conflicts of interest.

References

  1. Kellogg, R. The history of whales: Their adaptation to life in the water. Q. Rev. Biol. 1928, 3, 29–76. [Google Scholar] [CrossRef]
  2. Howell, A.B. Aquatic Mammals: Their Adaptations to Life in the Water; Charles C. Thomas: Springfield, IL, USA, 1930. [Google Scholar]
  3. Thewissen, J.G.M.; Cooper, L.N.; George, J.C.; Bajpai, S. From land to water: The origin of whales, dolphins, and porpoises. Evol. Educ. Outreach 2009, 2, 272–288. [Google Scholar] [CrossRef] [Green Version]
  4. Reynolds, J.E.; Rommel, S.A. Biology of Marine Mammals; Smithsonian: Washington, DC, USA, 1999. [Google Scholar]
  5. Reidenberg, J. Anatomical adaptations of aquatic mammals. Anat. Rec. 2007, 290, 507–513. [Google Scholar] [CrossRef] [PubMed]
  6. Thewissen, J.G.M.; Bajpai, S. Whale origins as a poster child for macroevolution. BioScience 2001, 51, 1037–1049. [Google Scholar] [CrossRef] [Green Version]
  7. Zimmer, C. At the Water’s Edge: Fish with Fingers, Whales with Legs, and How Life Came Ashore But then Went Back Again; Atria Books: New York, NY, USA, 1999. [Google Scholar]
  8. Berta, A. Return to the Sea: The Life and Evolutionary Times of Marine Mammals; University of California Press: Berkeley, CA, USA, 2012. [Google Scholar]
  9. Thewissen, J.G.M. The Walking Whales: From Land to Water in Eight Million Years; University of California Press: Berkeley, CA, USA, 2014. [Google Scholar]
  10. Pyenson, N.D. Spying on Whales: The Past, Present, and Future of Earth’s Most Awesome Creatures; Viking: New York, NY, USA, 2018. [Google Scholar]
  11. Mueller, T. Valley of the whales. Natl. Geogr. 2010, 218, 118–137. [Google Scholar]
  12. Giggs, R. Giants of the Deep. The Atlantic 2018, 9. Available online: https://www.theatlantic.com/magazine/archive/2018/09/whale-evolution/565760/ (accessed on 1 April 2020).
  13. Black, R. How Did Whales Evolve? Smithsonian Magazine. Available online: https://www.smithsonianmag.com/science-nature/how-did-whales-evolve-73276956/ (accessed on 1 April 2020).
  14. When Whales Walked on Four Legs. Natural History Museum of London. Available online: https://www.nhm.ac.uk/discover/when-whales-walked-on-four-legs.html (accessed on 1 April 2020).
  15. The Evolution of Whales. University of California Museum of Paleontology. Available online: https://evolution.berkeley.edu/evolibrary/article/evograms_03 (accessed on 1 April 2020).
  16. Whale Evolution. PBS WGBH Evolution Series Library. Available online: https://www.pbs.org/wgbh/evolution/library/03/4/l_034_05.html (accessed on 1 April 2020).
  17. Evolution of Whales Animation. Smithsonian Institution. Available online: https://ocean.si.edu/through-time/ancient-seas/evolution-whales-animation (accessed on 1 April 2020).
  18. Whale Evolution from Walking Whales to Janjucetus. Melbourne Museum-Museums Victoria. Available online: https://museumsvictoria.com.au/website/melbournemuseum/discoverycentre/600-million-years/videos/whale-evolution/index.html (accessed on 1 April 2020).
  19. Thewissen, J.G.M. The Emergence of Whales: Evolutionary Patterns in the Origins of Cetacea; Springer: New York, NY, USA, 1998. [Google Scholar]
  20. Mchedlidze, G.A. General Features of the Paleobiological Evolution of Cetacea; Oxonian: New Delhi, India, 1984. [Google Scholar]
  21. Marx, F.; Lambert, O.; Uhen, M.D. Cetacean Paleobiology; Wiley-Blackwell: London, UK, 2016. [Google Scholar]
  22. Berta, A.; Sumich, J.L.; Kovacs, K. Marine Mammals: Evolutionary Biology, 3rd ed.; Elsevier/Academic Press: San Diego, CA, USA, 2015. [Google Scholar]
  23. McGowen, M.R.; Gatesy, J.; Wildman, D.E. Molecular evolution tracks macroevolutionary transitions in Cetacea. Trends Ecol. Evol. 2014, 29, 336–346. [Google Scholar] [CrossRef]
  24. Gatesy, J.; O’Leary, M.A. Deciphering whale origins with molecules and fossils. Trends Ecol. Evol. 2001, 16, 562–571. [Google Scholar] [CrossRef]
  25. Gould, S.J. Hooking leviathan by its past. Nat. Hist. 1995, 94, 8–15. [Google Scholar]
  26. O’Leary, M.A.; Uhen, M.D. The time and origin of whales and the role of behavioral changes in the terrestrial-aquatic transition. Paleobiology 1999, 25, 534–556. [Google Scholar] [CrossRef]
  27. Gatesy, J.; Geiser, J.H.; Chang, J.; Buell, C.; Berta, A.; Meredith, R.W.; Spring, M.S.; McGowen, M.R. A phylogenetic blueprint for a modern whale. Mol. Phylogen. Evol. 2013, 66, 479–506. [Google Scholar] [CrossRef] [PubMed]
  28. Deméré, T.A.; McGowen, M.R.; Berta, A.; Gatesy, J. Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales. Syst. Biol. 2008, 57, 15–37. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  29. Thewissen, J.G.M.; Hussain, S.T.; Arif, M. Fossil evidence for the origin of aquatic locomotion in archaeocete whales. Science 1994, 263, 210–212. [Google Scholar] [CrossRef] [PubMed]
  30. Thewissen, J.G.M.; Cohn, M.J.; Stevens, L.S.; Bajpai, S.; Heyning, J.; Horton, W.E., Jr. Developmental basis for hind-limb loss in dolphins and origin of the cetacean bodyplan. Proc. Natl. Acad. Sci. USA 2006, 103, 8414–8418. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  31. Thewissen, J.G.M.; Williams, E.N. The early radiation of Cetacea (Mammalia): Evolutionary pattern and developmental correlations. Ann. Rev. Ecol. Syst. 2002, 33, 73–90. [Google Scholar] [CrossRef]
  32. Thewissen, J.G.M.; Cooper, L.N.; Clementz, M.T.; Bajpai, S.; Tiwari, B.N. Whales originated from aquatic artiodactyls in the Eocene epoch of India. Nature 2007, 450, 1190–1191. [Google Scholar] [CrossRef]
  33. Gingerich, P.D.; Wells, N.A.; Russell, D.E.; Ibrahim Shah, S.M. Origin of whales in epicontinental remnant seas: New evidence from the Early Eocene of Pakistan. Science 1983, 220, 403–406. [Google Scholar] [CrossRef]
  34. Roston, R.; Roth, V.L. Cetacean skull telescoping brings evolution of cranial sutures into focus: Telescoping and cranial suture evolution. Anat. Rec. 2019, 302, 1055–1073. [Google Scholar] [CrossRef]
  35. Geisler, J.H.; Colbert, M.W.; Carew, J.L. A new fossil species supports and early origin for toothed whale echolocation. Nature 2014, 508, 383–386. [Google Scholar] [CrossRef]
  36. Mourlam, M.J.; Orliac, M.J. Infrasonic and ultrasonic hearing evolved after the emergence of modern whales. Curr. Biol. 2017, 27, 1776–1781. [Google Scholar] [CrossRef] [Green Version]
  37. Pilleri, G. Adaptation to water and the evolution of echolocation in the Cetacea. Ethol. Ecol. Evol. 1990, 2, 135–163. [Google Scholar] [CrossRef]
  38. Ketten, D.R. The marine mammal ear: Specializations for aquatic audition and echolocation. In The Evolutionary Biology of Hearing; Webster, D.B., Popper, A.N., Fay, R.R., Eds.; Springer: New York, NY, USA, 1992; pp. 717–750. [Google Scholar]
  39. Werth, A.J. Adaptations of the cetacean hyolingual apparatus for aquatic feeding and thermoregulation. Anat. Rec. 2007, 290, 546–568. [Google Scholar] [CrossRef]
  40. Fasick, J.I.; Robinson, P.R. Adaptations of cetacean retinal pigments to aquatic environments. Front. Ecol. Evol. 2016, 4, e70. [Google Scholar] [CrossRef] [Green Version]
  41. Noren, S.R.; Williams, T.M. Body size and skeletal muscle myoglobin of cetaceans: Adaptations for maximizing dive duration. Comp. Biochem. Physiol. A 2000, 126, 181–191. [Google Scholar] [CrossRef]
  42. McClellan, D.A.; Palfreyman, E.J.; Smith, M.J.; Moss, J.L.; Christensen, R.G.; Sailsbery, J.K. Physicochemical evolution and molecular adaptation of the cetacean and artiodactyl cytochrome b proteins. Mol. Biol. Evol. 2005, 22, 437–455. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  43. Shen, T.; Xu, S.; Wang, X.; Yu, W.; Zhou, K.; Yang, G. Adaptive evolution and functional constraint at TLR4 during the secondary aquatic adaptation and diversification of cetaceans. BMC Evol. Biol. 2012, 12, e39. [Google Scholar] [CrossRef] [Green Version]
  44. Slater, G.J.; Price, S.A.; Santini, F.; Alfaro, M.E. Diversity versus disparity and the radiation of modern cetaceans. Proc. R. Soc. B 2010, 277, 3097–3104. [Google Scholar] [CrossRef] [Green Version]
  45. Steeman, M.E.; Hebagaard, M.B.; Fordyce, R.E.; Ho, S.Y.W.; Rabosky, D.L.; Nielsen, R.; Rahbek, C.; Glenner, H.; Sorensen, M.V.; Willersley, E. Radiation of extant cetaceans driven by restructuring of the oceans. Syst. Biol. 2009, 58, 573–585. [Google Scholar] [CrossRef] [Green Version]
  46. Marx, F.G.; Fordyce, R.E. Baleen boom and bust: A synthesis of mysticete phylogeny, diversity, and disparity. R. Soc. Open Sci. 2015, 2, e140434. [Google Scholar] [CrossRef] [Green Version]
  47. Sacher, G.A. Constitutional basis of longevity in the Cetacea: Do the whales and the terrestrial mammals obey the same laws? Rep. Int. Whal. Comm. Spec. Issue 1980, 3, 209–213. [Google Scholar]
  48. Seim, I.; Ma, S.; Zhou, X.; Gerashchenko, M.V.; Lee, S.G.; Suydam, R.; George, J.C.; Bickham, J.W.; Gladyshev, V.N. The transcriptome of the bowhead whale Balaena mysticetus reveals adaptations of the longest-lived mammal. Aging 2014, 6, 879–899. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  49. Fertl, D.; Rosen, P.E. Albinism. In Encyclopedia of Marine Mammals, 3rd ed.; Würsig, B., Thewissen, J.G.M., Kovacs, K.M., Eds.; Academic Press: San Diego, CA, USA, 2018; pp. 20–21. [Google Scholar]
  50. Gero, S.; Englhaupt, D.; Rendell, L.; Whitehead, H. Who Cares? Between-group variation in alloparental caregiving in sperm whales. Behav. Ecol. 2009, 20, 838–843. [Google Scholar] [CrossRef]
  51. Augusto, J.F.; Frasier, T.R.; Whitehead, H. Characterizing alloparental care in the pilot whale (Globicephala melas) population that summers off Cape Breton, Nova Scotia, Canada. Mar. Mamm. Sci. 2016, 33, 440–456. [Google Scholar] [CrossRef]
  52. Weinpress, M.R.; Herzing, D. Maternal and Alloparental Discipline in Atlantic Spotted Dolphins (Stenella frontalis) in the Bahamas. Anim. Behav. Cogn. 2015, 2, 348–364. [Google Scholar] [CrossRef]
  53. Hare, M.P.; Cipriano, F.; Palumbi, S.R. Genetic evidence on the demography of speciation in allopatric dolphin species. Evolution 2002, 56, 804–816. [Google Scholar] [CrossRef]
  54. Connor, R.; Norris, K.S. Are Dolphins Reciprocal Altruists? Am. Nat. 1982, 119, 358–374. [Google Scholar] [CrossRef]
  55. Pitman, R.L.; Deecke, V.B.; Gabriele, C.M.; Srinivasan, M.; Black, N.; Denkinger, J.; Durban, J.W.; Matthews, E.A.; Matkin, D.R.; Neilson, J.L.; et al. Humpback whales interfering when mammal-eating killer whales attack other species: Mobbing behavior and interspecific altruism? Mar. Mamm. Sci. 2017, 33, 7–58. [Google Scholar] [CrossRef]
  56. Fordyce, R.E. Cetacean evolution. In Encyclopedia of Marine Mammals, 3rd ed.; Würsig, B., Thewissen, J.G.M., Kovacs, K.M., Eds.; Academic Press: San Diego, CA, USA, 2018; pp. 180–185. [Google Scholar]
  57. Pyenson, N.D.; Sponberg, S.N. Reconstructing body size in extinct crown Cetacea (Neoceti) using allometry, phylogenetic methods and tests from the fossil record. J. Mamm. Evol. 2011, 18, e269. [Google Scholar] [CrossRef]
  58. Didier, G. Time-dependent-asymmetric-linear-parsimonious ancestral state reconstruction. Bull. Math. Biol. 2017, 79, 2334–2355. [Google Scholar] [CrossRef]
  59. Johnston, C.; Berta, A. Comparative anatomy and evolutionary history of suction feeding in cetaceans. Mar. Mamm. Sci. 2011, 27, 493–513. [Google Scholar] [CrossRef]
  60. Geisler, J.H.; Luo, Z. The petrosal and inner ear of Herpetocetus sp. (Mammalia: Cetacea) and their implications for the phylogeny and hearing of archaic mysticetes. J. Paleont. 1996, 70, 1045–1066. [Google Scholar] [CrossRef]
  61. Luo, Z. Homology and transformation of cetacean ectotympanic structures. In The Emergence of Whales: Advances in Vertebrate Paleobiology; Thewissen, J.G.M., Ed.; Springer: Boston, MA, USA, 1998; Volume 1, pp. 269–301. [Google Scholar]
  62. O’Leary, M.A. Phylogenetic and morphometric reassessment of the dental evidence for a mesonychian and cetacean clade. In The Emergence of Whales: Advances in Vertebrate Paleobiology; Thewissen, J.G.M., Ed.; Springer: Boston, MA, USA, 1998; Volume 1, pp. 133–161. [Google Scholar]
  63. Simões-Lopes, P.C.; Gutstein, C.S. Notes on the anatomy, positioning and homology of the pelvic bones in small cetaceans (Cetacea, Delphinidae, Pontoporiidae). Lat. Am. J. Aq. Mamm. 2004, 3, 157–162. [Google Scholar] [CrossRef] [Green Version]
  64. Davies, J.L. The antitropical factor in cetacean speciation. Evolution 1963, 17, 107–116. [Google Scholar] [CrossRef] [Green Version]
  65. Hamilton, H.; Caballero, S.; Collins, A.G.; Brownell, R.L. Evolution of river dolphins. Proc. R. Soc. Lond. B 2001, 268, 549–556. [Google Scholar] [CrossRef] [PubMed]
  66. Loseto, L.L.; Stern, G.A.; Ferguson, S.H. Size and biomagnification: How habitat selection explains beluga mercury levels. Environ. Sci. Technol. 2008, 42, 3982–3988. [Google Scholar] [CrossRef] [PubMed]
  67. Hoekstra, P.F.; O’Hara, T.M.; Pallant, S.J.; Solomon, K.R.; Muir, D.C. Bioaccumulation of Organochlorine Contaminants in Bowhead Whales (Balaena mysticetus) from Barrow, Alaska. Arch. Environ. Contam. Toxicol. 2002, 42, 497–507. [Google Scholar] [CrossRef]
  68. Baron, E.; Gimenez, J.; Verborgh, P.; Gauffier, P.; DeStephanis, R.; Eljarrat, E.; Barcelo, D. Bioaccumulation and biomagnification of classical flame retardants, related halogenated natural compounds and alternative flame retardants in three delphinids from Southern European waters. Environ. Pollut. 2015, 203, 107–115. [Google Scholar] [CrossRef]
  69. Alonso, M.B.; Azevedo, A.; Torres, J.P.; Dorneles, P.R.; Eljarrat, E.; Barcelo, D.; Lailson-Brito, J.; Malm, O. Anthropogenic (PBDE) and naturally-produced (MeO-PBDE) brominated compounds in cetaceans—A review. Sci. Total Environ. 2014, 481, 619–634. [Google Scholar] [CrossRef]
  70. Bianucci, G.; Sart, G.; Catanzariti, R.; Santini, U. Middle Pliocene cetaceans from Monte Voltraio (Tuscany, Italy) biostratigraphical, paleoecological, and paleoclimatic observations. Rev. Ital. Paleont. Strat. 1998, 104, 1. [Google Scholar]
  71. Pyenson, N.D.; Irmis, R.B.; Lipps, J.H.; Barnes, L.G.; Mitchell, E.D.; MacLeod, S.A. Origin of a widespread marine bonebed deposited during the middle Miocene Climatic Optimum. Geology 2009, 37, 519–522. [Google Scholar] [CrossRef]
  72. Borrell, A.; Gomez-Campos, E.; Aguilar, A. Influence of reproduction on stable-isotope rations: Nitrogen and carbon isotope discrimination between mothers, fetuses, and milk in the fin whale, a capital breeder. Physiol. Biochem. Zool. 2016, 89, e684632. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  73. Christiansen, F.; Vikingsson, G.A.; Rasmussen, M.H.; Lusseau, D. Female body condition affects foetal growth in a capital breeding mysticete. Funct. Ecol. 2014, 28, 579–588. [Google Scholar] [CrossRef]
  74. Roman, J.; McCarthy, J.J. The whale pump: Marine mammals enhance primary productivity in a coastal basin. PLoS ONE 2010, 5, e13255. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  75. Lavery, T.J.; Roudnew, B.; Gill, P.; Seymour, J.; Seuront, L.; Johnson, G.; Mitchell, J.G.; Smetacek, V. Iron defecation by sperm whales stimulates carbon export in the Southern Ocean. Proc. R. Soc. Lond. B 2010, 277, 3527–3531. [Google Scholar] [CrossRef] [Green Version]
  76. Werth, A.J. Odontocete suction feeding: Experimental analysis of water flow and head shape. J. Morph. 2006, 267, 1415–1428. [Google Scholar] [CrossRef]
  77. Werth, A.J. Mandibular and dental variation and the evolution of suction feeding in Odontoceti. J. Mamm. 2006, 87, 579–588. [Google Scholar] [CrossRef]
  78. Bisconti, M. Taxonomy and evolution of the Italian Pliocene Mysticeti (Mammalia, Cetacea): A state of the art. Bull. Soc. Paleont. Ital. 2009, 48, 147–156. [Google Scholar]
  79. Mead, J.G.; Mitchell, E.D. Atlantic gray whales. In The Gray Whale: Eschrichtius robustus; Jones, M.L., Swartz, S.L., Leatherwood, S., Eds.; Academic Press: San Diego, CA, USA, 1984; pp. 33–53. [Google Scholar]
  80. Balbuena, J.A.; Raga, J.A. Ecology and host relationships of the whale-louse Isocyamus delphini (Amphipoda: Cyamidae) parasitizing long-finned pilot whales (Globicephala melas) off the Faroe Islands (Northeast Atlantic). Can. J. Zool. 1991, 69, 141–145. [Google Scholar] [CrossRef]
  81. Mitchell, E. Pigmentation pattern evolution in delphinid cetaceans: An essay in adaptive coloration. Can. J. Zool. 1970, 48, 717–740. [Google Scholar] [CrossRef]
  82. MacLeod, C.D. The relationship between body mass and relative investment in testes mass in cetaceans: Implications for inferring interspecific variations in the extent of sperm competition. Mar. Mamm. Sci. 2010, 26, 370–380. [Google Scholar] [CrossRef]
  83. Ainley, D.G.; Ballard, G.; Dugger, K.M. Competition among penguins and cetaceans reveals trophic cascades in the Western Ross Sea, Antarctica. Ecology 2006, 87, 2080–2093. [Google Scholar] [CrossRef]
  84. Collin, R.; Janis, C.M. Morphological constraints on tetrapod feeding mechanisms: Why were there no suspension-feeding marine reptiles? In Ancient Marine Reptiles; Callaway, J.M., Nicholls, E.L., Eds.; Academic Press: San Diego, CA, USA, 1997; pp. 451–466. [Google Scholar]
  85. O’Keefe, F.R.; Otero, R.A.; Soto-Acuna, S.; O’Gorman, J.P.; Godfrey, S.J.; Chatterjee, S. Cranial anatomy of Morturnia seymourensis from Antarctica, and the evolution of filter feeding in plesiosaurts of the Austral Late Cretaceous. J. Vert. Paleontol. 2017, 37, e1347570. [Google Scholar] [CrossRef]
  86. Muizon, C. Walrus-like feeding adaptation in a new cetacean from the Pliocene of Peru. Nature 1993, 365, 745–748. [Google Scholar] [CrossRef]
  87. Muizon, C.; Domning, D.P.; Ketten, D.R. Odobenocetops peruvianus, the walrus-convergent delphinoid (Mammalia: Cetacea) from the early Pliocene of Peru. Smithson. Contrib. Paleobiol. 2002, 93, 223–261. [Google Scholar]
  88. Fordyce, R.E.; Quilty, P.G.; Daniels, J. Australodelphis mirus, a bizarre new toothless ziphiid-like fossil dolphin (Cetacea: Delphinidae) from the Pliocene of Vestfold Hills, East Antarctica. Antarct. Sci. 2002, 14, 37–54. [Google Scholar] [CrossRef]
  89. Ballance, L.T. Cetacean ecology. In Encyclopedia of Marine Mammals, 3rd ed.; Würsig, B., Thewissen, J.G.M., Kovacs, K.M., Eds.; Academic Press: San Diego, CA, USA, 2018; pp. 172–180. [Google Scholar]
  90. Caro, T.; Beeman, K.; Stankowich, T.; Whitehead, H. The functional significance of colouration in cetaceans. Evol. Ecol. 2011, 25, e1231. [Google Scholar] [CrossRef]
  91. Whitehead, H. Gene–culture coevolution in whales and dolphins. Proc. Natl. Acad. Sci. USA 2017, 114, 7814–7821. [Google Scholar] [CrossRef] [Green Version]
  92. Rendell, L.; Whitehead, H. Culture in whales and dolphins. Behav. Brain Sci. 2001, 24, 309–382. [Google Scholar] [CrossRef] [Green Version]
  93. Bearzi, G.; Eddy, L.; Piwetz, S.; Reggente, M.A.L.; Cozzi, B. Cetacean behavior toward the dead and dying. In Encyclopedia of Animal Cognition and Behavior; Vonk, J., Shackelford, T., Eds.; Springer Nature: Basel, Switzerland, 2017. [Google Scholar] [CrossRef]
  94. Ishiyama, M. Enamel structure in odontocete whales. Scanning Microsc. 1987, 1, 1071–1079. [Google Scholar]
  95. Werth, A.J.; Loch, C.; Fordyce, R.E. Enamel microstructure in Cetacea: A case study in evolutionary loss of complexity. J. Mammal Evol. 2019. [Google Scholar] [CrossRef]
  96. Johnstone, R.A.; Cant, M.A. The evolution of menopause in cetaceans and humans: The role of demography. Proc. R. Soc. Lond. B 2010, 277, 3765–3771. [Google Scholar] [CrossRef] [PubMed]
  97. Gladden, J.G.B.; Ferguson, M.M.; Clayton, J.W. Matriarchal genetic population structure of North American beluga whales Delphinapterus leucas (Cetacea: Monodontidae). Mol. Ecol. 1997, 6, 1033–1046. [Google Scholar] [CrossRef] [PubMed]
  98. Fontaine, M.C.; Tolley, K.A.; Michaux, J.R.; Birkun, A.; Ferreira, M.; Jauniaux, T.; Llavona, A.; Ozturk, B.; Ozturk, A.A.; Ridoux, V.; et al. Genetic and historic evidence for climate-driven population fragmentation in a top cetacean predator: The harbour porpoises in European water. Proc. R. Soc. B 2010, 277, 2829–2837. [Google Scholar] [CrossRef] [PubMed]
  99. Lopes-Marques, M.; Machado, A.M.; Barbosa, S.; Fonseca, M.M.; Ruiyo, R.; Castro, L.F.C. Cetacea are natural knockouts for IL20. Immunogenetics 2018, 70, 681–687. [Google Scholar] [CrossRef] [PubMed]
  100. Foote, A.D.; Vijay, N.; Avila-Arcos, M.C.; Baird, R.; Durban, J.W.; Fumagalli, M.; Gibbs, R.A.; Hanson, M.B.; Korneliussen, T.S.; Martin, M.D.; et al. Genome-culture coevolution promotes rapid divergence of killer whale ecotypes. Nat. Commun. 2016, 7, e11693. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  101. DeBruyn, P.J.; Tosh, C.A.; Terauds, A. Killer whale ecotypes: Is there a global model. Biol. Rev. Camb. Philos. Soc. 2013, 88, 62–80. [Google Scholar] [CrossRef] [Green Version]
  102. Thewissen, J.G.M.; Heyning, J. Embryogenesis and development in Stenella attenuata and other cetaceans. In Reproductive Biology and Phylogeny of Cetacea; Miller, D.L., Ed.; CRC Press: Boca Raton, FL, USA, 2007; pp. 307–329. [Google Scholar]
  103. Thewissen, J.G.M.; Hieronymus, T.L.; George, J.C.; Suydam, R.; Stimmelmayr, R.; McBurney, D. Evolutionary aspects of the development of teeth and baleen in the bowhead whale. J. Anat. 2017, 230, 549–566. [Google Scholar] [CrossRef] [Green Version]
  104. Marino, L. A comparison of encephalization between odontocete cetaceans and anthropoid primates. Brain. Behav. Evol. 1998, 51, 230–238. [Google Scholar] [CrossRef] [Green Version]
  105. Perez-Alvarez, M.J.; Olavarria, C.; Moraga, R.; Baker, C.S.; Hamner, R.M.; Poulin, E. Microsatellite markers reveal strong genetic structure in the endemic Chilean dolphin. PLoS ONE 2015, 10, e0123956. [Google Scholar] [CrossRef] [Green Version]
  106. Beal, A.P.; Kiszka, J.J.; Wells, R.S.; Eirin-Lopez, J.M. The bottlenose dolphin epigenetic aging tool (BEAT): A molecular age estimation tool for small cetaceans. Front. Mar. Sci. 2019, 6, e561. [Google Scholar] [CrossRef] [Green Version]
  107. Tejada-Martinez, D.; Magalhaes, J.P.; Opazo, J.C. Positive selection and fast turnover rate in tumor suppressor genes reveal how cetaceans resist cancer. bioRxiv 2020. [Google Scholar] [CrossRef] [Green Version]
  108. Bearzi, G.; Reggente, M.A.L. Epimeletic behavior. In Encyclopedia of Marine Mammals, 3rd ed.; Würsig, B., Thewissen, J.G.M., Kovacs, K.M., Eds.; Academic Press: San Diego, CA, USA, 2018; pp. 337–338. [Google Scholar]
  109. Caldwell, M.C.; Caldwell, D.K. Epimeletic (care-giving) behavior in Cetacea. In Whales, Dolphins, and Porpoises; Norris, K.N., Ed.; University of California Press: Berkeley, CA, USA, 1966; pp. 755–789. [Google Scholar]
  110. Berta, A.; Ekdale, E.G.; Cranford, T.W. Review of the cetacean nose: Form, function, and evolution. Anat. Rec. 2014, 297, 2205–2215. [Google Scholar] [CrossRef] [PubMed]
  111. Cranford, T.W. The sperm whale’s nose: Sexual selection on a grand scale? Mar. Mamm. Sci. 1999, 15, 1133–1157. [Google Scholar] [CrossRef]
  112. Marx, F.G.; Fitzgerald, E.M.G.; Fordyce, R.E. Like phoenix from the ashes: How modern baleen whales arose from a fossil “dark age.”. Acta Palaeontol. Pol. 2019, 64, 231–238. [Google Scholar] [CrossRef]
  113. Armfield, B.A.; Zheng, Z.; Bajpai, S.; Vinyard, C.J.; Thewissen, J.G.M. Development and evolution of the unique cetacean dentition. PeerJ 2013, 1, e24. [Google Scholar] [CrossRef] [Green Version]
  114. Bejder, L.; Hall, B.K. Limbs in whales and limblessness in other vertebrates: Mechanisms of evolutionary and developmental transformation and loss. Evol. Dev. 2002, 4, 445–458. [Google Scholar] [CrossRef] [Green Version]
  115. Fahlke, J.M.; Hampe, O. Cranial symmetry in baleen whales (Cetacea, Mysticeti) and the occurrence of cranial asymmetry throughout cetacean evolution. Sci. Nat. 2015, 102, e58. [Google Scholar] [CrossRef]
  116. Lindberg, D.R.; Pyenson, N.D. Things that go bump in the night: Evolutionary interactions between cephalopods and cetaceans in the tertiary. Lethaia 2007, 40, 335–343. [Google Scholar] [CrossRef]
  117. Geisler, J.H.; McGowen, M.R.; Yang, G.; Gatesy, J. A supermatrix analysis of genomic, morphological, and paleontological data from crown Cetacea. BMC Evol. Biol. 2011, 11, e112. [Google Scholar] [CrossRef] [Green Version]
  118. Turvey, S.T.; Pitman, R.L.; Taylor, B.L.; Barlow, J.; Akamatsu, T.; Barrett, L.A.; Zhao, X.; Reeves, R.R.; Stewart, B.S.; Wang, K.; et al. First human-caused extinction of a cetacean species? Biol. Lett. 2007, 3, 537–540. [Google Scholar] [CrossRef] [Green Version]
  119. Snively, E.; Fahlke, J.M.; Welsh, R.C. Bone-breaking bite force of Basilosaurus isis (Mammalia, Cetacea) from the Late Eocene of Egypt estimated by finite element analysis. PLoS ONE 2015, 10, e0118380. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  120. Parra, G.J.; Corkeron, P.J.; Arnold, P. Grouping and fission–fusion dynamics in Australian snubfin and Indo-Pacific humpback dolphins. Anim. Behav. 2011, 82, 1423–1433. [Google Scholar] [CrossRef]
  121. Best, R.C. The tusk of the narwhal (Monodon monoceros L.): Interpretation of its function (Mammalia: Cetacea). Can. J. Zool. 1981, 59, 2386–2393. [Google Scholar] [CrossRef]
  122. Nweeia, M.T.; Eichmiller, F.C.; Nutarak, C.; Eidelman, N.; Giuseppetti, A.A.; Quinn, J.; Mead, J.G.; K’issuk, K.; Hauschka, P.V.; Tyler, E.M. Considerations of anatomy, morphology, evolution, and function for narwhal dentition. In Smithsonian at the Poles: Contributions to International Polar Year Science; Lang, M.A., Miller, S.E., Eds.; Smithsonian Press: Washington, DC, USA, 2009; pp. 223–240. [Google Scholar]
  123. Kiel, S.; Goedert, J.L. Deep-sea food bonanzas: Early Cenozoic whale-fall communities resemble wood-fall rather than seep communities. Proc. R. Soc. Lond. B 2006, 273, 2625–2632. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  124. Fujiwara, Y.; Kawato, M.; Yamanaka, T.; Sato-Okoshi, W.; Noda, C.; Tsuchida, S.; Komai, T.; Cubelio, S.S. Three-year investigations into sperm whale-fall ecosystems in Japan. Mar. Ecol. 2007, 28, 219–232. [Google Scholar] [CrossRef]
  125. McGowen, M.R.; Spaulding, M.; Gatesy, J. Divergence date estimation and a comprehensive molecular tree of extant cetaceans. Mol. Phylogen. Evol. 2009, 53, 891–906. [Google Scholar] [CrossRef]
  126. Bosio, G.; Malinverno, E.; Villa, I.M.; DiCelma, C.; Gariboldi, K.; Gioncada, A.; Barberini, V.; Urbina, M.; Bianucci, G. Tephrochronology and chronostratigraphy of the Miocene Chilcatay and Pisco formations (East Pisco Basin, Peru). Newslett. Stratigr. 2020, 53, 213–247. [Google Scholar] [CrossRef]
  127. Gariboldi, K.; Bosio, G.; Malinverno, E.; Gioncada, A.; DiCelma, C.; Villa, I.M.; Urbina, M.; Bianucci, G. Biostratigraphy, geochronology, and sedimentation rates of the upper Miocene Pisco Formation at two important marine vertebrate fossil-bearing sites of southern Peru. Newslett. Stratigr. 2017, 50, 417–444. [Google Scholar] [CrossRef]
  128. Gioncada, A.; Collareta, A.; Gariboldi, K.; Lambert, O.; DiCelma, C.; Bonaccorsi, E.; Urbina, M.; Bianucci, G. Inside baleen: Exceptional microstructure preservation in a late Miocene whale skeleton from Peru. Geology 2016, 44, 839–842. [Google Scholar] [CrossRef]
  129. Brand, L.; Urbina, M.; Chadwick, A.; DeVries, T.J.; Esperante, R. A high resolution stratigraphic framework for the remarkable fossil cetacean assemblage of the Miocene/Pliocene Pisco Formation, Peru. J. S. Am. Earth Sci. 2011, 31, 414–425. [Google Scholar] [CrossRef]
  130. Marino, L.; Uhen, M.D.; Pyenson, N.D.; Frohlich, B. Reconstructing cetacean brain evolution using computed tomography. Anat. Rec. 2003, 272B, 107–117. [Google Scholar] [CrossRef] [PubMed]
  131. Buchholtz, E.A. Vertebral osteology and swimming style in living and fossil whales (Order: Cetacea). J. Zool. 2001, 253, 175–190. [Google Scholar] [CrossRef]
  132. Lee, K.; Lee, J.M.; Sohn, S.; Cho, Y.; Choi, Y.M.; Kim, H.K.; Kim, J.H.; Jeong, D.G. Complete mitochondrial genome of the Pacific white-sided dolphin Lagenorhynchus obliquidens (Cetacea: Delphinidae). Conserv. Gen. Res. 2018, 10, 201–204. [Google Scholar] [CrossRef]
  133. Rooney, A.; Honeycutt, R.; Davis, S.; Derr, J.N. Evaluating a putative bottleneck in a population of bowhead whales from patterns of microsatellite diversity and genetic disequilibria. J. Mol. Evol. 1999, 49, 682–690. [Google Scholar] [CrossRef]
  134. Amendola-Pimenta, M.; Camelo-Marrufo, M.; Zamora-Briseno, J.A.; Hernandez-Velazques, I.M.; Zamora-Bustillos, R.; Rodrigues-Canul, R. Genetic bottleneck and founder effect signatures in a captive population of common bottlenose dolphins Tursiops truncatus (Montagu 1821) in Mexico. PeerJ 2018, 6, e26891v1. [Google Scholar]
  135. Gaspari, S.; Airoldi, S.; Hoelzel, A.R. Risso’s dolphins (Grampus griseus) in UK waters are differentiated from a population in the Mediterranean Sea and genetically less diverse. Conserv. Gen. 2007, 8, 727–732. [Google Scholar] [CrossRef]
  136. McGowan, M.R.; Tsagkogeorga, G.; Alvarez-Carretero, S.; dos Reis, M.; Struebig, M.; Deaville, R.; Jepson, P.D.; Jarman, S.; Polanowski, A.; Morin, P.A. Phylogenomic resolution of the cetacean tree of life using target sequence capture. Syst. Biol. 2020, 69, 479–501. [Google Scholar] [CrossRef]
  137. Nery, M.F.; Gonzalez, D.J.; Opazo, J.C. How to Make a Dolphin: Molecular Signature of Positive Selection in Cetacean Genome. PLoS ONE 2013, 8, e65491. [Google Scholar] [CrossRef] [Green Version]
  138. Yim, H.S.; Cho, Y.S.; Guang, X.; Kang, S.G.; Jeong, J.Y.; Cha, S.S.; Oh, H.M.; Lee, J.H.; Yang, E.C.; Kwon, K.K.; et al. Minke whale genome and aquatic adaptation in cetaceans. Nat. Gen. 2014, 46, 88–92. [Google Scholar] [CrossRef] [Green Version]
  139. Goldbogen, J.A.; Madsen, P.T. The evolution of foraging capacity and gigantism in cetaceans. J. Exp. Biol. 2018, 221, e166033. [Google Scholar] [CrossRef] [Green Version]
  140. Fordyce, R.E.; Marx, F.G. Gigantism precedes filter feeding in baleen whale evolution. Curr. Biol. 2018, 28, 1670–1676. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  141. Bianucci, G.; Marx, F.G.; Collareta, A.; DiStefano, A.; Landini, W.; Morigi, C.; Varola, A. Rise of the titans: Baleen whales became giants earlier than thought. Biol. Lett. 2019, 15, e0175. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  142. Lambert, O.; Bianucci, G.; Post, K.; de Muizon, C.; Salas-Gismondi, R.; Urbina, M.; Reuner, J. The giant bite of a new raptorial sperm whale from the Miocene epoch of Peru. Nature 2010, 466, 105–108. [Google Scholar] [CrossRef] [PubMed]
  143. Whitehead, H. Cultural selection and genetic diversity in matrilineal whales. Science 1998, 28, 1708–1711. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  144. Huang, S.L.; Hao, Y.; Mei, Z.; Turvey, S.T.; Wang, D. Common pattern of population decline for freshwater cetacean species in deteriorating habitats. Freshw. Biol. 2012, 57, 1266–1276. [Google Scholar] [CrossRef]
  145. Gol’din, P. Naming an innominate: Pelvis and hindlimbs of Miocene whales give and insight into evolution and homology of cetacean pelvic girdle. Evol. Biol. 2017, 41, 473–479. [Google Scholar] [CrossRef]
  146. Peredo, C.M.; Pyenson, N.D.; Uhen, M.D.; Marshall, C.D. Alveoli, teeth, and tooth loss: Understanding the homology of internal mandibular structures in mysticete cetaceans. PLoS ONE 2017, 12, e0178243. [Google Scholar] [CrossRef]
  147. Fraija-Fernandez, N.; Olson, P.D.; Crespo, E.A.; Raga, J.A.; Aznar, F.J.; Fernandez, M. Independent host switching events by digenean parasites of cetaceans inferred from ribosomal DNA. Int. J. Parasitol. 2015, 45, 167–173. [Google Scholar] [CrossRef]
  148. Marx, F.G.; Uhen, M.D. Climate, critters, and cetaceans: Cenozoic drivers of the evolution of modern whales. Science 2010, 327, 993–996. [Google Scholar] [CrossRef] [Green Version]
  149. Simões-Lopes, P.C.; Daura-Jorge, F.G.; Cantor, M. Clues of cultural transmission in cooperative foraging between artisanal fishermen and bottlenose dolphins, Tursiops truncatus (Cetacea: Delphinidae). Zool. Curitiba 2016, 33. [Google Scholar] [CrossRef] [Green Version]
  150. Peterson, D.; Hanazaki, N.; Simões-Lopes, P.C. Natural resource appropriation in cooperative artisanal fishing between fishermen and dolphins (Tursiops truncatus) in Laguna, Brazil. Ocean Coast. Manag. 2008, 51, 469–475. [Google Scholar] [CrossRef]
  151. Crossman, C.A.; Taylor, E.B.; Barrett-Lennard, L.G. Hybridization in the Cetacea: Widespread occurrence and associated morphological, behavioral, and ecological factors. Ecol. Evol. 2016, 6, 1293–1303. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  152. Van Bressem, M.F.; Raga, J.A.; DiGuardo, G.; Jepson, P.D.; Duignan, P.J.; Siebert, U.; Barrett, T.; Santos, M.C.O.; Moreno, I.B.; Siciliano, S.; et al. Emerging infectious diseases in cetaceans worldwide and the possible role of environmental stressors. Dis. Aquat. Org. 2009, 86, 143–157. [Google Scholar] [CrossRef] [PubMed]
  153. Darwin, C.R. On the Origin of Species; John Murray: London, UK, 1859. [Google Scholar]
  154. Gillet, A.; Frederich, B.; Parmentier, E. Divergent evolutionary morphology of the axial skeleton as a potential key innovation in modern cetaceans. Proc. R. Soc. B 2019, 286, e20191771. [Google Scholar] [CrossRef] [PubMed]
  155. Estes, J.A.; Tinker, M.T.; Williams, T.M.; Doak, D.F. Killer whale predation on sea otters linking oceanic and nearshore ecosystems. Science 1998, 285, 473–476. [Google Scholar] [CrossRef] [Green Version]
  156. Parsons, K.M.; Durban, J.W.; Claridge, D.E.; Balcomb, K.C.; Noble, L.R.; Thompson, P.M. Kinship as a basis for alliance formation between male bottlenose dolphins, Tursiops truncatus, in the Bahamas. Anim. Behav. 2003, 66, 185–194. [Google Scholar] [CrossRef] [Green Version]
  157. Huang, S.L.; Ni, I.H.; Chou, L.S. Correlations in cetacean life history traits. Raffles Bull. Zool. 2008, 19, 285–292. [Google Scholar]
  158. Marx, F.G.; Fordyce, R.E. A link no longer missing: New evidence for the cetotheriid affinity for Caperea. PLoS ONE 2016, 11, e0164059. [Google Scholar] [CrossRef]
  159. Boessenecker, R.W. Pleistocene survival of an archaic dwarf baleen whale (Mysticeti: Cetotheriidae). Naturwissenschaften 2013, 100, 365–371. [Google Scholar] [CrossRef] [Green Version]
  160. Perrin, W.M.; Mesnick, S.L. Sexual ecology of the spinner dolphin, Stenella longirostris: Geographic variation in mating system. Mar. Mamm. Sci. 2003, 19, 462–483. [Google Scholar] [CrossRef]
  161. Schaeff, C.M. Courtship and mating behavior. In Reproductive Biology and Phylogeny of Cetacea; Miller, D.L., Ed.; CRC Press: Boca Raton, FL, USA, 2007; pp. 349–370. [Google Scholar]
  162. Hershkovitz, P. Catalogue of Living Whales; Smithsonian: Washington, DC, USA, 1966. [Google Scholar]
  163. Corkeron, P.J.; Connor, R.C. Why do baleen whales migrate? Mar. Mamm. Sci. 2006, 15, 1228–1245. [Google Scholar] [CrossRef]
  164. Churchill, M.; Miguel, J.; Beatty, B.L.; Gowami, A.; Geisler, J.H. Asymmetry drives modularity of the skull in the common dolphin (Delphinus delphis). Biol. J. Linn. Soc. 2018, 126, 225–239. [Google Scholar] [CrossRef]
  165. Jackson, J.A.; Baker, C.S.; Vant, M.; Steel, D.J.; Medrano-Gonzalez, L.; Palumbi, S.R. Big and slow: Phylogenetic estimates of molecular evolution in baleen whales (Suborder Mysticeti). Mol. Biol. Evol. 2009, 26, 2427–2440. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  166. Milinkovitch, M.C.; Mayer, A.; Powell, J.R. Phylogeny of all major groups of cetaceans based on DNA sequences from three mitochondrial genes. Mol. Biol. Evol. 1994, 11, 939–948. [Google Scholar] [PubMed] [Green Version]
  167. Theodor, J.M. Molecular clock divergence estimates and the fossil record of Cetartiodactyla. J. Paleont 2004, 78, 39–44. [Google Scholar] [CrossRef] [Green Version]
  168. Naylor, G.J.P.; Adams, D.C. Are the fossil data really at odds with the molecular data? Morphological evidence for Cetartiodactyl phylogeny reexamined. Syst. Biol. 2001, 50, 444–453. [Google Scholar]
  169. Messenger, S.L.; McGuire, J.A. Morphology, molecules, and the phylogenetics of cetaceans. Syst. Biol. 1998, 47, 90–124. [Google Scholar] [CrossRef] [Green Version]
  170. Amaral, A.R.; Coelho, M.M.; Marugan-Lobon, J.; Rohlf, F.J. Cranial shape differentiation in three closely related delphinid cetacean species: Insights into evolutionary history. Zoology 2009, 112, 38–47. [Google Scholar] [CrossRef]
  171. Falk, D.; Dudek, B. Mosaic evolution of the neocortex. Behav. Brain Sci. 1993, 16, 701–702. [Google Scholar] [CrossRef]
  172. Zhu, K.; Zhou, X.; Xu, S.; Sun, D.; Ren, W.; Zhou, K.; Yang, G. The loss of taste genes in cetaceans. BMC Evol. Biol. 2014, 14, e218. [Google Scholar] [CrossRef] [Green Version]
  173. Huelsmann, M.; Hecker, N.; Springer, M.S.; Gatesy, J.; Sharma, V.; Hiller, M. Genes lost during the transition from land to water in cetaceans highlight genomic changes associated with aquatic adaptations. Sci. Adv. 2019, 5, e6671. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  174. Chen, Z.; Wang, Z.; Xu, S.; Zhou, K.; Yang, G. Characterization of hairless (Hr) and FGF5 genes provides insights into the molecular basis of hair loss in cetaceans. BMC Evol. Biol. 2013, 13, e34. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  175. Katona, S.; Whitehead, H. Are Cetacea ecologically important? Oceanogr. Mar. Biol. Ann. Rev. 1988, 26, 553–568. [Google Scholar]
  176. Trull, P. Symbiotic Relationship between Humpback Whales and Marine Birds. Orleans Conservation Trust. Available online: https://orleansconservationtrust.org/symbiotic-relationship-between-humpback-whales-and-marine-birds-presentation-recap/ (accessed on 1 April 2020).
  177. Tsai, C.H.; Fordyce, R.E. Juvenile morphology in baleen whale phylogeny. Naturwissenschaften 2014, 101, 765–769. [Google Scholar] [CrossRef]
  178. Galatius, A. Paedomorphosis in two small species of toothed whales (Odontoceti): How and why? Biol. J. Linn. Soc. 2010, 99, 278–295. [Google Scholar] [CrossRef] [Green Version]
  179. Waples, R.S. Genetic methods for estimating the effective size of cetacean populations. Rep. Int. Whal. Comm. 1991, 13, 279–300. [Google Scholar]
  180. Friedlaender, A.; Lawon, G.L.; Halpin, P.N. Evidence of resource partitioning between humpback and minke whales around the western Antarctic Peninsula. Mar. Mamm. Sci. 2009, 25, 402–415. [Google Scholar] [CrossRef] [Green Version]
  181. Erwin, D.H. Macroevolution: Dynamics of diversity. Curr. Biol. 2011, 21, R1000–R1001. [Google Scholar] [CrossRef] [Green Version]
  182. Milinkovitch, M.C.; Berube, M.; Palsboll, P.J. Cetaceans are highly derived artiodactyls. In The Emergence of Whales; Thewissen, J.G.M., Ed.; Plenum: New York, NY, USA, 1998; pp. 113–131. [Google Scholar]
  183. Santos, M.; Rosso, S. Ecological aspects of marine tucuxi dolphins (Sotalia guianensis) based on group size and composition in the Cananéia Estuary, southeastern Brazil. Lat. Am. J. Aquat. Mamm. 2007, 6, 71–82. [Google Scholar] [CrossRef] [Green Version]
  184. Collareta, A.; Lambert, O.; Landini, W.; DiCelma, C.; Malinverno, E.; Varas-Malca, R.; Urbina, M.; Bianucci, G. Did the giant extinct shark Carcharocles megalodon target small prey? Bite marks on marine mammal remains from the late Miocene of Peru. Paleogeog. Paleoclimatol. Paleoecol. 2017, 469, 84–91. [Google Scholar] [CrossRef]
  185. Yablokov, A.V. Convergence or parallelism in the evolution of cetaceans. Int. Geol. Rev. 1965, 7, 1461–1468. [Google Scholar] [CrossRef]
  186. Perrin, W.F.; Rosel, P.E.; Cipriano, F. How to contend with paraphyly in the taxonomy of delphinine cetaceans? Mar. Mamm. Sci. 2013, 29, 567–588. [Google Scholar] [CrossRef]
  187. Arvy, L. Phoresies and parasitism in cetaceans: A review. Invest Cetacea 1982, 14, 233–335. [Google Scholar]
  188. Paulos, R.D.; Trone, M.; Kuczaj, S.A. Play in wild and captive cetaceans. Int. J. Comp. Psychol. 2010, 23, 701–722. [Google Scholar]
  189. Wang, J.Y.; Liao, W.B. Ontogenesis and evolutionary allometry shape divergent evolution of genitalia in female cetaceans. Evolution 2018, 72, 404–405. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  190. Loch, C.; Duncan, W.; Simões-Lopes, P.C.; Kieser, J.A.; Fordyce, R.E. Ultrastructure of enamel and dentine in extant dolphins (Cetacea: Delphinoidea and Inioidea). Zoomorphology 2013, 132, 215–225. [Google Scholar] [CrossRef]
  191. Cooper, L.N.; Berta, A.; Dawson, S.D.; Reidenberg, J.S. Evolution of hyperphalangy and digit reduction in the cetacean manus. Anat. Rec. 2007, 290, 654–672. [Google Scholar] [CrossRef]
  192. Cercio, S.; Jacobsen, J.K.; Cholewiak, D.M.; Falcone, E.A.; Merriwether, D.A. Paternity in humpback whales, Megaptera novaeangliae: Assessing polygyny and skew in male reproductive success. Anim. Behav. 2005, 70, 267–277. [Google Scholar] [CrossRef]
  193. Vachon, F.; Whitehead, H.; Frasier, T.R. What factors shape genetic diversity in cetaceans? Ecol. Evol. 2018, 8, 1554–1572. [Google Scholar] [CrossRef] [Green Version]
  194. Hoelzel, A.R.; Hancock, J.M.; Dover, G.A. Evolution of the cetacean mitochondrial D-loop region. Mol. Biol. Evol. 1991, 8, 475–493. [Google Scholar]
  195. Pagan, H.J.T.; Ferrer, T.; O’Corry-Crowe, G. Positive selection in coding regions and motif duplication in regulatory regions of bottlenose dolphin MHC class II genes. PLoS ONE 2018, 13, e0203450. [Google Scholar] [CrossRef] [PubMed]
  196. Kerem, D.; Goffman, O.; Elasar, M.; Hadar, N.; Scheinin, A.; Lewis, T. The rough-toothed dolphin, Steno bredanensis, in the Eastern Mediterranean Sea: A relict population? Adv. Mar. Biol. 2016, 75, 233–258. [Google Scholar] [PubMed]
  197. Riesch, R.; Barrett-Lennard, L.G.; Ellis, G.M.; Ford, J.K.B.; Deecke, V.B. Cultural traditions and the evolution of reproductive isolation: Ecological speciation in killer whales? Biol. J. Linn. Soc. 2012, 106, 1–17. [Google Scholar] [CrossRef] [Green Version]
  198. Marsh, H.; Kasuya, T. 1986 Evidence for reproductive senescence in female cetaceans. Rep. Int. Whal. Comm. 1986, 8, 57–73. [Google Scholar]
  199. Ellis, S.; Franks, D.W.; Nattrass, S.; Currie, T.E.; Cant, M.A.; Giles, D.; Balcomb, K.C.; Croft, D.P. Analyses of ovarian activity reveal repeated evolution of post-reproductive lifespans in toothed whales. Sci. Rep. 2018, 8, e12833. [Google Scholar] [CrossRef] [PubMed]
  200. Ansmann, I.C.; Lanyon, J.M.; Seddon, J.M.; Parra, G.J. Habitat and resource partitioning among Indo-Pacific bottlenose dolphins in Moreton Bay, Australia. Mar. Mamm. Sci. 2015, 31, 211–230. [Google Scholar] [CrossRef]
  201. Pyenson, N.D.; Goldbogen, J.A.; Shadwick, R.E. Mandible allometry in extant and fossil Balaenopteridae (Cetacea: Mammalia): The largest vertebrate skeletal element and its role in rorqual lunge feeding. Biol. J. Linn. Soc. 2013, 108, 586–599. [Google Scholar] [CrossRef] [Green Version]
  202. Alexander, A.; Steel, D.; Slikas, B.; Hoekzema, K.; Carraher, C.; Parks, M.; Cronn, R.; Baker, C.S. Low diversity in the mitogenome of sperm whales revealed by next generation sequencing. Genome Biol. Evol. 2013, 5, 113–129. [Google Scholar] [CrossRef] [Green Version]
  203. Zheng, R.; Karczmarki, L.; Lin, W.; Chan, S.C.Y.; Chang, W.L.; Wu, Y. Infanticide in the Indo-Pacific humpback dolphin (Sousa chinensis). J. Ethol. 2016, 34, 299–307. [Google Scholar] [CrossRef]
  204. Dalebout, M.L.; Steel, D.; Baker, C.S. Phylogeny of the beaked whale genus Mesoplodon (Ziphiidae: Cetacea) revealed by nuclear introns: Implications for the evolution of male tusks. Syst. Biol. 2008, 57, 857–875. [Google Scholar] [CrossRef] [Green Version]
  205. Borrell, A.; Velásquez Vacca, A.; Pinela, A.M.; Kinze, C.; Lockyer, C.H.; Vighi, M.; Aguilar, A. Stable isotopes provide insight into population structure and segregation in Eastern North Atlantic sperm whales. PLoS ONE 2013, 8, e82398. [Google Scholar] [CrossRef] [PubMed]
  206. Clementz, M.T.; Fordyce, R.E.; Peek, S.L.; Fox, D.L. Ancient marine isoscapes and isotopic evidence of bulk-feeding by Oligocene cetaceans. Paleogeogr. Paleoclimatol. Paleoecol. 2014, 400, 28–40. [Google Scholar] [CrossRef]
  207. Perrin, W.F.; Mead, J.G.; Brownell, R.L. Review of the Evidence Used in the Description of Currently Recognized Cetacean Subspecies. 2009. Available online: https://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1115&context=usdeptcommercepub (accessed on 21 May 2020).
  208. Yang, G.; Liu, S.; Ren, W.; Zhou, K.; Wei, F. Mitochondrial control region variability of baiji and the Yangtze finless porpoises, two sympatric small cetaceans in the Yangtze river. Acta Theriol. 2003, 48, 469–483. [Google Scholar] [CrossRef]
  209. Hoelzel, A.R. Genetic structure of cetacean populations in sympatry, parapatry, and mixed assemblages: Implications for conservation policy. J. Hered. 1998, 89, 451–458. [Google Scholar] [CrossRef] [Green Version]
  210. Boessenecker, R.W.; Perry, F.A.; Schmitt, J.G. Comparative taphonomy, taphofacies, and bonebeds of the Mio-Pliocene Purisima Formation, Central California: Strong physical control on marine vertebrate preservation in shallow marine settings. PLoS ONE 2014, 9, e91419. [Google Scholar] [CrossRef] [PubMed]
  211. Bianucci, G.; Collareta, A.; Bosio, G.; Landini, W.; Gariboldi, K.; Gioncada, A.; Lambert, O.; Malinverno, E.; de Muizon, C.; Varas-Malca, R.; et al. Taphonomy and palaeoecology of the lower Miocene marine vertebrate assemblage of Ullujaya (Chilcatay Formation, East Pisco Basin, southern Peru). Paleogeogr. Paleoclimatol. Paleoecol. 2018, 511, 256–279. [Google Scholar] [CrossRef]
  212. Esperante, R.; Brand, L.; Chadwick, A.; Poma, O. Taphonomy of fossil whales in the diatomaceous sediments of the Miocene/Pliocene Pisco Formation, Peru. Curr. Top. Taph. Fossil. 2002, 337–344. [Google Scholar]
  213. Krutzen, M.; Mann, J.; Heithaus, M.R.; Connor, R.C.; Bajder, L.; Sherwin, W.B. Cultural transmission of tool use in bottlenose dolphins. Proc. Natl. Acad. Sci. USA 2005, 102, 8939–8943. [Google Scholar] [CrossRef] [Green Version]
  214. Baum, J.K.; Worm, B. Cascading top-down effects of changing ocean predator abundances. J. Anim. Ecol. 2009, 78, 699–714. [Google Scholar] [CrossRef]
  215. Lynam, C.P.; Llope, M.; Mollman, C.; Helaouet, P.; Bayliss-Brown, G.A.; Stenseth, N.C. Interaction between top-down and bottom-up control in marine food webs. Proc. Natl. Acad. Sci. USA 2017, 114, 1952–1957. [Google Scholar] [CrossRef] [Green Version]
  216. Ainley, D.; Ballard, G.; Blight, L.K.; Ackley, S.; Emslie, S.D.; Lescroel, A.; Olmastroni, S.; Townsend, S.E.; Tynan, C.T.; Wilson, P.; et al. Impacts of cetaceans on the structure of Southern Ocean food webs. Mar. Mamm. Sci. 2010, 26, 482–498. [Google Scholar] [CrossRef]
  217. Braulik, G.; Barnett, R.; Odon, V.; Islas-Villaneuva, V.; Hoelzel, R.; Graves, J.A. One species or two? Vicariance, lineage divergence, and low mtDNA diversity in geographically isolated populations of South Asian river dolphin. J. Mamm. Evol. 2014, 22, 111–120. [Google Scholar] [CrossRef]
  218. Bisconti, M. Anatomy of a new cetotheriid genus and species from the Miocene of Herentals, Belgium, and the phylogenetic and palaeobiogeographical relationships of Cetotheriidae s.s. (Mammalia, Cetacea, Mysticeti). J. Syst. Palaeont. 2015, 13, 377–395. [Google Scholar] [CrossRef]

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Werth, A.J. Cetaceans as Exemplars of Evolution and Evolutionary Ecology: A Glossary. Oceans 2020, 1, 56-76. https://doi.org/10.3390/oceans1020006

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Werth AJ. Cetaceans as Exemplars of Evolution and Evolutionary Ecology: A Glossary. Oceans. 2020; 1(2):56-76. https://doi.org/10.3390/oceans1020006

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Werth, Alexander J. 2020. "Cetaceans as Exemplars of Evolution and Evolutionary Ecology: A Glossary" Oceans 1, no. 2: 56-76. https://doi.org/10.3390/oceans1020006

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