Delphinidae Research Articles

The amygdaloid body of the family Delphinidae: a morphological study of its central nucleus through calbindin-D28k.

The amygdala is a noticeable bilateral structure in the medial temporal lobe and it is composed of at least 13 different nuclei and cortical areas, subdivided into the deep nuclei, the superficial nuclei, and the remaining nuclei which contain the central nucleus (CeA). CeA mediates the behavioral and physiological responses associated with fear and anxiety through pituitary-adrenal responses by modulating the liberation of the hypothalamic Corticotropin Releasing Factor/Hormone. Five dolphins of three different species, belonging to the family Delphinidae (three striped dolphins, one common dolphin, and one Atlantic spotted dolphin), were used for this study. For a precise overview of the CeA's structure, thionine staining and the immunoperoxidase method using calbindin D-28k were employed. CeA extended mainly dorsal to the lateral nucleus and ventral to the striatum. It was medial to the internal capsule and lateral to the optic tract and the medial nucleus of the amygdala. The dolphin amygdaloid complex resembles that of primates, including the subdivision, volume, and location of the CeA.

Common dolphin whistle responses to experimental mid-frequency sonar.

Oceanic delphinids that occur in and around Navy operational areas are regularly exposed to intense military sonar broadcast within the frequency range of their hearing. However, empirically measuring the impact of sonar on the behavior of highly social, free-ranging dolphins is challenging. Additionally, baseline variability or the frequency of vocal state-switching among social oceanic dolphins during undisturbed conditions is lacking, making it difficult to attribute changes in vocal behavior to anthropogenic disturbance. Using a network of drifting acoustic buoys in controlled exposure experiments, we investigated the effects of mid-frequency (3-4 kHz) active sonar (MFAS) on whistle production in short-beaked (Delphinus delphis delphis) and long-beaked common dolphins (Delphinus delphis bairdii) in southern California. Given the complexity of acoustic behavior exhibited by these group-living animals, we conducted our response analysis over varying temporal windows (10 min- 5 s) to describe both longer-term and instantaneous changes in sound production. We found that common dolphins exhibited acute and pronounced changes in whistle rate in the 5 s following exposure to simulated Navy MFAS. This response was sustained throughout sequential MFAS exposures within experiments simulating operational conditions, suggesting that dolphins may not habituate to this disturbance. These results indicate that common dolphins exhibit brief yet clearly detectable acoustic responses to MFAS. They also highlight how variable temporal analysis windows-tuned to key aspects of baseline vocal behavior as well as experimental parameters related to MFAS exposure-enable the detection of behavioral responses. We suggest future work with oceanic delphinids explore baseline vocal rates a-priori and use information on the rate of change in vocal behavior to inform the analysis time window over which behavioral responses are measured.

Dawn of the Delphinidans: New Remains of Kentriodon from the Lower Miocene of Italy Shed Light on the Early Radiation of the Most Diverse Extant Cetacean Clade.

Nowadays, the infraorder Delphinida (oceanic dolphins and kin) represents the most diverse extant clade of Cetacea, with delphinids alone accounting for more than 40% of the total number of living cetacean species. As for other cetacean groups, the Early Miocene represents a key interval for the evolutionary history of Delphinida, as it was during this time span that the delphinidans became broadly distributed worldwide, first and foremost with the widespread genus Kentriodon and closely related forms. Here, we report on a new odontocete find from Burdigalian (20.4-19.0 Ma) deposits of the Friulian-Venetian Basin of northeastern Italy, consisting of the partial cranium of a small delphinidan with associated ear bones (right periotic, stapes, malleus and tympanic bulla). Osteoanatomical considerations and comparisons allow us to assign the studied specimen to the genus Kentriodon. This is the first confirmed record of Kentriodon from Europe as well as from the whole proto-Mediterranean region. Stratigraphic and phylogenetic considerations suggest that our new specimen may represent the geologically oldest member of Kentriodon. The evolutionary success of Kentriodon may correlate with the emergence of narrow-band high-frequency echolocation as a possible strategy to escape acoustic detection by large marine predators such as the squalodontids. In addition, the relatively high encephalization quotient of Kentriodon spp. may have provided these early dolphins with some kind of competitive advantage over the coeval non-delphinidan odontocetes.

Molecular Evidence for Relaxed Selection on the Enamel Genes of Toothed Whales (Odontoceti) with Degenerative Enamel Phenotypes.

Different species of toothed whales (Odontoceti) exhibit a variety of tooth forms and enamel types. Some odontocetes have highly prismatic enamel with Hunter-Schreger bands, whereas enamel is vestigial or entirely lacking in other species. Different tooth forms and enamel types are associated with alternate feeding strategies that range from biting and grasping prey with teeth in most oceanic and river dolphins to the suction feeding of softer prey items without the use of teeth in many beaked whales. At the molecular level, previous studies have documented inactivating mutations in the enamel-specific genes of some odontocete species that lack complex enamel. At a broader scale, however, it is unclear whether enamel complexity across the full diversity of extant Odontoceti correlates with the relative strength of purifying selection on enamel-specific genes. Here, we employ sequence alignments for seven enamel-specific genes (ACP4, AMBN, AMELX, AMTN, ENAM, KLK4, MMP20) in 62 odontocete species that are representative of all extant families. The sequences for 33 odontocete species were obtained from databases, and sequences for the remaining 29 species were newly generated for this study. We screened these alignments for inactivating mutations (e.g., frameshift indels) and provide a comprehensive catalog of these mutations in species with one or more inactivated enamel genes. Inactivating mutations are rare in Delphinidae (oceanic dolphins) and Platanistidae/Inioidea (river dolphins) that have higher enamel complexity scores. By contrast, mutations are much more numerous in clades such as Monodontidae (narwhal, beluga), Ziphiidae (beaked whales), Physeteroidea (sperm whales), and Phocoenidae (porpoises) that are characterized by simpler enamel or even enamelless teeth. Further, several higher-level taxa (e.g., Hyperoodon, Kogiidae, Monodontidae) possess shared inactivating mutations in one or more enamel genes, which suggests loss of function of these genes in the common ancestor of each clade. We also performed selection (dN/dS) analyses on a concatenation of these genes and used linear regression and Spearman's rank-order correlation to test for correlations between enamel complexity and two different measures of selection intensity (# of inactivating mutations per million years, dN/dS values). Selection analyses revealed that relaxed purifying selection is especially prominent in physeteroids, monodontids, and phocoenids. Linear regressions and correlation analyses revealed a strong negative correlation between selective pressure (dN/dS values) and enamel complexity. Stronger purifying selection (low dN/dS) is found on branches with more complex enamel and weaker purifying selection (higher dN/dS) occurs on branches with less complex enamel or enamelless teeth. As odontocetes diversified into a variety of feeding modes, in particular, the suction capture of prey, a reduced reliance on the dentition for prey capture resulted in the relaxed selection of genes that are critical to enamel development.

First report of false killer whale Pseudorca crassidens in Bizerte, Tunisia

False killer whale (Pseudorca crassidens) is a species of oceanic dolphin that is the only extant representative of the genus Pseudorca. It is found worldwide but mainly frequents tropical regions. This manuscript deals with the first sighting and stranding event recorded of the false killer whale in Mediterranean waters of Bizerte, northern Tunisia. The species has never been reported in the area despite years of collecting anecdotal and scientific information of cetaceans. This report extends previous knowledge of the false killer whale in the Mediterranean Sea.

Natural and anthropogenic organic brominated compounds in the southwestern Atlantic ocean: Bioaccumulation in coastal and oceanic dolphin species

Marine pollution is considered a current driver of change in the oceans and despite the urgency to develop more studies, there is limited information in the southern hemisphere. This study aimed to analyze the levels and profiles of natural (MeO-PBDEs) and anthropogenic (BFRs: PBDEs, HBB, PBEB) organic brominated compounds in adipose tissue of two species of dolphins with different distribution and trophic requirements from the Southwestern Atlantic Ocean; the short-beaked common dolphin (Delphinus delphis) and the Fraser's dolphin (Lagenodelphis hosei). In addition, we aim to investigate maternal transfer and biological pattern relationship (sex, age, sexual maturity) in short-beaked common dolphin bioaccumulation. The levels of both groups of contaminants were in the same order of magnitude as those reported for other marine mammals on both a regional and global scale. BFRs profiles were dominated by BDE 28 and BDE 47 in short-beaked common dolphin and Fraser's dolphin, respectively, whereas 2-MeO-BDE 68 was the most abundant natural compound in both species. Evidence of maternal transfer, temporary increase in BDE 154 levels and no influence of sex, age, or sexual maturity on brominated compound concentration was observed in short-beaked common dolphin. This study fills a gap in the knowledge of the Southwestern Atlantic Ocean providing new information on emerging organic pollutants bioavailability for dolphins and, therefore, for the different trophic webs. In addition, it serves as a baseline for further contamination assessments.

Dolphins from a prehistoric midden imply long-term philopatry of delphinids around Tokyo Bay

Abstract Cetaceans of the family Delphinidae are widely distributed throughout the global ocean, and it has been suggested that females exhibit philopatry. However, the duration of time that a maternal group of these highly mobile creatures spends inhabiting a particular region remains unknown. The objective of this study is to examine the genetic diversity of delphinids in the past and to shed light on the above question. In this study, we amplified and sequenced the mitochondrial DNA D-loop region from the skeletal remains of delphinids that were excavated from the Shomyoji Midden, a site situated on the western coast of Tokyo Bay and dating to 5000–3000 years before the present. We acquired prehistoric D-loop sequences of seven bottlenose dolphins, six Pacific white-sided dolphins, and a false killer whale. Notably, one or more haplotypes from the prehistoric individuals were detected in the geographically proximate modern conspecific populations across all three species. Whilst the genetic diversity of prehistoric and modern Pacific white-sided dolphin populations appears to be comparable, the modern bottlenose dolphin population demonstrates significantly lower genetic diversity when compared to its prehistoric counterpart in the vicinity of Tokyo Bay. The maternal groups of these species are speculated to have persisted in the same habitat for at least 4000 years.

First mercury and stable isotope assessment from an unusual mass stranding of rough-toothed dolphins (Steno bredanensis) (Artiodactyla: Delphinidae) in Azuero peninsula, Pacific coast of Panama

Introduction: Small cetaceans are good bioindicators of environmental contamination; however, knowledge about their ecotoxicological status in Central America is scarce. In Panama, access to samples from wild populations to determine the ecotoxicological status of oceanic dolphins is limited; therefore, stranding events provide an alternative for obtaining samples. In April 2016, a rare mass stranding event occurred in the Azuero Peninsula (Pacific coast of Panama), where 60 rough-toothed dolphins (Steno bredanensis) stranded, including ten which died on the beach.
 Objective: To assess total mercury (THg) concentrations, and δ13C and δ15N stable isotope values in rough-toothed dolphins for the first time in this region.
 Methods: Nine skin samples were collected from adults, stored in 70 % ethanol, and posteriorly analyzed to determine THg concentrations and stable isotope values.
 Results: THg concentrations ranged from 4 764 to 18 689 ng g-1 dry weight (dw) (mean = 12 841; SD = 5 083 ng g-1 dw), δ13C values ranged between −16.8 and −15.2 ‰ (mean = −16.2; SD = 0.6 ‰), and δ15N values ranged between 14.3 and 15.9 ‰ (mean = 15.0; SD = 0.5 ‰).
 Conclusions: High THg concentrations reported for this species in the Azuero Peninsula are consistent with values reported for rough-toothed dolphins in other areas worldwide, such as the central-northern Rio de Janeiro State in Brazil and La Guajira in the Colombian Caribbean. Elevated mercury (Hg) concentrations may be related to the rough-toothed dolphin diet, which according to δ15N values found here, appears to be based mainly on high trophic level prey that bioaccumulate more Hg in their tissues compared to lower trophic level organisms. However, additional dietary studies would be required to support these findings. Continuing monitoring of traditional dietary analysis, as well as contamination levels in fish and dolphins, is necessary to understand the dolphins’ ecotoxicology in Panama.

Differences in the whistles of two ecotypes of bottlenose dolphins from the Gulf of California.

Intraspecific whistle variation in dolphins has been explained in the context of geographical isolation, soundscape, and social structure. Whistles of two ecotypes of bottlenose dolphins from La Paz Bay, Gulf of California, were analyzed. Both ecotypes showed similar whistle contours. However, they could be identified by contour maximum frequency which was predominantly above 15 kHz for oceanic dolphins and below 15 kHz for coastal dolphins. The different whistle frequencies of the two ecotypes could result from differences in group size and acoustic characteristics of their respective habitats, suggesting potential future passive acoustic monitoring applications.

How does habitat influence metabolism? Clues from biomarker response and contaminant profile in Tursiops truncatus (Montagu, 1821) subspecies living in parapatry

In western South America (WSA) two subspecies of bottlenose dolphin are recognized: Tursiops truncatus gephyreus, predominantly found in estuaries and river mouths, and Tursiops truncatus truncatus, occurring along the continental shelf. Despite a partial spatial overlap, both subspecies are considered to occupy different habitats and ecological niches. In the present study, chemical analyzes as well as biochemical and molecular biomarkers were used to investigate the influence of niche partitioning over metabolic pathways associated with the detoxification of persistent organic pollutants (POPs), antioxidant metabolism, immune activity and lipid metabolism in Tursiops truncatus subspecies living in parapatry. Overall, the profile and levels of bioaccumulated PCBs, pesticides and PBDEs were similar between groups, with a greater variety of pesticides, such as γ-HCHs, heptachlor, oxychlordane and o,p'DDT, detected in T. truncatus gephyreus. Multivariate analysis of variance (MANOVA) and non-metric multidimensional scaling (NMDS) results indicated that glutathione reductase (GR) and superoxide dismutase (SOD) enzymatic activities were higher in coastal dolphins, as were the mRNA levels of metallothionein 2A (MT2A), interleukin-1α (IL-1α), ceramide synthase 3 (CERS3) and fatty acid elongase (ELOVL4). In parallel, mRNA levels of fatty acid synthase complex 1 (FASN 1) were higher in oceanic dolphins. These findings suggest that, due to their occurrence in coastal habitats, T. truncatus gephyreus is more exposed to environmental pollutants and pathogenic microorganisms. Likewise, niche partitioning may influence lipid biosynthesis, possibly due to differences on feeding habits, reflecting in an enhanced long chain ceramides biosynthesis in T. truncatus gephyreus. Collectively, these data reinforce the need to address habitat specificities in conservation efforts, since distinct groups can be facing different anthropogenic pressures in WSA.

Aquatic wild meat consumption of cetaceans in São Tomé and Príncipe (Gulf of Guinea)

Relatively little is known about cetaceans in São Tomé and Príncipe and in recent years, some episodes of cetacean bycatch and human consumption have been described. Aiming to provide insight into the reliance upon cetaceans as a source of meat, we present findings from complementary studies conducted in the islands of São Tomé and Príncipe from 2012 to 2021. Based on interviews with local residents, we assessed the prevalence of human consumption of cetaceans and identified potential sociodemographic drivers. This study provides the first comprehensive assessment of harvest (direct remove or incidental take) and consumption of cetaceans (mainly the family Delphinidae) in the country. These findings will be useful in the development of regional conservation measures and contribute to our understanding of anthropogenic activities affecting cetacean populations in the Gulf of Guinea.

Alzheimer's disease-like neuropathology in three species of oceanic dolphin.

Alzheimer's disease (AD) is the most common neurodegenerative disease and the primary cause of disability and dependency among elderly humans worldwide. AD is thought to be a disease unique to humans although several other animals develop some aspects of AD-like pathology. Odontocetes (toothed whales) share traits with humans that suggest they may be susceptible to AD. The brains of 22 stranded odontocetes of five different species were examined using immunohistochemistry to investigate the presence or absence of neuropathological hallmarks of AD: amyloid-beta plaques, phospho-tau accumulation and gliosis. Immunohistochemistry revealed that all aged animals accumulated amyloid plaque pathology. In three animals of three different species of odontocete, there was co-occurrence of amyloid-beta plaques, intraneuronal accumulation of hyperphosphorylated tau, neuropil threads and neuritic plaques. One animal showed well-developed neuropil threads, phospho-tau accumulation and neuritic plaques, but no amyloid plaques. Microglia and astrocytes were present as expected in all brain samples examined, but we observed differences in cell morphology and numbers between individual animals. The simultaneous occurrence of amyloid-beta plaques and hyperphosphorylated tau pathology in the brains of odontocetes shows that these three species develop AD-like neuropathology spontaneously. The significance of this pathology with respect to the health and, ultimately, death of the animals remains to be determined. However, it may contribute to the cause(s) of unexplained live-stranding in some odontocete species and supports the 'sick-leader' theory whereby healthy conspecifics in a pod mass strand due to high social cohesion.

Modeling the Evolution of Rates of Continuous Trait Evolution.

Rates of phenotypic evolution vary markedly across the tree of life, from the accelerated evolution apparent in adaptive radiations to the remarkable evolutionary stasis exhibited by so-called "living fossils". Such rate variation has important consequences for large-scale evolutionary dynamics, generating vast disparities in phenotypic diversity across space, time, and taxa. Despite this, most methods for estimating trait evolution rates assume rates vary deterministically with respect to some variable of interest or change infrequently during a clade's history. These assumptions may cause underfitting of trait evolution models and mislead hypothesis testing. Here, we develop a new trait evolution model that allows rates to vary gradually and stochastically across a clade. Further, we extend this model to accommodate generally decreasing or increasing rates over time, allowing for flexible modeling of "early/late bursts" of trait evolution. We implement a Bayesian method, termed "evolving rates" (evorates for short), to efficiently fit this model to comparative data. Through simulation, we demonstrate that evorates can reliably infer both how and in which lineages trait evolution rates varied during a clade's history. We apply this method to body size evolution in cetaceans, recovering substantial support for an overall slowdown in body size evolution over time with recent bursts among some oceanic dolphins and relative stasis among beaked whales of the genus Mesoplodon. These results unify and expand on previous research, demonstrating the empirical utility of evorates.

Changes in whistle parameters of two common bottlenose dolphin ecotypes as a result of the physical presence of the research vessel.

In the presence of vessels, dolphins have been found to change their habitat, behavior, group composition and whistle repertoire. The modification of the whistle parameters is generally considered to be a response to the engine noise. Little is known about the impact of the physical presence of vessels on dolphin acoustics. Whistle parameters of the coastal and oceanic ecotypes of common bottlenose dolphins in La Paz Bay, Mexico, were measured after the approach of the research vessel and its engine shutdown. Recordings of 10 min were made immediately after turning off the engine. For analysis, these recordings were divided from minute 0 to minute 5, and from minute 5:01 to minute 10. The whistles of the oceanic ecotype showed higher maximum, minimum and peak frequency in the second time interval compared to the first one. The whistle rate decreased in the second time interval. The whistles of the coastal ecotype showed no difference between the two time intervals. The physical presence of the research vessel could have induced a change in the whistle parameters of the oceanic dolphins until habituation to the vessel disturbance. The oceanic ecotype could increase the whistle rate and decrease the whistle frequencies to maintain acoustic contact more frequently and for longer distances. The coastal ecotype, showing no significant changes in the whistle parameters, could be more habituated to the presence of vessels and display a higher tolerance.

Toothed Whales Have Black Neurons in the Blue Spot.

Simple SummaryNeuromelanin is a dark pigment that is present in several types of neurons of the brain. The role of human neuromelanin is a matter of controversy and, over the past few years, has been attributed to having a dual nature, either in a protective role to shield neurons from toxic compounds, or as a trigger of neuroinflammation. This pigment has been researched mainly in the human brain, but it has also been found in the neurons of monkeys, horses, giraffes, cattle, sheep, goats, dogs, rats, and even in frogs and tadpoles. Even so, neuromelanin in humans and primates presents unique features that are not shown in other animals. A study on the morphology of the locus ceruleus (a key brain structure) of the family Delphinidae highlighted the presence of a large amount of neuromelanin within this brain area. In an attempt to better define the ultrastructure of neuromelanin in toothed whales, two brain specimens of the suborder Odontoceti were investigated. The two toothed whales that were examined presented melanin granules associated with lipid droplets and membranes that bore a striking resemblance with human neuromelanin. Its accumulation takes place over the entire life span, and appears to contain the story of one’s life exposure to several endogenous and environmental metals and/or compounds.Neuromelanin (NM) is a dark polymer pigment that is located mostly in the human substantia nigra, and in the locus ceruleus, referred to as “the blue spot”. NM increases linearly with age, and has been described mainly in the human brain; however, it also occurs in the neurons of monkeys, horses, giraffes, cattle, sheep, goats, dogs, rats, and even in frogs. While in most of these mammals NM shows the histochemical and ultrastructural features typical of lipofuscins, human NM is confined within cytoplasmic organelles that are surrounded by a double membrane, suggesting an autophagic origin. In a study on the morphology of the locus ceruleus of the family Delphinidae, the presence of a variable quantity of NM in the interior of locus ceruleus neurons was observed for the first time; meanwhile, nothing is known about its ultrastructure and composition. Transmission electron microscopy demonstrated in two toothed whales—an Atlantic spotted dolphin (Stenella frontalis; family Delphinidae) and a Blainville’s beaked whale (Mesoplodon densirostris; family Ziphiidae)—the presence of melanin granules associated with lipid droplets and membranes that were very similar to that of human NM. The relationship between NM and neuronal vulnerability must be studied in depth, and cetaceans may offer a new natural-spontaneous comparative model for the study of NM and its implication in neurodegenerative diseases.

The ontogeny of asymmetry in echolocating whales.

Extreme asymmetry of the skull is one of the most distinctive traits that characterizes toothed whales (Odontoceti, Cetacea). The origin and function of cranial asymmetry are connected to the evolution of echolocation, the ability to use high-frequency sounds to navigate the surrounding environment. Although this novel phenotype must arise through changes in cranial development, the ontogeny of cetacean asymmetry has never been investigated. Here we use three-dimensional geometric morphometrics to quantify the changes in degree of asymmetry and skull shape during prenatal and postnatal ontogeny for five genera spanning odontocete diversity (oceanic dolphins, porpoises and beluga). Asymmetry in early ontogeny starts low and tracks phylogenetic relatedness of taxa. Distantly related taxa that share aspects of their ecology overwrite these initial differences via heterochronic shifts, ultimately converging on comparable high levels of skull asymmetry. Porpoises maintain low levels of asymmetry into maturity and present a decelerated rate of growth, probably retained from the ancestral condition. Ancestral state reconstruction of allometric trajectories demonstrates that both paedomorphism and peramorphism contribute to cranial shape diversity across odontocetes. This study provides a striking example of how divergent developmental pathways can produce convergent ecological adaptations, even for some of the most unusual phenotypes exhibited among vertebrates.

The partial merger of two dolphin societies.

Interactions between mammalian social groups are generally antagonistic as individuals in groups cooperate to defend resources from non-members. Members of the family Delphinidae inhabit a three-dimensional habitat where resource defence is usually impractical. Here, we describe a long-term partial fusion of two communities of Atlantic spotted dolphins (Stenella frontalis). The northern community, studied for 30 years, immigrated 160 km to the range of the southern community, observed for 20 years. Both communities featured fission–fusion grouping patterns, strongest associations between adult males, and frequent affiliative contact between individuals. For the 5-year period following the immigration, we found members of all age classes and both sexes in mixed groups, but there was a strong bias toward finding immigrant males in mixed groups. Some association levels between males, and males and females, from different communities were as high as the highest within-community associations. Affiliative contacts indicate that these individuals were forming social relationships. The mixing of two separate social groups with new bond formation is rare in terrestrial mammal groups. Such mixing between spotted dolphin groups suggests that adaptations to respond aggressively to ‘outsiders’ are diminished in this species and possibly other ecologically similar dolphins.

A new early Miocene archaic dolphin (Odontoceti, Cetacea) from New Zealand, and brain evolution of the Odontoceti

ABSTRACT The major increase in cetacean brain size happened in the middle Miocene, about 15 million years ago, and involved the modern oceanic dolphin lineage Delphinoidea. In this paper, we describe a new specimen of an archaic dolphin, aff. Prosqualodon davidis, from the Gee Greensand, near Oamaru, New Zealand. The specimen is from the early Miocene, approximately 23–19–18 Ma, and includes an incomplete cranium, endocast, teeth, and some postcranial elements. Comparison of the median sulcus among previously reported endocasts and brains of the Cetacea suggests that the cerebrum was expanded incipiently in the Odontoceti from the early Miocene in the Eurhinodelphinidae + Delphinoidea lineage. Conversely, the olfactory fossa has been reduced in Odontoceti, and completely lost in at least two clades (Delphinoidea and Platanista gangetica), because olfaction was most likely no longer an essential sense for animals spending their life in water. Reduction and loss of the olfactory fossa saved energy, which may be related to the expansion of the cerebrum and cortex in an Odontoceti lineage that led to modern oceanic dolphins. The early Miocene appears to have been a transitional period for archaic and modern-type odontocetes, which were possibly and partly separated by these differences.

Vertebral Trabecular Bone Mechanical Properties Vary Among Functional Groups of Cetaceans.

SynopsisSince their appearance in the fossil record 34 million years ago, modern cetaceans (dolphins, whales, and porpoises) have radiated into diverse habitats circumglobally, developing vast phenotypic variations among species. Traits such as skeletal morphology and ecologically linked behaviors denote swimming activity; trade-offs in flexibility and rigidity along the vertebral column determine patterns of caudal oscillation. Here, we categorized 10 species of cetaceans (families Delphinidae and Kogiidae; N = 21 animals) into functional groups based on vertebral centra morphology, swimming speeds, diving behavior, and inferred swimming patterns. We quantified trabecular bone mechanical properties (yield strength, apparent stiffness, and resilience) among functional groups and regions of the vertebral column (thoracic, lumbar, and caudal). We extracted 6 mm3 samples from vertebral bodies and tested them in compression in 3 orientations (rostrocaudal, dorsoventral, and mediolateral) at 2 mm min−1. Overall, bone from the pre-fluke/fluke boundary had the greatest yield strength and resilience, indicating that the greatest forces are translated to the tail during caudal oscillatory swimming. Group 1, composed of 5 shallow-diving delphinid species, had the greatest vertebral trabecular bone yield strength, apparent stiffness, and resilience of all functional groups. Conversely, Group 3, composed of 2 deep-diving kogiid species, had the least strong, stiff, and resilient bone, while Group 2 (3 deep-diving delphinid species) exhibited intermediate values. These data suggest that species that incorporate prolonged glides during deep descents in the water column actively swim less, and place relatively smaller loads on their vertebral columns, compared with species that execute shallower dives. We found that cetacean vertebral trabecular bone properties differed from the properties of terrestrial mammals; for every given bone strength, cetacean bone was less stiff by comparison. This relative lack of material rigidity within vertebral bone may be attributed to the non-weight-bearing locomotor modes of fully aquatic mammals.

Extensive Interspecific Gene Flow Shaped Complex Evolutionary History and Underestimated Species Diversity in Rapidly Radiated Dolphins

Recently diverged taxa are often characterized by high rates of hybridization, which can complicate phylogenetic reconstruction. For this reason, the phylogenetic relationships and evolutionary history of dolphins are still not very well resolved; the question of whether the genera Tursiops and Stenella are monophyletic is especially controversial. Here, we performed re-sequencing of six dolphin genomes and combined them with eight previously published dolphin SRA datasets and six whole-genome datasets to investigate the phylogenetic relationships of dolphins and test the monophyly hypothesis of Tursiops and Stenella. Phylogenetic reconstruction with the maximum likelihood and Bayesian methods of concatenated loci, as well as with coalescence analyses of sliding window trees, produced a concordant and well-supported tree. Our studies support the non-monophyletic status of Tursiops and Stenella because the species referred these genera do not form exclusive monophyletic clades. This suggests that the current taxonomy of both genera might not reflect their evolutionary history and may underestimate their diversity. A four-taxon D-statistic (ABBA-BABA) test, five-taxon DFOIL test, and tree-based PhyloNet analyses all showed extensive gene flow across dolphin species, which could explain the instability in resolving phylogenetic relationship of oceanic dolphins with different and limited markers. This study could be a good case to demonstrate how genomic data can reveal complex speciation and phylogeny in rapidly radiating animal groups.