Distribution and development of the highly specialized lipids in the sound reception systems of dolphins

Fat bodies in the heads of toothed whales, which serve to transmit and receive sound, represent extraordinary examples of physiological specialization in adipose tissues among mammals, yet we know surprisingly little about their biochemical composition. We describe the spatial distributions and development of unusual endogenous lipids (branched-chain [“iso”] molecules and wax esters) in the mandibular fat bodies of bottlenose dolphins (Tursiops truncatus) using an ontogenetic series (fetus to adult; n = 10). Although concentrations of iso-acids, iso-alcohols and waxes were lower in younger dolphins than in adults, the same relative spatial arrangement was present in all age classes, implying a set “pattern” of acoustic lipid distribution that is established very early in life. In all age classes, a small region of blubber overlying the lateral region contained unusually high concentrations of iso-acids, exhibiting a tenfold increase over “normal” adjacent blubber. Being chemically more similar to the acoustic fat bodies, this region may serve as an entry point for sound into the head. Developmental accumulations of some iso-acids and iso-alcohols occurred more rapidly than others, implying that not only are the spatial distributions of branched-chain molecules under extremely fine-scale control, but the regulatory mechanisms controlling acoustic lipid synthesis are also highly complex.     

Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow,Acoustically Complex Habitat

Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPa_pp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPa_pp. These source levels are 1–2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes.     

Auditory temporal resolution and evoked responses to pulsed sounds for the Yangtze finless porpoises (<Emphasis Type="Italic">Neophocaena phocaenoides asiaeorientalis</Emphasis>)

Temporal cues are important for some forms of auditory processing, such as echolocation. Among odontocetes (toothed whales, dolphins, and porpoises), it has been suggested that porpoises may have temporal processing abilities which differ from other odontocetes because of their relatively narrow auditory filters and longer duration echolocation signals. This study examined auditory temporal resolution in two Yangtze finless porpoises (Neophocaena phocaenoides asiaeorientalis) using auditory evoked potentials (AEPs) to measure: (a) rate following responses and modulation rate transfer function for 100 kHz centered pulse sounds and (b) hearing thresholds and response amplitudes generated by individual pulses of different durations. The animals followed pulses well at modulation rates up to 1,250 Hz, after which response amplitudes declined until extinguished beyond 2,500 Hz. The subjects had significantly better hearing thresholds for longer, narrower-band pulses similar to porpoise echolocation signals compared to brief, broadband sounds resembling dolphin clicks. Results indicate that the Yangtze finless porpoise follows individual acoustic signals at rates similar to other odontocetes tested. Relatively good sensitivity for longer duration, narrow-band signals suggests that finless porpoise hearing is well suited to detect their unique echolocation signals.     

Living in the fast lane: rapid development of the locomotor muscle in immature harbor porpoises (Phocoena phocoena)

Cetaceans (dolphins and whales) are born into the aquatic environment and are immediately challenged by the demands of hypoxia and exercise. This should promote rapid development of the muscle biochemistry that supports diving, but previous research on two odontocete (toothed whales and dolphins) species showed protracted postnatal development for myoglobin content and buffering capacity. A minimum of 1 and 1.5 years were required for Fraser’s (Lagenodelphis hosei) and bottlenose (Tursiops truncatus) dolphins to obtain mature myoglobin contents, respectively; this corresponded to their lengthy 2 and 2.5-year calving intervals (a proxy for the dependency period of cetacean calves). To further examine the correlation between the durations for muscle maturation and maternal dependency, we measured myoglobin content and buffering capacity in the main locomotor muscle (longissimus dorsi) of harbor porpoises (Phocoena phocoena), a species with a comparatively short calving interval (1.5 years). We found that at birth, porpoises had 51 and 69 % of adult levels for myoglobin and buffering capacity, respectively, demonstrating greater muscle maturity at birth than that found previously for neonatal bottlenose dolphins (10 and 65 %, respectively). Porpoises achieved adult levels for myoglobin and buffering capacity by 9–10 months and 2–3 years postpartum, respectively. This muscle maturation occurred at an earlier age than that found previously for the dolphin species. These results support the observation that variability in the duration for muscular development is associated with disparate life history patterns across odontocetes, suggesting that the pace of muscle maturation is not solely influenced by exposure to hypoxia and exercise. Though the mechanism that drives this variability remains unknown, nonetheless, these results highlight the importance of documenting the species-specific physiological development that limits diving capabilities and ultimately defines habitat utilization patterns across age classes.     

High concentrations of isovaleric acid in the fats of odontocetes: variation and patterns of accumulation in blubber vs. stability in the melon

Isovaleric acid (iso5:0) is an unusual fatty acid that is important for echolocation and hearing in acoustic tissues of some odontocetes, but its functional significance in blubber is unknown. We examined patterns of accumulation of this compound in blubber in 30 species of odontocetes ( n=299). Iso5:0 concentrations in blubber varied with phylogeny, ontogeny and body topography. Iso5:0 accumulated in greater quantities in superficial/outer blubber than in deep/inner blubber. In the outer blubber of northern right whale and Hector's dolphins, iso5:0 accounted for one-third to one-half of all fatty acids. Total blubber burden of iso5:0 in harbour porpoises represented up to 15 times the amount deposited in the melon. The composition of the melon does not change during starvation in harbour porpoises, supporting the hypothesis that lipids in melon are conserved for a specific function. Some odontocetes continually deposit iso5:0 in blubber after levels in melon have reached asymptotic levels, suggesting independent control of iso5:0 synthesis and storage in these compartments. Dolphins and porpoises inhabiting cold waters possess higher concentrations of iso5:0 in their outer blubber layers than species from warmer regions. We propose that this relationship represents an adaptive secondary role for iso5:0 in maintaining blubber flexibility in cold environments.     

Microvascular characteristics of the acoustic fats: Novel data suggesting taxonomic differences between deep and shallow‐diving odontocetes

Odontocetes have specialized mandibular fats, the extramandibular (EMFB) and intramandibular fat bodies (IMFB), which function as acoustic organs, receiving and channeling sound to the ear during hearing and echolocation. Recent strandings of beaked whales suggest that these fat bodies are susceptible to nitrogen (N₂) gas embolism and empirical evidence has shown that the N₂ solubility of these fat bodies is higher than that of blubber. Since N₂ gas will diffuse from blood into tissue at any blood/tissue interface and potentially form gas bubbles upon decompression, it is imperative to understand the extent of microvascularity in these specialized acoustic fats so that risk of embolism formation when diving can be estimated. Microvascular density was determined in the EMFB, IMFB, and blubber from 11 species representing three odontocete families. In all cases, the acoustic tissues had less (typically 1/3 to 1/2) microvasculature than did blubber, suggesting that capillary density in the acoustic tissues may be more constrained than in the blubber. However, even within these constraints there were clear phylogenetic differences. Ziphiid (Mesoplodon and Ziphius, 0.9 ± 0.4% and 0.7 ± 0.3% for EMFB and IMFB, respectively) and Kogiid families (1.2 ± 0.2% and 1.0 ± 0.01% for EMFB and IMFB, respectively) had significantly lower mean microvascular densities in the acoustic fats compared to the Delphinid species (Tursiops, Grampus, Stenella, and Globicephala, 1.3 ± 0.3% and 1.3 ± 0.3% for EMFB and IMFB, respectively). Overall, deep‐diving beaked whales had less microvascularity in both mandibular fats and blubber compared to the shallow‐diving Delphinids, which might suggest that there are differences in the N₂ dynamics associated with diving regime, phylogeny, and tissue type. These novel data should be incorporated into diving physiology models to further understand potential functional disruption of the acoustic tissues due to changes in normal diving behavior.     

Paedomorphosis in two small species of toothed whales (Odontoceti): how and why?

To evaluate and assess the ontogenetic background for paedomorphosis in phocoenids, samples of 144 harbour porpoises, 81 white‐beaked dolphins, and 130 Commerson's dolphins were compared in terms of the development of epiphyseal fusion, cranial suture fusion, and ontogeny of cranial shape. Harbour porpoises and Commerson's dolphins terminated growth and development of all investigated traits sooner than white‐beaked dolphins, leading to lesser degrees of fusion of skeletal elements and less postnatal allometric development. The latter occurred even though shape in the two paedomorphic species developed at twice the rate relative to the size of white‐beaked dolphins. These observations imply that progenetic evolution has occurred convergently in phocoenid and Cephalorhynchus ancestors. The truncated ontogenies allow sexual maturity to be attained earlier and provide a greater reproductive potential. Both species inhabit similar temperate productive habitats and, hence, ecological factors are proposed to have supplied the selection pressures leading to progenesis. Constant prey availability must be a prerequisite for the observed phenomena because frequent food‐intake is necessitated by the limited capacity for energy storage and high heat‐loss entailed by the resulting small body sizes. Progenesis has rarely been proposed in mammal species. This may reflect rarity or that mammalian expressions of progenesis are less obvious. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 278–295.     

A New Acoustic Portal into the Odontocete Ear and Vibrational Analysis of the Tympanoperiotic Complex

Global concern over the possible deleterious effects of noise on marine organisms was catalyzed when toothed whales stranded and died in the presence of high intensity sound. The lack of knowledge about mechanisms of hearing in toothed whales prompted our group to study the anatomy and build a finite element model to simulate sound reception in odontocetes. The primary auditory pathway in toothed whales is an evolutionary novelty, compensating for the impedance mismatch experienced by whale ancestors as they moved from hearing in air to hearing in water. The mechanism by which high-frequency vibrations pass from the low density fats of the lower jaw into the dense bones of the auditory apparatus is a key to understanding odontocete hearing. Here we identify a new acoustic portal into the ear complex, the tympanoperiotic complex (TPC) and a plausible mechanism by which sound is transduced into the bony components. We reveal the intact anatomic geometry using CT scanning, and test functional preconceptions using finite element modeling and vibrational analysis. We show that the mandibular fat bodies bifurcate posteriorly, attaching to the TPC in two distinct locations. The smaller branch is an inconspicuous, previously undescribed channel, a cone-shaped fat body that fits into a thin-walled bony funnel just anterior to the sigmoid process of the TPC. The TPC also contains regions of thin translucent bone that define zones of differential flexibility, enabling the TPC to bend in response to sound pressure, thus providing a mechanism for vibrations to pass through the ossicular chain. The techniques used to discover the new acoustic portal in toothed whales, provide a means to decipher auditory filtering, beam formation, impedance matching, and transduction. These tools can also be used to address concerns about the potential deleterious effects of high-intensity sound in a broad spectrum of marine organisms, from whales to fish.     

The trouble with flippers: a report on the prevalence of digital anomalies in Cetacea

The forelimbs of cetaceans (whales, dolphins, and porpoises) are unique among mammals as the digits exhibit hyperphalangy, and the entire limb is encased in a soft tissue flipper that functions to generate lift. The typical morphology of cetacean digits has been well documented by detailed anatomical studies. This study however furthers our understanding of cetacean forelimb anatomy by conducting a taxonomically broad survey of cetacean digital anomalies. Forelimb radiographs from museum collections provided the basis upon which we calculated the prevalence and documented the morphology of cetacean digital abnormalities. Results indicated that 11% (n = 255) of toothed whales displayed some type of aberrant ossification: the majority of these cases displayed a fusion of elements within a single digital ray, whereas cases exhibiting branched digits were rare. A small sample of baleen whale radiographs (n = 6) contained the only documented case of baleen whale polydactyly in a specimen of the gray whale (Eschrichtius). Furthermore, some Balaenoptera specimens displayed ossified elements within the interdigital spaces that lacked attachment to the adjacent digits and carpus. In addition, we speculated on the role that several genes may have played in creating cetacean digital anomalies. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 722–735.     

Post-mortem investigations on stranded dolphins and porpoises from Hong Kong waters

Stranded cetaceans reported from the territorial waters of Hong Kong during the period May 1993 to March 1998 were examined to establish factors that may have contributed to their death. During the current study, 28 Indo-Pacific hump-backed dolphins (Sousa chinensis), 32 finless porpoises (Neophocaena phocaenoides), and four bottlenose dolphins (Tursiops truncatus) were necropsied. Bacteria (15 species) were isolated from nine animals. Of these bacteria, 47% were of possible fecal origin reflecting the high level of sewage contamination in Hong Kong's waters. One finless porpoise displayed wounds caused by a shark attack, and two female finless porpoises presented prolapsed uteri. At least 10 finless porpoises showed evidence of moderate to heavy lungworm infections (Halocercus pingi), and this appears to have been a factor contributing to death in at least six animals. Evidence suggesting blunt traumatic injury (probably caused by boat collisions) was found in six cetaceans (three finless porpoises and three hump-backed dolphins). Signs of fishery-related mortality were detected in at least nine animals (six hump-backed dolphins, two finless porpoises, and one bottlenose dolphin). Of these two human-caused mortality types, pre-existing disease or bacterial infection were detected in 29% of cases. Results indicate that human factors may have played a significant role in the death of at least 15 animals (32% of hump-backed dolphins, 15% of finless porpoises, and 25% of bottlenose dolphins).     

Evidence for infanticide in bottlenose dolphins: an explanation for violent interactions with harbour porpoises?

Most harbour porpoises found dead on the north-east coast of Scotland show signs of attack by sympatric bottlenose dolphins, but the reason(s) for these violent interactions remain(s) unclear. Post-mortem examinations of stranded bottlenose dolphins indicate that five out of eight young calves from this same area were also killed by bottlenose dolphins. These data, together with direct observations of an aggressive interaction between an adult bottlenose dolphin and a dead bottlenose dolphin calf, provide strong evidence for infanticide in this population. The similarity in the size range of harbour porpoises and dolphin calves that showed signs of attack by bottlenose dolphins suggests that previously reported interspecific interactions could be related to this infanticidal behaviour. These findings appear to provide the first evidence of infanticide in cetaceans (whales, dolphins and porpoises). We suggest that infanticide must be considered as a factor shaping sociality in this and other species of cetaceans, and may have serious consequences for the viability of small populations.     

Bayesian inference of cetacean phylogeny based on mitochondrial genomes

The phylogeny of Cetacea (whales, dolphins, porpoises) has long attracted the interests of biologists and has been investigated by many researchers based on different datasets. However, some phylogenetic relationships within Cetacea still remain controversial. In this study, Bayesian analyses were performed to infer the phylogeny of 25 representative species within Cetacea based on their mitochondrial genomes for the first time. The analyses recovered the clades resolved by the previous studies and strongly supported most of the current cetacean classifications, such as the monophyly of Odontoceti (toothed whales) and Mysticeti (baleen whales). The analyses provided a reliable and comprehensive phylogeny of Cetacea which can provide a foundation for further exploration of cetacean ecology, conservation and biology. The results also showed that: (i) the mitochondrial genomes were very informative for inferring phylogeny of Cetacea; and (ii) the Bayesian analyses outperformed other phylogenetic methods on inferring mitochondrial genome-based phylogeny of Cetacea.     

The extracranial arterial system in the heads of beaked whales,with implications on diving physiology and pathogenesis

Beaked whales are medium‐sized toothed whales that inhabit depths beyond the continental shelf; thus beaked whale strandings are relatively infrequent compared to those of other cetaceans. Beaked whales have been catapulted into the spotlight by their tendency to strand in association with naval sonar deployment. Studies have shown the presence of gas and fat emboli within the tissues and analysis of gas emboli is suggestive of nitrogen as the primary component. These findings are consistent with human decompression sickness (DCS) previously not thought possible in cetaceans. Because, tissue loading with nitrogen gas is paramount for the manifestation of DCS and nitrogen loading depends largely on the vascular perfusion of the tissues, we examined the anatomy of the extracranial arterial system using stranded carcasses of 16 beaked whales from five different species. Anatomic regions containing lipid and/or air spaces were prioritized as potential locations of nitrogen gas absorption due to the known solubility of nitrogen in adipose tissue and the nitrogen content of air, respectively. Attention was focused on the acoustic fat bodies and accessory sinus system on the ventral head. We found much of the arterial system of the head to contain arteries homologous to those found in domestic mammals. Robust arterial associations with lipid depots and air spaces occurred within the acoustic fat bodies of the lower jaw and pterygoid air sacs of the ventral head, respectively. Both regions contained extensive trabecular geometry with small arteries investing the trabeculae. Our findings suggest the presence of considerable surface area between the arterial system, and the intramandibular fat bodies and pterygoid air sacs. Our observations may provide support for the hypothesis that these structures play an important role in the exchange of nitrogen gas during diving. J. Morphol. 277:5–33, 2016. © 2015 Wiley Periodicals, Inc.     

Frequent Loss and Alteration of the MOXD2 Gene in Catarrhines and Whales: A Possible Connection with the Evolution of Olfaction

The MOXD2 gene encodes a membrane-bound monooxygenase similar to dopamine-β-hydroxylase, and has been proposed to be associated with olfaction. In this study, we analyzed MOXD2 genes from 64 mammalian species, and identified loss-of-function mutations in apes (humans, Sumatran and Bornean orangutans, and five gibbon species from the four major gibbon genera), toothed whales (killer whales, bottlenose dolphins, finless porpoises, baijis, and sperm whales), and baleen whales (minke whales and fin whales). We also identified a shared 13-nt deletion in the last exon of Old World cercopithecine monkeys that results in conversion of a membrane-bound protein to a soluble form. We hypothesize that the frequent inactivation and alteration of MOXD2 genes in catarrhines and whales may be associated with the evolution of olfaction in these clades.     

Living with dolphins: Local ecological knowledge and perceptions of small cetaceans along the Sindhudurg coastline of Maharashtra,India

Two near shore small cetaceans occur commonly along the Maharashtra coast, the Indian Ocean humpback dolphin and Indo Pacific finless porpoise. These cetaceans frequently interact with fisheries in this region due to overlap in space and resource use. Besides stranding records, little ecological information is available about these species from Maharashtra. We conducted 143 semistructured interviews to document local ecological knowledge and community perceptions of small cetaceans in 30 coastal fishing villages in Sindhudurg. Perceptions of finless porpoises were largely neutral, whereas humpback dolphins were negative. A classification regression tree (CART) analysis (root node error: 60%) showed that the annual cost of gear damage was an important predictor variable of humpback dolphin perceptions, followed by occupation (gear type) and age. Entanglements were reported for both species in large and small gill nets, and shore seines. Perceived net damage and catch loss due to humpback dolphins was six times greater than that of finless porpoises. However, finless porpoises were reportedly more frequently entangled in gear than humpback dolphins. We provide an insight into the perceptions of cetaceans in the local community and the fisheries‐cetacean interactions that shape them.     

Population biology, social behavior and communication in whales and dolphins

The baleen whales differ from the toothed whales and dolphins in life history and in social organization. Even though they grow to a larger size, young baleen whales tend to develop more rapidly than dolphins and toothed whales. Except for the mother-calf bond, most groups of baleen whales are short-lived, lasting only for hours, and individual-specific associations appear to be exceptions to the norm. Most toothed whales live in more structured groups, in which young animals have a long period of dependency and social learning. The communication signals described for different cetacean species have functions suited to the interactions that predominate in their societies.     

Evidence for infanticide in bottlenose dolphins of the western North Atlantic

Nine bottlenose dolphin (Tursiops truncatus) calves that stranded in Virginia in 1996 and 1997 died of severe blunt-force trauma. Injuries were concentrated on the head and chest and multiple rib fractures, lung lacerations, and soft tissue contusions were prominent. Skeletal and/or soft tissue trauma occurred bilaterally in all of the calves. One had a bite wound across the left mandible that exhibited deep punctures consistent with the tooth placement in an adult bottlenose dolphin. The lesions were not compatible with predation, boat strike, fisheries interactions, rough-surf injury, or blast injury. However, they were similar to traumatic injuries described in stranded bottlenose dolphin calves and harbor porpoises (Phocoena phocoena) in Great Britain attributed to violent dolphin interactions. The evidence suggests that violent dolphin behavior was the cause of the trauma in the nine calves reported here and that infanticide occurs in bottlenose dolphins of the western North Atlantic.     

Toothed whale monophyly reassessed by SINE insertion analysis: the absence of lineage sorting effects suggests a small population of a common ancestral species

Morphological data have indicated that toothed whales form a monophyletic group. However, research published in the last several years has made the issue of the monophyly or paraphyly of toothed whales a subject of debate. Our group previously characterized three independent loci in which SINE insertions were shared among dolphins and sperm whales, thus supporting the traditional, morphologically based hypothesis of toothed whale monophyly. Although in recent years a few additional molecular works proposed this topology, there is still skepticism over this monophyly from the view point of molecular systematics. When the phylogeny of rapidly radiated taxa is examined using the SINE method, it is important to consider the ascertainment bias that arises when choosing a particular taxon for SINE loci screening. To overcome this methodological problem specific to the SINE method, we examined all possible topologies among sperm whales, dolphins and baleen whales by extensively screening SINE loci from species of all three lineages. We characterized nine independent SINE loci from the genomes of sperm whales and dolphins, all of which cluster sperm whales and dolphins but exclude baleen whales. Furthermore, we characterized ten independent loci from baleen whales, all of which were amplified in a common ancestor of these whales. From these observations, we conclude that toothed whales form a monophyletic group and that no ancestral SINE polymorphisms hinder their phylogenetic assignment despite the short divergence times of the major lineages of extant whales during evolution. These results suggest that a small population of common ancestors of all toothed whales ultimately diverged into the lineages of sperm whales and dolphins.     

How Can Dolphins Recognize Fish According to Their Echoes? A Statistical Analysis of Fish Echoes

Echo-based object classification is a fundamental task of animals that use a biosonar system. Dolphins and porpoises should be able to rely on echoes to discriminate a predator from a prey or to select a desired prey from an undesired object. Many studies have shown that dolphins and porpoises can discriminate between objects according to their echoes. All of these studies however, used unnatural objects that can be easily characterized in human terminologies (e.g., metallic spheres, disks, cylinders). In this work, we collected real fish echoes from many angles of acquisition using a sonar system that mimics the emission properties of dolphins and porpoises. We then tested two alternative statistical approaches in classifying these echoes. Our results suggest that fish species can be classified according to echoes returning from porpoise- and dolphin-like signals. These results suggest how dolphins and porpoises can classify fish based on their echoes and provide some insight as to which features might enable the classification.     

Observation and analysis of sonar signal generation in the bottlenose dolphin (Tursiops truncatus): Evidence for two sonar sources

Indirect evidence for multiple sonar signal generators in odontocetes exists within the published literature. To explore the long-standing controversy over the site of sonar signal generation, direct evidence was collected from three trained bottlenose dolphins (Tursiops truncatus) by simultaneously observing nasal tissue motion, internal nasal cavity pressure, and external acoustic pressure. High-speed video endoscopy revealed tissue motion within both sets of phonic lips, while two hydrophones measured acoustic pressure during biosonar target recognition. Small catheters measured air-pressure changes at various locations within the nasal passages and in the basicranial spaces. Video and acoustic records demonstrate that acoustic pulses can be generated along the phonic fissure by vibrating the phonic labia within each set of phonic lips. The left and right phonic lips are capable of operating independently or simultaneously. Air pressure in both bony nasal passages rose and fell synchronously, even if the activity patterns of the two phonic lips were different. Whistle production and increasing sound pressure levels are generally accompanied by increasing intranarial air pressure. One acoustic “click” occurred coincident with one oscillatory cycle of the phonic labia. Changes in the click repetition rate and cycles of the phonic labia were simultaneous, indicating that these events are coupled. Structural similarity in the nasal apparatus across the Odontoceti suggests that all extant toothed whales generate sonar signals using the phonic lips and similar biomechanical processes.