Genetic homogeneity in the face of morphological heterogeneity in the harbor porpoise from the Black Sea and adjacent waters (Phocoena phocoena relicta).

Absence of genetic differentiation is usually taken as an evidence of panmixia, but can also reflect other situations, including even nearly complete demographic independence among large-sized populations. Deciphering which situation applies has major practical implications (e.g., in conservation biology). The endangered harbor porpoises in the Black Sea illustrates this point well. While morphological heterogeneity suggested that population differentiation may exist between individuals from the Black and Azov seas, no genetic study provided conclusive evidence or covered the entire subspecies range. Here, we assessed the genetic structure at ten microsatellite loci and a 3904 base-pairs mitochondrial fragment in 144 porpoises across the subspecies range (i.e., Aegean, Marmara, Black, and Azov seas). Analyses of the genetic structure, including FST, Bayesian clustering, and multivariate analyses revealed a nearly complete genetic homogeneity. Power analyses rejected the possibility of underpowered analyses (power to detect FST ≥ 0.008 at microsatellite loci). Simulations under various demographic models, evaluating the evolution of FST, showed that a time-lag effect between demographic and genetic subdivision is also unlikely. With a realistic effective population size of 1000 individuals, the expected "gray zone" would be at most 20 generations under moderate levels of gene flow (≤10 migrants per generation). After excluding alternative hypotheses, panmixia remains the most likely hypothesis explaining the genetic homogeneity in the Black Sea porpoises. Morphological heterogeneity may thus reflect other processes than population subdivision (e.g., plasticity, selection). This study illustrates how combining empirical and theoretical approaches can contribute to understanding patterns of weak population structure in highly mobile marine species.

Biosonar signal propagation in the harbor porpoise's (Phocoena phocoena) head: The role of various structures in the formation of the vertical beam.

Harbor porpoises (Phocoena phocoena) use narrow band echolocation signals for detecting and locating prey and for spatial orientation. In this study, acoustic impedance values of tissues in the porpoise's head were calculated from computer tomography (CT) scan and the corresponding Hounsfield Units. A two-dimensional finite element model of the acoustic impedance was constructed based on CT scan data to simulate the acoustic propagation through the animal's head. The far field transmission beam pattern in the vertical plane and the waveforms of the receiving points around the forehead were compared with prior measurement results, the simulation results were qualitatively consistent with the measurement results. The role of the main structures in the head such as the air sacs, melon and skull in the acoustic propagation was investigated. The results showed that air sacs and skull are the major components to form the vertical beam. Additionally, both beam patterns and sound pressure of the sound waves through four positions deep inside the melon were demonstrated to show the role of the melon in the biosonar sound propagation processes in the vertical plane.

Phenotypic and genetic divergence among harbour porpoise populations associated with habitat regions in the North Sea and adjacent seas.

Determining the mechanisms that generate population structure is essential to the understanding of speciation and the evolution of biodiversity. Here, we investigate a geographical range that transects two habitat gradients, the North Sea to North Atlantic transition, and the temperate to subpolar regions. We studied the harbour porpoise (Phocoena phocoena), a small odontocete inhabiting both subpolar and temperate waters. To assess differentiation among putative populations, we measured morphological variation at cranial traits (N = 462 individuals) and variation at eight microsatellite loci for 338 of the same individuals from Norwegian, British and Danish waters. Significant morphological differentiation reflected the size of the buccal cavity. Porpoises forage in relatively shallow waters preying mainly on benthic species in British and Danish waters, and on mesopelagic and pelagic fish off the coast of Norway. We suggest that the observed differentiation may be explained by resource specialization and either adaptation or developmental responses to different local habitats.

Stereology of the neocortex in Odontocetes: qualitative, quantitative, and functional implications.

We investigated the quantitative morphology of the neocortex (gray matter) in 2 toothed whale (odontocete) species (harbor porpoise, Phocoena phocoena; bottlenose dolphin, Tursiops truncatus) with stereological methods. The 4 primary projection areas (motor, somatosensory, auditory, and visual fields) are analyzed for their cell densities in layers III and V with standard design-based stereology methods. Along cortical areas M1, S1, A1, and V1 in Tursiops, neuron density is always higher in layer III than in layer V, whereas the data in Phocoena are variable. Moreover, neuron density in layer III is generally around 1.5 times higher in Tursiops than in Phocoena. Maximal density values are seen in layer III of A1 and V1 in Tursiops and the ratio of layer III/layer V density is maximal in A1 of this species. Thus, layer III could have a higher capacity in the bottlenose dolphin with regard to intrinsic connectivity. Extant knowledge on toothed whale neurobiology and behavior suggests that quantitative/stereological differences between the 2 odontocete species regarding the neuron density of standard cortical units may be correlated with specific adaptations to their respective habitats. In contrast to layers V and VI which mainly serve as an executive system, layer III could represent an intermediate level in sensory and premotor processing which works more tangentially in the cortices via horizontal connections with other cortical areas, respectively. The generally higher density of cortical layer III in Tursiops suggests a higher connectivity of this layer in view of the more agile and complicated behavior of these gregarious animals including versatile phonation by complex sound and ultrasound signals.

Vertebral pattern variation in the North Sea harbor porpoise (Phocoena phocoena) by computed tomography.

Vertebral series in the harbor porpoise (Phocoena phocoena) include cervical, thoracic, lumbar, and caudal. In contrast to studying skeletons from museums, in which small bones can be missed, evaluation of full body computed tomography (CT) scans provides an overview of the vertebral column, while maintaining interrelationship of all structures. The aim of this study was to document variations in vertebral patterning of the harbor porpoise via evaluation of CT images of intact stranded harbor porpoises. The harbor porpoises were divided into age classes, based on developmental stage of reproductive organs on postmortem examination and closure of proximal humeral physis on CT. Numbers of vertebrae per series, fusion state of the syncervical, type of first hemal arch, number of double articulating ribs, and floating ribs were recorded based on CT images. Included in the study were 48 harbor porpoises (27 males and 21 females), which were divided in two age classes (27 immatures and 21 adults). Total vertebral count varied from 63 to 68 with vertebral formula range C7T12-14L12-16Cd29-33. Twenty-five different vertebral formulas were found, of which C7T13L14Ca30 was the most common (n = 8, 17%). Thoracic vertebrae with six, seven, or eight double articulating ribs and zero, one, or two vertebrae with floating ribs were seen. Four different fusion states of the syncervical and four types of hemal arches were recognized. This study showed a great variation in vertebral patterning in the harbor porpoise, with homeotic and meristic variation in the thoracic, lumbar, and caudal vertebral series.

Single source sound production and dynamic beam formation in echolocating harbour porpoises (Phocoena phocoena).

Echolocating toothed whales produce high-powered clicks by pneumatic actuation of phonic lips in their nasal complexes. All non-physeteroid toothed whales have two pairs of phonic lips allowing many of these species to produce both whistles and clicks at the same time. That has led to the hypothesis that toothed whales can increase the power outputs and bandwidths of clicks, and enable fast clicking and beam steering by acutely timed actuation of both phonic lip pairs simultaneously. Here we test that hypothesis by applying suction cup hydrophones on the sound-producing nasal complexes of three echolocating porpoises (Phocoena phocoena) with symmetrical pairs of phonic lips. Using time of arrival differences on three hydrophones, we show that all recorded clicks from these three porpoises are produced by the right pair of phonic lips with no evidence of simultaneous or independent actuation of the left pair. It is demonstrated that porpoises, despite actuation of only one sound source, can change their output and sound beam probably through conformation changes in the sound-producing soft tissues and nasal sacs, and that the coupling of the phonic lips and the melon acts as a waveguide for sound energy between 100 and 160 kHz to generate a forward-directed sound beam for echolocation.

Asymmetric and Spiraled Genitalia Coevolve with Unique Lateralized Mating Behavior.

Asymmetric genitalia and lateralized mating behaviors occur in several taxa, yet whether asymmetric morphology in one sex correlates or coevolves with lateralized mating behavior in the other sex remains largely unexplored. While lateralized mating behaviors are taxonomically widespread, among mammals they are only known in the harbor porpoise (Phocoena phocoena). Males attempt copulation by approaching a female exclusively on her left side. To understand if this unusual lateralized behavior may have coevolved with genital morphology, we quantified the shape of female and male harbor porpoise reproductive tracts using 2D geometric morphometrics and 3D models of the vaginal lumen and inflated distal penis. We found that the vaginas varied individually in shape and that the vaginas demonstrated both significant directional and fluctuating asymmetry. This asymmetry resulted from complex 3D spirals and vaginal folds with deep recesses, which may curtail the depth or direction of penile penetration and/or semen movement. The asymmetric shapes of the vaginal lumen and penis tip were both left-canted with similar angular bends that mirrored one another and correspond with the left lateral mating approach. We suggest that the reproductive anatomy of both sexes and their lateral mating behavior coevolved.

Testicular morphology and spermatogenesis in harbour porpoises (Phocoena phocoena).

Knowledge about reproductive parameters in male harbour porpoises such as testicular histology and germ cell maturation as well as seasonal changes in spermatogenesis is scarce. Thus, the aim of the present study was to report changes in the histological appearance of the testicular morphology of neonatal and juvenile harbour porpoises during maturation, to identify stages of spermatogenesis in adult males and to detect seasonal modifications. The identification of these stages can be used to assess the developmental profile of gene expression during spermatogenesis and to identify defects in spermatogenesis arising in pathological conditions. Testes of adult male harbour porpoises from the North and Baltic Sea that became stranded or by-caught in the years 1998-2016 were histologically examined using Haematoxylin and Eosin - staining. The Periodic Acid Schiff (PAS) staining was used for spermatogenic staging and the evaluation of the development of the acrosomic cap. For the identification of changes in testes morphology and morphometry during the course of the year, histological characteristics like germ cell associations and diameter of the convoluted seminiferous tubules were noted for each month. The analysis showed that in adult males more than one stage of spermatogenesis could be found per cross section of the convoluted seminiferous tubules similar to findings in men and some ape species. This rare phenomenon is called multi-stage-arrangement. In sexually active males from the peak breeding season (June and July) eight stages of spermatogenesis were identified and all stages occurred simultaneously, while during the low breeding season (August to May) only residual spermatogenesis or constituent germ cell populations were found. Missing germ cell generations were recorded in specimens from July to September. Our investigations provide a detailed staging of spermatogenesis and give new insight into the reproductive biology of male harbour porpoises. With these new basic parameters, indicators for endocrine disruptors can be developed in the future, aiming to detect how environmental factors could affect male fertility in wildlife.

Life history constrains biochemical development in the highly specialized odontocete echolocation system.

The vertebrate head has undergone enormous modification from the features borne by early ancestors. The growth of skull bones has been well studied in many species, yet little is known about corresponding soft tissue development. Among mammals, some of the most unusual examples of cranial evolution exist in the toothed whales (odontocetes). Specialized fat bodies in toothed whale heads play important roles in sound transmission and reception. These fat bodies contain unique endogenous lipids, with favourable acoustic properties, arranged in highly organized, three-dimensional patterns. We link variation in developmental rates of acoustic fats with life-history strategy, using bottlenose dolphins and harbour porpoises. Porpoise acoustic fats attain adult configurations earlier (less than 1 year) and at a faster pace than dolphins. The accelerated lipid accumulation in porpoises reflects the earlier need for fully functional echolocation systems. Dolphins enjoy 3-6 years of maternal care; porpoises must achieve total independence by approximately nine months. Further, a stereotypic 'blueprint' for the spatial distribution of lipids is established prior to birth, demonstrating the highly conserved nature of the intricate biochemical arrangement in acoustic tissues. This system illustrates an unusual case of soft tissue development being constrained by life history, rather than the more commonly observed mechanistic or phyletic constraints.

Calcium-binding proteins in the cerebellar cortex of the bottlenose dolphin and harbour porpoise.

Studying the distribution of Ca2+-binding proteins allows one to discover specific neuron chemotypes involved in the regulation of the activity of various neural elements. While extensive data exist on Ca2+-binding proteins in the nervous system, in particular, in the cerebellar cortex of terrestrial mammals, the localization of these proteins in the cerebellar cortex of marine mammals has not been studied. We studied the localization of calretinin, calbindin, and parvalbumin immunoreactivity in the cerebellar cortex of the bottlenose dolphin Tursiops truncates and harbour porpoise Phocoena phocoena. In both species, most Purkinje cells were calbindin-immunoreactive, while calretinin and parvalbumin were expressed in a small portion of Purkinje cells. In addition, calretinin-immunoreactive unipolar brush and granule cells and calbindin- and parvalbumin-immunoreactive basket, stellate, and Golgi cells were observed. Calretinin-immunoreactive corticopetal (mossy and climbing) fibers were found. Based on the length of the primary dendrite, short-, middle-, and long-dendrite unipolar brush cells could be distinguished. The validity of this classification was supported using cluster analysis suggesting the presence of several natural types of these cells. The distribution of Ca2+-binding proteins in the cerebellar cortex of the cetaceans studied was generally similar to that reported for terrestrial mammals, suggesting that this trait is evolutionarily conservative in mammals.

Comparative physiology of vocal musculature in two odontocetes, the bottlenose dolphin (Tursiops truncatus) and the harbor porpoise (Phocoena phocoena).

The mechanism by which odontocetes produce sound is unique among mammals. To gain insight into the physiological properties that support sound production in toothed whales, we examined myoglobin content ([Mb]), non-bicarbonate buffering capacity (ß), fiber-type profiles, and myosin heavy chain expression of vocal musculature in two odontocetes: the bottlenose dolphin (Tursiops truncatus; n = 4) and the harbor porpoise (Phocoena phocoena; n = 5). Both species use the same anatomical structures to produce sound, but differ markedly in their vocal repertoires. Tursiops produce both broadband clicks and tonal whistles, while Phocoena only produce higher frequency clicks. Specific muscles examined in this study included: (1) the nasal musculature around the phonic lips on the right (RNM) and left (LNM) sides of the head, (2) the palatopharyngeal sphincter (PPS), which surrounds the larynx and aids in pressurizing cranial air spaces, and (3) the genioglossus complex (GGC), a group of muscles positioned ventrally within the head. Overall, vocal muscles had significantly lower [Mb] and ß than locomotor muscles from the same species. The PPS was predominately composed of small diameter slow-twitch fibers. Fiber-type and myosin heavy chain analyses revealed that the GGC was comprised largely of fast-twitch fibers (Tursiops: 88.6%, Phocoena: 79.7%) and had the highest ß of all vocal muscles. Notably, there was a significant difference in [Mb] between the RNM and LNM in Tursiops, but not Phocoena. Our results reveal shared physiological characteristics of individual vocal muscles across species that enhance our understanding of key functional roles, as well as species-specific differences which appear to reflect differences in vocal capacities.

Comparative anatomical study of sound production and reception systems in the common dolphin (Delphinus delphis) and the harbour porpoise (Phocoena phocoena) heads.

Magnetic resonance imaging (MRI) and computed tomography (CT) scans were used to analyse, respectively, the soft tissues and the bones of the heads of four common dolphins and three harbour porpoises. This imaging study was completed by an examination of anatomical sections performed on two odontocete heads (a subadult common dolphin and a subadult harbour porpoise). The three complementary approaches allowed to illustrate anatomical differences in the echolocation systems of the common dolphin and the harbour porpoise. We captured images confirming strong differences of symmetry of the melon and of its connexions to the MLDB (Monkeys Lips/Dorsal Bursae) between the common dolphin and the harbour porpoise. The melon of the common dolphin is asymmetrically directly connected to the right bursae cantantes at its right side, whereas the melon of the harbour porpoise is symmetrical, and separated from the two bursae cantantes by a set of connective tissues. Another striking difference comes from the bursae cantantes themselves, less deeply located in the head of the common dolphin than in the harbour porpoise.

Phenotypical characterization of changes in thymus and spleen associated with lymphoid depletion in free-ranging harbor porpoises (Phocoena phocoena).

Harbor porpoises from the North and Baltic Seas exhibit a higher incidence of bacterial infections compared to whales from less polluted arctic waters. Furthermore, thymic atrophy and splenic depletion, associated with elevated concentrations of environmental contaminants, such as polychlorinated biphenyls (PCB) and polybrominated diphenyl ether (PBDE) body burdens, have been found in wildlife harbor porpoises. Thus, there is currently a debate about the potential adverse effect of xenobiotics on the immune system and therefore on the health status of this and other marine mammal species. The aim of the present study was to characterize phenotypical changes in lymphoid organs of harbor porpoises and their possible association with increased disease susceptibility due to an impaired immune response in this marine mammal species. Therefore, 29 by-caught and stranded harbor porpoises were necropsied and the health status was evaluated based upon the severity of main pathological findings. In addition, the distribution of CD2-, CD3epsilon-, and CD45R-positive cells as well as B lymphocytes, MHC class II-expressing and antigen-presenting cells was determined in the thymus and spleen using immunohistochemistry. Thymic atrophy and splenic depletion were associated with an impaired health status in investigated whales. Phenotypical changes in atrophic thymuses were characterized by a depletion of immature cortical thymocytes and medullary B cells. Furthermore, findings in depleted spleens were consistent with a loss of peripheral T lymphocytes in the periarteriolar lymphoid sheath (PALS). Based upon the results, an altered thymopoiesis and impaired cellular immune function cannot be excluded in whales with evidence of lymphoid depletion.

Vertebral bone microarchitecture and osteocyte characteristics of three toothed whale species with varying diving behaviour.

Although vertebral bone microarchitecture has been studied in various tetrapods, limited quantitative data are available on the structural and compositional changes of vertebrae in marine mammals. Whales exhibit exceptional swimming and diving behaviour, and they may not be immune to diving-associated bone pathologies. Lumbar vertebral bodies were analysed in three toothed whale species: the sperm whale (Physeter macrocephalus), orca (Orcinus orca) and harbour porpoise (Phocoena phocoena). The bone volume fraction (BV/TV) did not scale with body size, although the trabeculae were thicker, fewer in number and further apart in larger whale species than in the other two species. These parameters had a negative allometric scaling relationship with body length. In sperm whales and orcas, the analyses revealed a central ossification zone ("bone-within-bone") with an increased BV/TV and trabecular thickness. Furthermore, a large number of empty osteocyte lacunae was observed in the sperm whales. Quantitative backscattered electron imaging showed that the lacunae were significantly smaller and less densely packed. Our results indicate that whales have a unique vertebral bone morphology with an inside-out appearance and that deep diving may result in a small number of viable osteocytes because of diving depth-related osteocyte death.

Biological variables and health status affecting inorganic element concentrations in harbour porpoises (Phocoena phocoena) from Portugal (western Iberian Peninsula).

The coastal preferences of harbour porpoise (Phocoena phocoena) intensify their exposure to human activities. The harbour porpoise Iberian population is presently very small and information about the threats it endures is vital for the conservation efforts that are being implemented to avoid local extinction. The present study explored the possible relation between the accumulation of trace elements by porpoises and their sex, body length, nutritional state, presence of parasites and gross pathologies. The concentrations of arsenic (As), cadmium (Cd), copper (Cu), mercury (Hg), manganese (Mn), nickel (Ni), lead (Pb), zinc (Zn) and selenium (Se) were evaluated in 42 porpoises stranded in Portugal between 2005 and 2013. Considering European waters, porpoises stranded in Portugal present the highest Hg concentrations and the lowest Cd concentrations, which may reflect dietary preferences and the geographic availability of these pollutants. While no effect of sex on trace element concentrations was detected, there was a positive relationship between porpoise body length and the concentration of Cd, Hg and Pb. Animals in worse nutritional condition showed higher levels of Zn. Harbour porpoises with high parasite burdens showed lower levels of Zn and As in all analysed tissues and also lower levels of renal Ni, while those showing gross pathologies presented higher Zn and Hg levels. This is the first data on the relationship between trace elements and health-related variables in porpoises from southern European Atlantic waters, providing valuable baseline information about the contamination status of this vulnerable population.

Organization of the sleep-related neural systems in the brain of the harbour porpoise (Phocoena phocoena).

The present study provides the first systematic immunohistochemical neuroanatomical investigation of the systems involved in the control and regulation of sleep in an odontocete cetacean, the harbor porpoise (Phocoena phocoena). The odontocete cetaceans show an unusual form of mammalian sleep, with unihemispheric slow waves, suppressed REM sleep, and continuous bodily movement. All the neural elements involved in sleep regulation and control found in bihemispheric sleeping mammals were present in the harbor porpoise, with no specific nuclei being absent, and no novel nuclei being present. This qualitative similarity of nuclear organization relates to the cholinergic, noradrenergic, serotonergic, and orexinergic systems and is extended to the γ-aminobutyric acid (GABA)ergic elements involved with these nuclei. Quantitative analysis of the cholinergic and noradrenergic nuclei of the pontine region revealed that in comparison with other mammals, the numbers of pontine cholinergic (126,776) and noradrenergic (122,878) neurons are markedly higher than in other large-brained bihemispheric sleeping mammals. The diminutive telencephalic commissures (anterior commissure, corpus callosum, and hippocampal commissure) along with an enlarged posterior commissure and supernumerary pontine cholinergic and noradrenergic neurons indicate that the control of unihemispheric slow-wave sleep is likely to be a function of interpontine competition, facilitated through the posterior commissure, in response to unilateral telencephalic input related to the drive for sleep. In addition, an expanded peripheral division of the dorsal raphe nuclear complex appears likely to play a role in the suppression of REM sleep in odontocete cetaceans. Thus, the current study provides several clues to the understanding of the neural control of the unusual sleep phenomenology present in odontocete cetaceans. J. Comp. Neurol. 524:1999-2017, 2016. © 2016 Wiley Periodicals, Inc.

Characterization of the temporomandibular joint of the harbour porpoise (Phocoena phocoena) and Risso's dolphin (Grampus griseus).

OBJECTIVES: The temporomandibular joint (TMJ) in cetaceans is largely uncharacterized. This study aims to describe the macroscopic, microscopic, biochemical and biomechanical features of the TMJ of two species of the suborder Odontoceti: the harbour porpoise (Phocoena phocoena) and Risso's dolphin (Grampus griseus). Furthermore, we aim to elucidate the structure-function relationship of their TMJs and their possible role in echolocation. DESIGN: The TMJs from fresh cadaver heads of harbour porpoise (n=4) and Risso's dolphin (n=2) acquired from stranding were examined. Following macroscopical evaluation, the TMJs were investigated for their histological, mechanical and biochemical properties. RESULTS: The TMJs of the studied odontocetes were found to be fundamentally different from other mammals. Macroscopically, the TMJ lacks the typical joint cavity found in most mammals and is essentially a syndesmosis. Histological and microstructural analysis revealed that the TMJ discs were composed of haphazardly intersecting fibrous-connective tissue bundles separated by adipose tissue globules and various calibre blood vessels and nerve fibres. The collagen fibre composition was primarily collagen type I with lesser amounts of collagen type II. Sulphated glycosaminoglycan (sGAG) content was lower compared to other studied mammals. Finally, mechanical testing demonstrated the disc was stronger and stiffer in the dorsoventral direction than in the mediolateral direction. CONCLUSION: The spatial position of the TMJ, the absence of an articulating synovial joint, and the properties of the TMJ discs all reflect the unique suction-feeding mechanism adopted by the harbour porpoise and Risso's dolphin for underwater foraging. In addition, the presence of unique adipose globules, blood vessels and nerves throughout the discs may indicate a functional need beyond food apprehension. Instead, the disc may play a role in neurological sensory functions such as echolocation.

Range-dependent flexibility in the acoustic field of view of echolocating porpoises (Phocoena phocoena).

Toothed whales use sonar to detect, locate, and track prey. They adjust emitted sound intensity, auditory sensitivity and click rate to target range, and terminate prey pursuits with high-repetition-rate, low-intensity buzzes. However, their narrow acoustic field of view (FOV) is considered stable throughout target approach, which could facilitate prey escape at close-range. Here, we show that, like some bats, harbour porpoises can broaden their biosonar beam during the terminal phase of attack but, unlike bats, maintain the ability to change beamwidth within this phase. Based on video, MRI, and acoustic-tag recordings, we propose this flexibility is modulated by the melon and implemented to accommodate dynamic spatial relationships with prey and acoustic complexity of surroundings. Despite independent evolution and different means of sound generation and transmission, whales and bats adaptively change their FOV, suggesting that beamwidth flexibility has been an important driver in the evolution of echolocation for prey tracking.

Morphology and variation in porpoise (Cetacea: Phocoenidae) cranial endocasts.

Evolution of endocranial anatomy in cetaceans is important from the perspective of echolocation ability, intelligence, social structure, and alternate pathways for circulation to the brain. Apart from the importance of studying brain shape and asymmetries as they relate to aspects of behavior and intelligence, cranial endocasts can show a close correspondence to the hydrostatic shape of the brain in life, and canals and grooves can preserve features of the circulatory system. Multiple samples are rarely available for studies of individual variation, especially in fossils, thus a first step in quantifying variation and making comparisons with fossils is made possible with CT scans of osteological specimens. This study presents a series of high-resolution X-ray CT-derived cranial endocasts of six extant species of Phocoenidae, a clade including some of the smallest and one of the rarest cetaceans. Degree of gyrification varies interspecifically and intraspecifically, possibly resulting from variation in preservation of the ossified meninges. Computed tomographic data show that visually assessed asymmetry in the cranial endocasts is not correlated with volumetric measurements, but nonetheless may reflect torsion in the skull's shape such that the right cerebral and cerebellar hemispheres extend rostrally and laterally more than the left. Vasculature and canals are similar to other described cetacean species, but the hypophyseal casts are unusual. Similarities between brain shape and volume measurements in the different species can be attributed to paedomorphism and concomitant variation in ecological preferences. This may explain similarities Neophocaena phocaenoides and Phocoena sinus share with the juvenile Phocoena phocoena specimen studied.