Advances in research on the impacts of anti-submarine sonar on beaked whales.

Mass stranding events (MSEs) of beaked whales (BWs) were extremely rare prior to the 1960s but increased markedly after the development of naval mid-frequency active sonar (MFAS). The temporal and spatial associations between atypical BW MSEs and naval exercises were first observed in the Canary Islands, Spain, in the mid-1980s. Further research on BWs stranded in association with naval exercises demonstrated pathological findings consistent with decompression sickness (DCS). A 2004 ban on MFASs around the Canary Islands successfully prevented additional BW MSEs in the region, but atypical MSEs have continued in other places of the world, especially in the Mediterranean Sea, with examined individuals showing DCS. A workshop held in Fuerteventura, Canary Islands, in September 2017 reviewed current knowledge on BW atypical MSEs associated with MFAS. Our review suggests that the effects of MFAS on BWs vary among individuals or populations, and predisposing factors may contribute to individual outcomes. Spatial management specific to BW habitat, such as the MFAS ban in the Canary Islands, has proven to be an effective mitigation tool and mitigation measures should be established in other areas taking into consideration known population-level information.

Novel locomotor muscle design in extreme deep-diving whales.

Most marine mammals are hypothesized to routinely dive within their aerobic dive limit (ADL). Mammals that regularly perform deep, long-duration dives have locomotor muscles with elevated myoglobin concentrations that are composed of predominantly large, slow-twitch (Type I) fibers with low mitochondrial volume densities (V(mt)). These features contribute to extending ADL by increasing oxygen stores and decreasing metabolic rate. Recent tagging studies, however, have challenged the view that two groups of extreme deep-diving cetaceans dive within their ADLs. Beaked whales (including Ziphius cavirostris and Mesoplodon densirostris) routinely perform the deepest and longest average dives of any air-breathing vertebrate, and short-finned pilot whales (Globicephala macrorhynchus) perform high-speed sprints at depth. We investigated the locomotor muscle morphology and estimated total body oxygen stores of several species within these two groups of cetaceans to determine whether they (1) shared muscle design features with other deep divers and (2) performed dives within their calculated ADLs. Muscle of both cetaceans displayed high myoglobin concentrations and large fibers, as predicted, but novel fiber profiles for diving mammals. Beaked whales possessed a sprinter's fiber-type profile, composed of ~80% fast-twitch (Type II) fibers with low V(mt). Approximately one-third of the muscle fibers of short-finned pilot whales were slow-twitch, oxidative, glycolytic fibers, a rare fiber type for any mammal. The muscle morphology of beaked whales likely decreases the energetic cost of diving, while that of short-finned pilot whales supports high activity events. Calculated ADLs indicate that, at low metabolic rates, both beaked and short-finned pilot whales carry sufficient onboard oxygen to aerobically support their dives.

Lipid class and depth-specific thermal properties in the blubber of the short-finned pilot whale and the pygmy sperm whale.

Blubber, the specialized hypodermis of cetaceans, provides thermal insulation through the quantity and quality of lipids it contains. Quality refers to percent lipid content; however, not all lipids are the same. Certain deep-diving cetacean groups possess blubber with lipids - wax esters (WE) - that are not typically found in mammals, and the insulative quality of 'waxy' blubber is unknown. Our study explored the influence of lipid storage class - specifically WE in pygmy sperm whales (Kogia breviceps; N=7) and typical mammalian triacylglycerols in short-finned pilot whales (Globicephala macrorhynchus; N=7) - on blubber's thermal properties. Although the blubber of both species had similar total lipid contents, the thermal conductivity of G. macrorhynchus blubber (0.20±0.01 W m(-1) °C(-1)) was significantly higher than that of K. breviceps (0.15±0.01 W m(-1) °C(-1); P=0.0006). These results suggest that lipid class significantly influences the ability of blubber to resist heat flow. In addition, because the lipid content of blubber is known to be stratified, we measured its depth-specific thermal conductivities. In K. breviceps blubber, the depth-specific conductivity values tended to vary inversely with lipid content. In contrast, G. macrorhynchus blubber displayed unexpected depth-specific relationships between lipid content and conductivity, which suggests that temperature-dependent effects, such as melting, may be occurring. Differences in heat flux measurements across the depth of the blubber samples provide evidence that both species are capable of storing heat in their blubber. The function of blubber as an insulator is complex and may rely upon its lipid class, stratified composition and dynamic heat storage capabilities.

Ontogenetic changes in the structural stiffness of the tailstock of bottlenose dolphins (Tursiops truncatus).

Late-term fetal bottlenose dolphins (Tursiops truncatus) are bent ventrolaterally en utero, requiring extreme flexibility of the axial skeleton and associated soft tissues. At birth, neonatal dolphins must immediately swim to the surface to breath, yet the dorsoventral oscillations used during locomotion may be compromised by the lateral flexibility evident in the fetus. The unique fetal position of dolphins, coupled with their need to swim at birth, places conflicting mechanical demands on the tailstock. Our previous research demonstrated that neonatal dolphins possess laterally placed, axial muscles that are functionally specialized to actively maintain the straightened posture of the tailstock. Here, we investigated the development of passive lateral stability in the tailstock of bottlenose dolphins by performing whole-body bending tests on an ontogenetic series of stranded dolphin specimens (N=15), including fetuses, neonates and juveniles (total length 58-171 cm). Structural stiffness increased, while overall body curvature decreased, with increasing body length. Scaling analyses suggest that increased structural stiffness is due to increases in size and probably changes in the passive material properties of the tailstock through ontogeny. The neutral zone was approximately constant with increasing size, while the relative neutral zone (neutral zone/total length) decreased. The lateral stability of the tailstock appears to be controlled by a combination of active and passive systems and the role of these systems varies through ontogeny. While neonates use active, muscular mechanisms to limit lateral deformations of the tailstock, the stability of the maturing tailstock is due primarily to its passive tissue properties.

Morphology of the melon and its tendinous connections to the facial muscles in bottlenose dolphins (Tursiops truncatus).

The melon is a lipid-rich structure located in the forehead of odontocetes that functions to propagate echolocation sounds into the surrounding aquatic environment. To date, the melon's ability to guide and impedance match biosonar sounds to seawater has been attributed to its unique fatty acid composition. However, the melon is also acted upon by complex facial muscles derived from the m. maxillonasolabialis. The goal of this study was to investigate the gross morphology of the melon in bottlenose dolphins (Tursiops truncatus) and to describe how it is tendinously connected to these facial muscles. Standard gross dissection (N = 8 specimens) and serial sectioning (N = 3 specimens) techniques were used to describe the melon and to identify its connections to the surrounding muscles and blubber in three orthogonal body planes. The dolphin forehead was also thin-sectioned in three body planes (N = 3 specimens), and polarized light was used to reveal the birefringent collagen fibers within and surrounding the melon. This study identified distinct regions of the melon that vary in shape and display locally specific muscle-tendon morphologies. These regions include the bilaterally symmetric main body and cone and the asymmetric right and left caudal melon. This study is the first to identify that each caudal melon terminates in a lipid cup that envelopes the echolocation sound generators. Facial muscles of the melon have highly organized tendon populations that traverse the melon and insert into either the surrounding blubber, the connective tissue matrix of the nasal plug, or the connective tissue sheath surrounding the sound generators. The facial muscles and tendons also lie within multiple orthogonal body planes, which suggest that the melon is capable of complex shape change. The results of this study suggest that these muscles could function to change the frequency, beam width, and directionality of the emitted sound beam in bottlenose dolphins. The echolocation sound propagation pathway within the dolphin forehead appears to be a tunable system.

Malignant seminoma with metastasis, Sertoli cell tumor, and pheochromocytoma in a spotted dolphin (Stenella frontalis) and malignant seminoma with metastasis in a bottlenose dolphin (Tursiops truncatus).

Seminoma with metastasis was diagnosed in a spotted dolphin (Stenella frontalis) and an Atlantic bottlenose dolphin (Tursiops truncatus). Sertoli cell tumor and pheochromocytoma were also diagnosed in the spotted dolphin. The spotted and bottlenose dolphins were adult males that stranded and died on the coasts of northwest Florida and southeast North Carolina, respectively. Neoplasia is infrequently reported in cetaceans. This is the first report of seminoma, Sertoli cell tumor, and pheochromocytoma in a dolphin, the first report of three distinct neoplasms in a dolphin, and one of the few reports of malignant neoplasia in dolphins.

Blubber development in bottlenose dolphins (Tursiops truncatus).

Blubber, the lipid-rich hypodermis of cetaceans, functions in thermoregulation, buoyancy control, streamlining, metabolic energy storage, and locomotion. This study investigated the development of this specialized hypodermis in bottlenose dolphins (Tursiops truncatus) across an ontogenetic series, including fetuses, neonates, juveniles, subadults, and adults. Blubber samples were collected at the level of the mid-thorax, from robust specimens (n = 25) that stranded along the coasts of North Carolina and Virginia. Blubber was dissected from the carcass and its mass, and the depth and lipid content at the sample site, were measured. Samples were prepared using standard histological methods, viewed by light microscopy, and digital images of blubber captured. Images were analyzed through the depth of the blubber for morphological and structural features including adipocyte size, shape, and numbers, and extracellular, structural fiber densities. From fetus to adult, blubber mass and depth increased proportionally with body mass and length. Blubber lipid content increased dramatically with increasing fetal length. Adult and juvenile blubber had significantly higher blubber lipid content than fetuses, and this increase was reflected in mean adipocyte size, which increased significantly across all robust life history categories. In juvenile, subadult, and adult dolphins, this increase in cell size was not uniform across the depth of the blubber, with the largest increases observed in the middle and deep blubber regions. Through-depth counts of adipocytes were similar in all life history categories. These results suggest that blubber depth is increased during postnatal growth by increasing cell size rather than cell number. In emaciated adults (n = 2), lipid mobilization, as evidenced by a decrease in adipocyte size, was localized to the middle and deep blubber region. Thus, in terms of both lipid accumulation and depletion, the middle and deep blubber appear to be the most metabolically dynamic. The superficial blubber likely serves a structural role important in streamlining the animal. This study demonstrates that blubber is not a homogeneous tissue through its depth, and that it displays life history-dependent changes in its morphology and lipid content.

Magnetic resonance images of the brain of a dwarf sperm whale (Kogia simus).

Cetacean (dolphin, whale and porpoise) brains are among the least studied mammalian brains because of the difficulty of collecting and histologically preparing such relatively rare and large specimens. Among cetaceans, there exist relatively few studies of the brain of the dwarf sperm whale (Kogia simus). Magnetic resonance imaging (MRI) offers a means of observing the internal structure of the brain when traditional histological procedures are not practical. Therefore, MRI has become a critical tool in the study of the brain of cetaceans and other large species. This paper represents the first MRI-based anatomically labelled three-dimensional description of the dwarf sperm whale brain. Coronal plane sections of the brain of a sub-adult dwarf sperm whale were originally acquired and used to produce virtual digital scans in the other two orthogonal spatial planes. A sequential set of images in all three planes has been anatomically labelled and displays the proportions and positions of major neuroanatomical features.

Changes in blubber distribution and morphology associated with starvation in the harbor porpoise (Phocoena phocoena): evidence for regional differences in blubber structure and function.

To examine patterns of blubber loss accompanying a decline in body condition, blubber thickness of juvenile harbor porpoises in normal/robust body condition (n=69) was compared with that of starved conspecifics (n=31). Blubber thickness in the thorax of starved porpoises (9-11 mm) was only 50%-60% of that of normal animals (18-20 mm); however, very little tailstock blubber was lost during starvation. Adipocytes in thorax and tailstock blubber were measured in both groups (n=5) to determine whether thickness changes were homogeneous throughout blubber depth. In the thorax of normal porpoises, adipocytes near the epidermis (outer blubber) were smaller (0.11 nL) than inner blubber adipocytes (0.17 nL). Conversely, the size of tailstock adipocytes was uniform. Starved animals had fewer, smaller adipocytes in the inner thorax blubber, suggesting a possible combination of adipocyte shrinkage and loss. Lipids were withdrawn only from the inner layer of thorax blubber during starvation, supporting a hypothesis of regional specialization of function in blubber. Blubber of the thorax serves as the site of lipid deposition and mobilization, while the tailstock is metabolically inert and likely important in locomotion and streamlining. Therefore, some proportion of the blubber of small odontocetes must be considered structural/mechanical rather than an energy reserve.

Functional morphology of venous structures associated with the male and female reproductive systems in Florida manatees (Trichechus manatus latirostris).

The reproductive organs of Florida manatees (Trichechus manatus latirostris) are surrounded by thermogenic locomotory muscles and insulating fat. Manatees are reported to maintain core body temperatures of 35.6 degrees -36.4 degrees C, temperatures known to interfere with production and maturation of viable sperm in terrestrial mammals. We describe two novel venous plexuses associated with the manatee epididymis. Each epididymis is located in a hypogastric fossa at the caudolateral extremity of the abdominal cavity. Each hypogastric fossa is lined by an inguinal venous plexus that receives cooled blood from a superficial thoracocaudal plexus. We conclude that male manatees may prevent hyperthermic insult to their reproductive tissues by feeding cooled superficial blood to venous plexuses deep within their bodies. Female manatees also possess hypogastric fossae and venous structures similar to those found in male manatees. The ovaries, uterine tubes, and distal tips of the uterine horns are located in the hypogastric fossae. We suggest that the thermovascular structures we describe also prevent hypothermic insult to female manatee reproductive tissues. The venous structures in manatees are functionally similar to structures associated with reproductive thermoregulation in cetaceans and phocid seals. Thus, these thermovascular structures appear to be convergent morphological adaptations that occur in three clades of diving mammals with independent evolutionary histories.

Anatomy and three-dimensional reconstructions of the brain of a bottlenose dolphin (Tursiops truncatus) from magnetic resonance images.

Cetacean (dolphin, whale, and porpoise) brains are among the least studied mammalian brains because of the formidable challenge of collecting and histologically preparing such relatively rare and large specimens. Magnetic resonance imaging offers a means of observing the internal structure of the brain when traditional histological procedures are not practical. Furthermore, internal structures can be analyzed in their precise anatomic positions, which is difficult to accomplish after the spatial distortions often accompanying histological processing. In this study, images of the brain of an adult bottlenose dolphin, Tursiops truncatus, were scanned in the coronal plane at 148 antero-posterior levels. From these scans a computer-generated three-dimensional model was constructed using the programs VoxelView and VoxelMath (Vital Images, Inc.). This model, wherein details of internal and external morphology are represented in three-dimensional space, was then resectioned in orthogonal planes to produce corresponding series of virtual sections in the horizontal and sagittal planes. Sections in all three planes display the sizes and positions of major neuroanatomical features such as the arrangement of cortical lobes and subcortical structures such as the inferior and superior colliculi, and demonstrate the utility of MRI for neuroanatomical investigations of dolphin brains.

Buoyant balaenids: the ups and downs of buoyancy in right whales.

A variety of marine mammal species have been shown to conserve energy by using negative buoyancy to power prolonged descent glides during dives. A new non-invasive tag attached to North Atlantic right whales recorded swim stroke from changes in pitch angle derived from a three-axis accelerometer. These results show that right whales are positively buoyant near the surface, a finding that has significant implications for both energetics and management. Some of the most powerful fluke strokes observed in tagged right whales occur as they counteract this buoyancy as they start a dive. By contrast, right whales use positive buoyancy to power glides during ascent. Right whales appear to use their positive buoyancy for more efficient swimming and diving. However, this buoyancy may pose added risks of vessel collision. Such collisions are the primary source of anthropogenic mortality for North Atlantic right whales, whose population is critically endangered and declining. Buoyancy may impede diving responses to oncoming vessels and right whales may have a reduced ability to manoeuvre during free ascents. These risk factors can inform efforts to avoid collisions.

The development of diving in marine endotherms: preparing the skeletal muscles of dolphins, penguins, and seals for activity during submergence.

Myoglobin is an important oxygen store for supporting aerobic diving in endotherms, yet little is known about its role during postnatal development. Therefore, we compared the postnatal development of myoglobin in marine endotherms that develop at sea (cetaceans) to those that develop on land (penguins and pinnipeds). We measured myoglobin concentrations in the major locomotor muscles of mature and immature bottlenose dolphins (Tursiops truncatus) and king penguins (Aptenodytes patagonicus) and compared the data to previously reported values for northern elephant seals (Mirounga angustirostris). Neonatal dolphins, penguins, and seals lack the myoglobin concentrations required for prolonged dive durations, having 10%, 9%, and 31% of adult values, respectively. Myoglobin contents increased significantly during subsequent development. The increases in myoglobin content with age may correspond to increases in activity levels, thermal demands, and time spent in apnea during swimming and diving. Across these phylogenetically diverse taxa (cetaceans, penguins, and pinnipeds), the final stage of postnatal development of myoglobin occurs during the initiation of independent foraging, regardless of whether development takes place at sea or on land.

Precocial development of axial locomotor muscle in bottlenose dolphins (Tursiops truncatus).

At birth, the locomotor muscles of precocial, terrestrial mammals are similar to those of adults in both mass, as a percent of total body mass, and fiber-type composition. It is hypothesized that bottlenose dolphins (Tursiops truncatus), marine mammals that swim from the instant of birth, will also exhibit precocial development of locomotor muscles. Body mass data from neonatal and adult dolphins are used to calculate Grand's (1992) Neural and Muscular Indices of Development. Using these indices, the bottlenose dolphin is a Condition "3.5" neonate, where Condition 4 is the documented extreme of precocial development in terrestrial mammals. Moreover, myosin ATPase (alkaline preincubation) analyses of the epaxial locomotor m. extensor caudae lateralis show that neonatal dolphins have fiber-type profiles very similar to those of adults. Thus, based on mass and myosin ATPase activity, muscle development in dolphins is precocial. However, succinic dehydrogenase and Nile red histochemistry demonstrate that neonatal dolphin muscle has mitochondrial and lipid distributions different from those found in adults. These data suggest that neonates have a lower aerobic capacity than adults. Dolphin neonates may compensate for an apparent lack of aerobic stamina in two ways: 1) by being positively buoyant, with a relatively increased investment of their total body mass in blubber, and 2) by "free-riding" off their mothers. This study investigates quantitatively the development of a dolphin locomotor muscle and offers suggestions about adaptations required for a completely aquatic existence.

Locomotor design of dolphin vertebral columns: bending mechanics and morphology of Delphinus delphis.

The primary skeletal structure used by dolphins to generate the dorsoventral bending characteristic of cetacean swimming is the vertebral column. In the vertebral column of the saddleback dolphin Delphinus delphis, we characterize the static and dynamic mechanical properties of the intervertebral joints, describe regional variation and dorsoventral asymmetries in mechanical performance, and investigate how the mechanical properties are correlated with vertebral morphologies. Using a bending machine that applies an external load (N m) to a single intervertebral segment, we measured the resulting angular deformation (rad) of the segment in both dorsal extension and ventral flexion. Intervertebral segments from the thoracic, lumbar and caudal regions of the vertebral column were tested from five individuals. Using quasi-static bending tests, we measured the initial (low-strain) bending stiffness (N m rad-1) as a function of segment position, direction of bending (extension and flexion) and sequential cutting of intervertebral ligaments. We found that initial bending stiffness was significantly greater in the lumbar region than in adjacent thoracic and caudal regions, and all joints were stiffer in extension than is predicted (r2 = 0.554) by the length and width of the intervertebral disc and the length of the cranial vertebral body in the segment. Stiffness in flexion is predicted (r2 = 0.400) by the width of the nucleus pulposus, the length of the caudal vertebral body in the segment and the height of the transverse processes from the ventral surface of the vertebral body. We also performed dynamic bending tests on intervertebral segments from the lumbo-caudal joint and the joint between caudal vertebrae 7 and 8. Dynamic bending stiffness (N m rad-1) increases with increasing bending amplitude and is independent of bending frequency. Damping coefficient (kg m2 rad-2 s-1) decreases with increasing bending amplitude and frequency. Resilience (% energy return) increases from approximately 20% at low bending amplitudes (+/-0.6 degree) to approximately 50% at high bending amplitudes (+/-2.9 degrees). Based on these findings, the dolphin's vertebral column has the mechanical capacity to help control the body's locomotor reconfigurations, to store elastic energy and to dampen oscillations.

Venous structures associated with thermoregulation of phocid seal reproductive organs.

BACKGROUND: Seal reproductive systems are surrounded by thermogenic muscle and insulating blubber, suggesting elevated temperatures at the gonads and uterus. In the limbs of terrestrial mammals, cooled blood returning from superficial veins is mixed proximally with warm blood returning from deep veins. Thus, mixed cool-superficial and warm-deep venous blood from the hind limbs is returned to the central circulation. METHODS: We describe structures observed in salvaged carcasses of harbor (Phoca vitulina), gray (Haliochoerus gryphus), harp (Phoca groendlandica), hooded (Cystophora cristata), and ringed (Phoca hispida) seals. Vessels were identified by dissection of injected and uninjected material. RESULTS: In contrast to terrestrial mammals, phocid seals have anastomoses between the veins of the distal hind limb and the pelvis which allow large volumes of cool blood returning from the skin surface of the flipper to enter the gluteal, pelvic, or pudendo-epigastric veins. This provides a cool-superficial venous return that remains separate from the warm-deep venous return of the femoral veins. The cooled venous blood from the hind flippers supplies venous plexuses lining the inguinal region and the abdominal and pelvic cavities. CONCLUSIONS: Cooled blood may prevent hyperthermic insult to seal reproductive systems.

Thermoregulation of the intra-abdominal testes of the bottlenose dolphin (Tursiops truncatus) during exercise.

Dolphins possess a vascular countercurrent heat exchanger (CCHE) that functions to cool their intra-abdominal testes. Spermatic arteries in the posterior abdomen are juxtaposed to veins returning cooled blood from the surfaces of the dorsal fin and tail flukes. In this study, we investigated the effect of exercise on CCHE function in the bottlenose dolphin. The CCHE flanks a region of the bowel in the posterior abdomen and influences colonic temperatures. A rectal probe housing a linear array of seven copper-constantan thermocouples was designed to measure colonic temperatures simultaneously at positions anterior to, within and posterior to the region of the colon flanked by the CCHE. Immediately after vigorous swimming, temperatures at the CCHE decreased relative to resting and pre-swim values: post-swim temperatures at the CCHE were maximally 0.5 degrees C cooler than pre-swim temperatures. These data suggest that the CCHE has an increased ability to cool the arterial blood supply to the testes when the dolphin is swimming. This ability could offset the increased thermal load on the testes is an exercising dolphin. To the best of our knowledge, this is the first report of deep body cooling in an exercising mammal that is not undertaking a dive.

Temperature regulation of the testes of the bottlenose dolphin (Tursiops truncatus): evidence from colonic temperatures.

Dolphins possess a countercurrent heat exchanger that functions to cool their intra-abdominal testes. Spermatic arteries in the posterior abdomen are juxtaposed to veins returning cooled blood from the surfaces of the dorsal fin and flukes. A rectal probe housing a linear array of five copper-constantan thermocouples was designed to measure colonic temperatures simultaneously at positions anterior to, within, and posterior to the region of the colon flanked by the countercurrent heat exchanger. Colonic temperatures adjacent to the countercurrent heat exchanger were maximally 1.3 degrees C cooler than temperatures measured outside this region. Temporary heating and cooling of the dorsal fin and flukes affected temperatures at the countercurrent heat exchanger, but had little or no effect on temperatures posterior to its position. These measurements support the hypothesis that cooled blood is introduced into the deep abdominal cavity and functions specifically to regulate the temperature of arterial blood flow to the dolphin testes.

Functional morphology of the vascular plexuses associated with the cetacean uterus.

The cetacean reproductive system is surrounded by thermogenic locomotory muscle and insulating blubber. This arrangement suggests elevated temperatures at the uterus that could induce detrimental effects on fetal development. We present anatomical evidence for a complex countercurrent heat exchange system that could function to regulate the thermal environment of the uterus and a developing fetus. Cooled venous blood from the surfaces of the dorsal fin and flukes enters the abdominal cavity via the lumbo-caudal venous plexus. This plexus is juxtaposed to the arterial and venous plexuses associated with the uterus. The morphology of the lumbo-caudal venous plexus suggests that it acts as a "heat sink" for the adjacent tissues. Heat may be transferred to the cool, lumbo-caudal venous plexus from the warm blood in the arterial and venous plexuses supplying the uterus. Heat may also be transferred from adjacent locomotory muscles to the cool lumbo-caudal venous plexus. The countercurrent heat exchanger created by the juxtaposition of the lumbo-caudal venous plexus with the uterovarian arterial plexus is similar in design to that of the countercurrent heat exchanger described for male cetaceans. The functional implications of introducing cool superficial blood into the abdominal cavity of a diving, and locomoting female cetacean are discussed.

Anatomical evidence for a countercurrent heat exchanger associated with dolphin testes.

Cetaceans possess cryptic testes that lie within the abdominal cavity, that are surrounded by primary locomotor muscles, and that are presumably exposed to core or above core body temperatures. It has remained a question as to how cetaceans produce and store viable sperm at these high temperatures. We offer anatomical evidence for a two layer arterio-venous countercurrent heat exchanger at the cetacean testis. Subcutaneous veins from the peripheral surfaces of the dorsal fin and flukes carry cool blood from the fins to the lumbo-caudal venous plexus. The lumbo-caudal venous plexus is juxtaposed to the spermatic arterial plexus, which supplies the testis. Venous plexus flow is form the ventro-lateral margins of the visceral cavity towards the vena cava. Arterial plexus flow is from the aorta towards the ventro-lateral margins of the visceral cavity and into the testis. The existence of a countercurrent heat exchanger suggests that cetaceans potentially compensate for detrimental effects of core temperatures on sperm viability and storage by regulating the temperature of blood flow to the testis.