Melon and rostral muscle morphology of Gervais' beaked whale (Mesoplodon europaeus): Alternating patterns of bilateral asymmetry.

Toothed whales utilize specialized nasal structures such as the lipid-rich melon to produce sound and propagate it into the aquatic environment. Very little nasal morphology of mesoplodont beaked whales has been described in the literature, and the anatomy of the melon and associated musculature of Gervais' beaked whale (Mesoplodon europaeus) remains undescribed. Heads of three (n = 3) Gervais' beaked whales were examined in detail via dissection as well as computed tomography (CT) and magnetic resonance imaging (MRI). Two additional Gervais' beaked whale individuals (n = 2) were studied via archived CT and MRI scans. Representative transverse dissection sections of the melon were processed for polarized light imaging to verify the presence of tendons inserting into the melon tissue. Three-dimensional (3D) CT reconstructions of the melon, rostral muscles, and associated structures were performed to assess morphology and spatial relationships. In all individuals, the melon's main body demonstrated a bilaterally asymmetrical, curvilinear geometry. This curvilinear shape was defined by a pattern of alternating asymmetry in the medial rostral muscles that projected into the melon's tissue. In transverse polarized light imaging, a network of tendons originating from these asymmetrical rostral muscle projections was observed permeating the melon's lipid tissue. This curvilinear melon morphology and associated asymmetrical musculature suggest a means of lengthening the lipid pathway within a relatively short dimensional footprint. In addition, the species-specific arrangement of muscular projections suggests complex fine-tuning of the melon's geometry during echolocation. Further studies may lend additional insight into the function of this unusual melon morphology.

Gross and histological morphology of the cervical gill slit gland of the pygmy sperm whale (Kogia breviceps).

Odontocete cetaceans have undergone profound modifications to their integument and sensory systems and are generally thought to lack specialized exocrine glands that in terrestrial mammals function to produce chemical signals (Thewissen & Nummela, 2008). Keenan-Bateman et al. (2016, 2018), though, introduced an enigmatic exocrine gland, associated with the false gill slit pigmentation pattern in Kogia breviceps. These authors provided a preliminary description of this cervical gill slit gland in their helminthological studies of the parasitic nematode, Crassicauda magna. This study offers the first detailed gross and histological description of this gland and reports upon key differences between immature and mature individuals. Investigation reveals it is a complex, compound tubuloalveolar gland with a well-defined duct that leads to a large, and expandable central chamber, which in turn leads to two caudally projecting diverticula. All regions of the gland contain branched tubular and alveolar secretory regions, although most are found in the caudal diverticula, where the secretory process is holocrine. The gland lies between slips of cutaneous muscle, and is innervated by lamellar corpuscles, resembling Pacinian's corpuscles, suggesting that its secretory product may be actively expressed into the environment. Mature K. breviceps display larger gland size, and increased functional activity in glandular tissues, as compared to immature individuals. These results demonstrate that the cervical gill slit gland of K. breviceps shares morphological features of the specialized, chemical signaling, exocrine glands of terrestrial members of the Cetartiodactyla.

Comparative morphology of the spinal cord and associated vasculature in shallow versus deep diving cetaceans.

The cetacean vertebral canal houses the spinal cord and arterial supply to and venous drainage from the entire central nervous system (CNS). Thus, unlike terrestrial mammals, the cetacean spinal cord lies within a highly vascularized space. We compared spinal cord size and vascular volumes within the vertebral canal across a sample of shallow and deep diving odontocetes. We predicted that the (a) spinal cord, a metabolically expensive tissue, would be relatively small, while (b) volumes of vascular structures would be relatively large, in deep versus shallow divers. Our sample included the shallow diving Tursiops truncatus (n = 2) and Delphinus delphis (n = 3), and deep diving Kogia breviceps (n = 2), Mesoplodon europaeus (n = 2), and Ziphius cavirostris (n = 1). Whole, frozen vertebral columns were cross-sectioned at each intervertebral disc, scaled photographs of vertebral canal contents acquired, and cross-sectional areas of structures digitally measured. Areas were multiplied by vertebral body lengths and summed to calculated volumes of neural and vascular structures. Allometric analyses revealed that the spinal cord scaled with negative allometry (b = 0.51 ± 0.13) with total body mass (TBM), and at a rate significantly lower than that of terrestrial mammals. As predicted, the spinal cord represented a smaller percentage of the total vertebral canal volume in the deep divers relative to shallow divers studied, as low as 2.8% in Z. cavirostris. Vascular volume scaled with positive allometry (b = 1.2 ± 0.22) with TBM and represented up to 96.1% (Z. cavirostris) of the total vertebral canal volume. The extreme deep diving beaked whales possessed 22-35 times more vascular volume than spinal cord volume within the vertebral canal, compared with the 6-10 ratio in the shallow diving delphinids. These data offer new insights into morphological specializations of neural and vascular structures that may contribute to differential diving capabilities across odontocete cetaceans.

Heat flux in manatees: an individual matter and a novel approach to assess and monitor the thermal state of Florida manatees (Trichechus manatus latirostris).

Florida manatees (Trichechus manatus latirostris) possess an unusual suite of adaptations to accommodate both a fully aquatic lifestyle and an herbivorous diet, including a low metabolic rate and a very limited thermoneutral zone. Their relatively high lower critical temperature of around 20 °C suggests strong sensitivity to cold, thereby limiting their distribution to tropical and subtropical waters. "Cold stress syndrome" affects and kills Florida manatees every year during intense or prolonged cold weather, posing one of the major threats to manatees. However, knowledge regarding manatee thermoregulation is sparse, but essential for effective conservation and management of this threatened species. We measured heat flux in two captive Florida manatees at multiple times of the year, at 41 sites distributed across the entire body surface of each manatee. Heat flux differed significantly between individuals, and among body sites and times of the year. The pectoral flippers and axillae were identified as areas with highest heat exchange. Despite exposure to constant water temperature throughout the year, the manatees in this study had significantly lower heat flux in winter than in summer. We used the measured heat flux values to calculate total heat dissipation in individual manatees. The values estimated this way correspond well with the low metabolic rates estimated in previous studies, confirming the reliability of our novel approach. Our method provides simple and useful options for enhancing manatee welfare by monitoring the animals' thermal state during potentially stressful activities such as during medical treatment, capture restraints and transportation.

Habitat use pattern of the giant parasitic nematode Crassicauda magna within the pygmy sperm whale Kogia breviceps.

The giant (>3 m) parasitic nematode Crassicauda magna infects kogiid whales, although only 3 studies to date have provided detailed descriptions of these worms, all based upon fragmented specimens. These fragments were found within the neck region of kogiids, an unusual anatomic site for this genus of parasites. C. magna is a species-specific parasite among kogiids, infecting only pygmy sperm whales Kogia breviceps, and with a primarily cervico-thoracic distribution. To date, however, the pattern of habitat use within the host and transmission path of this parasite remain unknown. We used detailed dissections (n = 12), histological examination of host tissues (n = 2), and scanning electron microscopy of excised nematodes (n = 7) to enhance our understanding of this host-parasite relationship. Results revealed that a critical habitat for the parasite is an exocrine gland in the whale's ventral cervical region. C. magna male and female tails were found intertwined within the glandular lumen, and eggs were observed within its presumed secretion, illuminating the transmission path out of the host. The cephalic ends of these worms were often meters away (curvilinearly), embedded deeply within epaxial muscle. A single worm's complete, tortuous 312 cm course, from the gland to its termination in the contralateral epaxial muscle, is described for the first time. This study also provides the first scanning electron micrographs of C. magna, which illustrate taxonomically important features of the heads and tails of both male and female worms.

How to Build a Deep Diver: The Extreme Morphology of Mesoplodonts.

Mesoplodont beaked whales are extreme divers, diving for over 45 mins and to depths of over 800 m. These dives are of similar depth and duration to those of the giant sperm whale (Physeter macrocephalus) whose body mass can be 50 times larger. Velten et al. (2013) provided anatomical data that demonstrated that on-board oxygen stores were sufficient to aerobically support the extreme dives of mesoplodonts if their diving metabolic rates are low. Because no physiological data yet exist, we utilized an anatomical approach-the body composition technique-to examine the relative metabolic rates of mesoplodonts. We utilized a systematic mass dissection protocol to compare the body composition of mesoplodonts with those of two short duration, shallow divers-the harbor porpoise (Phocoena phocoena) and bottlenose dolphin (Tursiops truncatus). We then investigated the body composition of two other extreme divers, the southern elephant seal (Mirounga leonina) and P. macrocephalus using data from the literature. Our results demonstrate that extreme divers invest a smaller percentage of their total body mass (TBM) in metabolically expensive brain and viscera, and a larger percent of their TBM in inexpensive integument, bone, and muscle, than do the shallow divers. Deep divers also share features of their locomotor muscle that contribute to relatively low tissue metabolic rates and high oxygen storage capacity, including large muscle fiber diameters, low mitochondrial volume densities, and high myoglobin concentrations. One feature of the locomotor muscle of mesoplodonts, though, is unique among deep divers investigated to date. Rather than having an endurance athlete's muscle fiber profile, dominated by slow oxidative fibers, mesoplodonts possess a sprinter's profile, dominated by fast glycolytic fibers. Velten et al. (2013) hypothesized that these fibers are likely inactive during routine swimming and provide a large, metabolically inexpensive oxygen store for the slow oxidative fibers to aerobically power swimming. We suggest that future anatomical analyses, coupled with performance data transduced through tagging studies, will enhance our understanding of the extreme diving capabilities of marine mammals.

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

Beaked whales are a poorly known but diverse group of whales that have received considerable attention due to strandings that have been temporally and spatially associated with naval sonar deployment. Postmortem studies on stranded carcasses have revealed lesions consistent with decompression sickness, including intravascular gas and fat emboli. These findings have been supported by analyses of intravascular gas emboli showing composition dominated by nitrogen gas. To increase our understanding of the pathophysiology of nitrogen bubble formation and intravascular embolization, we examined the gross and microscopic anatomy of the venous system in the head of beaked whales. Since the potential sources of intravascular fat and gas emboli were of greatest interest, focus was placed on the acoustic fat bodies and pneumatic accessory sinus system. Herein, we describe intimate arteriovenous associations with specialized adipose depots and air sinuses in beaked whales. These vascular structures comprise an extensive network of thin-walled vessels with a large surface area, which is likely to facilitate exchange of nitrogen gas and may, therefore, form anatomic regions that may be important in physiological management of diving gases. These structures may also be vulnerable to pathologic introduction of emboli into the vascular system. Expansive, thin-walled venous lakes are found within the pterygoid region, which suggest the potential for nitrogen exchange as well as for compensation of middle-ear pressures during descent on a dive. These findings warrant further research into the structure and function of this morphology as it relates to normal and pathologic physiology.

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.

Morphology of the Nasal Apparatus in Pygmy (Kogia Breviceps) and Dwarf (K. Sima) Sperm Whales.

Odontocete echolocation clicks are generated by pneumatically driven phonic lips within the nasal passage, and propagated through specialized structures within the forehead. This study investigated the highly derived echolocation structures of the pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales through careful dissections (N = 18 K. breviceps, 6 K. sima) and histological examinations (N = 5 K. breviceps). This study is the first to show that the entire kogiid sound production and transmission pathway is acted upon by complex facial muscles (likely derivations of the m. maxillonasolabialis). Muscles appear capable of tensing and separating the solitary pair of phonic lips, which would control echolocation click frequencies. The phonic lips are enveloped by the "vocal cap," a morphologically complex, connective tissue structure unique to kogiids. Extensive facial muscles appear to control the position of this structure and its spatial relationship to the phonic lips. The vocal cap's numerous air crypts suggest that it may reflect sounds. Muscles encircling the connective tissue case that surrounds the spermaceti organ may change its shape and/or internal pressure. These actions may influence the acoustic energy transmitted from the phonic lips, through this lipid body, to the melon. Facial and rostral muscles act upon the length of the melon, suggesting that the sound "beam" can be focused as it travels through the melon and into the environment. This study suggests that the kogiid echolocation system is highly tunable. Future acoustic studies are required to test these hypotheses and gain further insight into the kogiid echolocation system.

Vascularization of Air Sinuses and Fat Bodies in the Head of the Bottlenose Dolphin (Tursiops truncatus): Morphological Implications on Physiology.

Cetaceans have long been considered capable of limiting diving-induced nitrogen absorption and subsequent decompression sickness through a series of behavioral, anatomical, and physiological adaptations. Recent studies however suggest that in some situations these adaptive mechanisms might be overcome, resulting in lethal and sublethal injuries. Perhaps most relevant to this discussion is the finding of intravascular gas and fat emboli in mass-stranded beaked whales. Although the source of the gas emboli has as yet to been ascertained, preliminary findings suggest nitrogen is the primary component. Since nitrogen gas embolus formation in divers is linked to nitrogen saturation, it seems premature to dismiss similar pathogenic mechanisms in breath-hold diving cetaceans. Due to the various anatomical adaptations in cetacean lungs, the pulmonary system is thought of as an unlikely site of significant nitrogen absorption. The accessory sinus system on the ventral head of odontocete cetaceans contains a sizeable volume of air that is exposed to the changing hydrostatic pressures during a dive, and is intimately associated with vasculature potentially capable of absorbing nitrogen through its walls. The source of the fat emboli has also remained elusive. Most mammalian fat deposits are considered poorly vascularized and therefore unlikely sites of intravascular introduction of lipid, although cetacean blubber may not be as poorly vascularized as previously thought. We present new data on the vasculature of air sinuses and acoustic fat bodies in the head of bottlenose dolphins and compare it to published accounts. We show that the mandibular fat bodies and accessory sinus system are associated with extensive venous plexuses and suggest potential physiological and pathological implications.

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.

Fatty acid profiles as a potential lipidomic biomarker of exposure to brevetoxin for endangered Florida manatees (Trichechus manatus latirostris).

Fatty acid signature analysis (FASA) is an important tool by which marine mammal scientists gain insight into foraging ecology. Fatty acid profiles (resulting from FASA) represent a potential biomarker to assess exposure to natural and anthropogenic stressors. Florida manatees are well studied, and an excellent necropsy program provides a basis against which to assess this budding tool. Results using samples from 54 manatees assigned to four cause-of-death categories indicated that those animals exposed to or that died due to brevetoxin exposure (red tide, or RT samples) demonstrate a distinctive hepatic fatty acid profile. Discriminant function analysis indicated that hepatic fatty acids could be used to classify RT versus non-RT liver samples with reasonable certainty. A discriminant function was derived based on 8 fatty acids which correctly classified 100% of samples from a training dataset (10 RT and 25 non-RT) and 85% of samples in a cross-validation dataset (5 RT and 13 non-RT). Of the latter dataset, all RT samples were correctly classified, but two of thirteen non-RT samples were incorrectly classified. However, the "incorrect" samples came from manatees that died due to other causes during documented red tide outbreaks; thus although the proximal cause of death was due to watercraft collisions, exposure to brevetoxin may have affected these individuals in ways that increased their vulnerability. This use of FASA could: a) provide an additional forensic tool to help scientists and managers to understand cause of death or debilitation due to exposure to red tide in manatees; b) serve as a model that could be applied to studies to improve assessments of cause of death in other marine mammals; and c) be used, as in humans, to help diagnose metabolic disorders or disease states in manatees and other species.

Lung size and thoracic morphology in shallow- and deep-diving cetaceans.

Shallow-diving, coastal bottlenose dolphins (Tursiops truncatus) and deep-diving, pelagic pygmy and dwarf sperm whales (Kogia breviceps and K. sima) will experience vastly different ambient pressures at depth, which will influence the volume of air within their lungs and potentially the degree of thoracic collapse they experience. This study tested the hypotheses that lung size will be reduced and/or thoracic mobility will be enhanced in deeper divers. Lung mass (T. truncatus, n = 106; kogiids, n = 18) and lung volume (T. truncatus, n = 5; kogiids, n = 4), relative to total body mass, were compared. One T. truncatus and one K. sima were cross-sectioned to calculate lung, thoracic vasculature, and other organ volumes. Excised thoraxes (T. truncatus, n = 3; kogiids, n = 4) were mechanically manipulated to compare changes in thoracic cavity shape and volume. Kogiid lungs were half the mass and one-fifth the volume of those of similarly sized T. truncatus. The lungs occupied only 15% of the total thoracic cavity volume in K. sima and 37% in T. truncatus. The kogiid and dolphin thoraxes underwent similar changes in shape and volume, although the width of the thoracic inlet was relatively constrained in kogiids. A broader phylogenetic comparison demonstrated that the ratio of lung mass to total body mass in kogiids, physeterids, and ziphiids was similar to that of terrestrial mammals, while delphinids and phocoenids possessed relatively large lungs. Thus, small lung size in deep-diving odontocetes may be a plesiomorphic character. The relatively large lung size of delphinids and phocoenids appears to be a derived condition that may permit the lung to function as a site of respiratory gas exchange throughout a dive in these rapid breathing, short-duration, shallow divers.

The gross morphology and histochemistry of respiratory muscles in bottlenose dolphins, Tursiops truncatus.

Most mammals possess stamina because their locomotor and respiratory (i.e., ventilatory) systems are mechanically coupled. These systems are decoupled, however, in bottlenose dolphins (Tursiops truncatus) as they swim on a breath hold. Locomotion and ventilation are coupled only during their brief surfacing event, when they respire explosively (up to 90% of total lung volume in approximately 0.3 s) (Ridgway et al. 1969 Science 166:1651-1654). The predominantly slow-twitch fiber profile of their diaphragm (Dearolf 2003 J Morphol 256:79-88) suggests that this muscle does not likely power their rapid ventilatory event. Based on Bramble's (1989 Amer Zool 29:171-186) biomechanical model of locomotor-respiratory coupling in galloping mammals, it was hypothesized that locomotor muscles function to power ventilation in bottlenose dolphins. It was further hypothesized that these muscles would be composed predominantly of fast-twitch fibers to facilitate the bottlenose dolphin's rapid ventilation. The gross morphology of craniocervical (scalenus, sternocephalicus, sternohyoid), thoracic (intercostals, transverse thoracis), and lumbopelvic (hypaxialis, rectus abdominis, abdominal obliques) muscles (n = 7) and the fiber-type profiles (n = 6) of selected muscles (scalenus, sternocephalicus, sternohyoid, rectus abdominis) of bottlenose dolphins were investigated. Physical manipulations of excised thoracic units were carried out to investigate potential actions of these muscles. Results suggest that the craniocervical muscles act to draw the sternum and associated ribs craniodorsally, which flares the ribs laterally, and increases the thoracic cavity volume required for inspiration. The lumbopelvic muscles act to draw the sternum and caudal ribs caudally, which decreases the volumes of the thoracic and abdominal cavities required for expiration. All muscles investigated were composed predominantly of fast-twitch fibers (range 61-88% by area) and appear histochemically poised for rapid contraction. These combined results suggest that dolphins utilize muscles, similar to those used by galloping mammals, to power their explosive ventilation.

Nephrolithiasis and pyelonephritis in two West Indian manatees (Trichechus manatus spp.).

Two West Indian manatees (Trichechus manatus spp.) were reported with severe emaciation. One animal was a Florida manatee from the Everglades; the other was an Antillean manatee from Cuba. On necropsy, both animals had nephrolithiasis, pyelonephritis, and moderate to severe renomegaly. Histopathology revealed multifocal to diffuse pyelonephritis, interstitial nephritis, and nephrocalcinosis. The stones were analyzed and consisted primarily of calcium carbonate. Serum chemistry values for the Florida animal revealed no renal abnormalities. The mechanism of calculus formation remains unclear in manatees. In horses, another hindgut fermenter, the most common urolith is also calcium carbonate. Urinalyses performed on manatees are very similar to those of horses (i.e., alkaline urine, low specific gravity, and calcium carbonate crystals). Formation of uroliths in manatees may have a pathogenesis similar to equine urolithiasis.

CT scans and 3D reconstructions of Florida manatee (Trichechus manatus latirostris) heads and ear bones.

The auditory anatomy of the Florida manatee (Trichechus manatus latirostris) was investigated using computerized tomography (CT), three-dimensional reconstructions, and traditional dissection of heads removed during necropsy. The densities (kg/m3) of the soft tissues of the head were measured directly using the displacement method and those of the soft tissues and bone were calculated from CT measurements (Hounsfield units). The manatee's fatty tissue was significantly less dense than the other soft tissues within the head (p<0.05). The squamosal bone was significantly less dense than the other bones of the head (p<0.05). Measurements of the ear bones (tympanic, periotic, malleus, incus, and stapes) collected during dissection revealed that the ossicular chain was overly massive for the mass of the tympanoperiotic complex.

Methods used during gross necropsy to determine watercraft-related mortality in the Florida manatee (Trichechus manatus latirostris).

Between 1993 and 2003, 713 (24%) of 2,940 dead Florida manatees (Trichechus manatus latirostris) recovered from Florida waters and examined were killed by watercraft-induced trauma. It was determined that this mortality was the result of watercraft trauma because the external wound patterns and the internal lesions seen during gross necropsy are recognizable and diagnostic. This study documents the methods used in determining watercraft-related mortality during gross necropsy and explains why these findings are diagnostic. Watercraft can inflict sharp- and blunt-force trauma to manatees, and both types of trauma can lead to mortality. This mortality may be a direct result of the sharp and blunt forces or from the chronic effects resulting from either force. In cases in which death is caused by a chronic wound-related complication, the original incident is usually considered to be the cause of death. Once a cause of death is determined, it is recorded in an extensive database and is used by Federal and state managers in developing strategies for the conservation of the manatee. Common sequelae to watercraft-induced trauma include skin lesions, torn muscles, fractured and luxated bones, lacerated internal organs, hemothorax, pneumothorax, pyothorax, hydrothorax, abdominal hemorrhage and ascites, and pyoperitoneum.

Brevetoxicosis: red tides and marine mammal mortalities.

Potent marine neurotoxins known as brevetoxins are produced by the 'red tide' dinoflagellate Karenia brevis. They kill large numbers of fish and cause illness in humans who ingest toxic filter-feeding shellfish or inhale toxic aerosols. The toxins are also suspected of having been involved in events in which many manatees and dolphins died, but this has usually not been verified owing to limited confirmation of toxin exposure, unexplained intoxication mechanisms and complicating pathologies. Here we show that fish and seagrass can accumulate high concentrations of brevetoxins and that these have acted as toxin vectors during recent deaths of dolphins and manatees, respectively. Our results challenge claims that the deleterious effects of a brevetoxin on fish (ichthyotoxicity) preclude its accumulation in live fish, and they reveal a new vector mechanism for brevetoxin spread through food webs that poses a threat to upper trophic levels.

Vascular adaptations for heat conservation in the tail of Florida manatees (Trichechus manatus latirostris).

Although Florida manatees (Trichechus manatus latirostris) have relatively low basal metabolic rates for aquatic mammals of their size, they maintain normal mammalian core temperatures. We describe vascular structures in the manatee tail that permit countercurrent heat exchange (CCHE) to conserve thermal energy. Approximately 1000 arteries juxtaposed to 2000 veins are found at the cranial end of the caudal vascular bundle (CVB); these numbers decrease caudally, but the 1:2 ratio of arteries to veins persists. Arterial walls are relatively thin when compared to those previously described in vascular countercurrent heat exchangers in cetaceans. It is assumed that CCHE in the CVB helps manatees to maintain core temperatures. Activity in warm water, however, mandates a mechanism that prevents elevated core temperatures. The tail could transfer heat to the environment if arterial blood delivered to the skin were warmer than the surrounding water; unfortunately, CCHE prevents this heat transfer. We describe deep caudal veins that provide a collateral venous return from the tail. This return, which is physically outside the CVB, reduces the venous volume within the bundle and allows arterial expansion and increased arterial supply to the skin, and thus helps prevent elevated core temperatures.