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.

Fetal distress and in utero pneumonia in perinatal dolphins during the Northern Gulf of Mexico unusual mortality event.

An unusual mortality event (UME) involving primarily common bottlenose dolphins Tursiops truncatus of all size classes stranding along coastal Louisiana, Mississippi, and Alabama, USA, started in early 2010 and continued into 2014. During this northern Gulf of Mexico UME, a distinct cluster of perinatal dolphins (total body length <115 cm) stranded in Mississippi and Alabama during 2011. The proportion of annual dolphin strandings that were perinates between 2009 and 2013 were compared to baseline strandings (2000-2005). A case-reference study was conducted to compare demographics, histologic lesions, and Brucella sp. infection prevalence in 69 UME perinatal dolphins to findings from 26 reference perinates stranded in South Carolina and Florida outside of the UME area. Compared to reference perinates, UME perinates were more likely to have died in utero or very soon after birth (presence of atelectasis in 88 vs. 15%, p < 0.0001), have fetal distress (87 vs. 27%, p < 0.0001), and have pneumonia not associated with lungworm infection (65 vs. 19%, p = 0.0001). The percentage of perinates with Brucella sp. infections identified via lung PCR was higher among UME perinates stranding in Mississippi and Alabama compared to reference perinates (61 vs. 24%, p = 0.01), and multiple different Brucella omp genetic sequences were identified in UME perinates. These results support that from 2011 to 2013, during the northern Gulf of Mexico UME, bottlenose dolphins were particularly susceptible to late-term pregnancy failures and development of in utero infections including brucellosis.

Nitrogen solubility in odontocete blubber and mandibular fats in relation to lipid composition.

Understanding toothed whale (odontocete) diving gas dynamics is important given the recent atypical mass strandings of odontocetes (particularly beaked whales) associated with mid-frequency naval sonar. Some stranded whales have exhibited gas emboli (pathologies resembling decompression sickness) in their specialized intramandibular and extramandibular fat bodies used for echolocation and hearing. These tissues have phylogenetically unique, endogenous lipid profiles with poorly understood biochemical properties. Current diving gas dynamics models assume an Ostwald nitrogen (N2) solubility of 0.07 ml N2 ml(-1) oil in odontocete fats, although solubility in blubber from many odontocetes exceeds this value. The present study examined N2 solubility in the blubber and mandibular fats of seven species across five families, relating it to lipid composition. Across all species, N2 solubility increased with wax ester content and was generally higher in mandibular fats (0.083 ± 0.002 ml N2 ml(-1) oil) than in blubber (0.069 ± 0.007 ml N2 ml(-1) oil). This effect was more pronounced in mandibular fats with higher concentrations of shorter, branched fatty acids/alcohols. Mandibular fats of short-finned pilot whales, Atlantic spotted dolphins and Mesoplodon beaked whales had the highest N2 solubility values (0.097 ± 0.005, 0.081 ± 0.007 and 0.080 ± 0.003 ml N2 ml(-1) oil, respectively). Pilot and beaked whales may experience high N2 loads during their relatively deeper dives, although more information is needed about in vivo blood circulation to mandibular fats. Future diving models should incorporate empirically measured N2 solubility of odontocete mandibular fats to better understand N2 dynamics and potential pathologies from gas/fat embolism.

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.

Gut bacterial communities in Atlantic bottlenose dolphins (Tursiops truncatus) throughout a disease-driven (Morbillivirus) unusual mortality event.

Gut microbiomes are important determinants of animal health. In sentinel marine mammals where animal and ocean health are connected, microbiome impacts can scale to ecosystem-level importance. Mass mortality events affect cetacean populations worldwide, yet little is known about the contributory role of their gut bacterial communities to disease susceptibility and progression. Here, we characterized bacterial communities from fecal samples of common bottlenose dolphins, Tursiops truncatus, across an unusual mortality event (UME) caused by dolphin Morbillivirus (DMV). 16S rRNA gene sequence analysis revealed similar diversity and structure of bacterial communities in individuals stranding before, during, and after the 2013-2015 Mid-Atlantic Bottlenose Dolphin UME and these trends held in a subset of dolphins tested by PCR for DMV infection. Fine-scale shifts related to the UME were not common (10 of 968 bacterial taxa) though potential biomarkers for health monitoring were identified within the complex bacterial communities. Accordingly, acute DMV infection was not associated with a distinct gut bacterial community signature in T. truncatus. However, temporal stratification of DMV-positive dolphins did reveal changes in bacterial community composition between early and late outbreak periods, suggesting that gut community disruptions may be amplified by the indirect effects of accumulating health burdens associated with chronic morbidity.

Remarkable consistency of spinal cord microvasculature in highly adapted diving odontocetes.

Odontocetes are breath-hold divers with a suite of physiological, anatomical, and behavioral adaptations that are highly derived and vastly different from those of their terrestrial counterparts. Because of these adaptations for diving, odontocetes were originally thought to be exempt from the harms of nitrogen gas embolism while diving. However, recent studies have shown that these mammals may alter their dive behavior in response to anthropogenic sound, leading to the potential for nitrogen supersaturation and bubble formation which may cause decompression sickness in the central nervous system (CNS). We examined the degree of interface between blood, gases, and neural tissues in the spinal cord by quantifying its microvascular characteristics in five species of odontocetes (Tursiops truncatus, Delphinus delphis, Grampus griseus, Kogia breviceps, and Mesoplodon europaeus) and a model terrestrial species (the pig-Sus scrofa domesticus) for comparison. This approach allowed us to compare microvascular characteristics (microvascular density, branching, and diameter) at several positions (cervical, thoracic, and lumbar) along the spinal cord from odontocetes that are known to be either deep or shallow divers. We found no significant differences (p < 0.05 for all comparisons) in microvessel density (9.30-11.18%), microvessel branching (1.60-2.12 branches/vessel), or microvessel diameter (11.83-16.079 µm) between odontocetes and the pig, or between deep and shallow diving odontocete species. This similarity of spinal cord microvasculature anatomy in several species of odontocetes as compared to the terrestrial mammal is in contrast to the wide array of remarkable physio-anatomical adaptations marine mammals have evolved within their circulatory system to cope with the physiological demands of diving. These results, and other studies on CNS lipids, indicate that the spinal cords of odontocetes do not have specialized features that might serve to protect them from Type II DCS.

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.

Lipid signature of neural tissues of marine and terrestrial mammals: consistency across species and habitats.

Marine mammals are exposed to O2-limitation and increased N2 gas concentrations as they dive to exploit habitat and food resources. The lipid-rich tissues (blubber, acoustic, neural) are of particular concern as N2 is five times more soluble in lipid than in blood or muscle, creating body compartments that can become N2 saturated, possibly leading to gas emboli upon surfacing. We characterized lipids in the neural tissues of marine mammals to determine whether they have similar lipid profiles compared to terrestrial mammals. Lipid profiles (lipid content, lipid class composition, and fatty acid signatures) were determined in the neural tissues of 12 cetacean species with varying diving regimes, and compared to two species of terrestrial mammals. Neural tissue lipid profile was not significantly different in marine versus terrestrial mammals across tissue types. Within the marine species, average dive depth was not significantly associated with the lipid profile of cervical spinal cord. Across species, tissue type (brain, spinal cord, and spinal nerve) was a significant factor in lipid profile, largely due to the presence of storage lipids (triacylglycerol and wax ester/sterol ester) in spinal nerve tissue only. The stability of lipid signatures within the neural tissue types of terrestrial and marine species, which display markedly different dive behaviors, points to the consistent role of lipids in these tissues. These findings indicate that despite large differences in the level of N2 gas exposure by dive type in the species examined, the lipids of neural tissues likely do not have a neuroprotective role in marine mammals.

Myoglobin Concentration and Oxygen Stores in Different Functional Muscle Groups from Three Small Cetacean Species.

Compared with terrestrial mammals, marine mammals possess increased muscle myoglobin concentrations (Mb concentration, g Mb · 100g-1 muscle), enhancing their onboard oxygen (O2) stores and their aerobic dive limit. Although myoglobin is not homogeneously distributed, cetacean muscle O2 stores have been often determined by measuring Mb concentration from a single muscle sample (longissimus dorsi) and multiplying that value by the animal's locomotor muscle or total muscle mass. This study serves to determine the accuracy of previous cetacean muscle O2 stores calculations. For that, body muscles from three delphinid species: Delphinus delphis, Stenella coeruleoalba, and Stenella frontalis, were dissected and weighed. Mb concentration was calculated from six muscles/muscle groups (epaxial, hypaxial and rectus abdominis; mastohumeralis; sternohyoideus; and dorsal scalenus), each representative of different functional groups (locomotion powering swimming, pectoral fin movement, feeding and respiration, respectively). Results demonstrated that the Mb concentration was heterogeneously distributed, being significantly higher in locomotor muscles. Locomotor muscles were the major contributors to total muscle O2 stores (mean 92.8%) due to their high Mb concentration and large muscle masses. Compared to this method, previous studies assuming homogenous Mb concentration distribution likely underestimated total muscle O2 stores by 10% when only considering locomotor muscles and overestimated them by 13% when total muscle mass was considered.

Host phylogeny and life history stage shape the gut microbiome in dwarf (Kogia sima) and pygmy (Kogia breviceps) sperm whales.

Gut microbiomes perform crucial roles in host health and development, but few studies have explored cetacean microbiomes especially deep divers. We characterized the gut microbiomes of stranded dwarf (Kogia sima) and pygmy (K. breviceps) sperm whales to examine the effects of phylogeny and life stage on microbiome composition and diversity. 16S rRNA gene sequence analysis revealed diverse gut communities (averaging 674 OTUs) dominated by a few symbiont taxa (25 OTUs accounted for 64% of total relative abundance). Both phylogeny and life stage shaped community composition and diversity, with species-specific microbiome differences present early in life. Further analysis showed evidence of microbiome convergence with host maturity, albeit through different processes: symbiont 'accumulation' in K. sima and 'winnowing' in K. breviceps, indicating different methods of community assembly during host development. Furthermore, culture-based analyses yielded 116 pure cultures matching 25 OTUs, including one isolate positive for chitin utilization. Our findings indicate that kogiid gut microbiomes are highly diverse and species-specific, undergo significant shifts with host development, and can be cultivated on specialized media under anaerobic conditions. These results enhance our understanding of the kogiid gut microbiome and may provide useful information for symbiont assessment in host health.

Locomotor muscle morphology of three species of pelagic delphinids.

The locomotor muscle morphology of diving mammals yields insights into how they utilize their environment and partition resources. This study examined a primary locomotor muscle, the longissimus, in three closely related, similarly sized pelagic delphinids (n = 7-9 adults of each species) that exhibit different habitat and depth preferences. The Atlantic spotted dolphin (Stenella frontalis) is a relatively shallow diver, inhabiting continental shelf waters; the striped (Stenella coeruleoalba) and short-beaked common (Delphinus delphis) dolphins are sympatric, deep-water species that dive to different depths. Based upon comparative data from other divers, it was hypothesized that the locomotor muscle of the deepest-diving S. coeruleoalba would exhibit a higher percentage of slow oxidative fibers, larger fiber diameters, a higher myoglobin concentration [Mb], and a lower mitochondrial density than that of the shallow-diving S. frontalis, and that the muscle of D. delphis would display intermediate values for these features. As expected, the locomotor muscle of S. coeruleoalba exhibited a significantly higher proportion of slow (57.3 ± 3.9%), oxidative (51.7 ± 2.5%) fibers and higher [Mb] (8.2 ± 0.7 g/100 g muscle) than that of S. frontalis (41.3 ± 3.9%, 31.0 ± 3.2%, 4.7 ± 0.05 g/100 g muscle, respectively). There were no differences in fiber size or mitochondrial density among these species. Like other deep divers, S. coeruleoalba displayed locomotor muscle features that enhance oxygen storage capacity and metabolic efficiency but did not display features that limit aerobic capacity. These results suggest a previously undescribed muscle design for an active, small-bodied, deep-diving cetacean. HIGHLIGHTS: The locomotor muscle features displayed by the striped dolphin, which are unique among deep divers, enhance oxygen stores but do not limit aerobic capacity. This novel muscle design may facilitate the active lifestyle of this small-bodied deep diver.

Evaluation of hematologic and biochemical values for convalescing seals from the coast of Maine.

Marine mammal researchers and veterinarians evaluate the health of individuals and populations by using observed ranges for blood values. The goal of this study was to establish observed ranges for dependent (0-4 week) and weanling (>4 week) harbor seal (Phoca vitulina) pups that were admitted for rehabilitation at the Marine Animal Lifeline in Portland, Maine from 1998-2003 and to record changes in hematologic and biochemical values as they convalesced. Blood values, measured both at admission to and release from rehabilitation, were quantitatively compared between these two groups. Of the 29 blood analytes measured in this study between dependent and weanling pups upon admission to rehabilitation, 14 were significantly different including white blood cell (WBC), red blood cell (RBC), hemoglobin (Hb), hematocrit (HCT), platelets, lymphocytes, monocytes, potassium, chloride, CO2, total bilirubin, calcium, alanine transaminase (ALT), and alkaline phosphatase. Between admission and release, 14 analytes had significantly changed for dependent pups including RBC, Hb, HCT, mean cell volume (MCV), mean corpuscular hemoglobin (MCH), platelets, lymphocytes, monocytes, potassium, blood urea nitrogen (BUN), total protein, total bilirubin, alkaline phosphatase, and ALT. Between admission and release, 10 analytes had significantly changed for weanlings including RBC, MCV, neutrophils, eosinophils, potassium, chloride, anion gap, total protein, total bilirubin, and calcium. At the time of release, six parameters were significantly different between pups admitted as dependent and those admitted as weanlings including MCV, neutrophils, eosinophils, total protein, globulin, and albumin.

Running, swimming and diving modifies neuroprotecting globins in the mammalian brain.

The vulnerability of the human brain to injury following just a few minutes of oxygen deprivation with submergence contrasts markedly with diving mammals, such as Weddell seals (Leptonychotes weddellii), which can remain underwater for more than 90 min while exhibiting no neurological or behavioural impairment. This response occurs despite exposure to blood oxygen levels concomitant with human unconsciousness. To determine whether such aquatic lifestyles result in unique adaptations for avoiding ischaemic-hypoxic neural damage, we measured the presence of circulating (haemoglobin) and resident (neuroglobin and cytoglobin) oxygen-carrying globins in the cerebral cortex of 16 mammalian species considered terrestrial, swimming or diving specialists. Here we report a striking difference in globin levels depending on activity lifestyle. A nearly 9.5-fold range in haemoglobin concentration (0.17-1.62 g Hb 100 g brain wet wt(-1)) occurred between terrestrial and deep-diving mammals; a threefold range in resident globins was evident between terrestrial and swimming specialists. Together, these two globin groups provide complementary mechanisms for facilitating oxygen transfer into neural tissues and the potential for protection against reactive oxygen and nitrogen groups. This enables marine mammals to maintain sensory and locomotor neural functions during prolonged submergence, and suggests new avenues for averting oxygen-mediated neural injury in the mammalian brain.

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.

High diversity and unique composition of gut microbiomes in pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales.

Mammals host diverse bacterial and archaeal symbiont communities (i.e. microbiomes) that play important roles in digestive and immune system functioning, yet cetacean microbiomes remain largely unexplored, in part due to sample collection difficulties. Here, fecal samples from stranded pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales were used to characterize the gut microbiomes of two closely-related species with similar diets. 16S rRNA gene sequencing revealed diverse microbial communities in kogiid whales dominated by Firmicutes and Bacteroidetes. Core symbiont taxa were affiliated with phylogenetic lineages capable of fermentative metabolism and sulfate respiration, indicating potential symbiont contributions to energy acquisition during prey digestion. The diversity and phylum-level composition of kogiid microbiomes differed from those previously reported in toothed whales, which exhibited low diversity communities dominated by Proteobacteria and Actinobacteria. Community structure analyses revealed distinct gut microbiomes in K. breviceps and K. sima, driven by differential relative abundances of shared taxa, and unique microbiomes in kogiid hosts compared to other toothed and baleen whales, driven by differences in symbiont membership. These results provide insight into the diversity, composition and structure of kogiid gut microbiomes and indicate that host identity plays an important role in structuring cetacean microbiomes, even at fine-scale taxonomic levels.

A novel gammaherpesvirus associated with genital lesions in a Blainville's beaked whale (Mesoplodon densirostris).

An adult male Blainville's beaked whale (Mesoplodon densirostris) was found stranded on the Atlantic coast of the USA on 28 January 2004. Necropsy revealed a focal papilloma-like penile lesion, the cells from which revealed single 4-6 microm basophilic intranuclear inclusions. Total DNA extracted from lesion material was tested using a pan-herpes-virus PCR assay that targets the DNA polymerase gene and found to be positive. When the amplified DNA fragment was cloned, sequenced, and compared to GenBank-deposited herpesvirus DNA polymerase sequences, the detected virus was determined to be a distinct member of the Gammaherpesvirinae subfamily of herpesviruses. This new virus, tentatively named Ziphiid herpesvirus type 1, was associated with but not determined to be the cause of genital disease in the Blainville's beaked whale.

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.

Habitat-based cetacean density models for the U.S. Atlantic and Gulf of Mexico.

Cetaceans are protected worldwide but vulnerable to incidental harm from an expanding array of human activities at sea. Managing potential hazards to these highly-mobile populations increasingly requires a detailed understanding of their seasonal distributions and habitats. Pursuant to the urgent need for this knowledge for the U.S. Atlantic and Gulf of Mexico, we integrated 23 years of aerial and shipboard cetacean surveys, linked them to environmental covariates obtained from remote sensing and ocean models, and built habitat-based density models for 26 species and 3 multi-species guilds using distance sampling methodology. In the Atlantic, for 11 well-known species, model predictions resembled seasonal movement patterns previously suggested in the literature. For these we produced monthly mean density maps. For lesser-known taxa, and in the Gulf of Mexico, where seasonal movements were less well described, we produced year-round mean density maps. The results revealed high regional differences in small delphinoid densities, confirmed the importance of the continental slope to large delphinoids and of canyons and seamounts to beaked and sperm whales, and quantified seasonal shifts in the densities of migratory baleen whales. The density maps, freely available online, are the first for these regions to be published in the peer-reviewed literature.

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.

Postural role of lateral axial muscles in developing bottlenose dolphins (Tursiops truncatus).

Foetal dolphins (Tursiops truncatus) are bent ventrolaterally, such that the tailflukes and lower jaw are juxtaposed. The lateral flexibility required en utero may compromise the efficiency of the dorsoventral oscillations required of the swimming neonate. The m. intertransversarius caudae dorsalis (IT) is the most laterally placed epaxial muscle. Bilateral contractions of the IT could limit lateral deformations of the flexible tailstock of the early neonate. We test the hypothesis that the IT is functioning as a postural muscle in neonates by examining its morphological, histological and biochemical properties. The neonatal IT has a relatively large cross-sectional area and bending moment, as well as a large proportion of slow-twitch fibres and elevated myoglobin concentrations. Our results demonstrate that the IT is functionally capable of performing this specific postural function in neonatal dolphins. In later life-history stages, when postural control is no longer needed, the IT serves to fine-tune the position of the tailstock during locomotion. The changing function of the adult IT is concomitant with changes in morphology and biochemistry, and most notably, with an increase in the proportion of fast-twitch fibres. We suggest that these changes reflect strong selective pressure to improve locomotor abilities by limiting lateral deformations during this critical life-history stage.