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Between June 1987 and March 1988, bottlenose dolphins (Tursiops truncatus Montagu 1821) along the US Atlantic coast experienced an epizootic. Monthly interquartile ranges of strandings during the epizootic were used to propose the Coastal Migratory Stock (CMS) of bottlenose dolphins (Scott et al., 1988). To date, the hypothesised CMS remains poorly understood. The goal of this study was to use a 25-year database to compare stranding patterns during the epizootic to those before (1972-1986) and after (1989-1997) the event. These comparisons reveal that monthly interquartile ranges during the epizootic are dissimilar to those before and after the event. The frequency distribution of total monthly strandings during the epizootic is also significantly different from those observed outside the event. Seasonal stranding patterns from 1989-1997 suggest more complex movements of dolphins along the US Atlantic coast than those of a single group ranging seasonally from Florida to New Jersey. In winter, for example, when the current model for the CMS predicts dolphin distributions concentrated in central Florida, the highest number of strandings occurred in North Carolina. Thus, these comparative analyses suggest that the pattern observed during the epizootic was anomalous, and not representative of stranding distributions for any other time period of the study. During the 15 years before the epizootic, and the nine years following, there was no clear picture of ‘migration’ of mortality along the coast. This study demonstrates how long-term, systematic collection of strandings data can be useful in testing hypotheses regarding the complex stock structure of coastal bottlenose dolphins. This knowledge will greatly enhance the ability to conserve and manage these animals as they recover from historic (i.e. directed takes and epizootic) and current sources of mortality.

In recent years, humpback whales (Megaptera novaeangliae) have been observed in the waters of the US mid-Atlantic states (USMA; New Jersey to North Carolina), notably in winter. The level of the mortality in this area (52 recorded deaths from 1990-2000), makes it important to understand the nature and population identity of this aggregation. Of the approximately 100 humpback whales documented in this study, photographs of 41 (live or dead) were of sufficient quality to be compared to catalogues from the Gulf of Maine (GOM, the closest feeding ground) and elsewhere in the North Atlantic. Of 22 live whales, 10 (45.5%) matched to the GOM, 5 (22.7%) to Newfoundland and 1 (4.5%) to the Gulf of St Lawrence (GSL). Of 19 dead whales, 6 (31.6%) were known GOM whales. Although the population composition of the USMA is dominated by GOM whales, lack of recent photographic effort in Newfoundland makes it likely that the observed match rates under represent the true presence of Canadian whales in the region. Length data from 48 stranded whales (18 females, 22 males and 8 of unknown sex) suggest that 39 (81.2%) were first-year animals, 7 (14.6%) were immature and 2 (4.2%) were adults. However, sighting histories of five of the dead whales indicate that some were small for their age and histories of live whales further indicate that the population contains a greater percentage of mature animals than is suggested by the stranded sample. The authors suggest that the study area primarily represents a supplemental winter feeding ground that is used by humpbacks for more than one purpose. From a management perspective, although the only successful matches of mortalities to date have been to the GOM, the observed mixing of live whales from different summer stocks might suggest that the high numbers of mortalities occurring there may not be impacting this single stock alone. Although further data are required before conclusions can be drawn, the mortality rate may be significant for the GOM population and this warrants further investigation.

In the US mid-Atlantic, multi-disciplinary studies are underway to elucidate the complex stock structure of coastal bottlenose dolphins (Tursiops truncatus), as well as the degree of overlap between coastal and offshore ecotypes. In this study we use geo-referenced data, collected during aerial surveys in 2000-2002, to describe the distribution and relative abundance of bottlenose dolphins along the US midAtlantic coast. Two aerial survey designs were used: (1) onshore/offshore surveys out to 35 n.miles during winter from Georgia to Virginia; and (2) coastal surveys throughout the year along North Carolina (NC). The winter onshore/offshore surveys demonstrated that significantly more bottlenose dolphins occur in Raleigh Bay (between Cape Hatteras and Cape Lookout, NC), than in all other regions. Additionally, in winter most bottlenose dolphins occur in the coastal waters of NC; nearly half of all sightings occurred between the shoreline and 3km from shore. The year-round, coastal surveys demonstrated that this winter distribution pattern is the result of a distinct seasonal increase in the number of dolphins within the coastal waters of NC. Circular statistical analyses demonstrated a strong influence of season on dolphin abundance. Relatively few bottlenose dolphins were observed in late spring, summer, and early autumn, with increased numbers observed during winter. In all seasons but summer, dolphin numbers were highest in Raleigh Bay. Thus, the results of both surveys indicate the importance of the habitat surrounding Cape Hatteras to bottlenose dolphins. Dolphins may preferentially use these waters in response to changes in prey distribution and/or abiotic factors such as water temperature. These results reveal an overall seasonal movement pattern along the US Atlantic coast, which appears to be correlated, at least in part, to water temperature gradients and prey availability. Although the stock identity of dolphins sighted during these aerial surveys could not be ascertained, focused photo-identification efforts, together with enhanced genetic sampling, would provide insights into the movement patterns, and, thus, stock identity, of dolphins in this region.

In this paper the occurrence, distribution and abundance of cetaceans in offshore waters of Onslow Bay, North Carolina, USA is described. Between June 2007 and June 2010 monthly aerial and shipboard line-transect surveys were conducted along ten 74km transects placed perpendicular to the shelf break. In total 42,676km of aerial trackline (218 sightings) and 5,209km of vessel trackline (100 sightings) were observed. Seven species of cetaceans were observed, but the fauna was dominated strongly by common bottlenose and Atlantic spotted dolphins. Both species were present year-round in the study area. Using photo-identification techniques, five bottlenose dolphins and one spotted dolphin were resighted during the three-year period. In general, the abundance of cetaceans in Onslow Bay was low and too few sightings were made to estimate monthly abundances for species other than bottlenose and spotted dolphins. Maximum monthly abundances of bottlenose and spotted dolphins were 4,100 (95% CI: 1,300–9,400) in May 2010 and 6,000 (95% CI: 2,500–17,400) in March 2009, respectively. Bottlenose dolphins were found throughout the study area, although they were encountered most frequently just off the shelf break. In contrast, spotted dolphins exhibited a strong preference for waters over the continental shelf and were not encountered beyond the shelf break.

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.

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.

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.

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.

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.

Beaked whales are vulnerable to the impacts of disturbance from several sources of anthropogenic sound. Here we report the distribution and abundance of beaked whales off Cape Hatteras, North Carolina, USA, an area utilized by the U.S. Navy for training exercises, and of particular interest for seismic geophysical surveys. From May 2011 through November 2015, monthly aerial surveys were conducted at the site. Beaked whales were encountered 74 times (n= 205 individuals) during these surveys. Ziphius cavirostris, the most commonly encountered species, was observed in every month of the year. Mesoplodon spp. were encountered in ten months of the year. Photographs of adult males with erupted teeth permitted six sightings to be identified conclusively as M. europaeus; M. mirus was also photographed just outside the study area. Beaked whale surface densities stratified by depth (0.005 – 0.007/km2) were among the highest reported in the world for small ziphiids. A quantitative comparison of sightings and stranding records suggests that strandings do not accurately reflect the relative abundance of beaked whale species in this area. We conclude that Cape Hatteras, at the convergence of the Labrador Current and Gulf Stream, is a particularly important year-round habitat for several species of beaked whales. Postprint

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 parasitic nematode Crassicauda sp. was initially described in kogiid whales from specimens collected within cervical tissues, uncommon sites of infection for this parasite. Crassicauda sp. has only been reported in Kogia breviceps to date, but no study has yet investigated a large sample of both kogiid species. A 15 yr record of 104 kogiid strandings (K. sima, n = 40; K. breviceps, n = 64) in North Carolina and Virginia, U.S.A. was used to determine the prevalence of Crassicauda sp. across species, within species across sex, and within sex across length and life history categories. Crassicauda sp. was confirmed to be a species-specific parasite among kogiids infecting only K. breviceps (prevalence = 45%). Within K. breviceps, prevalence was similar (45%) in both immature and mature males, but increased from 10% in immature to 76% in mature females. This study confirmed the cervico-thoracic distribution of the parasite, and identified a novel site of infection in a previously undescribed exocrine gland associated with the pigmented “false gill slit.” The species-specific nature of Crassicauda sp. infection, the exocrine gland, and the distinct features of the false gill slit pigmentation associated with the gland, are all useful characters to identify kogiid species in the field.

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.

Growth in common bottlenose dolphins (Tursiops truncatus) was investigated through examination of sex-specific, ontogenetic changes in the mass of 38 discrete body compartments, utilizing stranded dolphins in good body condition (n = 145). Ontogenetic allometry and the body composition technique were used to quantitatively describe growth patterns. Although adult males were significantly larger than adult females in total body mass (TBM) and total length, overall patterns of growth were remarkably similar between sexes. The integument, locomotor muscle, and vertebral column together represented 50%–58% of TBM across all life history categories, although their relative contributions varied ontogenetically. Young dolphins invested the greatest percentage of TBM in integument, while locomotor muscle was the single largest body component in adults. In both sexes (1) most muscle groups displayed positive allometry, (2) most skeletal elements displayed negative allometric or isometric growth, (3) most abdominal viscera associated with digestion displayed positive allometry, and (4) the brain displayed negative allometric growth. Reproductive tissues exhibited the highest rates of growth in both sexes, and increased as a percentage of TBM with maturity. This study provides an integrated view of bottlenose dolphin growth and a quantitative baseline of body composition for future monitoring of this sentinel species of ecosystem health.

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. Anat Rec, 298:1301–1326, 2015. © 2015 Wiley Periodicals, Inc.

Cetaceans span a large range of body sizes and include species with the largest known locomotor muscles. To date, force output and thrust production have only been directly measured in the common bottlenose dolphin (Tursiops truncatus), although thrust forces have been hydrodynamically modeled for some whales. In this study, two metrics of epaxial muscle size—cross-sectional area (CSA) and mass—were used to estimate force output for 22 species (n = 83 specimens) ranging in size from bottlenose dolphins to blue whales (Balaenoptera musculus). Relative to total body length (TL), maximum force output estimated based upon both muscle CSA (TL1.56 ± 0.05) and mass (TL2.64 ± 0.07) scaled at rates lower than those predicted by geometric scaling, suggesting relative force output decreases with increasing body size in cetaceans. Estimated maximal force outputs were compared to both published drag forces and to the breaking strengths of commercial fishing lines known to entangle whales. The breaking strengths of these lines are within the same order of magnitude, and in some cases, exceed the estimated maximal force output of whales. These results suggest that while powerful animals, large whales may be unable to break the extremely strong fishing line used today.

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.

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.

Several species of odontocete cetaceans depredate bait and catch and, as a result, become hooked and entangled in pelagic longline fisheries. The present study measured how selected commercial longline hooks, including “weak hooks”, behaved within odontocete mouths. Five hooks (Mustad-16/0, Mustad-18/0, Mustad J-9/0, Korean 16, and Korean 18) were tested on three species of odontocetes known to interact with longline fisheries—short-finned pilot whales (Globicephala macrorhynchus), Risso's dolphins (Grampus griseus), and false killer whales (Pseudorca crassidens). Specimens were secured to a stanchion, hooks were placed in the mouth at multiple positions along the dorsal lip, and the force required to pull each hook free was measured. The soft tissue lips of these odontocetes were capable of resisting forces up to 250 kg before failing. The polished steel M-16, M-18, and J-9 hooks straightened at forces between 50 and 225 kg, depending on hook gauge. When straightened, these hooks exposed the sharpened barb, which sliced through the lip tissue, usually releasing the hook intact. The K-16 and K-18 hooks behaved very differently, breaking at higher forces (110–250 kg) and consistently just at the barb; usually, there was measurable soft-tissue loss and often shards of the hook were retained within those soft tissues. The different behaviours of these two hook types—the M and J type polished steel vs. the K type carbon steel—were consistent across all species tested. Mechanical tests were also conducted to determine if hooks could fracture the mandible of these same odontocetes. Only the M-18 and K-18 hooks had sufficiently large gapes to hook around the mandible, and both hook types fractured bone in short-finned pilot whales and Risso's dolphins. These results support other lines of evidence indicating that longline hooks can cause serious injury to these species, and suggest possible steps to mitigate these impacts.

Electronic tags have proven to be valuable tools in assessing small cetacean movement and behavior. However, problems associated with tag size and attachment have limited duration and damaged dorsal fins. These outcomes have motivated researchers to develop a new satellite-linked tag design that reduces detrimental effects to tagged animals, while increasing transmission durations. The goals of this study were to review previous studies that deployed single-pin transmitters and determine factors that influence transmission duration. Then, test these factors utilizing computational fluid dynamics (CFD) models to identify an optimal single-pin satellite-linked tag design, and evaluate this prototype through field studies. A review of four projects, which deployed 77 single-pin radio tags, determined that tags attached along the lower third of the dorsal fin and approximately 33 mm from the trailing edge resulted in longer transmission durations and reduced negative impacts to the dorsal fin. Based upon these results and CFD modeling, prototype, single-pin satellite-linked tags (n = 25) transmitted for 163 ± 22 d (mean ± 95% CI) which greatly exceeded transmissions for previous small cetacean telemetry studies. These results suggest that the newly developed single-pin satellite-linked tag design strikes a balance between reducing impacts to the individual while maximizing transmissions.

A complementary approach of stomach content and stable isotope analyses was used to characterize the foraging ecology and evaluate niche overlap between pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales stranded on the U.S. mid-Atlantic coast between 1998 and 2011. Food habits analysis demonstrated both species were primarily teuthophagous, with 35 species of cephalopods, and 2 species of mesopelagic fishes represented in their overall diets. Pianka's Index of niche overlap suggested high overlap between whale diets (On = 0.92), with squids from the families Histioteuthidae, Cranchidae, and Ommastrephidae serving as primary prey. Pygmy sperm whales consumed slightly larger prey sizes (mean mantle length [ML] = 10.8 cm) than dwarf sperm whales (mean ML = 7.8 cm). Mean prey sizes consumed by pygmy sperm whales increased with growth, but showed no trend in dwarf sperm whales. Significant differences were not detected in δ15N and δ13C values of muscle tissues from pygmy (10.8‰ ± 0.5‰, −17.1‰ ± 0.6‰), and dwarf sperm whales (10.7‰ ± 0.5‰, −17.0‰ ± 0.4‰), respectively. Isotopic niche widths also did not differ significantly and dietary overlap was high between the two species. Results suggest the feeding ecologies of the pygmy and dwarf sperm whales are similar and both species occupy equivalent trophic niches in the region.

Under US policy, a stock is a group of animals shown to be demographically independent from other such groups. Evidence of demographic delineations for marine mammal stocks is gained primarily through genetic analysis. However, additional techniques play an important role in determining fine-scale ranging patterns that can be used to define a stock's geographic boundaries. In cases where genetic studies have not yet been performed, movement and ranging pattern data are essential in identifying a targeted geographic region for tissue sampling and genetic studies. Photo-identification surveys, vessel-based radio telemetry, automated radio telemetry systems (ARTS), and satellite-linked telemetry are sampling techniques that have been used to determine common bottlenose dolphin ranging patterns and provide detailed insight into stock boundaries. The results of near simultaneous use of these sampling techniques are compared using data from a case study of bottlenose dolphins within the estuaries of southern Georgia. Satellite-linked and radio telemetry were determined to be useful sampling techniques for identification of short-term ranging patterns. Satellite-linked telemetry had the second lowest cost per location ($122) and identified dolphin ranging patterns within and outside of the study area boundaries. Vessel-based radio telemetry was more costly ($195 per location) and had relatively limited tracking coverage. However, this sampling technique permitted visual observations of animal and tag condition. The combination of vessel-based radio telemetry and ARTS, which had the lowest cost per location ($34), was an effective method for determining ranging patterns of tagged individuals within and outside of the study area. Photo-identification surveys, relative to satellite-linked and radio telemetry, were not as efficient for determination of targeted individuals’ short-term ranging patterns and had the highest cost per location of the four sampling techniques ($292). However, photo-identification is more effective than any other technique for compiling data on large numbers of individuals within a designated study area. Photo-identification surveys are essential for long-term monitoring and provide additional insight into dolphin stock structure that cannot be determined through telemetry alone. Copyright © 2013 John Wiley & Sons, Ltd.

When a marine mammal dives, breathing and locomotion are mechanically uncoupled, and its loco- motor muscle must power swimming when oxygen is lim- ited. The morphology of that muscle provides insight into both its oxygen storage capacity and its rate of oxygen consumption. This study investigated the m. longissimus dorsi, an epaxial swimming muscle, in the long duration, deep-diving pygmy sperm whale (Kogia breviceps) and the short duration, shallow-diving Atlantic bottlenose dol- phin (Tursiops truncatus). Muscle myoglobin content, fiber type profile (based upon myosin ATPase and succi- nate dehydrogenase assays), and fiber size were meas- ured for five adult specimens of each species. In addition, a photometric analysis of sections stained for succinate dehydrogenase was used to create an index of mitochon- drial density. The m. longissimus dorsi of K. breviceps dis- played significantly a) higher myoglobin content, b) larger proportion of Type I (slow oxidative) fibers by area, c) larger mean fiber diameters, and d) lower indices of mi- tochondrial density than that of T. truncatus. Thus, this primary swimming muscle of K. breviceps has greater ox- ygen storage capacity, reduced ATP demand, and likely a reduced rate of oxygen consumption relative to that of T. truncatus. The locomotor muscle of K. breviceps appears able to ration its high onboard oxygen stores, a feature that may allow this species to conduct relatively long du- ration, deep dives aerobically. J. Morphol. 274:663-675, 2013. ! 2013 Wiley Periodicals, Inc.

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.

The health of common bottlenose dolphins (Tursiops truncatus) within southern Georgia estuaries is of particular concern due to high levels of anthropogenic contaminants in their tissues. Dolphins in this region have the highest polychlorinated biphenyl (PCB) concentrations recorded for any marine mammal and these concentrations correlate to distance from a Superfund point-source in the Turtle/Brunswick River Estuary (TBRE). Currently, little is known about the population structure of dolphins in this region. This study identifies and compares baseline data on abundance, habitat use, site-fidelity, and ranging patterns of dolphins across two adjacent field sites; Brunswick, including the TBRE, and Sapelo, including the Sapelo Island National Estuarine Research Reserve. Sapelo is relatively undeveloped and was selected for comparison to the more contaminated TBRE. Dolphin densities increased with tributary size in both sites but dolphin density and total abundance were significantly higher in Sapelo than in Brunswick. Anthropogenic stressors within the TBRE may be an important factor contributing to the differences in abundance, density, and habitat use observed in this study.

Blubber, a specialized form of subdermal adipose tissue, surrounds marine mammal bodies. Typically, adipose tissue is perfused by capillaries but information on blubber vascularization is lacking. This study's goals were to: 1) describe and compare the microvasculature (capillaries, microarterioles, and microvenules) of blubber across odontocete species; 2) compare microvasculature of blubber to adipose tissue; and 3) examine relationships between blubber's lipid composition and its microvasculature. Percent microvascularity, distribution, branching pattern, and diameter of microvessels were determined from images of histochemically stained blubber sections from shallow-diving bottlenose dolphins (Tursiops truncatus), deeper-diving pygmy sperm whales (Kogia breviceps), deep-diving beaked whales (Mesoplodon densirostris; Ziphius cavirostris), and the subdermal adipose tissue of domestic pigs (Sus scrofa). Tursiops blubber showed significant stratification in percent microvascularity among the superficial, middle, and deep layers and had a significantly higher percent microvascularity than all other animals analyzed, in which the microvasculature was more uniformly distributed. The percent microvasculature of Kogia blubber was lower than that of Tursiops but higher than that of beaked whales and the subdermal adipose tissue of domestic pigs. Tursiops had the most microvascular branching. Microvessel diameter was relatively uniform in all species. There were no clear patterns associating microvascular and lipid characteristics. The microvascular characteristics of the superficial layer of blubber resembled the adipose tissue of terrestrial mammals, suggesting some conservation of microvascular patterns in mammalian adipose tissue. The middle and deep layers of blubber, particularly in Tursiops, showed the greatest departure from typical mammalian microvascular arrangement. Factors such as metabolics or thermoregulation may be influencing the microvasculature in these layers. © J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.

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.

To enhance our understanding of how aquatic endotherms respond thermally to their environment, we present two new designs for recording heat flux (HF) and skin temperature (Ts) from the dorsal fins of free-ranging dolphins. The first package, held on by a series of small suction cups, was designed for short-term deployments on the order of hours. The second package, a long-term attachment, was held on via two 5.0-mm -Delrin® pins and was designed for deployments on the order of days. The former version was configured to release remotely, by way of a galvanic linkage, removing the need to re-capture tagged dolphins. The latter was designed to be removed after re-capture of the individual but also included a fail–safe galvanic release. One of the major advances of our approach is that it allows both convective and conductive pathways of heat loss to be dynamically recorded from freeswimming wild dolphins, whereas previous approaches have recorded primarily conductive heat loss from trained captive or restrained wild dolphins. The short-term, re-usable package has been field-tested on 55 bottlenose dolphins (Tursiops truncatus )i n Sarasota Bay, Florida and the long-term device on two spotted dolphins (Stenella attenuata) in the eastern tropical Pacific Ocean. By also incorporating into each package a time–depth–velocity recorder we were able to examine patterns of heat flux in relation to changing behavioural states of the tagged individual. Heat flux recorded at the dorsal fin was found to be individually variable and dynamic. Water temperature (Tw) had a profound effect on patterns of heat loss in both species with HF increasing with even minor decreases in Tw. This device significantly improves our ability to understand the thermal biology of dolphins as they move about in their natural environment. © 2007 Published by Elsevier B.V.

Field necropsies of individual or multiple animals can be fraught with difficulty and frustration. Planning and preparation for these events, including development of required supplies and equipment checklists, necropsy forms, designated test sites, and a personnel pool, can alleviate many problems. Flexibility is paramount in field situations as is the need for individuals from multiple disciplines to collaboratively collect, record, disseminate, and interpret findings.

SUMMARYIn Atlantic bottlenose dolphins Tursiops truncatus, both the thickness and lipid content of blubber vary across ontogeny and across individuals of differing reproductive and nutritional status. This study investigates how these changes in blubber morphology and composition influence its thermal properties. Thermal conductivity (W m–1deg.–1, where deg. is °C) and thermal insulation(m2 deg. W–1) of dolphin blubber were measured in individuals across an ontogenetic series (fetus through adult, N=36),pregnant females (N=4) and emaciated animals (N=5). These thermal properties were determined by the simultaneous use of two common experimental approaches, the heat flux disc method and the standard material method. Thickness, lipid and water content were measured for each blubber sample. Thermal conductivity and insulation varied significantly across ontogeny. Blubber from fetuses through sub-adults was less conductive(range=0.11–0.13±0.02 W m–1deg.–1) than that of adults (mean=0.18 W m–1deg.–1). The conductivity of blubber from pregnant females was similar to non-adult categories, while that of emaciated animals was significantly higher (0.24 ± 0.04 W m deg.–1) than all other categories. Blubber from sub-adults and pregnant females had the highest insulation values while fetuses and emaciated animals had the lowest. In nutritionally dependant life history categories, changes in blubber's thermal insulation were characterized by stable blubber quality (i.e. conductivity)and increased blubber quantity (i.e. thickness). In nutritionally independent animals, blubber quantity remained stable while blubber quality varied. A final, unexpected observation was that heat flux measurements at the deep blubber surface were significantly higher than that at the superficial surface, a pattern not observed in control materials. This apparent ability to absorb heat, coupled with blubber's fatty acid composition, suggest that dolphin integument may function as a phase change material.

Cetacean brains are among the least studied mammalian brains because of the formidable histological preparations of such relatively rare and large specimens. Although the bottlenose dolphin, Tursiops truncatus, has been the most extensively studied cetacean species, there have been relatively few studies of the brain of the infant bottlenose dolphin. In this study, we present the first magnetic resonance imaging (MRI)-based study of the brain of an infant bottlenose dolphin. Magnetic resonance images in the coronal plane were originally acquired and used to digitally generate a set of resectioned virtual images in orthogonal planes. A sequential set of images in all three planes was anatomically labeled and reveals major neuroanatomical features. Some of the distinctive features of cetacean brains are already evident in the infant bottlenose dolphin brain, while other features may represent differences that deserve further study.

This study investigated the functional morphology of the blubber that forms the caudal keels of the harbor porpoise (Phocoena phocoena). Blubber is a pliant biocomposite formed by adipocytes and structural fibers composed of collagen and elastic fibers. Caudal keels are dorsally and ventrally placed triangular wedges of blubber that define the hydrodynamic profile of the porpoise tailstock. Mechanical tests on carcasses demonstrate that when keels are bent, they strain nonuniformly along their lengths, with highest strains just caudal to the dorsal fin and lowest at the insertion of the flukes. Therefore, caudal keels undergo nonuniform longitudinal deformation while maintaining a stable, triangular cross-sectional shape. Polarizing and transmitted light microscopy techniques were used to investigate blubber's 3D fiber architecture along the length of the dorsal keel. The triangular cross-sectional shape of the keel appears to be maintained by structural fibers oriented to act as tensile stays. The construction of the blubber composite is regionally specific :structural fiber densities and diameters are higher in the relatively stiff caudal region of the keel than in the more deformable cranial keel region. The orientations of structural fibers also change along the length of the keel. Cranially, no fibers are oriented along the long axis, whereas a novel population of longitudinally oriented fibers reinforces the keel at the insertion of the flukes. Thus, differences in the distribution and orientation of structural fibers contribute to the regionally specific mechanical properties of the dorsal keel.

We describe reproductive seasonality of bottlenose dolphins in North Carolina (NC), U.S.A., using strandings data from the entire coast of NC and sighting data from Beaufort, NC and by estimating dates of birth of known females. We found a strong peak of neonate strandings in the spring (April-May), and low levels of neonate strandings in the fall and winter. The distribution of neonate strandings was significantly different from a uniform distribution (P < 0.001, K = 3.8). We

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

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.

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°–36.4° 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. Anat Rec 264:339–347, 2001. Published 2001 Wiley-Liss, Inc.

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.

The similarity in swimming style and external body shape between dolphins and scombrid fishes, especially tunas, is a textbook example of evolutionary convergence. I identify additional morphological features of the musculoskeletal system shared by dolphins and tunas. Specifically, these swimmers share a pattern of force transmission through a complex, three-dimensional system of collagenous fabrics, which are stiffened by muscular hydrostatic pressure. This force transmission system increases both the displacement advantage and moment arm of contracting axial muscle. These features represent a functionally significant design for steady swimming vertebrates.

We investigated patterns of abundance and distribution for coastal migratory Atlantic bottlenose dolphins (Tursiops truncatus) that appear seasonally in the nearshore waters of Virginia Beach, Virginia. The study was conducted along 24 km of shoreline at the southern point of the Chesapeake Bay mouth from April 1994 to March 1995. This is the first study to investigate the relationship between the abundance of coastal migratory dolphins and factors that might affect their movement. A profile analysis of variance revealed significant differences in local abundance and distribution throughout the year. Dolphin number was positively correlated with water temperature and not correlated with photoperiod. Although prey distribution and abundance are two factors thought to affect dolphin presence, in this study the relationship between these two factors and dolphin abundance was unclear. Greater numbers of dolphins were found in the ocean section of the study area. However, significantly higher ratios of neonatal dolphins were observed in the bay section, suggesting the bay serves as a nursery area. The observed relationship between local dolphin abundance and environmental factors in Virginia may provide insight into dolphin distribution and migration along the Atlantic coast of the United States.