[Show abstract][Hide abstract] ABSTRACT: 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.
[Show abstract][Hide abstract] ABSTRACT: Most marine mammals are hypothesized to routinely dive within their aerobic dive limit (ADL). Mammals that regularly perform deep, long-duration dives have locomotor muscles with elevated myoglobin concentrations and are composed of predominantly large, slow-twitch (Type I) fibers with low mitochondrial volume densities (V(mt)). These features contribute to extending ADL by increasing oxygen stores and decreasing metabolic rate. Recent tagging studies, however, have challenged the view that two groups of extreme deep-diving cetaceans dive within their ADLs. Beaked whales (Ziphius cavirostris, Cuvier and Mesoplodon densirostris, Blainville) routinely perform the deepest and longest average dives of any air-breathing vertebrate, and short-finned pilot whales (Globicephala macrorhynchus, Gray) perform high-speed sprints at depth. We investigated the locomotor muscle morphology and estimated total body oxygen stores of these cetaceans to determine whether they (a) shared muscle design features with other deep-divers and (b) performed dives within their calculated ADLs. Muscle of both cetaceans displayed high myoglobin concentrations and large fibers, as predicted, but novel fiber profiles for diving mammals. Beaked whales possessed a sprinter's fiber-type profile, composed of approximately 80% fast-twitch (Type II) fibers with low V(mt). Approximately one-third of the muscle fibers of short-finned pilot whales were slow-twitch, oxidative, glycolytic fibers, a rare fiber-type for any mammal. The muscle morphology of beaked whales likely decreases the energetic cost of diving, while that of short-finned pilot whales supports high activity events. Calculated ADLs indicate that, at low metabolic rates, both cetaceans carry sufficient onboard oxygen to aerobically support their dives.
Journal of Experimental Biology 02/2013; · 3.24 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: The health of common bottlenose dolphins (Tursiops truncatus) within south-ern Georgia estuaries is of particular concern due to high levels of anthropo-genic contaminants in their tissues. Dolphins in this region have the highest 1 Corresponding author (e-mail: email@example.com).
[Show abstract][Hide abstract] ABSTRACT: 1. 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. 2. 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. 3. 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. 4. 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.
[Show abstract][Hide abstract] ABSTRACT: Blubber, the specialized hypodermis of cetaceans, provides thermal insulation through the quantity and quality of lipids it contains. Quality refers to % lipid content; however, not all lipids are the same. Certain deep-diving cetacean groups possess blubber with lipids - wax esters (WE) - that are not typically found in mammals, and the insulative quality of "waxy" blubber is unknown. Our study explored the influence of lipid storage class - specifically WE in pygmy sperm whales (Kogia breviceps; n=7) and typical mammalian triacylglycerols (TAG) in short-finned pilot whales (Globicephala macrorhynchus; n=7) - on blubber's thermal properties. Although the blubber of both species had similar total lipid contents, the thermal conductivity of G. macrorhynchus blubber (0.20 ± 0.01 Wm-1°C-1) was significantly higher than that of K. breviceps (0.15 ± 0.01 Wm-1°C-1; P=0.0006). These results suggest that lipid class significantly influences blubber's ability to resist heat flow. In addition, because blubber's lipid content is known to be stratified, we measured its depth-specific thermal conductivities. In K. breviceps blubber, the depth-specific conductivity values tended to vary inversely with lipid content. In contrast, G. macrorhynchus blubber displayed unexpected depth-specific relationships between lipid content and conductivity, which suggests that temperature-dependent effects, such as melting, may be occurring. Differences in heat flux measurements across the depth of the blubber samples provide evidence that both species are capable of storing heat in their blubber. The function of blubber as an insulator is complex and may rely upon its lipid class, stratified composition, and dynamic heat storage capabilities.
Journal of Experimental Biology 09/2012; · 3.24 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: 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.
Journal of Morphology 05/2012; 273(8):932-42. · 1.60 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Bottlenose dolphins (Tursiops truncatus) are apex predators in coastal southeastern U.S. waters; as such they are indicators of persistent organic pollutants (POPs) in coastal ecosystems. POP concentrations measured in a dolphin's blubber are influenced by a number of factors, including the animal's sex and ranging pattern in relation to POP point sources. This study examined POP concentrations measured in bottlenose dolphin blubber samples (n=102) from the Georgia, USA coast in relation to individual ranging patterns and specifically, distance of sightings from a polychlorinated biphenyl (PCB) point source near Brunswick, Georgia. Dolphin ranging patterns were determined based upon 5years of photo-identification data from two field sites approximately 40km apart: (1) the Brunswick field site, which included the Turtle/Brunswick River Estuary (TBRE), and (2) the Sapelo field site, which included the Sapelo Island National Estuarine Research Reserve (SINERR). Dolphins were categorized into one of three ranging patterns from photo-identification data. Individuals with sighting histories exclusively within one of the defined field sites were considered to have either Brunswick or Sapelo ranging patterns. Individuals sighted in both field sites were classified as having a Mixed ranging pattern. Brunswick males had the highest concentrations of PCBs reported for any marine mammal. The pattern of PCB congeners was consistent with Aroclor 1268, a highly chlorinated PCB mixture associated with a Superfund site in Brunswick. PCB levels in Sapelo males were lower than in Brunswick males, but comparable to the highest levels measured in other dolphin populations along the southeastern U.S. Female dolphins had higher Aroclor 1268 proportions than males, suggesting that the highly chlorinated congeners associated with Aroclor 1268 may not be offloaded through parturition and lactation, as easily as less halogenated POPs. Individuals sighted farther from the Superfund point source had lower Aroclor 1268 proportions.
Science of The Total Environment 02/2011; 409(11):2094-101. · 3.16 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Electronic tags using the ARGOS system (CLS, 2008) have proven to be valuable tools in assess-ing small cetacean movement patterns and habitat use (e.g., Read & Westgate, 1997; Corkeron & Martin, 2004; Klatsky et al., 2007; Balmer et al., 2008; Wells et al., 2008, 2009). While tag design and success rates have varied, problems associ-ated with package size, attachment position on the dorsal fin, and number of attachment pins have, in some cases, shortened the predicted attachment duration or caused adverse impacts to the dorsal fins of the animals (Read & Westgate, 1997; Scott et al., 1990; Balmer et al., 2010). One of the most recent iterations in the evolution of smaller sat-ellite-linked tags for dolphins used a 65 g, side-mounted tag, which attached to the upper third of the dorsal fin with three plastic pins (Balmer et al., 2010). This tag design has been used to determine movement patterns and dive durations in sev-eral species of small cetaceans, including bottle-nose dolphins (Tursiops truncatus) off Bermuda (Klatsky et al., 2007), Risso's dolphins (Grampus griseus) in the Gulf of Mexico and Atlantic Ocean (Wells et al., 2009; R. Wells, pers. obs.), rough-toothed dolphins (Steno bredanensis) in the Atlantic Ocean (Wells et al., 2008; R. Wells, pers. obs.), and Franciscana dolphins (Pontoporia blain-villei) in the Atlantic coastal waters off Argentina (R. Wells, pers. obs.). While this tag design appears to be relatively robust, a recent deployment of the tag demonstrated that the design is not well-suited
[Show abstract][Hide abstract] ABSTRACT: The understanding of a species' niche is fundamental to the concept of ecology, yet rela-tively little work has been done on niches in pelagic marine mammal communities. Data collection on the distribution and abundance of marine mammals is costly, time consuming and complicated by logis-tical difficulties. Here we take advantage of a data archive comprising many different datasets on the dis-tribution and abundance of cetaceans from Nova Scotia through the Gulf of Mexico in an effort to un-cover community structure at large spatial scales (1000s of km). We constructed a multivariate ordination of the species data, tested for group structure that might exist within the ordination space, and deter-mined how these groups might differ in environmental space. We examined 3 biogeographic regions: the oceanic waters north and south of Cape Hatteras, NC, and the Gulf of Mexico. North of Hatteras, we found 2 main groups split along a temperature and chlorophyll gradient, with most piscivores being found in cooler, more productive waters of the continental shelf, and most teuthivores being found far-ther offshore in warmer, less productive waters at the shelf break (200 m isobath). South of Hatteras, we found 3 groups, with the largest group being in warmer, lower chlorophyll waters that are closest to shore. In the Gulf of Mexico, we found 7 groups arrayed along a bottom depth gradient. We also tested the effect of taxonomically lumping different beaked whale species on ordination results. Results showed that when beaked whales were identified to the species level, they clustered out into distinct niches that are separate from those of other Odontocete groups. These results add to an increasing understanding of wildlife habitat associations and niche partitionings in the community structure of pelagic species, and provide important baseline information for future population monitoring efforts.
Marine Ecology Progress Series 01/2011; 434:165-181. · 2.64 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Most exposure assessments for free-ranging cetaceans focus on contaminant concentrations measured in blubber, and few data are available for other tissues or the factors governing contaminant distribution among tissues. The goal of this study was to provide a detailed description of the distribution of persistent organic pollutants (POPs) within the common bottlenose dolphin (Tursiops truncatus) body and assess the role of lipid dynamics in mediating contaminant distribution. Thirteen tissues (brain, blubber, heart, liver, lung, kidney, mammary gland, melon, skeletal muscle, spleen, thyroid, thymus, and testis/uterus) were sampled during necropsy from bottlenose dolphins (n = 4) and analyzed for lipid and 85 POPs, including polychlorinated biphenyls, organochlorine pesticides, and polybrominated diphenyl ethers. Significant correlations between tissue POP concentrations and lipid suggest that distribution of POPs is generally related to tissue lipid content. However, blubber:tissue partition coefficients ranged widely from 0.753 to 6.25, suggesting that contaminant distribution is not entirely lipid-dependent. Tissue-specific and whole-body contaminant burdens confirmed that blubber, the primary site of metabolic lipid storage, is also the primary site for POP accumulation, contributing >90% to the whole-body burdens. Observations also suggest that as lipid mobilizes from blubber, contaminants may redistribute, leading to elevated tissue concentrations. These results suggest that individuals with reduced blubber lipid may be at increased risk for exposure-related health effects. However, this study also provides evidence that the melon, a metabolically inert lipid-rich structure, may serve as an alternate depot for POPs, thus preventing the bulk of blubber contaminants from being directly available to other tissues. This unique physiological adaptation should be taken into consideration when assessing contaminant-related health effects in wild cetacean populations.
Environmental Toxicology and Chemistry 06/2010; 29(6):1263-73. · 2.62 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: Shallow-diving, coastal bottlenose dolphins (Tursiops truncatus) and deep-diving, pelagic pygmy and dwarf sperm whales (Kogia breviceps and K. sima) will experience vastly different ambient pressures at depth, which will influence the volume of air within their lungs and potentially the degree of thoracic collapse they experience. This study tested the hypotheses that lung size will be reduced and/or thoracic mobility will be enhanced in deeper divers. Lung mass (T. truncatus, n = 106; kogiids, n = 18) and lung volume (T. truncatus, n = 5; kogiids, n = 4), relative to total body mass, were compared. One T. truncatus and one K. sima were cross-sectioned to calculate lung, thoracic vasculature, and other organ volumes. Excised thoraxes (T. truncatus, n = 3; kogiids, n = 4) were mechanically manipulated to compare changes in thoracic cavity shape and volume. Kogiid lungs were half the mass and one-fifth the volume of those of similarly sized T. truncatus. The lungs occupied only 15% of the total thoracic cavity volume in K. sima and 37% in T. truncatus. The kogiid and dolphin thoraxes underwent similar changes in shape and volume, although the width of the thoracic inlet was relatively constrained in kogiids. A broader phylogenetic comparison demonstrated that the ratio of lung mass to total body mass in kogiids, physeterids, and ziphiids was similar to that of terrestrial mammals, while delphinids and phocoenids possessed relatively large lungs. Thus, small lung size in deep-diving odontocetes may be a plesiomorphic character. The relatively large lung size of delphinids and phocoenids appears to be a derived condition that may permit the lung to function as a site of respiratory gas exchange throughout a dive in these rapid breathing, short-duration, shallow divers.
Journal of Morphology 06/2010; 271(6):654-73. · 1.60 Impact Factor
[Show abstract][Hide abstract] ABSTRACT: In Atlantic bottlenose dolphins (Tursiops truncatus) the thickness and lipid content of blubber (the integument's specialized hypodermis) varies across ontogeny and with reproductive and nutritional state. Because the integument comprises up to 25% of total body mass in this species, ontogenetic changes in its lipid content may influence whole body buoyancy. The density and volume of the integument were measured and its buoyancy calculated across an ontogenetic series of dolphins and in pregnant and emaciated adults (total n= 45). Regional differences between the metabolically labile trunk integument and the structural tailstock integument were also investigated. Mean densities of both trunk and tailstock integument were similar across life history categories (trunk = 1,040.7 ± 14.1 kg/m3; tailstock = 1,077.1 ± 21.2 kg/m3) and were statistically similar to the density of seawater (1,026 kg/m3). The mean buoyant force of integument from the trunk (−1.01 ± 1.74 N) and tailstock (−0.30 ± 0.21 N) did not vary significantly across ontogeny. In contrast, pregnancy and emaciation did influence the integument's buoyancy, which ranged between 9 N and −45 N in these categories. Although neutral during growth, the integument's contribution to whole body buoyancy can be influenced by an individual's reproductive and nutritional status.
[Show abstract][Hide abstract] ABSTRACT: The temperature differential (ΔT) between a body surface and the environment influences an organism's heat balance. In Sarasota Bay, FL, where ambient water temperature (Tw) ranges annually from 11° to 33°C, ΔT was investigated in a resident community of bottlenose dolphins (Tursiops truncatus). Dorsal fin surface temperatures (Tdfin) were measured on wild, free-swimming dolphins using infrared thermography. Field and laboratory calibration studies were also undertaken to assess the efficacy of this non-invasive technology in the marine environment. The portability of infrared thermography permitted measurements of Tdfin across the entire range of environmental temperatures experienced by animals in this region. Results indicated a positive, linear relationship between Tdfin and Tw (r2= 0.978, P < 0.001). On average, Tdfin was 0.9°C warmer than Tw across seasons, despite the 22°C annual range in Tw. Changes in integumentary and vascular insulation likely account for the stability of ΔTdfin − w and the protection of core temperature (Tcore) across seasons. The high thermal conductivity of water may also influence this ΔT. The use of infrared thermography is an effective, non-invasive method of assessing dorsal fin skin surface temperatures (±1°C) across large numbers of wild, free-swimming dolphins throughout their thermally dynamic aquatic environment.
[Show abstract][Hide abstract] ABSTRACT: The subdermal connective tissue sheath (SDS) of dolphins is a fibre-reinforced membrane connected to other locomotor tissues, including blubber, axial muscles and tendons, and vertebral column. The complicated connections between the SDS and other locomotor tissues suggest that the SDS acts as a peripheral skeletal element for the axial locomotor muscles and as an anchor for a de novo dermal appendage, the dorsal fin. The morphology of the SDS suggests that the dolphin can be modelled as a fibre-wound, thin-walled, pressurized cylinder. Existing cylinder models predict that the SDS functions to resist torsional forces, prevent aneurysms, and limit wrinkling when the dolphin bends in locomotion. I present a new functional model that more accurately represents the morphology of the dolphin cylinder wrapped by the SDS. The new model predicts that the SDS: (1) acts as a retinaculum for the terminal tendons of the axial locomotor muscles; and (2) plays a role in maintaining the laterally flattened cross-sectional shape of the caudal peduncle. The model is based on external morphological features of dolphins shared by other steady swimming aquatic vertebrates, such as carangiform and thunniform fishes. These features, which include a streamlined body shape and narrow necking of the caudal peduncle have been identified as adaptions to reduce drag. The new model offers insight into some of the structural features of the body wall required to maintain the hydrodynamicallytuned, external morphology of steady-swimming vertebrates.
[Show abstract][Hide abstract] ABSTRACT: Late-term fetal bottlenose dolphins (Tursiops truncatus) are bent ventrolaterally en utero, requiring extreme flexibility of the axial skeleton and associated soft tissues. At birth, neonatal dolphins must immediately swim to the surface to breath, yet the dorsoventral oscillations used during locomotion may be compromised by the lateral flexibility evident in the fetus. The unique fetal position of dolphins, coupled with their need to swim at birth, places conflicting mechanical demands on the tailstock. Our previous research demonstrated that neonatal dolphins possess laterally placed, axial muscles that are functionally specialized to actively maintain the straightened posture of the tailstock. Here, we investigated the development of passive lateral stability in the tailstock of bottlenose dolphins by performing whole-body bending tests on an ontogenetic series of stranded dolphin specimens (N=15), including fetuses, neonates and juveniles (total length 58-171 cm). Structural stiffness increased, while overall body curvature decreased, with increasing body length. Scaling analyses suggest that increased structural stiffness is due to increases in size and probably changes in the passive material properties of the tailstock through ontogeny. The neutral zone was approximately constant with increasing size, while the relative neutral zone (neutral zone/total length) decreased. The lateral stability of the tailstock appears to be controlled by a combination of active and passive systems and the role of these systems varies through ontogeny. While neonates use active, muscular mechanisms to limit lateral deformations of the tailstock, the stability of the maturing tailstock is due primarily to its passive tissue properties.
[Show abstract][Hide abstract] ABSTRACT: Most mammals possess stamina because their locomotor and respiratory (i.e., ventilatory) systems are mechanically coupled. These systems are decoupled, however, in bottlenose dolphins (Tursiops truncatus) as they swim on a breath hold. Locomotion and ventilation are coupled only during their brief surfacing event, when they respire explosively (up to 90% of total lung volume in approximately 0.3 s) (Ridgway et al. 1969 Science 166:1651-1654). The predominantly slow-twitch fiber profile of their diaphragm (Dearolf 2003 J Morphol 256:79-88) suggests that this muscle does not likely power their rapid ventilatory event. Based on Bramble's (1989 Amer Zool 29:171-186) biomechanical model of locomotor-respiratory coupling in galloping mammals, it was hypothesized that locomotor muscles function to power ventilation in bottlenose dolphins. It was further hypothesized that these muscles would be composed predominantly of fast-twitch fibers to facilitate the bottlenose dolphin's rapid ventilation. The gross morphology of craniocervical (scalenus, sternocephalicus, sternohyoid), thoracic (intercostals, transverse thoracis), and lumbopelvic (hypaxialis, rectus abdominis, abdominal obliques) muscles (n = 7) and the fiber-type profiles (n = 6) of selected muscles (scalenus, sternocephalicus, sternohyoid, rectus abdominis) of bottlenose dolphins were investigated. Physical manipulations of excised thoracic units were carried out to investigate potential actions of these muscles. Results suggest that the craniocervical muscles act to draw the sternum and associated ribs craniodorsally, which flares the ribs laterally, and increases the thoracic cavity volume required for inspiration. The lumbopelvic muscles act to draw the sternum and caudal ribs caudally, which decreases the volumes of the thoracic and abdominal cavities required for expiration. All muscles investigated were composed predominantly of fast-twitch fibers (range 61-88% by area) and appear histochemically poised for rapid contraction. These combined results suggest that dolphins utilize muscles, similar to those used by galloping mammals, to power their explosive ventilation.
Journal of Morphology 10/2008; 269(12):1520-38. · 1.60 Impact Factor