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.
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.
Marine mammals are exposed to O
2
-limitation and increased N
2
gas concentrations as they dive to exploit habitat and food resources. The lipid-rich tissues (blubber, acoustic, neural) are of ...particular concern as N
2
is five times more soluble in lipid than in blood or muscle, creating body compartments that can become N
2
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 N
2
gas exposure by dive type in the species examined, the lipids of neural tissues likely do not have a neuroprotective role in marine mammals.
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.
How to Build a Deep Diver Pabst, D. Ann; McLellan, William A.; Rommel, Sentiel A.
Integrative and comparative biology,
12/2016, Letnik:
56, Številka:
6
Journal Article
Recenzirano
Odprti dostop
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.
Aim
The aim of this study was to determine if marine mammals follow ecogeographic rules. We examined Bergmann's rule and Allen's rule in two pilot whale species with contrasting latitudinal ...distributions.
Location
Northwest Atlantic Ocean.
Taxon
Globicephala spp.
Methods
We analysed morphometric data collected from strandings of short‐ and long‐finned pilot whales in the Northwest Atlantic Ocean to assess intraspecific and interspecific variation in surface area to volume ratios (SA:V) of the body core and appendage surface area relative to body core SA (normalized appendage SA) using a novel 3D modelling method.
Results
Our results suggest that ecogeographic variation in morphometrics between the two pilot whale species is consistent with morphological adaptations required to balance heat conservation and heat dissipation. Interspecific differences in morphology supported Bergmann's rule for fully grown individuals: the more temperate long‐finned pilot whale had a larger body size and lower body core SA:V than the short‐finned pilot whale, which has a more tropical distribution. Allen's rule was not supported; when all appendages were considered together, long‐finned pilot whales had larger normalized SA than short‐finned pilot whales. However, the pectoral flippers were the primary driver of this relationship; while long‐finned pilot whales had proportionally larger pectoral flippers, short‐finned pilot whales had proportionally larger dorsal fins and flukes. In addition, larger long‐finned pilot whales (i.e. males and mature individuals) had proportionally larger pectoral flippers than smaller long‐finned pilot whales.
Main Conclusions
Pilot whales follow Bergmann's rule but do not follow Allen's rule when fully mature. Thinly insulated appendages in marine mammals can be used to dissipate heat as the core warms, and larger and better insulated marine mammals may require relatively larger appendages in order to offload heat and thermoregulate effectively. Our results provide novel insight into ecogeographic rules and suggest that species in higher latitude climates towards the poles will demonstrate tradeoffs between core body heat conservation and appendage heat dissipation.
Most marine mammals are hypothesized to routinely dive within their aerobic dive limit (ADL). Mammals that regularly perform deep, long-duration dives have locomotor muscles with elevated myoglobin ...concentrations that are composed of predominantly large, slow-twitch (Type I) fibers with low mitochondrial volume densities (V(mt)). These features contribute to extending ADL by increasing oxygen stores and decreasing metabolic rate. Recent tagging studies, however, have challenged the view that two groups of extreme deep-diving cetaceans dive within their ADLs. Beaked whales (including Ziphius cavirostris and Mesoplodon densirostris) routinely perform the deepest and longest average dives of any air-breathing vertebrate, and short-finned pilot whales (Globicephala macrorhynchus) perform high-speed sprints at depth. We investigated the locomotor muscle morphology and estimated total body oxygen stores of several species within these two groups of cetaceans to determine whether they (1) shared muscle design features with other deep divers and (2) performed dives within their calculated ADLs. Muscle of both cetaceans displayed high myoglobin concentrations and large fibers, as predicted, but novel fiber profiles for diving mammals. Beaked whales possessed a sprinter's fiber-type profile, composed of ~80% fast-twitch (Type II) fibers with low V(mt). Approximately one-third of the muscle fibers of short-finned pilot whales were slow-twitch, oxidative, glycolytic fibers, a rare fiber type for any mammal. The muscle morphology of beaked whales likely decreases the energetic cost of diving, while that of short-finned pilot whales supports high activity events. Calculated ADLs indicate that, at low metabolic rates, both beaked and short-finned pilot whales carry sufficient onboard oxygen to aerobically support their dives.
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 (
and
) and a model terrestrial species (the pig-
) 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 (
< 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.
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.
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.