New views of humpback whale flow dynamics and oral morphology during prey engulfment

The rise of inexpensive, user‐friendly cameras and editing software promises to revolutionize data collection with minimal disturbance to marine mammals. Video sequences recorded by aerial drones and GoPro cameras provided close‐up views and unique perspectives of humpback whales engulfing juvenile salmon at or just below the water surface in Southeast Alaska and Prince William Sound. Although humpback feeding is famous for its flexibility, several stereotyped events were noted in the 47 lunges we analyzed. Engulfment was rapid (mean 2.07 s), and the entrance through which the tongue inverts into the ventral pouch was seen as water rushes in. Cranial elevation was a major contributor to gape, and pouch contraction sometimes began before full gape closure, with reverberating waves indicating rebounding flow of water within the expanded pouch. Expulsion of filtered water began with a small splash at the anterior of the mouth, followed by sustained excurrent flow in the mouth's central or posterior regions. Apart from a splash of rebounding water, water within the mouth was surprisingly turbulence‐free during engulfment, but submersion of the whale's head created visible surface whirlpools and vortices which may aggregate prey for subsequent engulfment.     

Anatomy of the laryngeal apparatus of the pygmy right whale, Caperea marginata (Gray 1846).

The laryngeal apparatus of Caperea marginata is described for the first time and proves to be significantly different from that of any mysticete previously described. This difference is especially noticeable in the position of the laryngeal sac, which is separate from the tracheal rings and lies to the right of the animal. Massive ontogenetic development of the sac in adult males is demonstrated, far greater than that seen in most other mysticetes. Histological analysis of the laryngeal sac shows the walls to be very muscular, abundantly innervated, and vascularized, indicating an active organ. Coiled blood vessels and nerves support the hypothesis that the sac undergoes extensive expansion and contraction. A possible association with the unique thoracic development of the species is suggested. The roles of the laryngeal sac and arytenoid cartilages in mysticete sound production are also discussed.     

Hydrodynamic performance of the flippers of large‐bodied cetaceans in relation to locomotor ecology

Cetaceans evolved flippers that are unique in both size and shape probably due to selection pressures associated with foraging and body size. Flippers function as control surfaces for maneuverability and stability. Flippers of cetaceans and engineered hydrofoils are similar with streamlined cross‐sections and wing‐like planforms, which affect lift, drag and hydrodynamic efficiency. Scale models of the flippers from large‐bodied (body length > 6 m) cetaceans (fin whale, killer whale, sperm whale) were constructed from computed tomography (CT) scans of flippers. Flipper planforms were highly tapered for the fin whale, a rounded, paddle‐like design for the killer whale, and a square geometry for the sperm whale. Hydrodynamic properties of the models at varying angles of attack (?40º to 40^o) were determined in a water tunnel with a multi‐axis load cell. The flippers were found to have hydrodynamic characteristics similar to engineered wings. Differences in flipper morphology of large‐bodied cetaceans and their hydrodynamic performance are associated with the requirements of aquatic locomotion involved with ecology of the whales. The flippers of the killer whale provided the greatest maneuverability, whereas the flippers of the fin whale had low drag for lunging and the flippers of the sperm whale provided lift for diving.     

STRUCTURE AND FUNCTION IN WHALE EARS

ABSTRACT

Ultrasonic echolocation abilities are well documented in several dolphin species, but hearing characteristics are unknown for most whales. Vocalization data suggest whale hearing spans infra- to ultrasonic ranges. This paper presents an overview of whale ear anatomy and analyzes 1) how whale ears are adapted for underwater hearing and 2) how inner ear differences relate to different hearing capacities among whales.

Whales have adaptations for rapid, deep diving and long submersion; e.g., broad- bore Eustachian tubes, no pinnae, and no air-filled external canals, that impact sound reception. In odontocetes, two soft tissue channels conduct sound to the ear. In mysticetes, bone and soft tissue conduction are likely. The middle ear is air-filled but has an extensible mucosa. Cochlear structures are hypertrophied and vestibular components are reduced. Auditory ganglion cell densities are double land mammal averages (2000–4000/mm). Basilar membrane lengths range 20–70 mm; gradients are larger than in terrestrial mammals. Odontocetes have 20–60% bony membrane support and basal ratios >0.6, consistent with hearing >150 kHz. Mysticetes have apical ratios <0.002 and no bony lateral support, implying acute infrasonic hearing. Cochlear hypertrophy may be adaptive for high background noise. Vestibular loss is consistent with cervical fusion. Exceptionally high auditory fiber counts suggest both mysticetes and odontocetes have ears “wired” for more complex signal processing mechanisms than most land mammals.     

Baleen wear reveals intraoral water flow patterns of mysticete filter feeding

A survey of macroscopic and microscopic wear patterns in the baleen of eight whale species (Cetacea: Mysticeti) discloses structural, functional, and life history properties of this neomorphic keratinous tissue, including evidence of intraoral water flow patterns involved in filter feeding. All baleen demonstrates wear, particularly on its medial and ventral edges, as flat outer layers of cortical keratin erode to reveal horn tubes, also of keratin, which emerge as hair‐like fringes. This study quantified five additional categories of specific wear: pitting of plates, scratching of plates, scuffing of fringes, shortening of fringes, and reorientation of fringes (including fringes directed between plates to the exterior of the mouth). Blue whale baleen showed the most pitting and sei whale baleen the most scratching; gray whale baleen had the most fringe wear. The location of worn baleen within the mouth suggests that direct contact with the tongue is not responsible for most wear, and that flowing water as well as abrasive prey or sediment carried by the flowing water likely causes pitting and scratching of plates as well as fringe fraying, scuffing, shortening, and reorientation. Baleen also has elevated vertical and horizontal ridges that are unrelated to wear; these are probably related to growth and may allow for age determination. J. Morphol. 277:453–471, 2016. © 2016 Wiley Periodicals, Inc.     

Whales from space: Four mysticete species described using new VHR satellite imagery

Large‐bodied animals such as baleen whales can now be detected with very high resolution (VHR) satellite imagery, allowing for scientific studies of whales in remote and inaccessible areas where traditional survey methods are limited or impractical. Here we present the first study of baleen whales using the WorldView‐3 satellite, which has a maximum spatial resolution of 31 cm in the panchromatic band, the highest currently available to nonmilitary professionals. We manually detected, described, and counted four different mysticete species: fin whales (Balaenoptera physalus) in the Ligurian Sea, humpback whales (Megaptera novaeangliae) off Hawaii, southern right whales (Eubalaena australis) off Península Valdés, and gray whales (Eschrichtius robustus) in Laguna San Ignacio. Visual and spectral analyses were conducted for each species, their surrounding waters, and nonwhale objects (e.g., boats). We found that behavioral and morphological differences made some species more distinguishable than others. Fin and gray whales were the easiest to discern due to their contrasting body coloration with surrounding water, and their prone body position, which is proximal to the sea surface (i.e., body parallel to the sea surface). These results demonstrate the feasibility of using VHR satellite technology for monitoring the great whales.     

MORPHOLOGY OF THE EYE AND SURROUNDING STRUCTURES OF THE BOWHEAD WHALE, BALAENA MYSTICETUS

Observations were made on eyes from 46 bowhead whales, Balaena mysticetus , taken in the subsistence harvest near Barrow, Point Hope, Savoonga, and Kaktovik, Alaska. Data reported here include palpebral, eyeball, corneal, scleral, pupillary, and lens dimensions. These quantitative data have allowed us to compare structures relative to one another and sometimes to compare them with similar structures in other species. We found, for example, that the cornea is almost three times as thick at its periphery as at its center; that when the ratio of scleral thickness and eyeball size are compared, the ratio, in the bowhead whale, is twice that of any other cetacean for which data were available; and that the corneal and pupillary width to height ratios indicate a less elongated cornea and pupil than has been reported in other cetaceans. We also found a strong correlation between body length and eyeball size indicating that within the species, unlike what is seen between species, larger animals have larger eyes. Novel observations include the presence of three periorbital fatty layers, 112 ciliary processes, the presence of scleral canals, the absence of an obvious fovea or macular region in the retina, a holangiotic pattern of fundic vessels, the presence of zonular fibers and a lens sheath, and the absence of an obvious pupillary operculum. Anatomical features like the wide angle of divergence and the palpebral dimensions suggest the absence of binocular vision while features like the size of the palpebral sac, abundant conjuctival fat, and the prominence of the retractor bulbi muscle suggest mechanisms for the protrusion and retraction of the eyeball.     

Multiaxial movements at the minke whale temporomandibular joint

Mandibular mobility accompanying gape change in Northern and Antarctic minke whales was investigated by manipulating jaws of carcasses, recording jaw movements via digital instruments (inclinometers, accelerometers, and goniometers), and examining osteological and soft tissue movements via computed tomography (CT)-scans. We investigated longitudinal (α) rotation of the mandible and mediolateral displacement at the symphysis (Ω₁) and temporomandibular joint (Ω₂) as the mouth opened (Δ). Results indicated three phases of jaw opening. In the first phase, as gape increased from zero to 8°, there was slight (<1°) α and Ω rotation. As gape increased between 20 and 30°, the mandibles rotated slightly laterally (Mean 3°), the posterior condyles were slightly medially displaced (Mean 4°), and the anterior ends at the symphysis were laterally displaced (Mean 3°). In the third phase of jaw opening, from 30° to full (≥90°) gape, these motions reversed: mandibles rotated medially (Mean 29°), condyles were laterally displaced (Mean 14°), and symphyseal ends were medially displaced (Mean 1°). Movements were observed during jaw manipulation and analyzed with CT-images that confirmed quantitative inclinometer/accelerometer data, including the unstable intermediate (Phase 2) position. Together these shifting movements maintain a constant distance for adductor muscles stretched between the skull's temporal fossa and mandible's coronoid process. Mandibular rotation enlarges the buccal cavity's volume as much as 36%, likely to improve prey capture in rorqual lunge feeding; it may strengthen and stabilize jaw opening or closure, perhaps via a simple locking or unlocking mechanism. Rotated lips may brace baleen racks during filtration. Mandibular movements may serve a proprioceptive mechanosensory function, perhaps via the symphyseal organ, to guide prey engulfment and water expulsion for filtration.     

BOOK REVIEWS

ABSTRACT

This paper provides our views on the areas of cetacean bioacoustics that are in the greatest need of study over the next several years. In doing this, we ask a number of questions we see as important to developing a better understanding of cetacean bioacoustics. The topics we will cover are: Auditory Capabilities, including hearing sensitivity, pathways of sound to the ear, intraspecific variation in hearing capabilities, and the effects of intense sound on hearing capabilities; Echolocation, including the information-bearing parameters exploited by dolphin sonar systems to discriminate and identify objects, and the functional characteristics of the internal representation generated by reflections from ensonified objects; and Acoustic Communication, including the nature of the cetacean sound generation mechanism, the behaviors associated with mysticete communication sounds, and the range over which mysticetes communicate. While other investigators may not fully agree with our suggestions as to which questions are most important for future studies of cetacean bioacoustics, it is clear that a considerable effort must still be made in order that we can better understand the bioacoustics and general behavior of these animals.     

Morphological specializations of baleen whales associated with hydrodynamic performance and ecological niche

Feeding behavior, prey type, and habitat appear to be associated with the morphological design of body, fluke, and flippers in baleen whales. Morphometric data from whaling records and recent stranding events were compiled, and morphometric parameters describing the body length, and fluke and flipper dimensions for an "average" blue whale Balaenoptera musculus, humpback whale Megaptera novaeangliae, gray whale Eschrichtius robustus, and right whale Eubalaena glacialis were determined. Body mass, body volume, body surface area, and fluke and flipper surface areas were estimated. The resultant morphological configurations lent themselves to the following classifications based on hydrodynamic principles: fast cruiser, slow cruiser, fast maneuverer, and slow maneuverer. Blue whales have highly streamlined bodies with small, high aspect ratio flippers and flukes for fast efficient cruising in the open ocean. On the other hand, the rotund right whale has large, high aspect ratio flukes for efficient slow speed cruising that is optimal for their continuous filter feeding technique. Humpbacks have large, high aspect ratio flippers and a large, low aspect ratio tail for quick acceleration and high-speed maneuvering which would help them catch their elusive prey, while gray whales have large, low aspect ratio flippers and flukes for enhanced low-speed maneuvering in complex coastal water habitats.