Baleen Hydrodynamics and Morphology of Cross-Flow Filtration in Balaenid Whale Suspension Feeding

The traditional view of mysticete feeding involves static baleen directly sieving particles from seawater using a simple, dead-end flow-through filtration mechanism. Flow tank experiments on bowhead (Balaena mysticetus) baleen indicate the long-standing model of dead-end filtration, at least in balaenid (bowhead and right) whales, is not merely simplistic but wrong. To recreate continuous intraoral flow, sections of baleen were tested in a flume through which water and buoyant particles circulated with variable flow velocity. Kinematic sequences were analyzed to investigate movement and capture of particles by baleen plates and fringes. Results indicate that very few particles flow directly through the baleen rack; instead much water flows anteroposteriorly along the interior (lingual) side of the rack, allowing items to be carried posteriorly and accumulate at the posterior of the mouth where they might readily be swallowed. Since water flows mainly parallel to rather than directly through the filter, the cross-flow mechanism significantly reduces entrapment and tangling of minute items in baleen fringes, obviating the need to clean the filter. The absence of copepods or other prey found trapped in the baleen of necropsied right and bowhead whales supports this hypothesis. Reduced through-baleen flow was observed with and without boundaries modeling the tongue and lips, indicating that baleen itself is the main if not sole agent of crossflow. Preliminary investigation of baleen from balaenopterid whales that use intermittent filter feeding suggests that although the biomechanics and hydrodynamics of oral flow differ, cross-flow filtration may occur to some degree in all mysticetes.     

Archaeocete-like jaws in a baleen whale

The titanic baleen whales (Cetacea, Mysticeti) have a bizarre skull morphology, including an elastic mandibular symphysis, which permits dynamic oral cavity expansion during bulk feeding. How this key innovation evolved from the sutured symphysis of archaeocetes has remained unclear. Now, mandibles of the Oligocene toothed mysticete Janjucetus hunderi show that basal mysticetes had an archaeocete-like sutured symphysis. This archaic morphology was paired with a wide rostrum typical of later-diverging baleen whales. This demonstrates that increased oral capacity via rostral widening preceded the evolution of mandibular innovations for filter feeding. Thus, the initial evolution of the mysticetes' unique cranial form and huge mouths was perhaps not linked to filtering plankton, but to enhancing suction feeding on individual prey.     

Developmental changes in the skull morphology of common minke whales Balaenoptera acutorostrata

We investigated growth‐related and sex‐related morphological changes in the skulls of 144 North Pacific common minke whales Balaenoptera acutorostrata. Measurement was conducted at 39 points on the skull and mandible to extract individual allometric equations relating the length and zygomatic width of the skull. The results revealed no significant differences in skull morphology by sex except for width of occipital bone. The size relative to the skull of the anatomical parts involved in feeding, such as the rostrum and mandible, increased after birth. In contrast, the sensory organs and the anatomical regions involved in neurological function, such as the orbit, tympanic bullae, and foramen magnum, were fully developed at birth, and their relative size reduced over the course of development. This is the first study to investigate developmental changes in the skull morphology using more than 100 baleen whale specimens, and we believe the results of this study will contribute greatly to multiple areas of baleen whale research, including taxonomy and paleontology. J. Morphol. 275:1113–1121, 2014. © 2014 Wiley Periodicals, Inc.     

Material and structural properties of fin whale (Balaenoptera physalus) Zwischensubstanz

The oral anatomy of the fin whale (Balaenoptera physalus) consists of several major structures crucial to its engulfment method of feeding, such as stiff keratinized baleen plates, a large flaccid tongue, and a prominent vomer. One under‐documented part of this anatomy is the cream white Zwischensubstanz that holds the baleen plates to the rostrum at their dorsal base. The mechanical and structural properties of Zwischensubstanz play a key role in baleen plate dynamics and, on the grand scale, contribute to baleen whales' filtration efficiency and attainment of large body size. Compression and tensile tests on the Zwischensubstanz sampled from an 18 m fin whale showed that this material unexpectedly exhibits linear isotropic behaviour with Elastic Modulus of 2.56 ± 0.60 MPa and hysteresis of 0.44 ± 0.02 in compression despite apparent unidirectional growth. Acting similar to a soft rubber, the Zwischensubstanz absorbs and dissipates the enormous forces acting on baleen plates during engulfment feeding while maintaining spacing between the plates to maximize filtration efficiency. Microscopic analysis provided images of connective tissue papillae penetrating the base of the Zwischensubstanz and developing within it to emerge as fully formed, keratinized baleen plates. The plates develop from the papillae and a connective tissue sheet within the 5–7 cm deep Zwischensubstanz. The Zwischensubstanz provides a keratin matrix of concentrically oriented fibers around each papilla forming the hard baleen plates and frayed fringes used for filter feeding. During this formation, the Zwischensubstanz remains unchanged and appears to slough away to allow the baleen plate to grow unhindered. J. Morphol. 274:947–955, 2013. © 2013 Wiley Periodicals, Inc.     

Exceptional occurrence of fossil baleen in shallow marine sediments of the Neogene Pisco Formation,Southern Peru

Fossil baleen is rare in the sedimentary record. This paper documents the exceptional occurrence of thirty seven fossil whale specimens with preserved baleen in the Neogene Pisco Formation during a transect survey in a limited area west of the Ica River Valley near the town of Ocucaje in southern Peru. The sedimentary layers consist of tuffaceous and diatomaceous sandstones, diatomaceous mudstones, and dolomites, deposited in a shallow marine embayment. Observations of modern whale carcasses on the seafloor and stranded individuals indicate that baleen detaches from the mouth of the whales very rapidly after death, and that bones deteriorate very rapidly as a result of scavenging activity and abrasion. In contrast, the bones of the Pisco Formation whales are exceptionally well preserved, and their baleen is often found in life position suspended from the rostrum. Sedimentary structures found associated with some skeletons indicate tidal and storm processes, suggesting that the environment was not anoxic. This exceptional occurrence of fossil baleen suggests early mineralization of the baleen attachment to the rostrum or rapid burial of the skeletons before any detachment or loss could occur.     

Feeding ecology of gray whales inferred from stable-carbon and nitrogen isotopic analysis of baleen plates

Stable-carbon and nitrogen isotopic compositions (δ¹³C, δ¹⁵N) of baleen plates of two juvenile and four adult gray whales (Eschrichtius robustus) were examined. High variance in isotopic composition of baleen plates was detected among individuals and, unlike other migratory species of baleen whales examined isotopically, δ¹³C and δ¹⁵N values of most whales showed no regular oscillations. Only one baleen plate reflected the assumed principal Arctic prey (ampeliscid amphipods) during growth on the summer grounds. The rate of baleen growth inferred for one of the juveniles in the last 5 months of life (4.7 mm/week) was similar to the rate calculated previously for a rehabilitating gray whale calf. Autumn corresponded to the timing of the formation of lowest δ¹⁵N values measured along plates. We estimate that the baleen length in all adult gray whales recorded around a year of information (1.3 ± 0.3; Mean ± SD). This short period of dietary integration precludes long time series analyses in this species and reflects the extensive wear on baleen plates due to benthic foraging. Handling editor: M. Power     

WHY DO BALEEN WHALES MIGRATE?1

The annual migrations of baleen whales are a conspicuous but unexplained feature of their behavioral repertoire. Some hypotheses offered to explain whale migration focus on direct benefits to the calf (thermoregulation, calm water) and some do not (resource tracking, and the “evolutionary holdover” hypothesis). Here, we suggest that a major selective advantage to migrating pregnant female baleen whales is a reduced risk of killer whale (Orcinus orca) predation on their newborn calves in low-latitude waters. Killer whale abundance in high latitudes is substantially greater than that in lower latitudes, and most killer whales do not appear to migrate with baleen whales. We suggest that the distribution of killer whales is determined more by their primary marine mammal prey, pinnipeds, and that following the baleen whale migrations would remove them from their pinniped prey. There are problems with all current hypotheses, most of which stem from a lack of directed research. We explore variation in migratory habits between species, populations, and individuals that may provide a “natural laboratory” for discriminating among the competing hypotheses.     

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.     

医院口腔颌面部骨折患者的临床特征分析及治疗效果调查

目的:调查研究医院口腔颌面部骨折(OMF)患者的临床特征及治疗效果。方法:选取我院2011年2月至2013年2月期间收治的OMF患者135例作为研究对象。对患者临床特征进行全面分析,并根据患者具体情况制定相应治疗方案,分析对比临床疗效情况。结果:患者男女比例为2.14:1,平均年龄为34.6±4.2岁,其中20~30岁所占比例最高,达42.96%。骨折原因主要为交通事故,占45.19%。骨折类型以单纯型下颌骨骨折最高发,占46.67%。切开复位并行钛板钛钉内固定较单纯接受钛钉颌间牵引内固定,前者临床有效率显著高于后者,差异有统计学意义(P0.05)。结论:OMF高发于男性青壮年,主要骨折原因为交通事故,且以单纯型下颌骨骨折类型居多,临床治疗切开复位并行钛板钛钉内固定疗效显著优于单纯接受钛钉颌间牵引内固定。     

A bizarre new toothed mysticete (Cetacea) from Australia and the early evolution of baleen whales

Extant baleen whales (Cetacea, Mysticeti) are all large filter-feeding marine mammals that lack teeth as adults, instead possessing baleen, and feed on small marine animals in bulk. The early evolution of these superlative mammals, and their unique feeding method, has hitherto remained enigmatic. Here, I report a new toothed mysticete from the Late Oligocene of Australia that is more archaic than any previously described. Unlike all other mysticetes, this new whale was small, had enormous eyes and lacked derived adaptations for bulk filter-feeding. Several morphological features suggest that this mysticete was a macrophagous predator, being convergent on some Mesozoic marine reptiles and the extant leopard seal (Hydrurga leptonyx). It thus refutes the notions that all stem mysticetes were filter-feeders, and that the origins and initial radiation of mysticetes was linked to the evolution of filter-feeding. Mysticetes evidently radiated into a variety of disparate forms and feeding ecologies before the evolution of baleen or filter-feeding. The phylogenetic context of the new whale indicates that basal mysticetes were macrophagous predators that did not employ filter-feeding or echolocation, and that the evolution of characters associated with bulk filter-feeding was gradual.     

Subunit principles in midface fractures: the importance of sagittal buttresses, soft-tissue reductions, and sequencing treatment of segmental fractures

The patterns of midface fractures were related to postoperative computed tomography scans and clinical results to assess the value of ordering fracture assembly in success of treatment methods. A total of 550 midface fractures were studied for their midface components and the presence of fractures in the adjacent frontal bone or mandible. Preoperative and postoperative computed tomography scans were analyzed to generate recommendations regarding exposure and postoperative stability related to fracture pattern and treatment sequence, both within the midface alone and when combined with frontal bone and mandibular fractures. Large segment (Le Fort I, II, and III) fractures were seen in 68 patients (12 percent); more comminuted midface fracture combinations were seen in 93 patients (17 percent). Midface and mandibular fractures were seen in 166 patients (30 percent). Midface, mandible, and nasoethmoid fractures were seen in 38 patients (7 percent). Frontal bone and midface fractures were seen in 131 patients (24 percent). Split-palate fractures accompanied 8 percent of midface fractures. Frontal bone, midface, and mandibular fractures were seen in 54 patients (10 percent). The midface, because of weak bone structure and comminuted fracture pattern, must therefore be considered a dependent, less stable structure. Its injuries more commonly occur with fractures of the frontal bone or mandible (two-thirds of cases) and, more often than not (>60 percent), are comminuted. Comminuted and pan-facial (multiple area) fractures deserve individualized consideration regarding the length of intermaxillary immobilization. Examples of common errors are described from this patient experience.     

Phylogenetic relationships among the baleen whales based on maternally and paternally inherited characters

Phylogenetic relationships in the Cetacean suborder Mysticeti (baleen whales) have recently been the focus of increased attention. Here, we examine the evolutionary history of this group by comparing genealogies derived from Y chromosome and mitochondrial DNA sequences. We generated topologies based on paternally and maternally inherited characters for males from nine baleen whale species, including representatives of three families (Balaenidae, Eschrichtiidae, and Balaenopteridae) and four genera (Balaena, Eschrichtius, Balaenoptera, and Megaptera). Divergence among species was fifteen times greater for mtDNA than for Y-specific DNA. Both mtDNA and yDNA topologies revealed the family Balaenopteridae to be paraphyletic, but this relationship was neither strongly supported nor consistent across phylogenetic analysis methodologies. Humpback and fin whales, representing different genera, were reciprocally monophyletic sister species according to mtDNA. Although the monophyly of fin whales decayed for yDNA, a close relationship between fin and humpback whales was retained in yDNA trees. The paraphyly of fin whales and the long branch leading to humpback whales for the yDNA marker may suggest life history differences between these species. Specifically, male humpback whales showed higher than average divergence from other baleen whales at yDNA, although not at mtDNA, suggesting a potential for smaller effective population sizes among male humpbacks on an evolutionary timescale. The observation that those species that have been found to hybridize in nature (blue/fin and blue/humpback) do not reveal evidence for paraphyly for either maternal or paternal markers suggests that introgressive hybridization has not historically been extensive and thus may not represent a substantial source of phylogenetic error for Mysticeti.     

Morphological and molecular evidence for a stepwise evolutionary transition from teeth to baleen in mysticete whales

The origin of baleen in mysticete whales represents a major transition in the phylogenetic history of Cetacea. This key specialization, a keratinous sieve that enables filter-feeding, permitted exploitation of a new ecological niche and heralded the evolution of modern baleen-bearing whales, the largest animals on Earth. To date, all formally described mysticete fossils conform to two types: toothed species from Oligocene-age rocks ( approximately 24 to 34 million years old) and toothless species that presumably utilized baleen to feed (Recent to approximately 30 million years old). Here, we show that several Oligocene toothed mysticetes have nutrient foramina and associated sulci on the lateral portions of their palates, homologous structures in extant mysticetes house vessels that nourish baleen. The simultaneous occurrence of teeth and nutrient foramina implies that both teeth and baleen were present in these early mysticetes. Phylogenetic analyses of a supermatrix that includes extinct taxa and new data for 11 nuclear genes consistently resolve relationships at the base of Mysticeti. The combined data set of 27,340 characters supports a stepwise transition from a toothed ancestor, to a mosaic intermediate with both teeth and baleen, to modern baleen whales that lack an adult dentition but retain developmental and genetic evidence of their ancestral toothed heritage. Comparative sequence data for ENAM (enamelin) and AMBN (ameloblastin) indicate that enamel-specific loci are present in Mysticeti but have degraded to pseudogenes in this group. The dramatic transformation in mysticete feeding anatomy documents an apparently rare, stepwise mode of evolution in which a composite phenotype bridged the gap between primitive and derived morphologies; a combination of fossil and molecular evidence provides a multifaceted record of this macroevolutionary pattern.     

A three-dimensional numerical simulation of mandible fracture reduction with screwed miniplates

A three-dimensional finite element model of a fractured human mandible treated with plating technique was developed to simulate and to study the biomechanical loads and the stress field distribution. Biomechanical properties of bone have been thoroughly investigated experimentally. In this work, using the finite element method, complete clinical conditions (after surgical reduction, post-operatory period, complete healing period) were simulated. The mandible fracture was located in the symphysis region and one or two titanium miniplates, fixed with monocortical screws, were evaluated. The behaviour of a reduced human mandible with screwed miniplates, as well as its complete healing, is investigated and described.     

Treatment of mandible fractures using bioabsorbable plates

This study evaluated the short-term outcome of treating fractures of the mandible with bioabsorbable plates. Sixty-nine fractures of the mandible in 49 patients were treated by open reduction and internal fixation using bioabsorbable plates and 2.4-mm, 2.0-mm, and 1.5-mm pretapped screws. The duration of intermaxillary fixation ranged from 0 to 23 days, with a mean of 4.6 days. Patients were evaluated for complications during the follow-up period, which ranged from 1 to 18 months. Six patients (12.2 percent) experienced complications. These included infection (four patients), premature occlusal contact (one patient), and temporomandibular disorder (one patient). Except for one case, all complications were minor and adequately managed with incision and drainage, elastic traction, physical therapy, and medication. Delayed infection (osteomyelitis) developed in a symphysis fracture and was treated by saucerization and antibiotics. The fracture line subsequently showed complete consolidation. Bioabsorbable plates can be selectively used for internal fixation in mandibular fractures, with the advantage that they do not need to be removed.     

Age estimation in bowhead whales using tympanic bulla histology and baleen isotopes

Tympanic bullae and baleen plates from bowhead whales of the Western Arctic population were examined. Growth layer groups (GLGs) in the involucrum of the tympanic bone were used to estimate age of the whales, and compared to stable isotope signatures along transects of baleen plates and the involucrum. The involucrum of the tympanic bone consists of three regions that form in utero, during nursing in the first year, and during the first decades of life, respectively. Life history events, such as annual migration, are recorded in the bowhead tympanic bulla. It is likely that bone growth in the bowhead tympanic occurs during periods of high food intake, while slow or arrested growth occurs during periods of low food intake. Comparisons between numbers of GLGs in the tympanic, number of isotopic oscillations in a baleen plate, length of the baleen plate, and total whale length show correlation coefficients as high as 0.97. The tympanic GLG method is particularly useful for estimating the age of whales up to 20 yr old.     

Lethal entanglement in baleen whales

Understanding the scenarios whereby fishing gear entanglement of large whales induces mortality is important for the development of mitigation strategies. Here we present a series of 21 cases involving 4 species of baleen whales in the NW Atlantic, describing the available sighting history, necropsy observations, and subsequent data analyses that enabled the compilation of the manners in which entanglement can be lethal. The single acute cause of entanglement mortality identified was drowning from entanglement involving multiple body parts, with the animal's inability to surface. More protracted causes of death included impaired foraging during entanglement, resulting in starvation after many months; systemic infection arising from open, unresolved entanglement wounds; and hemorrhage or debilitation due to severe gear-related damage to tissues. Serious gear-induced injury can include laceration of large vessels, occlusion of the nares, embedding of line in growing bone, and massive periosteal proliferation of new bone in an attempt to wall off constricting, encircling lines. These data show that baleen whale entanglement is not only a major issue for the conservation of some baleen whale populations, but is also a major concern for the welfare of each affected individual.     

Population biology, social behavior and communication in whales and dolphins

The baleen whales differ from the toothed whales and dolphins in life history and in social organization. Even though they grow to a larger size, young baleen whales tend to develop more rapidly than dolphins and toothed whales. Except for the mother-calf bond, most groups of baleen whales are short-lived, lasting only for hours, and individual-specific associations appear to be exceptions to the norm. Most toothed whales live in more structured groups, in which young animals have a long period of dependency and social learning. The communication signals described for different cetacean species have functions suited to the interactions that predominate in their societies.     

Mechanics of the right whale mandible: Full scale testing and finite element analysis

In an effort to better understand the mechanics of ship-whale collision and to reduce the associated mortality of the critically endangered North Atlantic right whale, a comprehensive biomechanical study has been conducted by the Woods Hole Oceanographic Institution and the University of New Hampshire. The goal of the study is to develop a numerical modeling tool to predict the forces and stresses during impact and thereby the resulting mortality risk to whales from ship strikes.Based on post-mortem examinations, jaw fracture was chosen as a fatal endpoint for the whales hit by a vessel. In this paper we investigate the overall mechanical behavior of a right whale mandible under transverse loading and develop a finite element analysis model of the bone. The equivalent elastic modulus of the cortical component of right whale mandible is found by comparing full-scale bending tests with the results of numerical modeling. The finite element model of the mandible can be used in conjunction with a vessel-whale collision event model to predict bone fracture for various ship strike scenarios.     

Mandibular shape correlates of tooth fracture in extant Carnivora: implications to inferring feeding behaviour of Pleistocene predators

Percentages of tooth fracture and mandible shape are robust predictors of feeding habits in Carnivora. If these parameters co‐vary above the species level, more robust palaeobiological inferences could be made on fossil species. A test of association is presented between mandible shape and tooth fracture in a subset of extant carnivorans together with large Pleistocene fossil predators from Rancho La Brea (Canis dirus, Panthera atrox, and Smilodon fatalis). Partial least square (PLS) and comparative methods are employed to validate co‐variation of these two parameters in extant carnivorans. Association between mandible shape and percentage of tooth fracture is strongly supported, even if both blocks of data exhibit a phylogenetic signal to a different degree. Dietary adaptations drive shape/fracture co‐variation in extant species, although no significant differences occur in the PLS scores between carnivores and bone/hard food consumers. The fossil species project into PLS morphospace as outliers. Their position suggests a unique feeding behaviour. The increase in the size of prey, together with consumption of skin and hair from carcasses in a cold environment, might have generated unusual tooth breakage patterns in large predators from Rancho La Brea. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 70–80.