Cetacean Bones Research Articles

Effects of pre-treatment, historical age, and sample characteristics on the stable isotope analyses of killer whale (Orcinus orca) bone.

Stable isotope analysis of bone provides insight into animal foraging and allows for ecological reconstructions over time, however pre-treatment is required to isolate collagen. Pre-treatments typically consist of demineralization to remove inorganic components and/or lipid extraction to remove fats, but these protocols can differentially affect stable carbon (δ13C) and nitrogen (δ15N) isotope values depending on the chemicals, tissues, and/or species involved. Species-specific methodologies create a standard for comparability across studies and enhance understanding of collagen isolation from modern cetacean bone. Elemental analyzers coupled to isotope ratio mass spectrometers were used to measure the δ13C and δ15N values of powdered killer whale (Orcinus orca) bone that was intact (control) or subjected to one of three experimental conditions: demineralized, lipid-extracted, and both demineralized and lipid-extracted. Additionally, C:N ratios were evaluated as a proxy for collagen purity. Lastly, correlations were examined between control C:N ratios vs. historical age and control C:N ratios vs. sample characteristics. No significant differences in the δ15N values were observed for any of the experimental protocols. However, the δ13C values were significantly increased by all three experimental protocols: demineralization, lipid extraction, and both treatments combined. The most influential protocol was both demineralization and lipid extraction. Measures of the C:N ratios were also significantly lowered by demineralization and both treatments combined, indicating the material was closer to pure collagen after the treatments. Collagen purity as indicated via C:N ratio was not correlated with historical age nor sample characteristics. If only the δ15N values from killer whale bone are of interest for analysis, no pre-treatment seems necessary. If the δ13C values are of interest, samples should be both demineralized and lipid-extracted. As historical age and specimen characteristics are not correlated with sample contamination, all samples can be treated equally.

Whale fall chemosymbiotic communities in a southwest Australian submarine canyon fill a distributional gap

A whale fall community of chemosymbiotic invertebrates living on cetacean bones has been identified off southwestern Australia during a Remotely Operated Vehicle (ROV) survey at bathyal depths within the Bremer Marine Park, which is part of important marine mammal areas (IMMA) of the Albany Canyon Region. Cetacean bones on the seafloor of the Hood Canyon, consisted of isolated skulls of three species of beaked whales (family Ziphiidae): Mesoplodon cf. layardii, M. grayi, and M. hectori, a few vertebrae, and lower jaws. One of the beaked whale skulls (Mesoplodon cf. layardii) was sampled and found to be intensely colonised by hundreds of specimens of a bathymodilinae mussel (“Adipicola” s.l.). Live polychaetes (Phyllochaetopterus?), skeneimorph gastropods, and amphipods (Seba, Leptamphopus) colonised the skull bone, which represent a later stage (sulfophilic) of carcass decomposition. The reducing sediment below the skull was inhabited by lucinid (Lucinoma) and vesicomyid (Calyptogena) chemosymbiotic bivalves. Additionally, the sediment thanatocoenosis comprised shells of various other chemosymbiotic bivalves, such as Acharax, thyasirids, lucinids, vesicomyids, and limpets, representing the complex ecological turnover phases through time in this whale fall chemosynthetic habitat. With one exception, all bones recovered were colonized by bathymodiolin mussels. This is the first documented case of a chemosynthetic community and associated chemosymbiotic fauna relating to beaked whales, and the first fully documented record of a whale fall community within the Australian Southern Ocean region.

The first record of a Konservat-Lagerstätten in which early post-settlement stages of fossil archaeobalanids (Cirripedia: Balanomorpha) are preserved.

A diverse sessile barnacle fauna from a Miocene shallow-water deposit at Dolnja Stara vas in Slovenia is described. It includes the first descriptions of early post settlement juveniles of Actinobalanus sloveniensis attached to mangrove leaves. These represent three distinct growth phases, the earliest being interpreted as being less than 24 h post settlement, the others being 1 to 2 days post settlement. An assessment of their taphonomy is provided. Associated adult balanomorphs are attached to a variety of organic substrates, including mangrove leaves and branches, fragments of the conifers ?Taxodioxylon, Carapoxylon, pine cones, molluscs, and cetacean bones. The barnacles include A. sloveniensis, Amphibalanus venustus, and Perforatus perforatus-many with opercula retained within the shells. A. venustus retains some of the original shell color. This is the second record of barnacle-plant associations from the Central Paratethys from Kamnik and Trbovlje. The paleoecology and paleogeography of the site are discussed.

Active Whaling, Opportunistic Scavenging or Long-Distance Trading: Zooarchaeological, Palaeoproteomic, and Historical Analyses on Whale Exploitation and Bone Working in Anglo-Saxon Hamwic

THE ANGLO-SAXON SITE OF HAMWIC (modern Southampton, Hampshire, UK) has been identified as a major bone-working centre. Besides antler and terrestrial mammal bone, cetacean bone has been recovered in high quantities. These specimens primarily represent working waste. Using peptide mass fingerprinting of bone collagen (ZooMS), it was determined that the majority of these specimens derive from the currently highly endangered population of North Atlantic right whale (Eubalaena glacialis). Limited historical sources appear to suggest that whaling was undertaken by the Anglo-Saxons, or by the Normans on the other side of the English Channel prior to the eleventh century ad. Nevertheless, the primary method of acquisition for whale bone was through opportunistic scavenging and trading.

Late Iron Age Whaling in Scandinavia

The use of marine mammal bone as a raw material in the manufacturing of gaming pieces in the Scandinavian late Iron Age has been observed and discussed in recent years. New empirical studies have created a chronology as well as a typology showing how the design of the gaming pieces is tightly connected to different choices of raw material; from antler in the Roman and Migration periods, to whale bone in the sixth century, and walrus in the tenth century. Macroscopic examination can, however, rarely go beyond determining that the material is ‘cetacean bone’. The following article presents the taxonomic identifications of 68 samples of whale bone gaming pieces, determined using Zooarchaeology by mass spectrometry analysis. The results demonstrate the consistent use of bones from Balaenidae sp. most probably the North Atlantic right whale (Eubalaena glacialis). This paper presents strong evidence for active, large-scale hunting of whales in Scandinavia, starting in the sixth century. The manufacture of gaming pieces was probably not the driver for the hunt, but merely a by-product that has survived in the archaeological record. Of greater importance were probably baleen, meat, and blubber that could be rendered into oil. This oil might have been an additional trading product in the far-reaching trade networks that were developing during the period. This study supports previous studies suggesting that Iron Age and medieval trade and resource exploitation had a much more severe influence on ecosystems than previously expected. It adds additional insights into anthropogenic impact on mammal populations in prehistory.

Vertebral Trabecular Bone Mechanical Properties Vary Among Functional Groups of Cetaceans.

SynopsisSince their appearance in the fossil record 34 million years ago, modern cetaceans (dolphins, whales, and porpoises) have radiated into diverse habitats circumglobally, developing vast phenotypic variations among species. Traits such as skeletal morphology and ecologically linked behaviors denote swimming activity; trade-offs in flexibility and rigidity along the vertebral column determine patterns of caudal oscillation. Here, we categorized 10 species of cetaceans (families Delphinidae and Kogiidae; N = 21 animals) into functional groups based on vertebral centra morphology, swimming speeds, diving behavior, and inferred swimming patterns. We quantified trabecular bone mechanical properties (yield strength, apparent stiffness, and resilience) among functional groups and regions of the vertebral column (thoracic, lumbar, and caudal). We extracted 6 mm3 samples from vertebral bodies and tested them in compression in 3 orientations (rostrocaudal, dorsoventral, and mediolateral) at 2 mm min−1. Overall, bone from the pre-fluke/fluke boundary had the greatest yield strength and resilience, indicating that the greatest forces are translated to the tail during caudal oscillatory swimming. Group 1, composed of 5 shallow-diving delphinid species, had the greatest vertebral trabecular bone yield strength, apparent stiffness, and resilience of all functional groups. Conversely, Group 3, composed of 2 deep-diving kogiid species, had the least strong, stiff, and resilient bone, while Group 2 (3 deep-diving delphinid species) exhibited intermediate values. These data suggest that species that incorporate prolonged glides during deep descents in the water column actively swim less, and place relatively smaller loads on their vertebral columns, compared with species that execute shallower dives. We found that cetacean vertebral trabecular bone properties differed from the properties of terrestrial mammals; for every given bone strength, cetacean bone was less stiff by comparison. This relative lack of material rigidity within vertebral bone may be attributed to the non-weight-bearing locomotor modes of fully aquatic mammals.

First records of grey whales, Eschrichtius robustus, from Scotland.

The grey whale, Eschrichtius robustus, occurred in the Atlantic Ocean until c.300 years ago, having been extirpated probably owing to hunting by humans. Three records of fossil grey whale bones were known previously from Cornwall and Devon in the 18th and 19th centuries. Morphological identification to species of cetacean remains from palaeontological and archaeological sites is problematic, because the bones are often fragmentary or have been modified by humans. The application of ancient DNA analysis and collagen fingerprinting (ZooMS) allows the identification of fragmentary and modified cetacean bone to species and/or generic level, thus offering new opportunities to understand the exploitation of cetaceans by human coastal communities. Analyses of cetacean bone from Orkney and Shetland have identified for the first time the presence of grey whale at archaeological sites in Scotland, extending the geographical range of records in Britain. These findings confirm for the first time that the seas around Scotland were part of the range of the grey whale in the eastern North Atlantic, thereby linking records of this species (from southern Scandinavia, the Northern Isles of and southwest Britain, the southern North Sea and Netherlands, to the Iberian Peninsula and Morocco) over a similar latitudinal range that the species still occupies in the Pacific Ocean.

Photogrammetric Three-dimensional Modeling and Printing of Cetacean Skeleton using an Omura's Whale Stranded in Hong Kong Waters as an Example.

The preparation of cetacean, in particular baleen whale, skeletons presents a great challenge due to their high lipid content and uncommon size. Documentation of the skeletal morphology is important to produce accurate and reliable models for both research and educational purposes. In this paper, we used a 10.8-meter long Omura's whale stranded in Hong Kong waters in 2014 as an example for the illustration. This rare and enormous specimen was defleshed, macerated, and sun-dried to yield the skeleton for research and public display. Morphology of each bone was then documented by photogrammetry. The complex contour of the skeleton made automated photoshoot inadequate and 3 manual methods were used on bones of different sizes and shapes. The captured photos were processed to generate three-dimensional (3D) models of 166 individual bones. The skeleton was printed half-size with polylactic acid for display purposes, which was easier to maintainthan the actual cetacean bones with high residual fat content. The printed bones reflected most anatomical features of the specimen, including the bowing out rostral region and the caudal condylar facet that articulated with Ce1, yet the foramina on the parieto-squamosal suture, which are diagnostic character of Balaenoptera omurai, and an indented groove on the frontal bone at the posterior end of the lateral edge were not clearly presented. Extra photoshoots or 3D surface scanning should be performed on areas with meticulous details to improve precision of the models. The electronic files of the 3D skeleton were published online to reach a global audience and facilitate scientific collaboration among researchers worldwide.

Humans, Dolphins, and Porpoises: Investigations at the Par-Tee Site, Seaside, Oregon, AD 100–800

Small cetaceans are understudied compared to whales and pinnipeds even though they represent a high-ranking prey choice when available in the environment. Building upon previous faunal analyses at the Par-Tee site, Seaside, Oregon that investigated whaling, this analysis of dolphin and porpoise remains suggests that people were hunting small cetaceans between AD 100–800 on the Oregon coast, especially harbor porpoise, which was found significantly more than any other cetacean species at the site. The quantity of small cetacean bone is unlikely to be the result of only acquiring stranded individuals. While there is no direct evidence of hunting, ethnographic literature and archaeologically recovered hunting technologies like harpoons provide insight into the means by which these species may have been hunted.

Une baleine grise dans le Périgord magdalénien : identification taxinomique d’une pointe de projectile en os de La Madeleine (Dordogne, France) par empreinte peptidique de masse du collagène

The Cetacean bone artifact described in this paper is the result of excavations carried out in La Madeleine (Tursac, Dordogne) under the direction of D. Peyrony, between 1910-1913. It was discovered in the middle layer (“couche moyenne”) of the stratigraphy described by Capitan and Peyrony and subsequently attributed to the “Magdalenian V” by H. Breuil (i.e the beginning of the Upper Magdalenian). This layer corresponds to layers C25 to C20 of the stratigraphy established later by J.-M. Bouvier. Two radiocarbon dates are currently available (C21: 12750 ± 240 BP, Ly 920; and C25: 13070 ± 190 BP, Ly 921). The artifact was recently identified as Cetacean bone during an integrated study of the bones and reindeer antlers kept at the Musee National de Prehistoire (Peyrony and Bouvier collections). It is a fully worked object, elongated in shape (129 X 13.2 X 12.8 mm; 18.6 g) with straight outline and profile, a circular cross-section and parallel sides. It is interpreted as a mesio-proximal fragment of projectile point. Collagen peptide mass fingerprinting (PMF) was applied to taxonomically identify the bone artefact, as a part of the PaleoCet project. Also known as ZooMS, collagen PMF discriminates taxonomic groups based on differences in the collagen protein sequence observed through Matrix-assisted laser desorption/ionization time-of-flight (MALDI-TOF) mass spectrometry. The bone subsamples from the exterior and interior of the artifact produced similar MALDI-TOF spectra, both of which could be confidently identified as gray whale (Eschrichtius robustus). The artifact from La Madeleine is the first evidence of the presence of this species in Atlantic waters at the end of MIS2, suggesting continuity between the Pleistocene and Holocene populations of this species. Moreover, to our knowledge, this gray whale bone artifact represents the oldest identified gray whale in an archaeological context, thus first witness of the millennia-old interactions between humans and this highly coastal whale species. The manufacturing of the La Madeleine artifact on the Basque coast, followed by its circulation through the western Pyrenees and then towards the north at the east of Landes and Gironde up to Perigord (a travel distance of roughly 350 km), might be the most plausible route taken by this artifact to reach La Madeleine.

Accelerated evolution and diversifying selection drove the adaptation of cetacean bone microstructure

BackgroundThe transition from land to sea by the ancestor of cetaceans approximately 50 million years ago was an incredible evolutionary event that led to a series of morphological, physiological, and behavioral adaptations. During this transition, bone microstructure evolved from the typical terrestrial form to the specialized structure found in modern cetaceans. While the bone microstructure of mammals has been documented before, investigations of its genetic basis lag behind. The increasing number of cetaceans with whole-genome sequences available may shed light on the mechanism underlying bone microstructure evolution as a result of land to water transitions.ResultsCetacean bone microstructure is consistent with their diverse ecological behaviors. Molecular evolution was assessed by correlating bone microstructure and gene substitution rates in terrestrial and aquatic species, and by detecting genes under positive selection along ancestral branches of cetaceans. We found that: 1) Genes involved in osteoclast function are under accelerated evolution in cetaceans, suggestive of important roles in bone remodeling during the adaptation to an aquatic environment; 2) Genes in the Wnt pathway critical for bone development and homeostasis show evidence of divergent evolution in cetaceans; 3) Several genes encoding bone collagens are under selective pressure in cetaceans.ConclusionsOur results suggest that evolutionary pressures have shaped the bone microstructure of cetaceans, to facilitate life in diverse aquatic environments.

Shark–Cetacean trophic interaction, Duinefontein, Koeberg, (5 Ma), South Africa

This study forms part of a larger project to reconstruct the Mio-Pliocene marine palaeoenvironment along South Africa’s west coast. It documents the shark–cetacean trophic interaction during the Zanclean (5 Ma) at Duinefontein (Koeberg). The damage described on the fragmentary cetacean bones was compared with similar damage observed on fossils from Langebaanweg, a Mio-Pliocene site on the west coast of South Africa, and data present in the literature. This comparison showed that the damage was the result of shark bites. The state of preservation makes it difficult to determine if the shark bite marks were the cause of death or as a result of scavenging. The presence of the bite marks on the bone would, however, indicate some degree of skeletonisation. Bite marks on some cranial fragments would suggest that the cetacean’s body was in an inverted position typical of a floating carcass. The preservation of the material suggests that the bones were exposed to wave action resulting in their fragmentation as well as abrasion, polishing and rolling. It also suggests that the cetacean skeletons were exposed for a long time prior to burial. The morphology of the bites suggests that the damage was inflicted by sharks with serrated and unserrated teeth. Shark teeth collected from the deposit include megalodon (Carcharodon megalodon), white (Carcharodon carcharias) as well as mako (Isurus sp. and Cosmopolitodus hastalis) sharks, making these sharks the most likely predators/scavengers.

Extensively remodeled, fractured cetacean tympanic bullae show that whales can survive traumatic injury to the ears.

Underwater human activities and anthropogenic noise in our oceans may be a major source of habitat degradation for marine life. This issue was highlighted by the opening of the United States Eastern Seaboard for seismic oil and gas exploration in 2014, which generated massive media coverage and widespread concern that seismic surveys could kill or deafen whales. We discovered 11 new specimens of fractured and healed cetacean ear bones, out of a survey of 2127 specimens housed in museum collections. This rare condition has been previously reported only in two specimens of blue whales (Balaenoptera musculus) from the early 1900s, summarized by Fraser & Purves (1953). All of our new specimens are represented by species for which this condition had never been reported previously, including both baleen and toothed whales. The baleen whale specimens (Balaenoptera physalus, Balaenoptera borealis, Balaenoptera acutorostrata) were collected during Canadian commercial whaling operations in the Atlantic Ocean in the 1970s; the specimens include ear bones with well-healed fractures, demonstrating that baleen whales are capable of overcoming traumatic injury to the ears. The toothed whale specimens (Delphinus sp., Berardius bairdii) were found dead on beaches in 1972 and 2001, respectively, with less remodeled fractures. Thus, ear injuries may be more lethal to the echolocating toothed whales, which rely on hearing for navigation and foraging. We explore several hypotheses regarding how these injuries could have occurred, and conclude that the most parsimonious explanations appear to be both direct and indirect effects of lytic processes from disease or calcium depletion, or damage from external pressure waves. Although further research is required to confirm whether the fractures resulted from natural or human-induced events, this study underscores the importance of museum collections and the work of stranding networks in understanding the potential effects of modern human activities on marine mammal health.

EARLY PLIOCENE (5 MA) SHARK-CETACEAN TROPHIC INTERACTION FROM LANGEBAANWEG, WESTERN COAST OF SOUTH AFRICA

We document an aspect of the marine paleoecology at Langebaanweg, a site that has produced an abundance of vertebrate fossils. Damage to the bone surfaces of cetacean fossils was not pathological as evident on the fossil seals from this site; the current study documents the damage and attempts to provide a parsimonious explanation. Literature reviews identified similar damage described elsewhere to be the result of shark feeding activity. Comparison of this material with Langebaanweg cetacean bones supports the interpretation that the damage resulted from shark teeth. Damage on the various skeletal elements appears to have been inflicted postmortem or, if they were made while the animal was alive, the whales did not survive the attack. Postmortem damage is also supported by the presence of bites on the dorsal, ventral, lateral, and medial surfaces of a pair of dentaries. Bites were inflicted by sharks with serrated teeth, as well as by sharks with unserrated teeth. Potential predators identified from the marks include white (Carcharodon spp.), Zambezi (bull) (Carcharhinus leucas), tiger (Galeocerdo sp.) and mako (Isurus sp.) sharks.

Long range inland–coastal networks during the Late Magdalenian: Evidence for individual acquisition of marine resources at Andernach-Martinsberg, German Central Rhineland

Recent re-examination of the osseous material assemblage from Andernach-Martinsberg, Central Rhineland, has resulted in the identification of an implement manufactured from cetacean bone (probably whale). Argued to be the proximal half of a foreshaft, this artefact is not only one of few such projectile elements to be identified in Magdalenian deposits in northern Europe, but also demonstrates that the exploitation of marine raw materials for use in manufacturing projectile elements is not restricted to southern France, instead extending to at least inland Germany. Additionally, in conjunction with the appearance of marine molluscs and engravings of seals at Andernach, it can now be forcefully argued that this region formed part of an inland–coastal network during the Late Magdalenian and allows us for the first time to suggest that we can identify the movements of individuals transporting valued marine sourced raw materials and their personal experiences across this vast region.

FT-Raman spectroscopy: A positive means of evaluating the impact of whale bone preservation treatment

The degreasing methods currently used for osteological collections are not always completely satisfactory. Numerous natural history museums encounter the problem of grease seeping to the surface of bones. FT-Raman spectroscopy was used to characterise cetacean bones, before and after degreasing treatment, in order to evaluate the efficacy of treatment and the impact thereof on bone constituents. The Raman spectra made it possible to monitor the changes in the main bone constituents: the mineral component with the apatite band at 960 cm −1, the organic component with the collagen amide III band at 1270 cm −1 and fat with the lipid CH 2 band at 2850 cm −1. The band associated with lipids decreased and even disappeared with degreasing treatment containing chlorinated solvents. This type of treatment enables fat to be extracted both from the surface and from the bone core; however, it debases the organic component of bone by denaturing collagen. Alternative types of treatment (acetone or enzyme baths) were tested over a limited period, which did not enable their true efficacy to be demonstrated. During the alternative treatments, no bone degradation was observed. Only the acetone solution was able to extract fat, though only from the surface.

First evidence of a whale-bone industry in the western European Upper Paleolithic: Magdalenian artifacts from Isturitz (Pyrénées-Atlantiques, France)

Recent reexamination of osseous material from the Magdalenian layers of the Isturitz cave (Pyrenees-Atlantiques, France) by the author and F. Poplin has shown that several artifacts once considered to be made of antler have now been identified deriving from cetacean bone, most probably whale bone. These artifacts demonstrate the existence of technical exploitation of whale bone, proving that the now submerged seashore was also a place where Paleolithic people gathered specific materials for producing weapons and tools.

The Exploitation of Sea-Mammals in Medieval England: Bones and their Social Context

There is little evidence for the practice of whaling in Anglo-Saxon or later medieval England, even though whales were caught on the opposite side of the English Channel in Normandy and Flanders. Stranded whales were exploited by coastal communities, but from the eleventh century whales and other cetaceans were claimed by the king as ‘royal fish’. The difficulties of enforcing this claim against, on the one hand the holders of coastal lordships, and on the other against local inhabitants, led to recognition by the king of seigneurial claims. Whales and porpoises were a high-status food, though by the end of the Middle Ages whales may have been declining in popularity. It is argued that sodai aspirations and tensions were expressed in the possession and consumption of cetaceans. The finds of cetacean bone on archaeological sites are interpreted against the changes in their social value. Between about A.D. 1000 and 1300 bones are mainly discovered on high-status sites.

An Odontocete (Cetacea) Meat Utility Index

A meat utility index for odontocetes (Cetacea), based on average flesh weight per skeletal portion of a harbour porpoise ( Phocoena phocoena), is presented. The highest ranked portions are associated with the middle and posterior vertebral column, while the lowest ranked portions are associated with the flippers, head, and anterior part of the vertebral column. The index (including both meat utility values and modified meat utility values) is applied to archaeologically- and ethnoarchaeologically-derived cetacean bone assemblages. Results suggest that while the ease of edible sculp and meat removal may significantly skew expected bone element frequencies, the index (especially that based on modified meat utility values) is nevertheless a useful predictive tool for small, and to some extent medium-sized, cetaceans.

Paleotemperatures versus sea level: Oxygen isotope signal from fish bone phosphate of the Miocene Calvert Cliffs, Maryland

The use of skeletal oxygen isotopic records for use in paleotemperature reconstruction has been hampered by the lack of independent evidence for ocean water oxygen isotopic composition. The δ18O record from homeothermic cetaceans has provided an independent estimate of ocean δ18O values represented by the Calvert and Choptank formations of Maryland. Fish teeth and bones (especially shark and ray teeth) were also collected from these sediments and provide the basis for paleotemperature estimates for represented time slices of the middle Miocene. Trends in δ18O values of the fish phosphate throughout the Calvert Formation are of opposite polarity to the trends from the cetacean bone phosphate. Paleotemperatures calculated using the cetacean proxies for ocean δ18O values sharpen the already present trend, indicating that ocean temperatures increased during episodes of greater glaciation and decreased during periods of lesser or no glaciation. When using modern average ocean values of 0‰ SMOW in the paleotemperature calculation, however, corrected paleotemperatures for the Choptank Formation do not alter the existing pattern of temperatures.