1

Introduction

1.1. Marine Mammals—“What Are They?”

Some 100 living species of mammals (listed in the Appendix) depend on the ocean formost or all of their life needs. Living marine mammals include a diverse assemblage ofspecies that have representatives in three mammalian orders. Within the order Carnivoraare the pinnipeds (i.e., seals, sea lions, walruses), the sea otter, and the polar bear. Theorder Cetacea includes whales, dolphins, and porpoises, and the order Sirenia is com-posed of sea cows (manatees and dugongs). Marine mammals were no less diverse in thepast and include extinct groups such as the hippopotamus-like desmostylians, the bizarrebear-like carnivore Kolponomos, and the aquatic sloth Thalassocnus.

1.2. Adaptations for Aquatic Life

Marine mammals are well adapted for life in the water though they differ in the degreeto which they are adapted to this habitat. Pinnipeds, sea otters, and polar bears areamphibious, spending some time on land or ice to give birth and to molt, whereascetaceans and sirenians are fully aquatic. A few major aquatic adaptations are brieflyreviewed in this chapter and are covered in greater detail in subsequent chapters.Adaptations of the skin, specifically its increased insulation (through development ofblubber or a dense fur layer) and countercurrent heat exchange systems, help them copewith the cold. Similarly, the eyes, nose, ears, and limbs of marine mammals havechanged in association with their ability to live in a variety of aquatic environments,which include saltwater, brackish, and freshwater. Perhaps the most notable amongsensory adaptations are the high frequency sounds produced by some whales for use innavigation and foraging. Other marine mammals (e.g., pinnipeds, polar bears, and seaotters) have an acute sense of smell; these same groups also possess well-developedwhiskers with sensitive nerve fibers that serve as tactile sense organs. Pinnipeds havefront and hind limbs modified as flippers that propel them both in the water and onland. In cetaceans and sirenians, the hind limbs are virtually absent and locomotion isaccomplished by vertical movement of the tail. Most marine mammals cope with

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living in salt water by conserving water in their heavily lobulated kidneys, which areefficient at concentrating urine.

Many marine mammals are capable of prolonged and deep dives. Adaptations of therespiratory system, such as flexible ribs that allow the lungs to collapse and thickenedtissue in the middle ear of pinnipeds and cetaceans, enable them to withstand the tremen-dous pressures encountered at great depths. The long dives of these animals are accom-plished by a variety of circulatory changes including a slowed heart rate, reduced oxygenconsumption, and shunting blood to only essential organs and tissues.

1.3. Scope and Use of This Book

Our goal for this second edition remains the same as for the first edition: to provide anoverview of the biology of marine mammals with emphasis on their evolution, anatomy,behavior, and ecology. These topics are presented and discussed using, in so far aspossible, an explicit phylogenetic context. In doing so we consider different ways ofincorporating evolutionary history into comparative analyses of marine mammal bio-logy. The phylogenetic approach advocated in this book is a young but vigorously devel-oping research field that we believe has much to offer marine mammal science. Over thepast six years, interest in this approach has grown and we are pleased to offer a numberof new case studies that integrate a phylogenetic approach into studies of marine mam-mal biodiversity.

The book is divided into two major sections: Part I: Evolutionary History (Chapters2–6) is where the origin and diversity of marine mammals are revealed, and Part II:Evolutionary Biology, Ecology, and Behavior (Chapters 7–15) is where we attempt toexplain how this diversity arose by examining patterns of morphological, behavioral,and ecologic diversity. We have intended to explain these concepts, wherever possible, byexample and with a minimum of professional jargon. Words and phrases included in theglossary appear in boldface type at their first appearance in the text. “Further reading”sections have been placed at the end of each chapter and are intended to guide the readerto more detailed information about a particular topic.

1.4. Time Scale

A historical discussion of marine mammals requires a standard time framework forrelating evolutionary events. Figure 1.1 presents the geologic time scale that is usedthroughout this book (based on Harland et al., 1990). Our interest lies in the CenozoicEra, the last 65 million years of earth history, during which time all marine mammalsmade their first appearance. Whales and sirenians were the first to appear, beginningapproximately 50 million years ago (Ma) during the early Eocene. Pinnipeds trace theirancestry back between 29 and 23 Ma to the late Oligocene. The sea otter lineage goesback approximately 7 Ma to the late Miocene, although the modern sea otter is known inthe fossil record only as far back as the early Pleistocene (1.6 Ma). Polar bears appeareven later, during the late Pleistocene (0.5 Ma). The desmostylians, extinct relatives ofsirenians, range from the early Oligocene through the late Miocene. The extinct carnivo-ran Kolponomos is known from a brief time interval during the early Miocene, and theextinct marine sloth Thalassocnus lived during the late Miocene–late Pliocene (7–3 Ma).

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1.5. Early Observations of Marine Mammals

The study of marine mammals probably began with casual observations of the appear-ance and behavior of whales in the 4th century B.C. Still, the knowledge and history ofthese animals themselves go much further back. Drawings of seals and dolphins on piecesof reindeer antler and in caves have been found from Paleolithic times. The Greek philo-sopher Aristotle (384–322 B.C.) in his Historia Animalium describes dolphins, killerwhales, and baleen whales, noting that “the [latter] has no teeth but does have hair thatresemble hog bristles.” Unfortunately, Aristotle’s observations were dismissed by manylater workers because of his misclassification of dolphins as fish. Following Aristotle, theonly other authority on whales in ancient times was Pliny the Elder (24–79 A.D.). In his37-volume Naturalis Historia, he included a book on whales and dolphins in which heprovided accounts based on Aristotle’s findings and his own observations. Knowledge ofmarine mammals languished for a thousand years after Aristotle and Pliny during theDark Ages. During the Renaissance, a rapid increase in exploration of the oceans was fol-lowed by the publication of scientific reports from various expeditions. The earliest ofthese was the Speculum Regale, an account of Iceland in the 13th century that consideredwhales the only truly interesting sight the island had to offer. Its author correctly distin-guished between northern right whales and bowhead whales, which were still confused bymany naturalists five centuries later. In the 16th century, explorers discovered the richfeeding grounds in the high Arctic and the large whale populations that these supported.In the mid-1500s, Konrad Gesner in his Historia Animalium presented illustrations ofwhales; among them was one so large that sailors mistook it for an island (Figure 1.2).

A walrus is also illustrated in Gesner’s work (Figure 1.3a). Among the earliest draw-ings of seals, Vitulus marinus (Figure 1.3b) in Pierre Belon’s De Aquatilibus (1553)is most remarkable for its accuracy, particularly in the detail of the hind limbs.In Guillaume Rondelet’s De Piscibus (1554), two seals are illustrated, one probably

1.5. Early Observations of Marine Mammals 3

Plio-ceneMioceneOligoceneEocene

Early MiddleLate Late LateMiddle Early Early

Kolponomos †

Sirenia

Desmostylia †

Cetacea

Pinnipedia

Thalassocnus †

510152025303540455055

Ma

Pleisto

E L

Ursus

EnhydraCarnivora

Edentata

0

Figure 1.1. Chronologic ranges of marine mammal taxa. Solid bars show reported maximum ranges.Ma = million years ago. (Time scale and correlations are from Harland et al., 1990, andBerggren et al., 1995.)

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representing the common seal and the other the Mediterranean monk seal (Figure 1.3c,d; King, 1983). In another book, The Natural History of Quadrupeds (1763) byR. Brookes, it is obvious from the illustration and description of the male with a largesnout or trunk that the elephant seal is depicted as a cheerful “sea lion”with a “seaweedtail” (Figure 1.3e; King, 1983).

4 1. Introduction

Figure 1.2. Woodcut by Conrad Gesner, from Historia Animalium, first published between 1551 and 1558,shows a whale so large that sailors mistook it for an island.

Figure 1.3. Early illustrations of pinnipeds. (a) Walrus from Conrad Gesner’s Historia Animalium, proba-bly taken from a drawing by Albert Dürer. (b) Seal from P. Belon, De Aquatilibus (1553). (c) Sealfrom Guillaume Rondelet, De Piscibus (1554). (d) Seal from Guillaume Rondelet, De Piscibus(1554). (e) “Sea lion”from R. Brookes, The Natural History of Quadrupeds (1763).

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In 1596, the Dutch navigator Wilhelm Barents discovered Spitzbergen (the largestisland in the Svalbard Archipelago, north of Norway) and early in the 17th century com-mercial whalers were sent there by Dutch and English companies to establish a whalingtown. Although these expeditions were concerned primarily with whale products, theyalso produced a number of publications that provided reasonably accurate descriptionsof the external appearance of the most common kinds of whales. The best of these arefound in Spitzbergische oder Groenlandische Reisen Beschreibung (1675) by FrederichMartens and Bloyeyende Opkomst der Aloude en Hedendaagsche GroenlandscheVisschery (1720) by C. G. Zorgdrager, both of which contained engravings that contin-ued to be reproduced in books until the early 19th century. Georg Wilhem Steller, ship’snaturalist and physician for Vitus Bering’s second expedition to North America, wasamong the first Europeans to explore Alaska and the Aleutian and Commander Islands.His notes of marine mammals living in the Bering Sea, The Beasts of the Sea (1751), con-tained a natural history account of the sea otter, sea lion, fur seal, and the now extinctSteller sea cow, the only first-hand scientific observation of this species.

Another naturalist, Lacépéde, compiled a volume on whales (1804), in which mostof the illustrations were copied from previous publications (Figure 1.4). Lacépédeacknowledged that not having ever seen a whale, he had made his descriptions fromthose of other naturalists. In the first half of the 19th century, additions to the

1.5. Early Observations of Marine Mammals 5

Figure 1.4. Woodcut of baleen whales from Lacépéde (1804).

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literature included Peter Camper’s Observations Anatomiques sur Plusiers Especes deCétacés (1820). The foremost European cetologist of the second half of the 19th cen-tury was P.-J. Van Beneden, a Belgian zoologist whose many monographs on whalesand pinnipeds (including Histoire Naturelle des Cétacés des Mers d’Europe, 1889) werepublished in Brussels between 1867 and 1892. John Edward Gray, who became Keeperof the Zoology Department at the British Museum of Natural History, published hisCatalogue of Seals and Whales in the British Museum in 1866. John Allen (1880), in hiscomprehensive monograph of North American pinnipeds, provided keys to the fami-lies and genera, described the North American species, and gave accounts of pinnipedspecies in other parts of the world.

Meanwhile, the whaling industries of several countries were making other contribu-tions to the study of whales. Whaling captains such as William Scoresby and CharlesScammon made their own observations in the field or collected those of their colleagues.Scoresby published An Account of the Arctic Regions (1820), which is still a valuablesource of information on the northern right whale. Scammon’s book, The MarineMammals of the North-Western Coast of North America, was published in 1874 and hasbecome a classic, particularly valued for its description of the natural history of the graywhale in California.

Land-based whaling stations used in more modern whaling provided the material forFrederick True’s 1904 monograph The Whalebone Whales of the Western North Atlanticand Roy Chapman Andrews’s 1916 monograph on the Sei whale in the Pacific.

Apart from whalers, the only people seriously interested in the study of whales (cetology)at this time were comparative anatomists (for a more detailed account of the beginnings ofcetology see Matthews, 1978). Among their ranks were Rondelet, Bartholin, Camper,Cuvier, Hunter, and Owen. These pioneers in the study of cetacean anatomy made themost of specimens that came their way and the writings that many of them produced showthat they made accurate observations. Cuvier in particular made several fundamentaladvances in cetology.His Le Régne Animal (1817) and Recherches sur les Ossemens Fossiles(1823) contain the original descriptions and illustrations of the three species of cetaceanthat he named (Cuvier’s beaked whale, Risso’s dolphin, and the spotted dolphin).

During this time, confusion over the affinities of another marine mammal group, thedugongs, led some to consider them an unusual tropical form of walrus. In a publicationfrom 1800, the manatee is inaccurately shown as hog-nosed (Figure 1.5a). The earliestillustration of a sirenian to be published, the West Indian manatee from the 1535 editionof La Historia General de la Indias by Gonzalo Fernandez de Oviedo y Valdés, is littlechanged from this depiction more than two centuries later (Figure 1.5b).

6 1. Introduction

Figure 1.5. Early illustrations of manatees. (a) An “American manatee” (species, unknown) from a litho-graph (Reynolds and Odell, 1991). (b) West Indian manatee from the 1535 edition of LaHistoria General de la Indias by Gonzalo Fernandez de Oviedo y Valdes.

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Descriptions of the anatomy of various pinnipeds followed including the walrus(Murie, 1870) and the Steller sea lion (Murie, 1872, 1874). Another accomplishedanatomist, W. C. S. Miller (1888), dissected a variety of pinnipeds including the southernfur seal and southern elephant seal, recovered on the H.M.S. Challenger expedition tothe Antarctic during the years 1873–1875. Thompson (1915) published the first accountof the osteology of Antarctic seals including the Ross seal, the Weddell seal, and theleopard seal. Howell (1929) published his well-known comparative study of both pho-cids and otariids based on the California sea lion and the ringed seal. He followed thiswith a book on aquatic adaptations in mammals (Howell, 1930).

1.6. Emergence of Marine Mammal Science

Marine mammal science has emerged as a discipline in its own right only in the last 20 – 30years. This increasing interest in marine mammals is clearly shown by the expansion ofthe literature dealing with these animals. J. A. Allen’s bibliography of cetaceans andsirenians (1882), covering the 350 years from 1495 to 1840, contains 1014 titles, justunder three publications per year. In the period from 1845 to 1960, between 3000 and4000 articles were published, with a conservative estimate of about 28 titles a year(Matthews, 1966). By comparison, c. 24,000 papers on marine mammals were publishedbetween 1961 and 1998 according to the Zoological Record, a rate of 646 per year. From1999 to 2004, marine mammal publications increased to a rate of more than 856 per year.Among the major influences that contributed to the birth of marine mammal sciencewas the growing recognition that marine mammal populations were limited in numbersand that their exploitation had to be regulated (Boyd, 1993). The aim of many early stud-ies was to obtain accurate information about the biology of these animals for use inestablishing an effective management policy for sustainable exploitation. It is ironic thatthe decline in whale stocks heralded the beginning of the scientific study of marine mam-mals. As a result of concerns regarding stock viability, the Discovery investigations(1925–1951) were undertaken to examine the biology of whale stocks in the SouthernOcean. Not only was the biology of whales examined but also their food supplies andtheir distributions and abundances in relation to oceanographic conditions. For exam-ple, British scientists N. A. Mackintosh and J. F. G. Wheeler (1929) examined 1600 car-casses for gut contents in order to produce their report on blue and fin whales. LeonardHarrison-Matthews had comparable samples in his reports on the humpback whale,sperm whale, and southern right whale in 1938 (Watson, 1981).

In the 1950s, the theme of the Discovery investigations was continued by the FalklandIslands Dependencies Survey (later known as the British Antarctic Survey) when it estab-lished a research program on the southern elephant seal on South Georgia Island under thedirectorship of R. M. Laws. In parallel with these and other studies, with a focus on popula-tion ecology, there also was growing interest in the anatomy and physiology of marine mam-mals (Irving, 1939; Scholander, 1940; Slijper, 1962; Norris, 1966; Andersen, 1969; Ridgway,1972; Harrison, 1972–1977). The establishment of various scientific committees (e.g., theInternational Whaling Commission’s Scientific Committee in 1946 and the U.S. MarineMammal Commission in 1972) to provide advice about the status of various marine mam-mal populations also required knowledge and data on the general biology of these animalsand thus served to stimulate research. Since the early 1980s, the biology of various marinemammal species has been the subject of many notable books, beginning with Ridgway and

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Harrison’s series entitled Handbook of Marine Mammals (1981–1998). This has been fol-lowed by detailed separate accounts of the biology of the Pacific walrus (Fay, 1982), graywhale (Jones et al., 1984), bowhead whale (Burns et al., 1993), bottlenose dolphin(Leatherwood and Reeves, 1990; Reynolds et al., 2000), Hawaiian spinner dolphin (Norriset al., 1994), harbor porpoise (Read et al., 1997) sperm whale (Whitehead, 2003), harp andhooded seals (Lavigne and Kovacs, 1988), elephant seals (Le Boeuf and Laws, 1994), andthe northern fur seal (Gentry, 1998). Comprehensive treatments of marine mammalgroups are available for pinnipeds (King, 1983; Bonner, 1990; Riedman, 1990; Renouf,1991), for whales (Matthews,1978;Gaskin,1982;Evans,1987;Mann et al.,2000), for man-atees and dugongs (Hartman, 1979; Reynolds and Odell, 1991), and for sea otters(Kenyon, 1969; Riedman and Estes, 1990). Valuable field identification guides for allmarine mammals are found in Reeves et al. (2002), for pinnipeds and sirenians in Reeveset al. (1992), and for whales and dolphins in Leatherwood and Reeves (1983) andCarwardine (1995).Recent additions to the growing literature on marine mammal biologyinclude edited books on health and medicine (Dierauf etal.,2001),cell and molecular biol-ogy (Pfeiffer, 2002), conservation biology (Evans and Raga, 2001), evolutionary biology(Hoelzel, 2002), and even an encyclopedia on marine mammals (Perrin et al., 2002).

Matthews (1966) wrote “the greatest revolution in the study of the Cetacea . . . hascome with the possibility of keeping living cetaceans in oceanariums.” However, one ofthe most significant advances in marine mammal science in recent years has undoubt-edly been the move toward studying animals under wild, unrestrained conditions at sea.This is in large part the result of technological advances in microelectronics (e.g., satel-lite telemetry and time-depth recorders). For example, the application of microelectron-ics led to the discovery that elephant seals regularly dive to depths of 1000 m withconsistently long dive durations, typically lasting 15 to 45 minutes. This feature ofelephant seal biology, in addition to studies on a variety of other species, has forcedphysiologists to reexamine our understanding of the biochemical pathways used bythese animals to maximize the efficiency of oxygen utilization. Studies with crittercamsprovide a visual record of everything that a marine mammal sees. For example, critter-cams have revealed Wedell seals flushing prey from crevices in the ice.

Technological advances in molecular biology (e.g., analysis of DNA variation) havealso provided unparalleled opportunities to examine interactions among populationsand the roles of individuals within those populations. For example, using DNA finger-printing and other techniques, it is possible to assess paternity and kinship amongwhales, animals for which this has previously been virtually impossible owing to thedifficulty of observing them mating underwater. These techniques have also made itpossible to measure effective population sizes and interpret historical events such as pop-ulation bottlenecks. Molecular techniques also have contributed to our knowledge ofthe systematics and taxonomy of various marine mammal groups.

As pointed out by Watkins and Wartzok (1985), information and research aboutmarine mammals range “from intensive to eclectic.” Much of the available data is diffi-cult to synthesize because techniques vary widely and sample sizes often are necessarilysmall. This is not a reflection of poor science but rather the environmental, practical, andlegal complications implicit in marine mammal research. It is apparent that the databasemust be expanded. Even within a relatively homogeneous group like odontocete whales,one well-known species (the bottlenose dolphin, Tursiops truncatus) cannot be usedreliably to characterize all toothed whales. With this in mind, we hope that as readers ofthis book you will be able to identify areas in which research must be done. We encourage

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you to pursue research on marine mammals—there are still many gaps in our knowledgeof this diverse and unique assemblage of mammals.

1.7. Further Reading and Resources

There are a large number of Internet addresses with information about marine mammalprograms and organizations; a few that we consider the most useful are listed here:http://www.marinemammalogy.org—Society for Marine Mammalogy (SMM), a pro-fessional international organization of marine mammal scientists, publishes a journal(quarterly) of original research on marine mammals: Marine Mammal Science.http://web.inter.NL.net/users/J.W.Broekema/ecs/index.htm—European CetaceanSociety (ECS), professional biologists and others interested in whales and dolphins.http://www.earthwatch.org—Earthwatch Institute, offers opportunities for marinemammal enthusiasts to work as volunteers with research scientists.

Also, for career and hobbyist information about marine mammals see books by Glen(1997) The Dolphin and Whale Career Guide, Samansky (2002) Starting Your Career as aMarine Mammal Trainer, and Strategies for Pursuing a Career in Marine MammalScience published by SMM and available online.

References

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Allen, J. A. (1882). “Preliminary List of Works and Papers Relating to the Mammalian Orders Cete andSirenia.”Bull. U.S. Geol. Geogr Surv. of the Territories 6(3) (Art. 18): 399–562.

Andersen, H. T. (ed.) (1969). The Biology of Marine Mammals. Academic Press, New York.Andrew, R. C. (1916). “Monographs of the Pacific Cetacea 2: The Sei Whale.”Mem. Amer. Mus. Nat. Hist. 1:

291–388.Belon, P. (1553). Petri Bellonii Cenomani De aquatilibus: libro duo cum conibus ad viuam ipsorum effigiem,

quoad eius fieri potuit, expressis. Apud Carolum Stephanum, Typographum Regium, Paris.Berggren, W. A., D. V. Kent, C. C. Swisher, Jr., and M. P. Aubry (1995). A Revised Cenozoic Geochronology

and Chronostratigraphy. In “Geochronology, Time Scales and Global Stratigraphic Correlations”(W. A. Berggren et al., eds.), pp. 129–212. SEPM Special Publication, No. 54.

Bonner, W. N. (1990). The Natural History of Seals. Christopher Helm, London.Boyd, I. L. (1993). “Introduction: Trends in Marine Mammal Science.”Symp. Zool. Soc. London 66: 1–12.Brookes, R. (1763). A New and Accurate System of Natural History (6 vols.) Vol. 1 “The Natural History of

Quadrupeds.”Printed for J. Newbery, London.Burns, J. J., J. J. Montague, and C. J. Cowles (1993). The Bowhead Whale. Special Publication, No. 2. Soc. Mar.

Mammal. Allen Press, KS.Camper, P. (1820). Observations anatomiques sur la structure intèrieure et le squelette de plusieurs espèces de

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Carwardine, M. (1995). Whales, Dolphins, and Porpoises. D. K. Publishing, New York.Cuvier, G. (1817). Le regne animal distribue d’apres son organisation, pour servir de basea l’histoire naturelle des

animaux et d’introduction a l’anatomie comparee. Deterville, Paris.Cuvier, G. (1823). Recherches sur les ossemens fossiles: ou l’on rétablit les caractères deplusieurs animaux dont

les révolutions du globe ont détruit les espèces. Nouvelle Édition, entirement refondue, et considérablementaugmentée. Dufour et d’Ocagne, 1821–1825. Paris.

Dierauf, L., and F. M. D. Gulland (eds.) (2001). CRC Handbook of Marine Mammal Medicine. CRC Press,Boca Raton, FL.

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Evans, P. G. H. (1987). The Natural History of Whales and Dolphins. Christopher Helm, London/Facts on File,New York.

Evans, P. G. H., and J. A. Raga (eds.) (2001). Marine Mammals: Biology and Conservation. KluwerAcademic/Plenum Publishers, New York.

Fay, F. H. (1982). “Ecology and Biology of the Pacific Walrus, Odobenus rosmarus divergens Illiger.”U. S. Dept.Int. Fish Wild. Serv. North American Fauna, No. 74.

Fernandez de Oviedo y Valdes, G. (1535). Historia general y natural de la Indias. Edición y estudio preliminar deJuan Pérez de Tudela Bueso. Ediciones Atlas, Madrid.

Gaskin, D. E. (1982). The Ecology of Whales and Dolphins. Heinemann, London.Gentry, R. L. (1998). Behavior and Ecology of the Northern Fur Seal. Princeton University Press, Princeton, NJ.Gesner, K. (1551–1587). Conradi Gesneri Historiæ animalium. C. Froschouerum, Tiguri.Glen, T. B. (1997). The Dolphin and Whale Career Guide. Omega Publishing Company, Chicago.Gray, J. E. (1866). Catalogue of Seals and Whales in the British Museum. 2nd ed. British Museum, London.Hamilton, R. (1839). “The Naturalists Library (conducted by W. Jardine).” Mammalian. Vol. 8. Amphibious

Carnivora, Including the Walrus and Seals, also of the Herbivorous Cetacea. W. H. Lizars, Edinburgh andW. Curry, Jun. and Co., Dublin.

Harland, W. B., R. L. Armstrong, A. V. Cox, L. E. Craig, A. G. Smith, and D. G. Smith (1990). A Geologic TimeScale-1989. Cambridge University Press, New York.

Harrison, R. J. (1972–1977). Functional Anatomy of Marine Mammals, Vols. 1–3. Academic Press, London.Hartman, D. S. (1979). “Ecology and Behavior of the Manatee (Trichechus manatus) in Florida.” Am. Soc.

Mammal. Special Publication, No. 5, 1–153.Hoelzel, A. R. (2002). Marine Mammal Biology, Blackwell Science, Oxford.Howell, A. B. (1929). “Contributions to the Comparative Anatomy of the Eared and Earless Seals (Genera

Zalophus and Phoca).”Proc. U. S. Natl. Mus. 73: 1–143.Howell, A. B. (1930). Aquatic Mammals. Thomas, Springfield, IL.Irving, L. (1939). “Respiration in Diving Mammals.” Physiol. Rev. 19 : 112–134.Jones, M. L., S. L. Swartz, and S. Leatherwood (eds.) (1984). The Gray Whale. Academic Press, New York.Kenyon, K. (1969). “The Sea Otter in the Eastern Pacific Ocean,” North American Fauna No. 68, Bur. Sport

Fish. Wild. U.S. Government Printing Office, Washington, DC.King, J. E. (1983). Seals of the World, 2nd ed., British Museum of Natural History, London, and Cornell

University Press, Ithaca, NY.Lacépéde, B. (1804). Histoire naturelle de Lacépède: comprenant les cétacés, les quadrupèdes ovipares, les ser-

pents et les poissons. Furne et cie, Paris.Larson, L.M. Speculum Regale (Iceland 13th Century). The King’s Mirror: Translated from the Old

Norwegian. (Scandinavian monographs: 3). American-Norwegian Foundation, 1917, New York.Lavigne, D. M., and K. M. Kovacs (1988). Harps and Hoods. University of Waterloo Press, Ontario, Canada.Leatherwood, S., and R. R. Reeves (1983). The Sierra Club Handbook of Whales and Dolphins. Sierra Club

Books, San Francisco, CA.Leatherwood, S., and R. R. Reeves (eds.) (1990). The Bottlenose Dolphin. Academic Press, San Diego, CA.Le Boeuf, B.J., and R. M. Laws (eds.) (1994). Elephant Seals. University of California Press, Berkeley.Mackintosh, N. A., and J. F. G. Wheeler (1929). “Southern Blue and Fin Whales.” Discovery Report 1:

257–540.Mann, J., R. C. Connor, P. L. Tyack, and H. Whitehead (eds.) (2000). Cetacean Societies: Field Studies of

Dolphins and Whales. University of Chicago Press, Chicago.Martens, F. (1675). Spitzbergische oder Groenlandische Reise Beschreibung gethan im Jahr 1671: aus eigner

Erfahrunge beschrieben, die dazu erforderte Figuren nach dem Leben selbst abgerissen (so hierbey in Kupfferzu sehen) und jetzo durch den Druck mitgetheilet. Auff Gottfried Schultzens Kosten gedruckt, Hamburg.

Matthews, L. H. (1966). Chairman’s Introduction to First Session of the International Symposium onCetacean Research. In “Whales, Dolphins and Porpoises.” (K. S. Norris, ed.), pp. 3–6. University ofCalifornia Press, Berkeley.

Matthews, L. H. (1978). The Natural History of the Whales. Columbia University Press, New York.Miller, W. C. G. (1888). The myology of the Pinnipedia. In “Report on the Scientific Results of the Voyage of

H.M.S. Challenger.” 26(2): 139–240; appendix to Turner’s report. Challenger Office, 1880–1895,Edinburgh.

Murie, J. (1870). “Researches Upon the Anatomy of the Pinnipedia. Part I. On the Walrus (Trichechus rosmarusLinn.).”Trans. Zool. Soc. London 7: 411–464.

Murie, J. (1872). “Researches Upon the Anatomy of the Pinnipedia. Part 2. Descriptive Anatomy of theSea-Lion (Otaria jubata).” Trans. Zool. Soc. London 7: 527–596.

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