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Overview

Brief Summary

Fossil species

recent & fossil

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Cetaceans is the official name for the whale family. When you hear the word 'whale', you usually picture a giant animal. However, porpoises which are only 1.5 meters long also fall under cetaceans. The predecessors of the cetaceans lived on land. During the Tertiary period (around 50 million years ago), a group of predator-like mammals moved to coastal regions. They slowly adapted to swimming in the sea. They were animals without back legs, but had fins and a wide flattened tail.
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Comprehensive Description

The order Cetacea comprises two extant sub-orders and one extinct sub-order. The extant sub-orders are Mysticeti (baleen whales) and Odontoceti (toothed whales). Both mysticetes and odontocetes are thought to be descendants of archaeocetes (Archaeoceti, ancient whales), an extinct sub-order. There are at least 83 living species of cetaceans, with 46 genera in 14 families. Of the two extant suborders, Odontoceti is larger and more diverse, with at least 70 species, 40 genera, and 10 families. Cetaceans, along with bats, are considered some of the most derived mammals on the planet. They evolved from terrestrial animals to an entirely aquatic life form that is completely separated from the land in all aspects of biology. Cetaceans live, breed, rest, and carry out all of their life functions in the water.

  • Reeves, R., B. Stewart, P. Clapham, J. Powell. 2002. National Audobon Society Guide to Marine Mammals of the World. New York: Alfred A. Knopf.
  • Gingerich, P., M. ul Haq, I. Zalmout, I. Khan, M. Malkani. 2001. Origin of whales from early artiodactyls: Hands and feet of Eocene Protocetidae from Pakistan. Science, 293: 2239-2242.
  • Rice, D. 1984. Cetaceans. Pp. 447-490 in S Anderson, J Jones Jr., eds. Orders and Families of Recent Mammals of the World. New York: John Wiley and Sons.
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Distribution

Cetaceans inhabit all of the world's oceans, as well as some freshwater lakes and rivers in South America, North America, and Asia. Some species, such as killer whales (Orcinus orca) are found in all of the world's oceans. Others are limited to one hemisphere (Antarctic minke whales) or ocean (Pacific white-sided dolphins). Still others have highly restricted ranges. For example, vaquitas (Phocoena sinus) only occur in the northern part of the Gulf of California.

Biogeographic Regions: arctic ocean (Native ); indian ocean (Native ); atlantic ocean (Native ); pacific ocean (Native ); mediterranean sea (Native )

Other Geographic Terms: cosmopolitan

  • Nowak, R. 1999. Walker's Mammals of the World, vol. II. Baltimore and London: The Johns Hopkins University Press.
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Physical Description

Morphology

All cetaceans share a number of similarities: they have a fusiform, or streamlined body shape; paddle-shaped front limbs; vestigial hind limbs (which are within the body wall); no external digits or claws; tail flattened laterally and bearing horizontal flukes at the tip; vestigial ear pinnae; basically hairless body (some young have hair on their snouts); thick subcutaneous blubber layer filled with fat and oil; telescoped skull bones; external nares (blowhole) on the top of the head; addition of compressed vertebrae; shortening of the neck; lack of sweat glands; internal reproductive organs; 3-chambered stomach; and an airway reinforced with cartilage to the alveoli. Many of these characteristics are adaptations to reduce drag for fast swimming in an aquatic environment. Protuberances such as external ears or genitals would create turbulence and would be inefficient for an animal in the water.

Cetaceans are white, black, gray, bluish-gray, or pink in color, and many are spotted, mottled, streaked, or boldly patterned. Most exhibit some countershading, tending to be lighter ventrally than dorsally. They are large animals, ranging in size from 20 to 180,000 kilograms and from 1.2 to 30 meters. Blue whales (Balaenoptera musculus) are the largest animals that have ever existed. Some species are sexually dimorphic in size. For example, female blue whales are larger than males and male bottlenose dolphins (Tursiops truncatus) are larger than female bottlenose dolphins. In a few species, such as narwhals (Monodon monoceros) and beaked whales (Ziphiidae), males have enlarged, protruding teeth that may be used in aggressive male-male encounters.

Some cetaceans are thought to be the most intelligent non-primates and many have proportionately large brains. They also have remarkably efficient lungs and circulatory systems, allowing them to dive for extended periods of time. Cetaceans use about 12% of the oxygen that they inhale, compared to 4% used by terrestrial mammals. They also have at least twice as many erythrocytes and myoglobin molecules in their blood, for efficient capture and transport of oxygen. When cetaceans dive, their heart rates slow by as much as 80 beats per minute, so their bodies use less oxygen than they would otherwise.

Cetaceans are found in all climates, including regions where sea water is near freezing. Small cetaceans can cope with cold temperatures because they have high metabolic rates. Also, their flippers and flukes have a countercurrent heat exchange system, wherein heat from arterial blood warms venous blood as it returns to the heart. Large cetaceans have a small surface area to volume ratio, so they lose little heat to the surrounding environment. Both small and large cetaceans are insulated by their thick blubber layer.

Cetaceans have diploid chromosome numbers of 42 to 44.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: female larger; male larger; ornamentation

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Ecology

Habitat

Cetaceans are exclusively aquatic. Most are species are marine, inhabiting coastal areas as well as open ocean. A few species inhabit freshwater rivers and lakes. Others live in the brackish waters of estuaries and coastal marshes.

Habitat Regions: temperate ; tropical ; polar ; saltwater or marine ; freshwater

Aquatic Biomes: pelagic ; lakes and ponds; rivers and streams; coastal ; brackish water

Other Habitat Features: riparian ; estuarine

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Trophic Strategy

Mysticetes are filter feeders, using their baleen to strain plankton and other tiny organisms from the water. Odontocetes primarily feed on fish, squid, and crustaceans, though the larger species also eat aquatic birds and mammals (including other cetaceans).

Foraging Behavior: filter-feeding

Primary Diet: carnivore (Eats terrestrial vertebrates, Piscivore , Eats non-insect arthropods, Molluscivore ); planktivore

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Associations

Cetaceans have vital ecosystem roles as consumers of plankton, fish, crustaceans, cephalopods, and other aquatic animals. They are host to a range of internal parasites, including cestodes in their intestines (Tetrabothrium and Diplogonoporus), plerocercoids in their blubber and peritonea (Phyllobothrium and Monorygma), trematodes in their stomachs, livers, intestines, and sinuses, acanthocephalans in their intestines (Bolbosoma and Corynosoma), and nematodes in their stomachs and urinogenital tracts (Anisakis, Crassicauda, and Placentonema). In addition, odontocetes are parasitized by cetacean lungworms in the family Pseudaliidae.

Cetaceans are not free from external parasites, either. Cookie-cutter sharks (Isistius brasiliensis) attack them and bite off chunks of blubber, and lampreys (Petromyzon marinus) have also been known to attack them. Cetaceans are host to small parasites such as barnacles, which live on or in their skin, and copepods, such as Balaenophilus unisetus and Pennella, which live on their skin or in their blubber. Diatoms (Cocconeis ceticola) coat the skin of mysticetes with a greenish slime. Nematodes (Odontobius ceti), protozoans, and whale lice (Cyamidae) also inhabit the skin. Some cetacean species are mutualists with animals that feed on these ectoparasites; for example, topsmelt (Atherinops affinis) consume whale lice that live on the skin of gray whales (Eschrichtius robustus). Some birds have commensal relationships with cetaceans. Seagulls often follow schools of dolphins and consume small fish stirred up by the feeding cetaceans. Remoras (Remora australis) sometimes attach themselves to the skin of cetaceans and hang on for the ride, and pilotfish (Naucrates ductor) sometimes accompany killer whales and eat scraps from their kills.

Mutualist Species:

Commensal/Parasitic Species:

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Cetaceans have few natural predators, save other cetaceans (killer whales, Orcinus orca), sharks, and occasionally walruses (Odobenus rosmarus) and polar bears (Ursus maritimus), which feed on belugas trapped in ice. Small odontocetes rely on their speed and agility to escape predators. Humans prey on cetaceans throughout the world.

Known Predators:

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Animal / carrion / dead animal feeder
fruitbody of Schizophyllum commune feeds on dead dead whale bone of Cetacea
Other: unusual host/prey

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Known predators

Cetacea (whales & dolphins) is prey of:
Chondrichthyes

Based on studies in:
South Africa, Southwest coast (Marine)

This list may not be complete but is based on published studies.
  • Yodzis P (2000) Diffuse effects in food webs. Ecology 81:261–266
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Known prey organisms

Cetacea (whales & dolphins) preys on:
Euphausia superba
Euphausia crystallorophias
zooplankton
Actinopterygii
Cephalopoda
Phocidae
macrozooplankton
Engraulidae
Clupeidae
Etrumeus teres
Diaphus splendidus
Callogobius atratus
Trachurus
Merluccius

Based on studies in:
Antarctic (Estuarine)
Southern Ocean (Marine, Tropical)
South Africa, Southwest coast (Marine)

This list may not be complete but is based on published studies.
  • G. A. Knox, Antarctic marine ecosystems. In: Antarctic Ecology, M. W. Holdgate, Ed. (Academic Press, New York, 1970) 1:69-96, from p. 87.
  • B. C. Patten and J. T. Finn, Systems approach to continental shelf ecosystems. In: Theoretical Systems Ecology, E. Halfon, Ed. (Academic Press, New York, 1979) pp. 183-212 from p. 202.
  • Yodzis P (2000) Diffuse effects in food webs. Ecology 81:261–266
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Life History and Behavior

Behavior

Cetaceans produce a variety of sounds. Baleen whales moan, grunt, chirp, whistle, and click to communicate; these sounds are made by the larynx. Male humpback whales (Megaptera novaeangliae) "sing" for up to 40 minutes at a time, presumably to attract females. Low-pitched moans produced by some baleen whales may be the loudest sounds produced by any animal; they carry underwater for hundreds of kilometers. Odontocetes communicate with whistles; these sounds are most likely produced by opening and closing nasal plugs. Odonotocetes also use clicks for echolocation, to navigate and to find food. They have a fatty organ called a melon on the forehead that focuses acoustic signals as they are emitted, and they receive sounds in the middle ear via the mandible. Odonocetes have a hearing range that greatly exceeds that of humans; they can perceive ultrasounds up to 120 kHz.

Though they rely mainly on sound to communicate, most cetaceans are able to see fairly well in both water and air. River dolphins (Inia, Lipotes, Platanista, Pontoporia) are the exception; there is little use for vision in the turbid waters where they reside, so their eyes are greatly reduced and some are nearly blind. Social odontocetes use touch extensively with pod-mates, which may be an important form of communication.

Communication Channels: tactile ; acoustic

Perception Channels: visual ; tactile ; acoustic ; ultrasound ; echolocation ; chemical

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Life Expectancy

Because of the difficulties involved with tracking and studying cetaceans, lifespans are difficult to estimate. Available estimates suggest that most species live at least two decades, and some live much longer than that. One-hundred and sixteen-year-old fin whales (Balaenoptera physalus) have been reported from the wild and bowhead whales (Balaena mysticetus) may live up to 200 years.

  • George, J., J. Bada, J. Zeh, L. Scott, S. Brown. 1999. Age and growth estimates of bowhead whales (Balaena mysticetus) via aspartic acid racemization. Canadian Journal of Zoology, 78: 1182-1198.
  • Carey, J., D. Judge. 2002. "Longevity Records: Life Spans of Mammals, Birds, Amphibians, Reptiles, and Fish" (On-line). Accessed November 28, 2005 at http://www.demogr.mpg.de/.
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Reproduction

Cetaceans are difficult creatures to study, and for this reason, the behavior and biology of many species remains a mystery. Most species that have been studied exhibit polyandry, polygyny or polygynandry. The large testis size of some (such as bowhead whales, Balaena mysticetus) indicates sperm competition. Sometimes, as with northern right whales (Eubalaena glacialis), courtship behavior can be observed at the surface. In this species, multiple males congregate around a single female and jostle for position in a frenzy of activity. The female may mate with several males in succession, or even with two at the same time.

Mating System: polyandrous ; polygynous ; polygynandrous (promiscuous)

Most cetacean species have just one mating season per year. Females each give birth to a single calf every one to six years, after a 10 to 17 month gestation. Minke whales (Balaenoptera acutorostrata) have a postpartum estrus. Calving usually occurs in the warm season. Calves are born tail-first and must swim from the moment of birth. Female cetaceans nurse their calves for anywhere from six months (many mysticetes) to over two years (many odontocetes: 13-year-old sperm whales have been found with milk in their stomachs). Calves grow quickly (nursing blue whale calves gain 90 kg per day) but do not reach sexual maturity for at least two years, and mysticetes do not reach sexual maturity until they are at least 10 years old. Even after sexual maturity, cetaceans may have to wait several years before they are socially mature enough to breed.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous ; post-partum estrous

Female cetaceans nurture their young inside their bodies for at least ten months, and then they provide them with milk extremely rich in protein and fat for at least six months. They are attentive mothers, helping their calves reach the surface to take their first breaths after they are born and keeping an eye out for predators. In some species, such as belugas (Delphinapterus leucas) mothers and calves form associations that last long after calves are weaned. Male parental care is rare, but males have been reported anecdotally to show interest in their offspring and bring them food. Calves are precocial, as they must be able to swim from birth.

Parental Investment: precocial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female); post-independence association with parents; extended period of juvenile learning

  • Nowak, R. 1999. Walker's Mammals of the World, vol. II. Baltimore and London: The Johns Hopkins University Press.
  • Reeves, R., B. Stewart, P. Clapham, J. Powell. 2002. National Audobon Society Guide to Marine Mammals of the World. New York: Alfred A. Knopf.
  • Rice, D. 1984. Cetaceans. Pp. 447-490 in S Anderson, J Jones Jr., eds. Orders and Families of Recent Mammals of the World. New York: John Wiley and Sons.
  • baiji.org Foundation, 2005. "World’s First Freshwater Porpoise Is Born In Captivity!" (On-line). Accessed November 28, 2005 at http://www.baiji.org/in_depth/porpoise_is_born.html.
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Evolution and Systematics

Functional Adaptations

Functional adaptation

Organs aid navigation: dolphin
 

The larynx and an organ on the head of dolphins detect and identify shapes of objects by sending and detecting sonar.

       
  "Moving at such speeds, navigation becomes critically important. Fish are helped by their lateral line system, but mammals lost that far back in their ancestry and the toothed whales have instead a system based on the sounds used by shrews and elaborated by bats, sonar. Dolphins produce the ultra-sound with larynx and maybe an organ in the front of the head, the melon. The frequencies they use are around 200,000 vibrations a second, which is comparable to those used by bats. With its aid, they can not only sense obstacles in their path, but identify from the quality of the echo, the nature of the objects ahead. This can be demonstrated easily enough for dolphins flourish in oceanaria and eagerly cooperate in training. Blindfolded dolphins demonstrate that they can, without difficulty, pick out particular shapes of floating rings and will swiftly swim through the water, with blindfolds on their eyes, and exultantly collect on their snout the one shape that they know will bring a reward." (Attenborough 1979:243)

Watch Video
  Learn more about this functional adaptation.
  • Attenborough, D. 1979. Life on earth. Boston, MA: Little, Brown and Company. 319 p.
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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
Specimen Records: 781
Specimens with Sequences: 665
Specimens with Barcodes: 594
Species: 71
Species With Barcodes: 59
Public Records: 589
Public Species: 59
Public BINs: 59
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Barcode data

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Conservation

Conservation Status

The IUCN lists 28 cetacean species as lower risk, 5 as vulnerable, 7 as endangered, 2 as critically endangered, and 39 as data deficient. All cetaceans are listed in CITES Appendix II, except for Lipotes vexillifer, Platanista spp., Berardius spp., Hyperoodon spp., Physeter catodon, Orcaella brevirostris, Sotalia fluviatilis, Sousa spp., Neophocaena phocaenoides, Phocoena sinus, Eschrichtius robustus, Balaenoptera acutorostrata, Balaenoptera bonaerensis, Balaenoptera borealis, Balaenoptera edeni, Balaenoptera musculus, Balaenoptera physalus, Megaptera novaeangliae, Balaena mysticetus, Eubalaena spp., and Caperea marginata, which are listed in Appendix I. Commercial whaling in the 18th, 19th, and 20th centuries took a huge toll on many populations of mysticetes, which have low reproductive rates and cannot recover quickly from overexploitation. Since whales were given legal protections in the latter half of the 20th century and commercial whaling was banned in 1986, some populations seem to be recovering, albeit slowly. Continued whaling by some nations as "scientific whaling" may still threaten some species. Many species of small odontocetes are threatened by commercial fishing operations. They become entangled in nets and drown or they are killed by explosives. Sometimes they are killed on purpose by fishermen who see them as competitors. They also risk being struck by vessels, and some wild populutions have been depleted by people capturing live animals for oceanariums. Captive breeding programs may be the only hope for such critically endangered odontocetes as Yangtze river dolphins (Lipotes vexillifer). The use of military sonar in the oceans, as well as the increase of ocean noise, is becoming more widely recognized as a serious and growing threat to cetaceans. Finally, all cetaceans face the threats of pollution and global climate change.

  • IUCN, 2004. "2004 IUCN Red List of Threatened Species" (On-line). Accessed November 28, 2005 at www.redlist.org.
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Relevance to Humans and Ecosystems

Benefits

Cetaceans may negatively impact commercial fisheries, because they compete for fish and become entangled in fishing nets.

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For centuries, cetaceans have been hunted for their meat, blubber, and oil. The late 19th and 20th centuries saw a boom in commercial whaling to provide oil for lighting and heating and facilitated by newer and deadlier technologies for hunting whales. However, in 1986 a moratorium on commercial whaling was enacted by the International Whaling Commission, and today only Norway continues a commercial harvest. Harvesting for subsistence and scientific purposes continues, however, and several countries, including Iceland and Japan, continue to hunt whales under the name of "scientific" whaling. Currently cetaceans are important for entertainment and tourist industries: captive odontocetes are trained to perform tricks for large crowds of spectators, and whale-watching boats are popular attractions for tourists wishing to catch a glimpse of cetaceans in the wild. Whales are also important cultural and mythological icons for peoples around the world.

Positive Impacts: food ; body parts are source of valuable material; ecotourism ; research and education

  • Price, S., O. Bininda-Emonds, J. Gittleman. 2005. A complete phylogeny of the whales, dolphins and even-toed hoofed mammals (Cetartiodactyla). Biological Reviews, 80: 445-473.
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