Toothed whale
The toothed whales are a parvorder of cetaceans that includes dolphins, porpoises, and all other whales with teeth, such as beaked whales and the sperm whales. 73 species of toothed whales are described. They are one of two living groups of cetaceans, with the other being the baleen whales, which have baleen instead of teeth. The two groups are thought to have diverged around 34 million years ago.
Toothed whales range in size from the and vaquita to the and sperm whale. Several species of odontocetes exhibit sexual dimorphism, in that there are size or other morphological differences between females and males. They have streamlined bodies and two limbs that are modified into flippers. Some can travel at up to 30 knots. Odontocetes have conical teeth designed for catching fish or squid. They have well-developed hearing that is adapted for both air and water, so much so that some can survive even if they are blind. Some species are well adapted for diving to great depths. Almost all have a layer of fat, or blubber, under the skin to keep warm in the cold water, with the exception of river dolphins.
Toothed whales consist of some of the most widespread mammals, but some, as with the vaquita, are restricted to certain areas. Odontocetes feed largely on fish and squid, but a few, like the orca, feed on mammals, such as pinnipeds. Males typically mate with multiple females every year, making them polygynous. Females mate every two to three years. Calves are typically born in the spring and summer, and females bear the responsibility for raising them, but more sociable species rely on the family group to care for calves. Many species, mainly dolphins, are highly sociable, with some pods reaching over a thousand individuals.
Once hunted for their products, cetaceans are now protected by international law. Some species are very intelligent. At the 2012 meeting of the American Association for the Advancement of Science, support was reiterated for a cetacean bill of rights, listing cetaceans as nonhuman persons. Besides whaling and drive hunting, they also face threats from bycatch and marine pollution. The baiji, for example, is considered functionally extinct by IUCN, with the last sighting in 2004, due to heavy pollution to the Yangtze River. Whales sometimes feature in literature and film, as in the great white sperm whale of Herman Melville's Moby-Dick. Small odontocetes, mainly dolphins, are kept in captivity and trained to perform tricks. Whale watching has become a form of tourism around the world.
Taxonomy
Research history
In Aristotle's time, the fourth century BC, whales were regarded as fish due to their superficial similarity. Aristotle, however, could already see many physiological and anatomical similarities with the terrestrial vertebrates, such as blood, lungs, uterus, and fin anatomy. His detailed descriptions were assimilated by the Romans, but mixed with a more accurate knowledge of the dolphins, as mentioned by Pliny the Elder in his Natural history. In the art of this and subsequent periods, dolphins are portrayed with a high-arched head and a long snout. The harbor porpoise is one of the most accessible species for early cetologists, because it could be seen very close to land, inhabiting shallow coastal areas of Europe. Many of the findings that apply to all cetaceans were therefore first discovered in the porpoises. One of the first anatomical descriptions of the airways of the whales on the basis of a harbor porpoise dates from 1671 by John Ray. It nevertheless referred to the porpoise as a fish.Evolution
Toothed whales, along with baleen whales, are descendants of land-dwelling mammals of the artiodactyl order. They are closely related to the hippopotamus, sharing a common ancestor that lived around 54 million years ago.The primitive cetaceans, or archaeocetes, first took to the sea approximately 49 mya and became fully aquatic within 5–10 million years. The ancestors of toothed whales and baleen whales diverged in the early Oligocene. This was due to a change in the climate of the southern oceans that affected the environment of the plankton that these whales ate.
The adaptation of echolocation and enhanced fat synthesis in blubber occurred when toothed whales split from baleen whales, and distinguishes modern toothed whales from fully aquatic archaeocetes. This happened around 34 mya. Unlike toothed whales, baleen whales do not have wax ester deposits nor branched fatty chain acids in their blubber. Thus, more recent evolution of these complex blubber traits occurred after baleen whales and toothed whales split, and only in the toothed whale lineage.
Modern toothed whales do not rely on their sense of sight, but on their sonar, to hunt prey. Echolocation also allowed toothed whales to dive deeper in search of food, with light no longer necessary for navigation, which opened up new food sources. Toothed whales echolocate by creating a series of clicks emitted at various frequencies. Sound pulses are emitted through the melon-shaped forehead, reflected off objects, and retrieved through the lower jaw. Skulls of Squalodon show evidence for the first hypothesized appearance of echolocation. Squalodon lived from the early to middle Oligocene to the middle Miocene, around 33–14 mya. Squalodon featured several commonalities with modern Odontocetes. The cranium was well compressed, the rostrum telescoped outward, giving Squalodon an appearance similar to that of modern toothed whales. However, it is thought unlikely that squalodontids are direct ancestors of living dolphins.
Biology
Anatomy
Toothed whales have torpedo-shaped bodies with usually inflexible necks, limbs modified into flippers, no outer ears, a large tail fin, and bulbous heads. Their skulls have small eye orbits, long beaks, and eyes placed on the sides of their heads. Toothed whales range in size from the and vaquita to the and sperm whale. Overall, they tend to be dwarfed by their relatives, the baleen whales. Several species have sexual dimorphism, with the females being larger than the males. One exception is with the sperm whale, which has males larger than the females.Odontocetes possess teeth with cementum cells overlying dentine cells. Unlike human teeth, which are composed mostly of enamel on the portion of the tooth outside of the gum, whale teeth have cementum outside the gum. Only in larger whales, where the cementum is worn away on the tip of the tooth, does enamel show. There is only a single set of functional teeth. Except for the sperm whale, most toothed whales are smaller than the baleen whales. The teeth differ considerably among the species. They may be numerous, with some dolphins bearing over 100 teeth in their jaws. At the other extreme are the narwhals with their single long tusks and the almost toothless beaked whales with tusk-like teeth only in males. In most beaked whales the teeth are seen to erupt in the lower jaw, and primarily occurs at the males sexual maturity. Not all species are believed to use their teeth for feeding. For instance, the sperm whale likely uses its teeth for aggression and showmanship.
Breathing involves expelling stale air from their one blowhole, forming an upward, steamy spout, followed by inhaling fresh air into the lungs. Spout shapes differ among species, which facilitates identification. The spout only forms when warm air from the lungs meets cold air, so it does not form in warmer climates, as with river dolphins.
Almost all cetaceans have a thick layer of blubber, except for river dolphins. In species that live near the poles, the blubber can be as thick as. This blubber can help with buoyancy, protection to some extent as predators would have a hard time getting through a thick layer of fat, energy for fasting during leaner times, and insulation from the harsh climate. Calves are born with only a thin layer of blubber, but some species compensate for this with thick lanugos.
Toothed whales have also evolved the ability to store large amounts of wax esters in their adipose tissue as an addition to or in complete replacement of other fats in their blubber. They can produce isovaleric acid from branched chain fatty acids. These adaptations are unique, are only in more recent, derived lineages and were likely part of the transition for species to become deeper divers as the families of toothed whales that have the highest quantities of wax esters and BCFAs in their blubber are also the species that dive the deepest and for the longest amount of time.
Toothed whales have a two-chambered stomach similar in structure to terrestrial carnivores. They have fundic and pyloric chambers.
Locomotion
Cetaceans have two flippers on the front, and a tail fin. These flippers contain four digits. Although toothed whales do not possess fully developed hind limbs, some, such as the sperm whale, possess discrete rudimentary appendages, which may contain feet and digits. Toothed whales are fast swimmers in comparison to seals, which typically cruise at 5–15 knots, or ; the sperm whale, in comparison, can travel at speeds of up to. The fusing of the neck vertebrae, while increasing stability when swimming at high speeds, decreases flexibility, rendering them incapable of turning their heads; river dolphins, however, have unfused neck vertebrae and can turn their heads. When swimming, toothed whales rely on their tail fins to propel them through the water. Flipper movement is continuous. They swim by moving their tail fin and lower body up and down, propelling themselves through vertical movement, while their flippers are mainly used for steering. Some species log out of the water, which may allow them to travel faster. Their skeletal anatomy allows them to be fast swimmers. Most species have a dorsal fin.Most toothed whales are adapted for diving to great depths, porpoises are one exception. In addition to their streamlined bodies, they can slow their heart rate to conserve oxygen; blood is rerouted from tissue tolerant of water pressure to the heart and brain among other organs; haemoglobin and myoglobin store oxygen in body tissue; and they have twice the concentration of myoglobin than haemoglobin. Before going on long dives, many toothed whales exhibit a behaviour known as sounding; they stay close to the surface for a series of short, shallow dives while building their oxygen reserves, and then make a sounding dive.