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The term "fish" most precisely describes any non-tetrapod craniate (i.e. an animal with a skull and in most cases a backbone) that has gills throughout life and whose limbs, if any, are in the shape of fins. Fish are gill-bearing aquatic craniate animals that lack limbs with digits. They form a sister group to the tunicates, together forming the olfactores. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Tetrapods emerged within lobe-finned fishes, so cladistically they are fish as well. However, traditionally fish are rendered paraphyletic by excluding the tetrapods (i.e., the amphibians, reptiles, birds and mammals which all descended from within the same ancestry). Because in this manner the term "fish" is defined negatively as a paraphyletic group, it is not considered a formal taxonomic grouping in systematic biology, unless it is used in the cladistic sense, including tetrapods. The earliest organisms that can be classified as fish were soft-bodied chordates that first appeared during the Cambrian period. Although they lacked a true spine, they possessed notochords which allowed them to be more agile than their invertebrate counterparts. Fish would continue to evolve through the Paleozoic era, diversifying into a wide variety of forms. Many fish of the Paleozoic developed external armor that protected them from predators. The first fish with jaws appeared in the Silurian period, after which many (such as sharks) became formidable marine predators rather than just the prey of arthropods. Most fish are ectothermic ("cold-blooded"), allowing their body temperatures to vary as ambient temperatures change, though some of the large active swimmers like white shark and tuna can hold a higher core temperature. Fish can communicate in their underwater environments through the use of acoustic communication. Acoustic communication in fish involves the transmission of acoustic signals from one individual of a species to another. The production of sounds as a means of communication among fish is most often used in the context of feeding, aggression or courtship behavior. The sounds emitted by fish can vary depending on the species and stimulus involved. They can produce either stridulatory sounds by moving components of the skeletal system, or can produce non-stridulatory sounds by manipulating specialized organs such as the swimbladder. Fish are abundant in most bodies of water. They can be found in nearly all aquatic environments, from high mountain streams (e.g., char and gudgeon) to the abyssal and even hadal depths of the deepest oceans (e.g., gulpers and anglerfish), although no species has yet been documented in the deepest 25% of the ocean. With 33,600 described species, fish exhibit greater species diversity than any other group of vertebrates. Fish are an important resource for humans worldwide, especially as food. Commercial and subsistence fishers hunt fish in wild fisheries (see fishing) or farm them in ponds or in cages in the ocean (see aquaculture). They are also caught by recreational fishers, kept as pets, raised by fishkeepers, and exhibited in public aquaria. Fish have had a role in culture through the ages, serving as deities, religious symbols, and as the subjects of art, books and movies. Fish, as vertebrata, developed as sister of the tunicata. As the tetrapods emerged deep within the fishes group, as sister of the lungfish, characteristics of fish are typically shared by tetrapods, including having vertebrae and a cranium. Drawing of animal with large mouth, long tail, very small dorsal fins, and pectoral fins that attach towards the bottom of the body, resembling lizard legs in scale and development.[9] Dunkleosteus was a gigantic, 10-metre (33 ft) long prehistoric fish of class Placodermi. Early fish from the fossil record are represented by a group of small, jawless, armored fish known as ostracoderms. Jawless fish lineages are mostly extinct. An extant clade, the lampreys may approximate ancient pre-jawed fish. The first jaws are found in Placodermi fossils. The diversity of jawed vertebrates may indicate the evolutionary advantage of a jawed mouth. It is unclear if the advantage of a hinged jaw is greater biting force, improved respiration, or a combination of factors. Fish may have evolved from a creature similar to a coral-like sea squirt, whose larvae resemble primitive fish in important ways. Many types of aquatic animals commonly referred to as "fish" are not fish in the sense given above; examples include shellfish, cuttlefish, starfish, crayfish and jellyfish. In earlier times, even biologists did not make a distinction – sixteenth century natural historians classified also seals, whales, amphibians, crocodiles, even hippopotamuses, as well as a host of aquatic invertebrates, as fish.