Humans make use of many other animal species for food, including meat, milk, and eggs; for materials, such as leather and wool; as pets; and as working animals for power and transport. Dogs have been used in hunting, while many terrestrial and aquatic animals are hunted for sport. Non-human animals have appeared in art from the earliest times and are featured in mythology and religion.
The word "animal" comes from the Latin animalis, meaning having breath, having soul or living being. The biological definition includes all members of the kingdom Animalia. In colloquial usage, as a consequence of anthropocentrism, the term animal is sometimes used nonscientifically to refer only to non-human animals.
Animals are unique in having the ball of cells of the early embryo (1) develop into a hollow ball or blastula (2).
All animals are composed of cells, surrounded by a characteristic extracellular matrix composed of collagen and elastic glycoproteins. During development, the animal extracellular matrix forms a relatively flexible framework upon which cells can move about and be reorganised, making the formation of complex structures possible. This may be calcified, forming structures such as shells, bones, and spicules. In contrast, the cells of other multicellular organisms (primarily algae, plants, and fungi) are held in place by cell walls, and so develop by progressive growth. Animal cells uniquely possess the cell junctions called tight junctions, gap junctions, and desmosomes.
With few exceptions--in particular, the sponges and placozoans--animal bodies are differentiated into tissues. These include muscles, which enable locomotion, and nerve tissues, which transmit signals and coordinate the body. Typically, there is also an internal digestive chamber with either one opening (in Ctenophora, Cnidaria, and flatworms) or two openings (in most bilaterians).
Nearly all animals make use of some form of sexual reproduction. They produce haploidgametes by meiosis; the smaller, motile gametes are spermatozoa and the larger, non-motile gametes are ova. These fuse to form zygotes, which develop via mitosis into a hollow sphere, called a blastula. In sponges, blastula larvae swim to a new location, attach to the seabed, and develop into a new sponge. In most other groups, the blastula undergoes more complicated rearrangement. It first invaginates to form a gastrula with a digestive chamber and two separate germ layers, an external ectoderm and an internal endoderm. In most cases, a third germ layer, the mesoderm, also develops between them. These germ layers then differentiate to form tissues and organs.
Animals originally evolved in the sea. Lineages of arthropods colonised land around the same time as land plants, probably between 510-471 million years ago during the Late Cambrian or Early Ordovician.Vertebrates such as the lobe-finned fishTiktaalik started to move on to land in the late Devonian, about 375 million years ago. Animals occupy virtually all of earth's habitats and microhabitats, including salt water, hydrothermal vents, fresh water, hot springs, swamps, forests, pastures, deserts, air, and the interiors of animals, plants, fungi and rocks. Animals are however not particularly heat tolerant; very few of them can survive at constant temperatures above 50 °C (122 °F). Only very few species of animals (mostly nematodes) inhabit the most extreme cold deserts of continental Antarctica.
The blue whale is the largest animal that has ever lived.
Largest and smallest
The blue whale (Balaenoptera musculus) is the largest animal that has ever lived, weighing up to 190 metric tonnes and measuring up to 33.6 metres (110 ft) long. The largest extant terrestrial animal is the African bush elephant (Loxodonta africana), weighing up to 12.25 tonnes and measuring up to 10.67 metres (35.0 ft) long. The largest terrestrial animals that ever lived were titanosaursauropod dinosaurs such as Argentinosaurus, which may have weighed as much as 73 tonnes. Several animals are microscopic; some Myxozoa (obligate parasites within the Cnidaria) never grow larger than 20 µm, and one of the smallest species (Myxobolus shekel) is no more than 8.5 µm when fully grown.
Numbers and habitats
The following table lists estimated numbers of described extant species for the animal groups with the largest numbers of species, along with their principal habitats (terrestrial, fresh water, and marine), and free-living or parasitic ways of life. Species estimates shown here are based on numbers described scientifically; much larger estimates have been calculated based on various means of prediction, and these can vary wildly. For instance, around 25,000-27,000 species of nematodes have been described, while published estimates of the total number of nematode species include 10,000-20,000; 500,000; 10 million; and 100 million. Using patterns within the taxonomic hierarchy, the total number of animal species--including those not yet described--was calculated to be about 7.77 million in 2011.[a]
The oldest animals are found in the Ediacaran biota, towards the end of the Precambrian, around 610 million years ago. It had long been doubtful whether these included animals, but the discovery of the animal lipid cholesterol in fossils of Dickinsonia establishes that these were indeed animals. Animals are thought to have originated under low-oxygen conditions, suggesting that they were capable of living entirely by anaerobic respiration, but as they became specialized for aerobic metabolism they became fully dependent on oxygen in their environments.
Some palaeontologists have suggested that animals appeared much earlier than the Cambrian explosion, possibly as early as 1 billion years ago.Trace fossils such as tracks and burrows found in the Tonian period may indicate the presence of triploblastic worm-like animals, roughly as large (about 5 mm wide) and complex as earthworms. However, similar tracks are produced today by the giant single-celled protist Gromia sphaerica, so the Tonian trace fossils may not indicate early animal evolution. Around the same time, another line of evidence may indicate the appearance of grazing animals: the layered mats of microorganisms called stromatolites decreased in diversity, perhaps due to grazing.
Non-bilaterians include sponges (centre) and corals (background).
Several animal phyla lack bilateral symmetry. Among these, the sponges (Porifera) probably diverged first, representing the oldest animal phylum. Sponges lack the complex organization found in most other animal phyla; their cells are differentiated, but in most cases not organised into distinct tissues. They typically feed by drawing in water through pores.
The Ctenophora (comb jellies) and Cnidaria (which includes jellyfish, sea anemones, and corals) are radially symmetric and have digestive chambers with a single opening, which serves as both mouth and anus. Animals in both phyla have distinct tissues, but these are not organised into organs. They are diploblastic, having only two main germ layers, ectoderm and endoderm. The tiny placozoans are similar, but they do not have a permanent digestive chamber.
Idealised bilaterian body plan.[c] With an elongated body and a direction of movement the animal has head and tail ends. Sense organs and mouth form the basis of the head. Opposed circular and longitudinal muscles enable peristaltic motion.
The remaining animals, the great majority--comprising some 29 phyla and over a million species--form a clade, the Bilateria. The body is triploblastic, with three well-developed germ layers, and their tissues form distinct organs. The digestive chamber has two openings, a mouth and an anus, and there is an internal body cavity, a coelom or pseudocoelom. Animals with this bilaterally symmetric body plan and a tendency to move in one direction have a head end (anterior) and a tail end (posterior) as well as a back (dorsal) and a belly (ventral); therefore they also have a left side and a right side.
Having a front end means that this part of the body encounters stimuli, such as food, favouring cephalisation, the development of a head with sense organs and a mouth. Many bilaterians have a combination of circular muscles that constrict the body, making it longer, and an opposing set of longitudinal muscles, that shorten the body; these enable soft-bodied animals with a hydrostatic skeleton to move by peristalsis. They also have a gut that extends through the basically cylindrical body from mouth to anus. Many bilaterian phyla have primary larvae which swim with cilia and have an apical organ containing sensory cells. However, there are exceptions to each of these characteristics; for example, adult echinoderms are radially symmetric (unlike their larvae), while some parasitic worms have extremely simplified body structures.
The bilaterian gut develops in two ways. In many protostomes, the blastopore develops into the mouth, while in deuterostomes it becomes the anus.
Protostomes and deuterostomes differ in several ways. Early in development, deuterostome embryos undergo radial cleavage during cell division, while many protostomes (the Spiralia) undergo spiral cleavage.
Animals from both groups possess a complete digestive tract, but in protostomes the first opening of the embryonic gut develops into the mouth, and the anus forms secondarily. In deuterostomes, the anus forms first while the mouth develops secondarily. Most protostomes have schizocoelous development, where cells simply fill in the interior of the gastrula to form the mesoderm. In deuterostomes, the mesoderm forms by enterocoelic pouching, through invagination of the endoderm.
The Ecdysozoa are protostomes, named after their shared trait of ecdysis, growth by moulting. They include the largest animal phylum, the Arthropoda, which contains insects, spiders, crabs, and their kin. All of these have a body divided into repeating segments, typically with paired appendages. Two smaller phyla, the Onychophora and Tardigrada, are close relatives of the arthropods and share these traits. The ecdysozoans also include the Nematoda or roundworms, perhaps the second largest animal phylum. Roundworms are typically microscopic, and occur in nearly every environment where there is water; some are important parasites. Smaller phyla related to them are the Nematomorpha or horsehair worms, and the Kinorhyncha, Priapulida, and Loricifera. These groups have a reduced coelom, called a pseudocoelom.
Jean-Baptiste de Lamarck led the creation of a modern classification of invertebrates, breaking up Linnaeus's "Vermes" into 9 phyla by 1809.
In the classical era, Aristotle divided animals,[d] based on his own observations, into those with blood (roughly, the vertebrates) and those without. The animals were then arranged on a scale from man (with blood, 2 legs, rational soul) down through the live-bearing tetrapods (with blood, 4 legs, sensitive soul) and other groups such as crustaceans (no blood, many legs, sensitive soul) down to spontaneously-generating creatures like sponges (no blood, no legs, vegetable soul). Aristotle was uncertain whether sponges were animals, which in his system ought to have sensation, appetite, and locomotion, or plants, which did not: he knew that sponges could sense touch, and would contract if about to be pulled off their rocks, but that they were rooted like plants and never moved about.
In 1758, Carl Linnaeus created the first hierarchical classification in his Systema Naturae. In his original scheme, the animals were one of three kingdoms, divided into the classes of Vermes, Insecta, Pisces, Amphibia, Aves, and Mammalia. Since then the last four have all been subsumed into a single phylum, the Chordata, while his Insecta (which included the crustaceans and arachnids) and Vermes have been renamed or broken up. The process was begun in 1793 by Jean-Baptiste de Lamarck, who called the Vermes une espèce de chaos (a chaotic mess)[e] and split the group into three new phyla, worms, echinoderms, and polyps (which contained corals and jellyfish). By 1809, in his Philosophie Zoologique, Lamarck had created 9 phyla apart from vertebrates (where he still had 4 phyla: mammals, birds, reptiles, and fish) and molluscs, namely cirripedes, annelids, crustaceans, arachnids, insects, worms, radiates, polyps, and infusorians.
In 1874, Ernst Haeckel divided the animal kingdom into two subkingdoms: Metazoa (multicellular animals, with five phyla: coelenterates, echinoderms, articulates, molluscs, and vertebrates) and Protozoa (single-celled animals), including a sixth animal phylum, sponges. The protozoa were later moved to the former kingdom Protista, leaving only the Metazoa as a synonym of Animalia.
^The application of DNA barcoding to taxonomy further complicates this; a 2016 barcoding analysis estimated a total count of nearly 100,000 insect species for Canada alone, and extrapolated that the global insect fauna must be in excess of 10 million species, of which nearly 2 million are in a single fly family known as gall midges (Cecidomyiidae).
^Castro, Peter; Huber, Michael E. (2007). Marine Biology (7th ed.). McGraw-Hill. p. 376. ISBN978-0-07-722124-9.
^Rota-Stabelli, Omar; Daley, Allison C.; Pisani, Davide (2013). "Molecular Timetrees Reveal a Cambrian Colonization of Land and a New Scenario for Ecdysozoan Evolution". Current Biology. 23 (5): 392-8. doi:10.1016/j.cub.2013.01.026. PMID23375891.
^Stork, Nigel E. (January 2018). "How Many Species of Insects and Other Terrestrial Arthropods Are There on Earth?". Annual Review of Entomology. 63 (1): 31-45. doi:10.1146/annurev-ento-020117-043348. PMID28938083. Stork notes that 1m insects have been named, making much larger predicted estimates.
^Sluys, R. (1999). "Global diversity of land planarians (Platyhelminthes, Tricladida, Terricola): a new indicator-taxon in biodiversity and conservation studies". Biodiversity and Conservation. 8 (12): 1663-1681. doi:10.1023/A:1008994925673.
^ abBobrovskiy, Ilya; Hope, Janet M.; Ivantsov, Andrey; Nettersheim, Benjamin J.; Hallmann, Christian; Brocks, Jochen J. (20 September 2018). "Ancient steroids establish the Ediacaran fossil Dickinsonia as one of the earliest animals". Science. 361 (6408): 1246-1249. Bibcode:2018Sci...361.1246B. doi:10.1126/science.aat7228. PMID30237355.
^Maloof, Adam C.; Rose, Catherine V.; Beach, Robert; Samuels, Bradley M.; Calmet, Claire C.; Erwin, Douglas H.; Poirier, Gerald R.; Yao, Nan; Simons, Frederik J. (17 August 2010). "Possible animal-body fossils in pre-Marinoan limestones from South Australia". Nature Geoscience. 3 (9): 653-659. Bibcode:2010NatGe...3..653M. doi:10.1038/ngeo934.
^Zimorski, Verena; Mentel, Marek; Tielens, Aloysius G.M.; Martin, William F. (2019). "Energy metabolism in anaerobic eukaryotes and Earth's late oxygenation". Free Radical Biology and Medicine. 140: 279-294. doi:10.1016/j.freeradbiomed.2019.03.030. PMID30935869.
^Maloof, A.C.; Porter, S.M.; Moore, J.L.; Dudas, F.O.; Bowring, S.A.; Higgins, J.A.; Fike, D.A.; Eddy, M.P. (2010). "The earliest Cambrian record of animals and ocean geochemical change". Geological Society of America Bulletin. 122 (11-12): 1731-1774. Bibcode:2010GSAB..122.1731M. doi:10.1130/B30346.1.
^Peters, Kenneth E.; Walters, Clifford C.; Moldowan, J. Michael (2005). The Biomarker Guide: Biomarkers and isotopes in petroleum systems and Earth history. 2. Cambridge University Press. p. 717. ISBN978-0-521-83762-0.
^Giribet, G.; Distel, D.L.; Polz, M.; Sterrer, W.; Wheeler, W.C. (2000). "Triploblastic relationships with emphasis on the acoelomates and the position of Gnathostomulida, Cycliophora, Plathelminthes, and Chaetognatha: a combined approach of 18S rDNA sequences and morphology". Syst Biol. 49 (3): 539-562. doi:10.1080/10635159950127385. PMID12116426.
^Kim, Chang Bae; Moon, Seung Yeo; Gelder, Stuart R.; Kim, Won (September 1996). "Phylogenetic Relationships of Annelids, Molluscs, and Arthropods Evidenced from Molecules and Morphology". Journal of Molecular Evolution. 43 (3): 207-215. Bibcode:1996JMolE..43..207K. doi:10.1007/PL00006079. PMID8703086.
^Pearnchob, N.; Siepmann, J.; Bodmeier, R. (2003). "Pharmaceutical applications of shellac: moisture-protective and taste-masking coatings and extended-release matrix tablets". Drug Development and Industrial Pharmacy. 29 (8): 925-938. doi:10.1081/ddc-120024188. PMID14570313.
^Munro, John H. (2003). Jenkins, David (ed.). Medieval Woollens: Textiles, Technology, and Organisation. The Cambridge History of Western Textiles. Cambridge University Press. pp. 214-215. ISBN978-0-521-34107-3.