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Animals
Temporal range:Cryogenian– present,665–0 Ma
EchinodermCnidariaTardigradeCrustaceanArachnidSpongeInsectBryozoaRotiferFlatwormMolluscaAnnelidVertebrateTunicatePhoronida
Scientific classificationEdit this classification
Domain: Eukaryota
Clade: Amorphea
Clade: Obazoa
(unranked): Opisthokonta
(unranked): Holozoa
(unranked): Filozoa
Clade: Choanozoa
Kingdom: Animalia
Linnaeus,1758
Subdivisions
Synonyms
  • MetazoaHaeckel 1874[1]
  • ChoanoblastaeaNielsen 2008[2]
  • GastrobiontaRothm. 1948[3]
  • ZooaeaBarkley 1939[3]
  • EuanimaliaBarkley 1939[3]

Animalsaremulticellular,eukaryoticorganismsin thebiological kingdomAnimalia(/ˌænɪˈmeɪliə/[4]). With few exceptions, animalsconsume organic material,breathe oxygen,havemyocytesand areable to move,canreproduce sexually,and grow from a hollow sphere of cells, theblastula,duringembryonic development.Animals form aclade,meaning that they arose from a single common ancestor.

Over 1.5 millionlivinganimalspecieshave beendescribed,of which around 1.05 million areinsects,over 85,000 aremolluscs,and around 65,000 arevertebrates.It has been estimated there are as many as 7.77 million animal species on Earth. Animal body lengths range from 8.5 μm (0.00033 in) to 33.6 m (110 ft). They have complexecologiesandinteractionswith each other and their environments, forming intricatefood webs.The scientific study of animals is known aszoology,and the study of animal behaviors is known asethology.

Most living animal species belong to the infrakingdomBilateria,a highly proliferativecladewhose members have abilaterally symmetricbody plan.The vast majority belong to two largesuperphyla:theprotostomes,which includes organisms such as thearthropods,molluscs,flatworms,annelidsandnematodes;and thedeuterostomes,which include theechinoderms,hemichordatesandchordates,the latter of which contains the vertebrates. The simpleXenacoelomorphahave an uncertain position within Bilateria.

Animals first appear in the fossil record in the lateCryogenianperiod, and diversified in the subsequentEdiacaran.Earlier evidence of animals is still controversial; thesponge-like organismOtaviahas been dated back to theTonianperiod at the start of theNeoproterozoic,but its identity as an animal is heavily contested.[5]Nearly all modern animal phyla became clearly established in the fossil record asmarine speciesduring theCambrian explosion,which began around 539million years ago(Mya), and mostclassesduring theOrdovician radiation485.4 Mya. 6,331 groups ofgenescommon to all living animals have been identified; these may have arisen from a singlecommon ancestorthat lived about 650 Mya during theCryogenianperiod.

Historically,Aristotledivided animalsinto those with blood and those without.Carl Linnaeuscreated the first hierarchicalbiological classificationfor animals in 1758 with hisSystema Naturae,whichJean-Baptiste Lamarckexpanded into 14 phyla by 1809. In 1874,Ernst Haeckeldivided the animal kingdom into the multicellularMetazoa(nowsynonymouswith Animalia) and theProtozoa,single-celled organisms no longer considered animals. In modern times, the biological classification of animals relies on advanced techniques, such asmolecular phylogenetics,which are effective at demonstrating theevolutionaryrelationships betweentaxa.

Humansmakeuse ofmany other animal species forfood(includingmeat,eggs,anddairy products), formaterials(such asleather,fur,andwool), aspetsand asworking animalsfortransportation,andservices.Dogs,the firstdomesticatedanimal, have been usedin hunting,in securityandin warfare,as havehorses,pigeonsandbirds of prey;while otherterrestrialandaquatic animalsarehuntedfor sports, trophies or profits. Non-human animals are also an importantculturalelement ofhuman evolution,having appeared incave artsandtotemssince the earliest times, and are frequently featured inmythology,religion,arts,literature,heraldry,politics,andsports.

Etymology

The wordanimalcomes from the Latin nounanimalof the same meaning, which is itself derived from Latinanimalis'having breath or soul'.[6]The biological definition includes all members of the kingdom Animalia.[7]In colloquial usage, the termanimalis often used to refer only to nonhuman animals.[8][9][10][11]The termmetazoais derived from Ancient Greek μετα (meta) 'after' (in biology, the prefixmeta-stands for 'later') and ζῷᾰ (zōia) 'animals', plural of ζῷονzōion'animal'.[12][13]

Characteristics

Animals are unique in having the ball of cells of the earlyembryo(1) develop into a hollow ball orblastula(2).

Animals have several characteristics that set them apart from other living things. Animals areeukaryoticandmulticellular.[14]Unlike plants andalgae,whichproduce their own nutrients,[15]animals areheterotrophic,[16][17]feeding on organic material and digesting it internally.[18]With very few exceptions, animalsrespire aerobically.[a][20]All animals aremotile[21](able to spontaneously move their bodies) during at least part of theirlife cycle,but some animals, such assponges,corals,mussels,andbarnacles,later becomesessile.Theblastulais a stage inembryonic developmentthat is unique to animals, allowingcells to be differentiatedinto specialised tissues and organs.[22]

Structure

All animals are composed of cells, surrounded by a characteristicextracellular matrixcomposed ofcollagenand elasticglycoproteins.[23]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 asshells,bones,andspicules.[24]In contrast, the cells of other multicellular organisms (primarily algae, plants, andfungi) are held in place by cell walls, and so develop by progressive growth.[25]Animal cells uniquely possess thecell junctionscalledtight junctions,gap junctions,anddesmosomes.[26]

With few exceptions—in particular, the sponges andplacozoans—animal bodies are differentiated intotissues.[27]These includemuscles,which enable locomotion, andnerve tissues,which transmit signals and coordinate the body. Typically, there is also an internaldigestivechamber with either one opening (in Ctenophora, Cnidaria, and flatworms) or two openings (in most bilaterians).[28]

Reproduction and development

See also:Sexual reproduction § Animals,andAsexual reproduction § Examples in animals
Sexual reproductionis nearly universal in animals, such as thesedragonflies.

Nearly all animals make use of some form of sexual reproduction.[29]They producehaploidgametesbymeiosis;the smaller, motile gametes arespermatozoaand the larger, non-motile gametes areova.[30]These fuse to formzygotes,[31]which develop viamitosisinto 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.[32]In most other groups, the blastula undergoes more complicated rearrangement.[33]It firstinvaginatesto form agastrulawith a digestive chamber and two separategerm layers,an externalectodermand an internalendoderm.[34]In most cases, a third germ layer, themesoderm,also develops between them.[35]These germ layers then differentiate to form tissues and organs.[36]

Repeated instances ofmating with a close relativeduring sexual reproduction generally leads toinbreeding depressionwithin a population due to the increased prevalence of harmfulrecessivetraits.[37][38]Animals have evolved numerous mechanisms foravoiding close inbreeding.[39]

Some animals are capable ofasexual reproduction,which often results in a genetic clone of the parent. This may take place throughfragmentation;budding,such as inHydraand othercnidarians;orparthenogenesis,where fertile eggs are produced withoutmating,such as inaphids.[40][41]

Ecology

Predators,such as thisultramarine flycatcher(Ficedula superciliaris), feed on other animals.

Animals are categorised into ecological groups depending on theirtrophic levelsandhow they consume organic material.Such groupings includecarnivores(further divided into subcategories such aspiscivores,insectivores,ovivores,etc.),herbivores(subcategorized intofolivores,graminivores,frugivores,granivores,nectarivores,algivores,etc.),omnivores,fungivores,scavengers/detritivores,[42]andparasites.[43]Interactionsbetween animals of eachbiomeform complexfood webswithin thatecosystem.In carnivorous or omnivorous species,predationis aconsumer–resource interactionwhere the predator feeds on another organism, itsprey,[44]who often evolvesanti-predator adaptationsto avoid being fed upon.Selective pressuresimposed on one another lead to anevolutionary arms racebetween predator and prey, resulting in various antagonistic/competitivecoevolutions.[45][46]Almost all multicellular predators are animals.[47]Someconsumersuse multiple methods; for example, inparasitoid wasps,the larvae feed on the hosts' living tissues, killing them in the process,[48]but the adults primarily consume nectar from flowers.[49]Other animals may have very specificfeeding behaviours,such ashawksbill sea turtleswhich mainlyeat sponges.[50]

Hydrothermal ventmussels and shrimps

Most animals rely onbiomassandbioenergyproduced byplantsandphytoplanktons(collectively calledproducers) throughphotosynthesis.Herbivores, asprimary consumers,eat the plant material directly to digest and absorb the nutrients, while carnivores and other animals on highertrophic levelsindirectly acquire the nutrients by eating the herbivores or other animals that have eaten the herbivores. Animals oxidizecarbohydrates,lipids,proteinsand other biomolecules, which allows the animal to grow and to sustainbasal metabolismand fuel other biological processes such aslocomotion.[51][52][53]Somebenthicanimals living close tohydrothermal ventsandcold seepson the darksea floorconsume organic matter produced throughchemosynthesis(viaoxidizinginorganic compoundssuch ashydrogen sulfide) byarchaeaandbacteria.[54]

Animals evolved in the sea. Lineages of arthropods colonised land around the same time asland plants,probably between 510 and 471 million years ago during theLate Cambrianor EarlyOrdovician.[55]Vertebratessuch as thelobe-finned fishTiktaalikstarted to move on to land in the lateDevonian,about 375 million years ago.[56][57]Animals occupy virtually all of earth'shabitatsand microhabitats, withfaunasadapted to salt water, hydrothermal vents, fresh water, hot springs, swamps, forests, pastures, deserts, air, and the interiors of other organisms.[58]Animals are however not particularlyheat tolerant;very few of them can survive at constant temperatures above 50 °C (122 °F)[59]or in the most extreme cold deserts of continentalAntarctica.[60]

Diversity

Size

Further information:Largest organismsandSmallest organisms
Theblue whaleis the largest animal that has ever lived.

Theblue whale(Balaenoptera musculus) is the largest animal that has ever lived, weighing up to 190tonnesand measuring up to 33.6 metres (110 ft) long.[61][62][63]The largest extant terrestrial animal is theAfrican bush elephant(Loxodonta africana), weighing up to 12.25 tonnes[61]and measuring up to 10.67 metres (35.0 ft) long.[61]The largest terrestrial animals that ever lived weretitanosaursauropod dinosaurssuch asArgentinosaurus,which may have weighed as much as 73 tonnes, andSupersauruswhich may have reached 39 meters.[64][65]Several animals are microscopic; someMyxozoa(obligate parasiteswithin the Cnidaria) never grow larger than 20μm,[66]and one of the smallest species (Myxobolus shekel) is no more than 8.5 μm when fully grown.[67]

Numbers and habitats of major phyla

The following table lists estimated numbers of described extant species for the major animal phyla,[68]along with their principal habitats (terrestrial, fresh water,[69]and marine),[70]and free-living or parasitic ways of life.[71]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.[72]Using patterns within thetaxonomichierarchy, the total number of animal species—including those not yet described—was calculated to be about 7.77 million in 2011.[73][74][b]

Phylum Example Described species Land Sea Freshwater Free-living Parasitic
Arthropoda wasp 1,257,000[68] Yes 1,000,000
(insects)[76] 		Yes >40,000
(Malac-
ostraca)[77] 		Yes 94,000[69] Yes[70] Yes >45,000[c][71]
Mollusca snail 85,000[68]
107,000[78] 		Yes 35,000[78] Yes 60,000[78] Yes 5,000[69]
12,000[78] 		Yes[70] Yes >5,600[71]
Chordata green spotted frog facing right >70,000[68][79] Yes 23,000[80] Yes 13,000[80] Yes 18,000[69]
9,000[80] 		Yes Yes 40
(catfish)[81][71]
Platyhelminthes 29,500[68] Yes[82] Yes[70] Yes 1,300[69] Yes[70]

3,000–6,500[83]

Yes >40,000[71]

4,000–25,000[83]

Nematoda 25,000[68] Yes (soil)[70] Yes 4,000[72] Yes 2,000[69] Yes
11,000[72] 		Yes 14,000[72]
Annelida 17,000[68] Yes (soil)[70] Yes[70] Yes 1,750[69] Yes Yes 400[71]
Cnidaria Table coral 16,000[68] Yes[70] Yes (few)[70] Yes[70] Yes >1,350
(Myxozoa)[71]
Porifera 10,800[68] Yes[70] 200–300[69] Yes Yes[84]
Echinodermata 7,500[68] Yes 7,500[68] Yes[70]
Bryozoa 6,000[68] Yes[70] Yes 60–80[69] Yes
Rotifera 2,000[68] Yes >400[85] Yes 2,000[69] Yes
Nemertea 1,350[86][87] Yes Yes Yes
Tardigrada 1,335[68] Yes[88]
(moist plants) 		Yes Yes Yes
Total number of described extant species as of 2013:1,525,728[68]

Evolutionary origin

Further information:Urmetazoan

Evidence of animals is found as long ago as theCryogenianperiod.24-Isopropylcholestane(24-ipc) has been found in rocks from roughly 650 million years ago; it is only produced by sponges andpelagophytealgae. Its likely origin is from sponges based onmolecular clockestimates for the origin of 24-ipc production in both groups. Analyses of pelagophyte algae consistently recover aPhanerozoicorigin, while analyses of sponges recover aNeoproterozoicorigin, consistent with the appearance of 24-ipc in the fossil record.[89][90]

The first body fossils of animals appear in theEdiacaran,represented by forms such asCharniaandSpriggina.It had long been doubted whether these fossils truly represented animals,[91][92][93]but the discovery of the animal lipidcholesterolin fossils ofDickinsoniaestablishes their nature.[94]Animals are thought to have originated under low-oxygen conditions, suggesting that they were capable of living entirely byanaerobic respiration,but as they became specialized for aerobic metabolism they became fully dependent on oxygen in their environments.[95]

Many animal phyla first appear in thefossilrecord during theCambrian explosion,starting about 539 million years ago, in beds such as theBurgess shale.[96]Extant phyla in these rocks includemolluscs,brachiopods,onychophorans,tardigrades,arthropods,echinodermsandhemichordates,along with numerous now-extinct forms such as thepredatoryAnomalocaris.The apparent suddenness of the event may however be an artifact of the fossil record, rather than showing that all these animals appeared simultaneously.[97][98][99][100]That view is supported by the discovery ofAuroralumina attenboroughii,the earliest known Ediacaran crown-group cnidarian (557–562 mya, some 20 million years before the Cambrian explosion) fromCharnwood Forest,England. It is thought to be one of the earliestpredators,catching small prey with itsnematocystsas modern cnidarians do.[101]

Some palaeontologists have suggested that animals appeared much earlier than the Cambrian explosion, possibly as early as 1 billion years ago.[102]Early fossils that might represent animals appear for example in the 665-million-year-old rocks of theTrezona FormationofSouth Australia.These fossils are interpreted as most probably being earlysponges.[103] Trace fossilssuch as tracks and burrows found in theTonianperiod (from 1 gya) may indicate the presence oftriploblasticworm-like animals, roughly as large (about 5 mm wide) and complex as earthworms.[104]However, similar tracks are produced by the giant single-celled protistGromia sphaerica,so the Tonian trace fossils may not indicate early animal evolution.[105][106]Around the same time, the layered mats ofmicroorganismscalledstromatolitesdecreased in diversity, perhaps due to grazing by newly evolved animals.[107]Objects such as sediment-filled tubes that resemble trace fossils of the burrows of wormlike animals have been found in 1.2 gya rocks in North America, in 1.5 gya rocks in Australia and North America, and in 1.7 gya rocks in Australia. Their interpretation as having an animal origin is disputed, as they might be water-escape or other structures.[108][109]

Phylogeny

Further information:Lists of animals

External phylogeny

Animals aremonophyletic,meaning they are derived from a common ancestor. Animals are the sister group to thechoanoflagellates,with which they form theChoanozoa.[110] The dates on thephylogenetic treeindicate approximately how many millions of years ago (mya) the lineages split.[111][112][113][114][115]

Ros-Rocher and colleagues (2021) trace the origins of animals to unicellular ancestors, providing the external phylogeny shown in the cladogram. Uncertainty of relationships is indicated with dashed lines.[116]

Opisthokonta
1300 mya

Internal phylogeny

The most basal animals, thePorifera,Ctenophora,Cnidaria,andPlacozoa,have body plans that lackbilateral symmetry.Their relationships are still disputed; the sister group to all other animals could be the Porifera or the Ctenophora,[117]both of which lackhox genes,which are important forbody plan development.[118]

Hox genes are found in the Placozoa,[119][120]Cnidaria,[121]and Bilateria.[122][123]6,331 groups ofgenescommon to all living animals have been identified; these may have arisen from a singlecommon ancestorthat lived650 million years agoin thePrecambrian.25 of these are novel core gene groups, found only in animals; of those, 8 are for essential components of theWntandTGF-betasignalling pathways which may have enabled animals to become multicellular by providing a pattern for the body's system of axes (in three dimensions), and another 7 are fortranscription factorsincludinghomeodomainproteins involved in thecontrol of development.[124][125]

Giribet and Edgecombe (2020) provide what they consider to be a consensus internal phylogeny of the animals, embodying uncertainty about the structure at the base of the tree (dashed lines).[126]

Animalia
multicellular

An alternative phylogeny, from Kapli and colleagues (2021), proposes a cladeXenambulacrariafor the Xenacoelamorpha + Ambulacraria; this is either within Deuterostomia, as sister to Chordata, or the Deuterostomia are recovered as paraphyletic, and Xenambulacraria is sister to the proposed cladeCentroneuralia,consisting of Chordata + Protostomia.[127]

Non-bilateria

Non-bilaterians include sponges (centre) and corals (background).

Several animal phyla lack bilateral symmetry. These are thePorifera(sea sponges),Placozoa,Cnidaria(which includesjellyfish,sea anemones,and corals), andCtenophora(comb jellies).

Sponges are physically very distinct from other animals, and were long thought to have diverged first, representing the oldest animal phylum and forming asister cladeto all other animals.[128]Despite their morphological dissimilarity with all other animals, genetic evidence suggests sponges may be more closely related to other animals than the comb jellies are.[129][130]Sponges lack the complex organization found in most other animal phyla;[131]their cells are differentiated, but in most cases not organised into distinct tissues, unlike all other animals.[132]They typically feed by drawing in water through pores, filtering out small particles of food.[133]

The comb jellies and Cnidaria are radially symmetric and have digestive chambers with a single opening, which serves as both mouth and anus.[134]Animals in both phyla have distinct tissues, but these are not organised into discreteorgans.[135]They arediploblastic,having only two main germ layers, ectoderm and endoderm.[136]

The tiny placozoans have no permanent digestive chamber and no symmetry; they superficially resemble amoebae.[137][138]Their phylogeny is poorly defined, and under active research.[129][139]

Bilateria

Main articles:BilateriaandSymmetry (biology) § Bilateral symmetry
Idealisedbilaterianbody plan.[d]With an elongated body and a direction of movement the animal has head and tail ends. Sense organs and mouth form thebasis of the head.Opposed circular and longitudinal muscles enableperistaltic motion.

The remaining animals, the great majority—comprising some 29 phyla and over a million species—form aclade,the Bilateria, which have a bilaterally symmetricbody plan.The Bilateria aretriploblastic,with three well-developed germ layers, and their tissuesform distinct organs.The digestive chamber has two openings, a mouth and an anus, and there is an internal body cavity, acoelomor pseudocoelom. These animals have a head end (anterior) and a tail end (posterior), a back (dorsal) surface and a belly (ventral) surface, and a left and a right side.[140][141]

Having a front end means that this part of the body encounters stimuli, such as food, favouringcephalisation,the development of a head withsense organsand a mouth. Many bilaterians have a combination of circularmusclesthat constrict the body, making it longer, and an opposing set of longitudinal muscles, that shorten the body;[141]these enable soft-bodied animals with ahydrostatic skeletonto move byperistalsis.[142]They also have a gut that extends through the basically cylindrical body from mouth to anus. Many bilaterian phyla have primarylarvaewhich swim withciliaand have an apical organ containing sensory cells. However, over evolutionary time, descendant spaces have evolved which have lost one or more of each of these characteristics. For example, adult echinoderms are radially symmetric (unlike their larvae), while someparasitic wormshave extremely simplified body structures.[140][141]

Genetic studies have considerably changed zoologists' understanding of the relationships within the Bilateria. Most appear to belong to two major lineages, theprotostomesand thedeuterostomes.[143]It is often suggested that the basalmost bilaterians are theXenacoelomorpha,with all other bilaterians belonging to the subcladeNephrozoa[144][145][146]However, this suggestion has been contested, with other studies finding that xenacoelomorphs are more closely related to Ambulacraria than to other bilaterians.[127]

Protostomes and deuterostomes

Further information:Embryological origins of the mouth and anus
Main articles:ProtostomeandDeuterostome
The bilaterian gut develops in two ways. In manyprotostomes,the blastopore develops into the mouth, while indeuterostomesit becomes the anus.

Protostomes and deuterostomes differ in several ways. Early in development, deuterostome embryos undergo radialcleavageduring cell division, while many protostomes (theSpiralia) undergo spiral cleavage.[147] Animals from both groups possess a complete digestive tract, but in protostomes the first opening of theembryonic gutdevelops into the mouth, and the anus forms secondarily. In deuterostomes, the anus forms first while the mouth develops secondarily.[148][149]Most protostomes haveschizocoelous development,where cells simply fill in the interior of the gastrula to form the mesoderm. In deuterostomes, the mesoderm forms byenterocoelic pouching,through invagination of the endoderm.[150]

The main deuterostome phyla are the Echinodermata and the Chordata.[151]Echinoderms are exclusively marine and includestarfish,sea urchins,andsea cucumbers.[152]The chordates are dominated by thevertebrates(animals withbackbones),[153]which consist offishes,amphibians,reptiles,birds,andmammals.[154]The deuterostomes also include theHemichordata(acorn worms).[155][156]

Ecdysozoa
Main article:Ecdysozoa
Ecdysis:adragonflyhas emerged from its dryexuviaeand is expanding its wings. Like otherarthropods,its body isdivided into segments.

The Ecdysozoa are protostomes, named after their sharedtraitofecdysis,growth by moulting.[157]They include the largest animal phylum, theArthropoda,which contains insects, spiders, crabs, and their kin. All of these have a body divided intorepeating segments,typically with paired appendages. Two smaller phyla, theOnychophoraandTardigrada,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;[158]some are important parasites.[159]Smaller phyla related to them are theNematomorphaor horsehair worms, and theKinorhyncha,Priapulida,andLoricifera.These groups have a reduced coelom, called a pseudocoelom.[160]

Spiralia
Main article:Spiralia
Spiral cleavagein a sea snail embryo

The Spiralia are a large group of protostomes that develop by spiral cleavage in the early embryo.[161]The Spiralia's phylogeny has been disputed, but it contains a large clade, the superphylumLophotrochozoa,and smaller groups of phyla such as theRouphozoawhich includes thegastrotrichsand theflatworms.All of these are grouped as thePlatytrochozoa,which has a sister group, theGnathifera,which includes therotifers.[162][163]

The Lophotrochozoa includes themolluscs,annelids,brachiopods,nemerteans,bryozoaandentoprocts.[162][164][165]The molluscs, the second-largest animal phylum by number of described species, includessnails,clams,andsquids,while the annelids are the segmented worms, such asearthworms,lugworms,andleeches.These two groups have long been considered close relatives because they sharetrochophorelarvae.[166][167]

History of classification

Further information:Taxonomy (biology),History of zoology through 1859,andHistory of zoology since 1859
Jean-Baptiste de Lamarckled the creation of a modern classification ofinvertebrates,breaking up Linnaeus's "Vermes" into 9 phyla by 1809.[168]

In theclassical era,Aristotledivided animals,[e]based on his own observations, into those with blood (roughly, the vertebrates) and those without. The animals were thenarranged on a scalefrom 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).Aristotlewas 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.[169]

In 1758,Carl Linnaeuscreated the firsthierarchicalclassification in hisSystema Naturae.[170]In his original scheme, the animals were one of three kingdoms, divided into the classes ofVermes,Insecta,Pisces,Amphibia,Aves,andMammalia.Since then the last four have all been subsumed into a single phylum, theChordata,while his Insecta (which included the crustaceans and arachnids) and Vermes have been renamed or broken up. The process was begun in 1793 byJean-Baptiste de Lamarck,who called the Vermesune espèce de chaos(a chaotic mess)[f]and split the group into three new phyla: worms, echinoderms, and polyps (which contained corals and jellyfish). By 1809, in hisPhilosophie Zoologique,Lamarck had created 9 phyla apart from vertebrates (where he still had 4 phyla: mammals, birds, reptiles, and fish) and molluscs, namelycirripedes,annelids, crustaceans, arachnids, insects, worms,radiates,polyps, andinfusorians.[168]

In his 1817Le Règne Animal,Georges Cuvierusedcomparative anatomyto group the animals into fourembranchements( "branches" with different body plans, roughly corresponding to phyla), namely vertebrates, molluscs, articulated animals (arthropods and annelids), andzoophytes (radiata)(echinoderms, cnidaria and other forms).[172]This division into four was followed by the embryologistKarl Ernst von Baerin 1828, the zoologistLouis Agassizin 1857, and the comparative anatomistRichard Owenin 1860.[173]

In 1874,Ernst Haeckeldivided 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.[174][173]The protozoa were later moved to the former kingdomProtista,leaving only the Metazoa as a synonym of Animalia.[175]

In human culture

Practical uses

Main article:Animals in culture
Sides ofbeefin aslaughterhouse

The human population exploits a large number of other animal species for food, both ofdomesticatedlivestock species inanimal husbandryand, mainly at sea, by hunting wild species.[176][177]Marine fish of many species arecaught commerciallyfor food. A smaller number of species arefarmed commercially.[176][178][179]Humans and theirlivestockmake up more than 90% of the biomass of all terrestrial vertebrates, and almost as much as all insects combined.[180]

Invertebratesincludingcephalopods,crustaceans,andbivalveorgastropodmolluscs are hunted or farmed for food.[181]Chickens,cattle,sheep,pigs,and other animals are raised as livestock for meat across the world.[177][182][183]Animal fibres such as wool are used to make textiles, while animalsinewshave been used as lashings and bindings, and leather is widely used to make shoes and other items. Animals have been hunted and farmed for their fur to make items such as coats and hats.[184]Dyestuffs includingcarmine(cochineal),[185][186]shellac,[187][188]andkermes[189][190]have been made from the bodies of insects.Working animalsincluding cattle and horses have been used for work and transport from the first days of agriculture.[191]

Animals such as the fruit flyDrosophila melanogasterserve a major role in science asexperimental models.[192][193][194][195]Animals have been used to createvaccinessince their discovery in the 18th century.[196]Some medicines such as the cancer drugtrabectedinare based ontoxinsor other molecules of animal origin.[197]

Agun dogretrieving a duck during a hunt

People have usedhunting dogsto help chase down and retrieve animals,[198]andbirds of preyto catch birds and mammals,[199]while tetheredcormorantshave beenused to catch fish.[200]Poison dart frogshave been used to poison the tips ofblowpipe darts.[201][202] A wide variety of animals are kept as pets, from invertebrates such as tarantulas, octopuses, andpraying mantises,[203]reptiles such assnakesandchameleons,[204]and birds includingcanaries,parakeets,andparrots[205]all finding a place. However, the most kept pet species are mammals, namelydogs,cats,andrabbits.[206][207][208]There is a tension between the role of animals as companions to humans, and their existence asindividuals with rightsof their own.[209]

A wide variety of terrestrial and aquatic animals are huntedfor sport.[210]

Symbolic uses

Thesigns of the WesternandChinese zodiacsare based on animals.[211][212]In China and Japan, thebutterflyhas been seen as thepersonificationof a person'ssoul,[213]and in classical representation the butterfly is also the symbol of the soul.[214][215]

Artistic vision:Still LifewithLobsterandOystersbyAlexander Coosemans,c. 1660

Animals have been thesubjects of artfrom the earliest times, both historical, as in ancient Egypt, and prehistoric, as in thecave paintings at Lascaux.Major animal paintings includeAlbrecht Dürer's 1515The Rhinoceros,andGeorge Stubbs'sc. 1762horse portraitWhistlejacket.[216]Insects,birds and mammals play roles in literature and film,[217]such as ingiant bug movies.[218][219][220]

Animals includinginsects[213]and mammals[221]feature in mythology and religion. Thescarab beetlewas sacred inancient Egypt,[222]and thecow is sacred in Hinduism.[223]Among other mammals,deer,[221]horses,[224]lions,[225]bats,[226]bears,[227]andwolves[228]are the subjects of myths and worship.

See also

Notes

  1. ^Henneguya zschokkeidoes not have mitochondrial DNA or utilize aerobic respiration.[19]
  2. ^The application ofDNA barcodingto taxonomy further complicates this; a 2016 barcoding analysis estimated a total count of nearly 100,000insectspecies forCanadaalone, 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).[75]
  3. ^Not includingparasitoids.[71]
  4. ^CompareFile:Annelid redone w white background.svgfor a more specific and detailed model of a particular phylum with this general body plan.
  5. ^In hisHistory of AnimalsandParts of Animals.
  6. ^The French prefixune espèce deis pejorative.[171]

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