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Panarthropoda

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46-408: Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda , Tardigrada (water bears) and Onychophora (velvet worms). Panarthropods also include extinct marine legged worms known as lobopodians (" Lobopodia "), a paraphyletic group where the last common ancestor and basal members ( stem-group ) of each extant panarthropod phylum are thought to have risen. However

92-399: 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. Repeated instances of mating with a close relative during sexual reproduction generally leads to inbreeding depression within

138-561: A clade Xenambulacraria for 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 clade Centroneuralia , consisting of Chordata + Protostomia. Eumetazoa , a clade which contains Ctenophora and ParaHoxozoa , has been proposed as a sister group to Porifera . A competing hypothesis

184-1109: A population due to the increased prevalence of harmful recessive traits. Animals have evolved numerous mechanisms for avoiding close inbreeding . Some animals are capable of asexual reproduction , which often results in a genetic clone of the parent. This may take place through fragmentation ; budding , such as in Hydra and other cnidarians ; or parthenogenesis , where fertile eggs are produced without mating , such as in aphids . Animals are categorised into ecological groups depending on their trophic levels and how they consume organic material . Such groupings include carnivores (further divided into subcategories such as piscivores , insectivores , ovivores , etc.), herbivores (subcategorized into folivores , graminivores , frugivores , granivores , nectarivores , algivores , etc.), omnivores , fungivores , scavengers / detritivores , and parasites . Interactions between animals of each biome form complex food webs within that ecosystem . In carnivorous or omnivorous species, predation

230-464: Is a consumer–resource interaction where the predator feeds on another organism, its prey , who often evolves anti-predator adaptations to avoid being fed upon. Selective pressures imposed on one another lead to an evolutionary arms race between predator and prey, resulting in various antagonistic/ competitive coevolutions . Almost all multicellular predators are animals. Some consumers use multiple methods; for example, in parasitoid wasps ,

276-433: Is a stage in embryonic development that is unique to animals, allowing cells to be differentiated into specialised tissues and organs. 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

322-463: Is disputed, as they might be water-escape or other structures. Animals are monophyletic , meaning they are derived from a common ancestor. Animals are the sister group to the choanoflagellates , with which they form the Choanozoa . The dates on the phylogenetic tree indicate approximately how many millions of years ago ( mya ) the lineages split. Ros-Rocher and colleagues (2021) trace

368-450: Is known as zoology , and the study of animal behaviour is known as ethology . The animal kingdom is divided into five infrakingdoms / superphyla , namely Porifera , Ctenophora , Placozoa , Cnidaria and Bilateria . Most living animal species belong to the infrakingdom Bilateria, a highly proliferative clade whose members have a bilaterally symmetric body plan , and the vast majority of bilaterians belong to two large superphyla :

414-808: Is often used to refer only to nonhuman animals. The term metazoa is derived from Ancient Greek μετα ( meta ) 'after' (in biology, the prefix meta- stands for 'later') and ζῷᾰ ( zōia ) 'animals', plural of ζῷον zōion 'animal'. Animals have several characteristics that set them apart from other living things. Animals are eukaryotic and multicellular . Unlike plants and algae , which produce their own nutrients , animals are heterotrophic , feeding on organic material and digesting it internally. With very few exceptions, animals respire aerobically . All animals are motile (able to spontaneously move their bodies) during at least part of their life cycle , but some animals, such as sponges , corals , mussels , and barnacles , later become sessile . The blastula

460-422: Is only produced by sponges and pelagophyte algae. Its likely origin is from sponges based on molecular clock estimates for the origin of 24-ipc production in both groups. Analyses of pelagophyte algae consistently recover a Phanerozoic origin, while analyses of sponges recover a Neoproterozoic origin, consistent with the appearance of 24-ipc in the fossil record. The first body fossils of animals appear in

506-547: Is the Benthozoa clade, which would consist of Porifera and ParaHoxozoa as a sister group of Ctenophora . Several animal phyla lack bilateral symmetry. These are the Porifera (sea sponges), Placozoa , Cnidaria (which includes jellyfish , sea anemones , and corals), and Ctenophora (comb jellies). Sponges are physically very distinct from other animals, and were long thought to have diverged first, representing

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552-622: The Cambrian explosion , which began around 539  million years ago (Mya), and most classes during the Ordovician radiation 485.4 Mya. 6,331 groups of genes common to all living animals have been identified; these may have arisen from a single common ancestor that lived about 650 Mya during the Cryogenian period. Historically, Aristotle divided animals into those with blood and those without . Carl Linnaeus created

598-619: The Ediacaran , represented by forms such as Charnia and Spriggina . It had long been doubted whether these fossils truly represented animals, but the discovery of the animal lipid cholesterol in fossils of Dickinsonia establishes their nature. 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. Many animal phyla first appear in

644-485: The Wnt and TGF-beta signalling 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 for transcription factors including homeodomain proteins involved in the control of development . Giribet and Edgecombe (2020) provide what they consider to be a consensus internal phylogeny of the animals, embodying uncertainty about

690-1042: The annelids , grouped together as the Articulata (animals with body segments), but subsequent phylogenomic studies consistently place them closer to cycloneuralians (nematodes, nematomorphs , loriciferans , kinorhynchas & priapulids ), grouped together as Ecdysozoa . While annelids are placed among the Spiralia (making them more closely related to mollusks , flatworms and such), having evolved their segmented bodies convergently . (Lobopodian taxa controversial)* (Lobopodian taxa controversial)* Antennacanthopodia [REDACTED] * Crown-group Onychophora [REDACTED] (Lobopodian taxa controversial)* Crown-group Tardigrada [REDACTED] Siberiid lobopodians [REDACTED] * Gilled lobopodians [REDACTED] * Opabinia [REDACTED] Radiodonta [REDACTED] Euarthropoda [REDACTED] There are three competing hyphotheses for

736-893: The biological kingdom Animalia ( / ˌ æ n ɪ ˈ m eɪ l i ə / ). With few exceptions, animals consume organic material , breathe oxygen , have myocytes and are able to move , can reproduce sexually , and grow from a hollow sphere of cells, the blastula , during embryonic development . Animals form a clade , meaning that they arose from a single common ancestor . Over 1.5 million living animal species have been described , of which around 1.05 million are insects , over 85,000 are molluscs , and around 65,000 are vertebrates . 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 complex ecologies and interactions with each other and their environments, forming intricate food webs . The scientific study of animals

782-665: The evolutionary relationships between taxa . Humans make use of many other animal species for food (including meat , eggs , and dairy products ), for materials (such as leather , fur , and wool ), as pets and as working animals for transportation , and services . Dogs , the first domesticated animal, have been used in hunting , in security and in warfare , as have horses , pigeons and birds of prey ; while other terrestrial and aquatic animals are hunted for sports, trophies or profits. Non-human animals are also an important cultural element of human evolution , having appeared in cave arts and totems since

828-473: The fossil record during the Cambrian explosion , starting about 539 million years ago, in beds such as the Burgess shale . Extant phyla in these rocks include molluscs , brachiopods , onychophorans , tardigrades , arthropods , echinoderms and hemichordates , along with numerous now-extinct forms such as the predatory Anomalocaris . The apparent suddenness of the event may however be an artifact of

874-510: The last common ancestor of Panarthropoda or Ecdysozoa . While most phylogenomic analyses support the monophyly of Panarthropoda, the results of interrelationship between the three phyla are less correlated - some of them inconsistently placing Tardigrada within Arthropoda, while the others mostly recovering either Antennopoda or Onychophora+Tardigrada. Within extinct lobopodians, at least Antennacanthopodia are widely accepted as part of

920-497: The lobe-finned fish Tiktaalik 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, with faunas adapted to salt water, hydrothermal vents, fresh water, hot springs, swamps, forests, pastures, deserts, air, and the interiors of other organisms. Animals are however not particularly heat tolerant ; very few of them can survive at constant temperatures above 50 °C (122 °F) or in

966-429: The monophyly of Panarthropoda, but most do, including neuroanatomical , phylogenomic and palaeontological studies. At least a close relationship between onychophorans and arthropods is widely agreed upon, but the position of tardigrades is more controversial. Some phylogenomic studies have found tardigrades to be more closely related to nematodes . Traditionally, panarthropods were considered to be closely related to

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1012-500: The protostomes , which includes organisms such as arthropods , molluscs , flatworms , annelids and nematodes ; and the deuterostomes , which include echinoderms , hemichordates and chordates , the latter of which contains the vertebrates . The much smaller basal phylum Xenacoelomorpha have an uncertain position within Bilateria. Animals first appear in the fossil record in the late Cryogenian period and diversified in

1058-715: The Cambrian explosion, possibly as early as 1 billion years ago. Early fossils that might represent animals appear for example in the 665-million-year-old rocks of the Trezona Formation of South Australia . These fossils are interpreted as most probably being early sponges . Trace fossils such as tracks and burrows found in the Tonian period (from 1 gya) 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 by

1104-717: 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. The following table lists estimated numbers of described extant species for the major animal phyla, 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

1150-463: The dark sea floor consume organic matter produced through chemosynthesis (via oxidizing inorganic compounds such as hydrogen sulfide ) by archaea and bacteria . Animals evolved in the sea. Lineages of arthropods colonised land around the same time as land plants , probably between 510 and 471 million years ago during the Late Cambrian or Early Ordovician . Vertebrates such as

1196-454: The earliest times, and are frequently featured in mythology , religion , arts , literature , heraldry , politics , and sports . The word animal comes from the Latin noun animal of the same meaning, which is itself derived from Latin animalis 'having breath or soul'. The biological definition includes all members of the kingdom Animalia. In colloquial usage, the term animal

1242-570: The first hierarchical biological classification for animals in 1758 with his Systema Naturae , which Jean-Baptiste Lamarck expanded into 14 phyla by 1809. In 1874, Ernst Haeckel divided the animal kingdom into the multicellular Metazoa (now synonymous with Animalia) and the Protozoa , single-celled organisms no longer considered animals. In modern times, the biological classification of animals relies on advanced techniques, such as molecular phylogenetics , which are effective at demonstrating

1288-448: 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,

1334-520: The fossil record, rather than showing that all these animals appeared simultaneously. That view is supported by the discovery of Auroralumina attenboroughii , the earliest known Ediacaran crown-group cnidarian (557–562 mya, some 20 million years before the Cambrian explosion) from Charnwood Forest , England. It is thought to be one of the earliest predators , catching small prey with its nematocysts as modern cnidarians do. Some palaeontologists have suggested that animals appeared much earlier than

1380-737: The giant single-celled protist Gromia sphaerica , so the Tonian trace fossils may not indicate early animal evolution. Around the same time, the layered mats of microorganisms called stromatolites decreased in diversity, perhaps due to grazing by newly evolved animals. 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

1426-534: The interrelationship between the extant panarthropod phyla, each known as Tactopoda (Arthropoda+Tardigrada), Antennopoda (Arthropoda+Onychophora), and the sister relationship between Onychophora and Tardigrada ( Lobopodia sensu Smith & Goldstein 2017). Onychophora Tardigrada Arthropoda Tardigrada Onychophora Arthropoda Arthropoda Tardigrada Onychophora Tactopoda had been supported by mitochondrial gene arrangements, palaeontological and neuroanatomical evidences, specifically

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1472-441: The larvae feed on the hosts' living tissues, killing them in the process, but the adults primarily consume nectar from flowers. Other animals may have very specific feeding behaviours , such as hawksbill sea turtles which mainly eat sponges . Most animals rely on biomass and bioenergy produced by plants and phytoplanktons (collectively called producers ) through photosynthesis . Herbivores, as primary consumers , eat

1518-513: The lobopodian members of tardigrade stem-group (represented by Onychodictyon ferox or Aysheaia ) are more controversial. Sialomorpha , a genus of microinvertebrate discovered in Dominican amber in 2019, is also considered to be a panarthropod. However, due to the unusual combination of tardigrade and mite -like characteristics, its exact placement is uncertain. Animal Animals are multicellular , eukaryotic organisms in

1564-753: The most extreme cold deserts of continental Antarctica . The blue whale ( Balaenoptera musculus ) is the largest animal that has ever lived, weighing up to 190 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 titanosaur sauropod dinosaurs such as Argentinosaurus , which may have weighed as much as 73 tonnes, and Supersaurus which may have reached 39 meters. Several animals are microscopic; some Myxozoa ( obligate parasites within

1610-620: The oldest animal phylum and forming a sister clade to all other animals. 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. Sponges lack the complex organization found in most other animal phyla; their cells are differentiated, but in most cases not organised into distinct tissues, unlike all other animals. They typically feed by drawing in water through pores, filtering out small particles of food. The comb jellies and Cnidaria are radially symmetric and have digestive chambers with

1656-434: The onychophoran stem-group. On the other hand, siberiids ( Siberion , Megadictyon and Jianshanopodia ) and gilled lobopodians ( Pambdelurion and Kerygmachela ) represent transitional forms between typical lobopodians and basal arthropods (e.g. Opabinia and Radiodonta ). The positions of most other lobopodians (e.g. Hallucigenia and luolishaniids as stem onychophorans or stem panarthropods), including

1702-525: The origins of animals to unicellular ancestors, providing the external phylogeny shown in the cladogram. Uncertainty of relationships is indicated with dashed lines. Holomycota (inc. fungi) [REDACTED] Ichthyosporea [REDACTED] Pluriformea [REDACTED] Filasterea [REDACTED] [REDACTED] [REDACTED] The most basal animals, the Porifera , Ctenophora , Cnidaria , and Placozoa , have body plans that lack bilateral symmetry . Their relationships are still disputed;

1748-519: The plant material directly to digest and absorb the nutrients, while carnivores and other animals on higher trophic levels indirectly acquire the nutrients by eating the herbivores or other animals that have eaten the herbivores. Animals oxidize carbohydrates , lipids , proteins and other biomolecules, which allows the animal to grow and to sustain basal metabolism and fuel other biological processes such as locomotion . Some benthic animals living close to hydrothermal vents and cold seeps on

1794-537: The presence of segmented ganglia shared by arthropods and tardigrades. Antennopodia united by the presence of specialized head appendages and deutocerebrum (additional second section of the brain), but subsequent anatomical studies suggest these features were convergently evolved between onychophoran and arthropod lineages. Onychophorans and tardigrades shared some lobopodian traits (e.g. soft cuticle, lobopodous appendages and peripheral nerve roots), but these were generally considered to be plesiomorphies traced back to

1840-753: The sister group to all other animals could be the Porifera or the Ctenophora, both of which lack hox genes , which are important for body plan development . Hox genes are found in the Placozoa, Cnidaria, and Bilateria. 6,331 groups of genes common to all living animals have been identified; these may have arisen from a single common ancestor that lived 650 million years ago in the Precambrian . 25 of these are novel core gene groups, found only in animals; of those, 8 are for essential components of

1886-407: 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

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1932-474: 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 haploid gametes by meiosis ;

1978-411: The structure at the base of the tree (dashed lines). Porifera [REDACTED] Ctenophora [REDACTED] Placozoa [REDACTED] Cnidaria [REDACTED] Xenacoelomorpha [REDACTED] Ambulacraria [REDACTED] Chordata [REDACTED] Ecdysozoa [REDACTED] Spiralia [REDACTED] An alternative phylogeny, from Kapli and colleagues (2021), proposes

2024-522: The subsequent Ediacaran period in what is known as the Avalon explosion . Earlier evidence of animals is still controversial; the sponge -like organism Otavia has been dated back to the Tonian period at the start of the Neoproterozoic , but its identity as an animal is heavily contested. Nearly all modern animal phyla became clearly established in the fossil record as marine species during

2070-543: The term "Lobopodia" is sometimes expanded to include tardigrades and onychophorans as well. Common characteristics of the Panarthropoda include a segmented body , paired ladder-like ventral nervous system , and the presence of paired appendages correlated with body segments. Echinodermata , Chordata and relatives Annelida , Mollusca and relatives Priapulida , Nematoda and relatives Onychophora , Tardigrada and Arthropoda Not all studies support

2116-467: 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. 3,000–6,500 4,000–25,000 Evidence of animals is found as long ago as the Cryogenian period. 24-Isopropylcholestane (24-ipc) has been found in rocks from roughly 650 million years ago; it

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