Panarthropoda is a proposed

animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

containing the extant phyla

Arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...

Tardigrada Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbä ...

(water bears) and

Onychophora Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...

(velvet worms). Panarthropods also include

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military * ...

legged worms known as lobopodians ("

Lobopodia The lobopodians, members of the informal group Lobopodia (from the Greek language, Greek, meaning "blunt feet"), or the formally erected phylum Lobopoda Cavalier-Smith (1998), are panarthropods with stubby legs called lobopods, a term which may ...

"), a

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

group where the

last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...

and basal members (

stem-group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

) of each extant panarthropod phylum are thought to have risen. However 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 The ventral nerve cord is a major structure of the invertebrate central nervous system. It is the functional equivalent of the vertebrate spinal cord. The ventral nerve cord coordinates neural signaling from the brain to the body and vice versa, in ...

, and the presence of paired

appendages An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including anten ...

correlated with body segments.

Taxonomy

Not all studies support the

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...

of Panarthropoda, but most do, including neuroanatomical,

phylogenomic Phylogenomics is the intersection of the fields of evolution and genomics. The term has been used in multiple ways to refer to analysis that involves genome data and evolutionary reconstructions. It is a group of techniques within the larger fields ...

and

palaeontological Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (geology), epoch (roughly 11,700 years before present). It includes t ...

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

nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...

s. Traditionally, panarthropods were considered to be closely related to the

annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...

s, grouped together as the Articulata (animals with body segments), but subsequent

studies consistenly place them closer to

cycloneuralia Cycloneuralia is a clade of ecdysozoan animals including the Scalidophora (Kinorhynchans, Loriciferans, Priapulids) and the Nematoida ( nematodes, Nematomorphs). It may be paraphyletic, or may be a sister group to Panarthropoda. Or perhaps P ...

ns (nematodes, nematomorphs,

loricifera Loricifera (from Latin, '' lorica'', corselet (armour) + ''ferre'', to bear) is a phylum of very small to microscopic marine cycloneuralian sediment-dwelling animals that had been determined to be 37 described species, in 9 genera, bu ...

ns,

kinorhyncha Kinorhyncha ( grc, κινέω, kīnéō, I move, ' "snout") is a phylum of small marine invertebrates that are widespread in mud or sand at all depths as part of the meiobenthos. They are also called mud dragons. Modern species are or less, ...

s &

priapulid Priapulida (priapulid worms, from Gr. πριάπος, ''priāpos'' 'Priapus' + Lat. ''-ul-'', diminutive), sometimes referred to as penis worms, is a phylum of unsegmented marine worms. The name of the phylum relates to the Greek god of fertility ...

s), grouped together as

Ecdysozoa Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerata, crustaceans, and myriapods), Nematoda, and several smaller phyla. They were first defined by Aguinaldo ''et al.'' in 1997, based mainly on phylogenetic tr ...

. While annelids are placed among the

Spiralia The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other taxa. The term ''Spiralia'' is applied to those phyla that exhibit canonical spiral cleavage, a ...

(making them closer related to

mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is e ...

flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegment ...

s and such), having evolved their segmented bodies convergently.

Interrelationship

There are three competing hyphothesis for the interrelationship between the extant panarthropod phyla, each known as Tactopoda (Arthropoda+Tardigrada), Antennopoda (Arthropoda+Onychophora), and the

sister A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to refer to ...

relationship between Onychophora and Tardigrada (

''sensu'' Smith & Goldstein 2017). Tactopoda had been supported by

mitochondrial gene Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...

arrangements, palaeontological and neuroanatomical evidences, specifically the presence of segmented

ganglia A ganglion is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system there are both sympatheti ...

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 The lobopodians, members of the informal group Lobopodia (from the Greek language, Greek, meaning "blunt feet"), or the formally erected phylum Lobopoda Cavalier-Smith (1998), are panarthropods with stubby legs called lobopods, a term which may ...

traits (e.g. soft cuticle, lobopodous appendages and peripheral nerve roots), but these were generally considered to be

plesiomorphies In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

traced back to the

of Panarthropoda or

. While most phylogenomic analysis 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. File:Britannica 1911 Arthropoda - Peripatus head.png, Anatomy of

onychophoran Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...

anterior region, showing specialized appendages (ant, md) and deutocerebrum (D). File:Tardigrade zenuw.jpg,

Nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...

of a

tardigrade Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbär ...

, showing ventral nerve cord with segmented ganglia (ga1-4). File:20200213 Remipedia Nectiopoda central nervous system.png,

of a

remipede Remipedia is a class of blind crustaceans found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Sea, and the Atlantic Ocean. The ...

arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...

, showing the presence of both deutocerebrum (dc) and ventral nerve cord (vnc) organized by segmented ganglia. Within extinct lobopodians, at least ''

Antennacanthopodia ''Antennacanthopodia'' is a rare unarmoured lobopodian from the Chengjiang biota, with prickly legs, a pair of 'antennae', and an onychophora Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms ...

'' are widely accepted as part of the onychophoran stem-group. On the other hand, siberiids (''

Siberion ''Siberion'' is an extinct genus of lobopodian from the Sinsk biota of Russia. Its anatomy, including the proboscis-like organ projecting from the face and prominent grasping first pair of appendages, suggests that xenusians like this organism ma ...

'', ''

Megadictyon ''Megadictyon'' is a genus of Cambrian lobopodian with similarities to ''Jianshanopodia'' and '' Siberion''. Occasionally mis-spelt ''Magadictyon''. Megadictyon is a large lobopodian, with body length (excluding appendages) possibly up to 20 c ...

'' and ''

Jianshanopodia ''Jianshanopodia decora'' is a Cambrian lobopodian. Its frontal, grasping appendages bear wedge-shaped plates. Its limbs branch, instead of being tipped with claws as many lobopods' are. It has a sediment-filled gut surrounded by serially repea ...

'') and gilled lobopodians (''

Pambdelurion ''Pambdelurion'' is an extinct genus of panarthropod from the Cambrian aged Sirius Passet site in northern Greenland. Like the morphologically similar ''Kerygmachela'' from the same locality, ''Pambdelurion'' is thought to be closely related to ar ...

'' and ''

Kerygmachela ''Kerygmachela kierkegaardi'' is a gilled lobopodian from the Cambrian Stage 3 aged Sirius Passet Lagerstätte in northern Greenland. Its anatomy strongly suggests that it, along with its relative '' Pambdelurion whittingtoni'', was a close rela ...

'') represent transitional forms between typical lobopodians and basal arthropods (e.g. ''

Opabinia ''Opabinia regalis'' is an extinct, stem group arthropod found in the Middle Cambrian Burgess Shale Lagerstätte (505 million years ago) of British Columbia. ''Opabinia'' was a soft-bodied animal, measuring up to 7 cm in body length, and it ...

'' and

Radiodonta Radiodonta is an extinct Order (biology), order of stem-group arthropods that was successful worldwide during the Cambrian period. They may be referred to as radiodonts, radiodontans, radiodontids, anomalocarids, or anomalocaridids, although the ...

). The position of most other lobopodians (e.g. ''

Hallucigenia ''Hallucigenia'' is a genus of Cambrian animal resembling worms, known from articulated fossils in Burgess Shale-type deposits in Canada and China, and from isolated spines around the world. The generic name reflects the type species' unusual ...

'' and luolishaniids as stem onychophorans or stem panarthropods), including the lobopodian members of tardigrade stem-group (represented by ''

Onychodictyon ''Onychodictyon'' is a genus of extinct lobopodian known from the Lower Cambrian Chengjiang Maotianshan Shales in the Yunnan Province in China. It was characterized by a stout body covered by fleshy papillae and pairs of sclerotized plates with s ...

ferox'' or ''

Aysheaia ''Aysheaia'' is an extinct genus of soft-bodied lobopod, known from the middle Cambrian of North America, with an average body length of 1–6 cm. Anatomy ''Aysheaia'' has ten body segments, each of which has a pair of spiked, annulate le ...

'') are more controversial. '' Sialomorpha'', a genus of microinvertebrate discovered in

Dominican amber Dominican amber is amber from the Dominican Republic derived from resin of the extinct tree '' Hymenaea protera''. Dominican amber differentiates itself from Baltic amber by being nearly always transparent, and it has a higher number of fossil inc ...

in 2019, also considered to be a panarthropod. However, due to the unusual combination of tardigrade and

mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...

-like characters, its exact placement is uncertain.

References

{{Taxonbar, from=Q134870 Extant Cambrian first appearances Ecdysozoa taxa Protostome unranked clades

Taxonomy

Interrelationship

See also

References