Tactopoda is a proposed clade of

protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryogenesis, embryonic development. This nature has since been discovered to be extremely variable among Pro ...

animals that includes the phyla

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 ...

and

Euarthropoda Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, oft ...

, supported by various morphological observations. The

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...

below shows the relationships implied by this hypothesis. The competing hypothesis is that

Arthropoda Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, ...

sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...

''J. Ortega-Hernández, "Making sense of 'lower' and 'upper' stem-group Euarthropoda, with comments on the strict use of the name Arthropoda von Siebold, 1848," Biol. Rev., vol. 91, no. 1, pp. 255–273, 2016. (=

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 ...

, the arthropods and the velvet worms) is monophyletic, and tardigrades lie outside this grouping. Anatomic arguments for the tactopoda monophyly include similarities in the anatomies of head, legs, and muscles between the arthropods and the tardigrades. Anatomic arguments against it include that tardigrades lack the kind of circulatory system (including a

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...

heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide to t ...

) which the arthropods and the velvet worms share.

Graham Budd Graham Edward Budd is a British palaeontologist. He is Professor and head of palaeobiology at Uppsala University. Budd's research focuses on the Cambrian explosion and on the evolution and development, anatomy, and patterns of diversificati ...

argued that the lack of this system in recent tardigrades is due to their miniature size, which makes a complex circulatory system superfluous; thus, the loss of this feature would be a secondary property, acquired as the tardigrade

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. ...

turned smaller, and both the Euarthropoda+Onychophora circulatory system and a relatively large size should be a feature of the last common ancestor of all three groups. However, Gregory Edgecombe also invoked

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 ...

evidence in favour of the alternative Euarthropoda+Onychophora grouping.

Etymology

Budd formed the suggested clade name 'tactopoda' from Greek ''taktos'', ordered, and ''poda'', feet, "with reference to the alleged well-formed stepping motion that characterises the group".

Proposed classification

*Phylum

**Class

Eutardigrada Eutardigrada are a class of tardigrades (Tardigrada) without lateral appendices. Primarily freshwater bound, some species have secondarily gained the ability to live in marine environments (''Halobiotus''). By cryptobiosis many species are able t ...

**Class Heterotardigrada **Class

Mesotardigrada Mesotardigrada is one of three classes of tardigrades, consisting of a single species, ''Thermozodium esakii''. The animal reportedly has six claws of equal length at each foot. This species was described in 1937 by German zoologist Gilbert Rah ...

*Phylum

**Class

Pycnogonida Sea spiders are marine arthropods of the order Pantopoda ( ‘all feet’), belonging to the class Pycnogonida, hence they are also called pycnogonids (; named after ''Pycnogonum'', the type genus; with the suffix '). They are cosmopolitan, foun ...

(?) **Clade

Arachnomorpha Arachnomorpha is a proposed subdivision or clade of Arthropoda, comprising the group formed by the trilobites and their close relatives (Artiopoda), Megacheira (which may be paraphyletic) and chelicerates. Under this proposed classification sch ...

***Class †

Trilobita Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...

***Order † Aglaspida ***Order †

Strabopida Strabopidae is the only family of the order Strabopida, a group of arachnomorph arthropods. The order Strabopida contains three valid genera and one doubtful, ''Khankaspis'', '' Paleomerus'', '' Parapaleomerus'' and '' Strabops'', all assigned in ...

***Class

(?) ***Order † Cheloniellida ***Subphylum

Chelicerata The subphylum Chelicerata (from New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks, and mi ...

****Class

Xiphosura Xiphosura () is an order of arthropods related to arachnids. They are more commonly known as horseshoe crabs (a name applied more specifically to the only extant family, Limulidae). They first appeared in the Hirnantian ( Late Ordovician). Curr ...

****Class †

Eurypterida Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 million years ago. The group is l ...

****Class † Chasmataspidida ****Class

(?) ****Class Arachnida **Clade

Mandibulata Mandibulata, termed "mandibulates", is a clade of arthropods that comprises the extant subphyla Myriapoda (millipedes and others), Crustacea and Hexapoda (insects and others). Mandibulata is currently believed to be the sister group of the clade ...

***Order † Euthycarcinoidea ***Subphylum

Myriapoda Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian ...

****Class

Chilopoda Centipedes (from New Latin , "hundred", and Latin , "foot") are predatory arthropods belonging to the class Chilopoda ( Ancient Greek , ''kheilos'', lip, and New Latin suffix , "foot", describing the forcipules) of the subphylum Myriapoda, a ...

****Class

Symphyla Symphylans, also known as garden centipedes or pseudocentipedes, are soil-dwelling arthropods of the class Symphyla in the subphylum Myriapoda. Symphylans resemble centipedes, but are very small, non-venomous, and only distantly related to both ...

****Class

Diplopoda Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...

****Class

Pauropoda Pauropods are small, pale, millipede-like arthropods. Around 830 species in twelve families are found worldwide, living in soil and leaf mold. They look rather like centipedes, or millipedes, and may be a sister group of the latter. However, this ...

***Clade Pancrustacea ****Clade Oligostraca *****Class

Ostracoda Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typical ...

*****Class Ichthyostraca ****Clade Altocrustacea *****Clade Multicrustacea ******Class

Hexanauplia The Hexanauplia is a clade proposed by Oakley et al. (2013) that constitutes a class of crustaceans, comprising the Copepoda and Thecostraca. A number of recent phylogenomic studies have not found support for this clade. Classification This t ...

******Class

Malacostraca Malacostraca (from New Latin; ) is the largest of the six classes of crustaceans, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crabs, l ...

******Class

Thecostraca Thecostraca is a Class (biology), class of marine invertebrates containing over 2,200 described species. Many species have planktonic larvae which become Sessility (zoology), sessile or parasite, parasitic as adults. The most important subgroup ...

*****Clade Allotriocarida ******Class

Branchiopoda Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca (or Cladocera), Notostraca and the Devonian ''Lepidocaris''. They are mostly small, freshwater animals that feed on plankton and detritus. Description ...

******Clade Miracrustacea *******Superclass

Xenocarida Xenocarida (from the Greek language, Greek for ''strange shrimp'') is a proposed clade inside the subphylum Crustacean, Crustacea that comprises two classes that were discovered in the 20th century: Remipedia and Cephalocarida. The clade was recov ...

********Class

Cephalocarida The Cephalocarida are a class in the subphylum Crustacea comprising only 12 benthic species. They were discovered in 1955 by Howard L. Sanders, and are commonly referred to as horseshoe shrimp. They have been grouped together with the Remipedi ...

********Class

Remipedia 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 ...

*******Subphylum

Hexapoda The subphylum Hexapoda (from Greek for 'six legs') comprises most species of arthropods and includes the insects as well as three much smaller groups of wingless arthropods: Collembola, Protura, and Diplura (all of these were once considered ins ...

******** Paraphyletic class

Entognatha The Entognatha are a class of wingless and ametabolous arthropods, which, together with the insects, makes up the subphylum Hexapoda. Their mouthparts are entognathous, meaning that they are retracted within the head, unlike the insects. Entog ...

********Clade Ectognatha *********Class

Insecta Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs o ...

Phylogeny

References

Ecdysozoa taxa Protostome unranked clades {{Protostome-stub