The Articulata is a proposed higher taxon of animals with segmented bodies, consisting of
Annelida
The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to variou ...
and
Panarthropoda
Panarthropoda is a clade comprising the greatest diversity of animal groups. It contains the extant phyla Arthropoda (Euarthropoda), Tardigrada (water bears) and Onychophora (velvet worms), although the precise relationships among these remained ...
. This theory states that these groups are descended from a common segmented ancestor. The Articulata hypothesis is an alternative to the hypothesis that
ecdysis
Ecdysis is the moulting of the cuticle in many invertebrates of the clade Ecdysozoa. Since the cuticle of these animals typically forms a largely inelastic exoskeleton, it is shed during growth and a new, larger covering is formed. The remnant ...
(the shedding of outer cuticle) is a primitive characteristic – this would place Panarthropoda in the group
Ecdysozoa
Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerates (including arachnids), crustaceans, and myriapods), Nematoda, and several smaller phylum (biology), phyla. The grouping of these animal phyla into a single ...
.
Origins
The Articulata hypothesis originates from the
phylogenetic
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analyses of
Georges Cuvier
Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...
in his 1817 published work ''
Le Règne animal, distribué après son organization''. In this work, Cuvier theorized that all
organisms
An organism is any living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have been pr ...
exist as a functional whole, meaning that all of the physiological structures of an organism are important for survival. By studying these physiological structures, Cuvier was able to group the known
animal
Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
kingdom according to structural similarities resulting from what he referred to as special "ground-plans", which are analogous to blueprints. Each of these ground-plans, he further argued, evolved separately from the others and structural similarities were due to common function and not to common ancestry. From these ground-plans, Cuvier separated the known animal kingdom into four branches or ':
Vertebrata
Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain.
The vertebrates make up the subphylum Verte ...
, Articulata,
Mollusca
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
and
Radiata
Radiata or Radiates is a historical taxonomic rank that was used to classify animals with Symmetry (biology)#Radial symmetry, radially symmetric body plans. The term Radiata is no longer accepted, as it united several different groupings of anim ...
. From this phylogenetic grouping, the Articulata hypothesis was born.
The Articulata hypothesis, simply stated, is the phylogenetic grouping of the phylum Annelida (which includes
polychaete
Polychaeta () is a paraphyletic class of generally marine Annelid, annelid worms, common name, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called c ...
s,
oligochaetes, and
leech
Leeches are segmented parasitism, parasitic or Predation, predatory worms that comprise the Class (biology), subclass Hirudinea within the phylum Annelida. They are closely related to the Oligochaeta, oligochaetes, which include the earthwor ...
es) together with the phylum Arthropoda (
arachnid
Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
s,
insect
Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s and
crustacean
Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s) into the common
taxon
In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
Articulata. Cuvier grouped these diverse phyla together according to the common structural feature: the segmented body plan. This hypothesis further implies that all segmented organisms have a common ancestral origin.
Development
Since its original formulation in 1817, there have been significant challenges and modifications to the Articulata hypothesis as new theories have been accepted (
Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
's
theory of evolution
Evolution is the change in the heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certai ...
) and new technologies have become available (
confocal microscopy
Confocal microscopy, most frequently confocal laser scanning microscopy (CLSM) or laser scanning confocal microscopy (LSCM), is an optical imaging technique for increasing optical resolution and contrast (vision), contrast of a micrograph by me ...
,
DNA sequencing
DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, thymine, cytosine, and guanine. The ...
,
genomics
Genomics is an interdisciplinary field of molecular biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, ...
). Additionally, the discovery of
Onychophora
Onychophora (from , , "claws"; and , , "to carry"), commonly known as velvet worms (for their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, ''Peripatus''), is a phylum of el ...
as its own phylum was incorporated into the theory by H. Bruce Bordreaux
Theory of evolution
Darwin's theory of evolution had a large, yet often understated impact of the Articulata hypothesis. Cuvier's original Articulata hypothesis was based on his assumption that current species no longer
evolved
Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
because to evolve would cause loss of integral structures necessary for the survival of the
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. While the general acceptance of the theory of evolution weakened Cuvier's general theory of the unique ground-plans as the origin of modern
taxa
In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
, it strengthened the Articulata hypothesis by organizing annelids and arthropods into a
clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
descended from a common segmented ancestor.
Confocal microscopy and modern molecular biology techniques

While each advance in modern molecular biology has shaken the
phylogenetic
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
tree of
Bilateria
Bilateria () is a large clade of animals characterised by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front (or "head") and a rear (or "tail") end, as well as a left� ...
, advances in
molecular biology
Molecular biology is a branch of biology that seeks to understand the molecule, molecular basis of biological activity in and between Cell (biology), cells, including biomolecule, biomolecular synthesis, modification, mechanisms, and interactio ...
techniques led to further data supporting the Articulata hypothesis but also led to the development of conflicting theories. Advances in
confocal microscopy
Confocal microscopy, most frequently confocal laser scanning microscopy (CLSM) or laser scanning confocal microscopy (LSCM), is an optical imaging technique for increasing optical resolution and contrast (vision), contrast of a micrograph by me ...
technology led to the discovery of
embryo
An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
nic
cleavage
Cleavage may refer to:
Science
* Cleavage (crystal), the way in which a crystal or mineral tends to split
* Cleavage (embryo), the division of cells in an early embryo
* Cleavage (geology), foliation of rock perpendicular to stress, a result of ...
patterns, which differs between the annelids and arthropods.
Annelids show spiral cleavage, meaning that each embryonic cleavage occurs at progressive 90-degree angles with respect to the animal–vegetal axis. Arthropods, on the other hand, display a heterogeneous mix of embryonic cleavage patterns including spiral-like cleavage and radial cleavage patterns. This led researchers to two theories: The first was that the arthropods lineage must have lost the ability to spiral cleave since differentiating from the last common ancestor between annelids and arthropods. The second is that this showed similarities between annelids and
mollusks
Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The num ...
who also spirally cleaves but lacks that the segmented body plan. This was not the only interpretation of this data but other hypotheses were seen to have less data or merit. Other studies such as those looking a neural patterns within the Articulata
clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
showed mixed patterns and thus mixed results.
DNA sequencing and genomics
The advancements in DNA sequencing techniques and the development of phylogenetic analysis algorithms led to the splitting of the Articulata clade. Original phylogenetic studies on the sequences of 18S and 28S
ribosomal DNA
The ribosomal DNA (rDNA) consists of a group of ribosomal RNA encoding genes and related regulatory elements, and is widespread in similar configuration in all domains of life. The ribosomal DNA encodes the non-coding ribosomal RNA, integral struc ...
sequence led to the suggestions that the annelids and arthropods had evolutionarily diverged much earlier than was previously thought but such limited genetic studies led to limited and often mixed results. As more genes were added to the studies, it became apparent that arthropods were genetically closer to
nematode
The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
s and other molting organisms whereas the annelids were closer evolutionary to mollusks. This
Ecdysozoa
Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerates (including arachnids), crustaceans, and myriapods), Nematoda, and several smaller phylum (biology), phyla. The grouping of these animal phyla into a single ...
hypothesis is generally accepted today as the best supported evolutionary hypothesis for annelids and arthropods.
However, a 2025 paper still suggests Articulata is monophyletic, alongside
Atelocerata, despite the vast majority of papers favouring Mandibulata and Ecdysozoa instead. While Ecdysozoa ''is'' recovered within the paper, cycloneuralians are instead found to be simplified lobopodians. Alongside this, Articulata is hypothesised to have descended from
spintherid-like crawling annelids.
See also
*
Proarticulata
Proarticulata is a proposed phylum of extinct, near-bilaterally symmetrical animals known from fossils found in the Ediacaran (Vendian) marine deposits, and dates to approximately . The name comes from the Greek () = "before" and Articulata, i. ...
References
{{Taxonbar, from=Q1526314
Phylogenetics