Articulata | |
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Extinct and modern arthropods | |
Glycera sp. (Annelida) | |
Scientific classification | |
Kingdom: | |
(unranked): | |
Superphylum: | Articulata H. Bruce Bordreaux, 1979 [1] |
Subordinate taxa | |
The Articulata hypothesis is the grouping in a higher taxon of animals with segmented bodies, consisting of Annelida and Panarthropoda. This theory states that these groups are descended from a common segmented ancestor. The Articulata hypothesis is an alternative to the hypothesis that ecdysis (the shedding of outer cuticle) is a primitive characteristic – this would place Panarthropoda in the group Ecdysozoa.
The Articulata hypothesis originates from the phylogenetic analyses of Georges Cuvier in his 1817 published work Le Règne animal, distribué après son organization . In this work, Cuvier theorized that all organisms 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 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 embranchements: Vertebrata, Articulata, Mollusca and Radiata. From this phylogenetic grouping, the Articulata hypothesis was born. [2]
The Articulata hypothesis, simply stated, is the phylogenetic grouping of the phylum Annelida (which includes polychaetes, oligochaetes, and leeches) together with the phylum Arthropoda (arachnids, insects and crustaceans) into the common taxon 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. [3]
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's theory of evolution) and new technologies have become available (confocal microscopy, DNA sequencing, genomics). Additionally, the discovery of Onychophora as its own phylum was incorporated into the theory by H. Bruce Bordreaux [1]
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 because to evolve would cause loss of integral structures necessary for the survival of the species. 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, it strengthened the Articulata hypothesis by organizing annelids and arthropods into a clade descended from a common segmented ancestor. [4]
While each advance in modern molecular biology has shaken the phylogenetic tree of Bilateria, advances in molecular biology techniques led to further data supporting the Articulata hypothesis but also led to the development of conflicting theories. Advances in confocal microscopy technology led to the discovery of embryonic cleavage patterns, which differs between the annelids and arthropods. [5] 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 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 showed mixed patterns and thus mixed results. [4] [6]
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 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 nematodes and other molting organisms whereas the annelids were closer evolutionary to mollusks. This Ecdysozoa hypothesis is generally accepted today as the best supported evolutionary hypothesis for annelids and arthropods. [7] [8] [9]
Invertebrates is an umbrella term describing animals that neither develop nor retain a vertebral column, which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum Vertebrata, i.e. vertebrates. Well-known phyla of invertebrates include arthropods, mollusks, annelids, echinoderms, flatworms, cnidarians, and sponges.
Onychophora, commonly known as velvet worms or more ambiguously as peripatus, is a phylum of elongate, soft-bodied, many-legged animals. In appearance they have variously been compared to worms with legs, caterpillars, and slugs. They prey upon other invertebrates, which they catch by ejecting an adhesive slime. Approximately 200 species of velvet worms have been described, although the true number of species is likely greater. The two extant families of velvet worms are Peripatidae and Peripatopsidae. They show a peculiar distribution, with the peripatids being predominantly equatorial and tropical, while the peripatopsids are all found south of the equator. It is the only phylum within Animalia that is wholly endemic to terrestrial environments, at least among extant members. Velvet worms are generally considered close relatives of the Arthropoda and Tardigrada, with which they form the proposed taxon Panarthropoda. This makes them of palaeontological interest, as they can help reconstruct the ancestral arthropod. Only two fossil species are confidently assigned as onychophorans: Antennipatus from the Late Carboniferous, and Cretoperipatus from the Late Cretaceous, the latter belonging to Peripatidae. In modern zoology, they are particularly renowned for their curious mating behaviours and the bearing of live young in some species.
Bilateria is a large clade or infrakingdom of animals called bilaterians, characterized by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front and a rear end, as well as a left–right–symmetrical belly (ventral) and back (dorsal) surface. Nearly all bilaterians maintain a bilaterally symmetrical body as adults; the most notable exception is the echinoderms, which extend to pentaradial symmetry as adults, but are only bilaterally symmetrical as an embryo. Cephalization is also a characteristic feature among most bilaterians, where the special sense organs and central nerve ganglia become concentrated at the front/rostral end.
In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of primates, the front flippers of whales, and the forelegs of four-legged vertebrates like dogs and crocodiles are all derived from the same ancestral tetrapod structure. Evolutionary biology explains homologous structures adapted to different purposes as the result of descent with modification from a common ancestor. The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843. Homology was later explained by Charles Darwin's theory of evolution in 1859, but had been observed before this, from Aristotle onwards, and it was explicitly analysed by Pierre Belon in 1555.
Ecdysozoa is a group of protostome animals, including Arthropoda, Nematoda, and several smaller phyla. The grouping of these animal phyla into a single clade was first proposed by Eernisse et al. (1992) based on a phylogenetic analysis of 141 morphological characters of ultrastructural and embryological phenotypes. This clade, that is, a group consisting of a common ancestor and all its descendants, was formally named by Aguinaldo et al. in 1997, based mainly on phylogenetic trees constructed using 18S ribosomal RNA genes.
Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada and Onychophora. 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 the term "Lobopodia" is sometimes expanded to include tardigrades and onychophorans as well.
Segmentation in biology is the division of some animal and plant body plans into a linear series of repetitive segments that may or may not be interconnected to each other. This article focuses on the segmentation of animal body plans, specifically using the examples of the taxa Arthropoda, Chordata, and Annelida. These three groups form segments by using a "growth zone" to direct and define the segments. While all three have a generally segmented body plan and use a growth zone, they use different mechanisms for generating this patterning. Even within these groups, different organisms have different mechanisms for segmenting the body. Segmentation of the body plan is important for allowing free movement and development of certain body parts. It also allows for regeneration in specific individuals.
Lophotrochozoa is a clade of protostome animals within the Spiralia. The taxon was established as a monophyletic group based on molecular evidence. The clade includes animals like annelids, molluscs, bryozoans, and brachiopods.
Cycloneuralia is a proposed clade of ecdysozoan animals including the Scalidophora, the Nematoida, and the extinct Palaeoscolecid. It may be paraphyletic, or may be a sister group to Panarthropoda. Or perhaps Panarthropoda is paraphyletic with respect to Cycloneuralia. The group has also been considered a single phylum, sometimes given the old name Nemathelminthes. The uniting character is the nervous system organization with a circumpharyngeal brain and somata–neuropil–somata pattern. The name derives from the position of the brain around the pharynx.
Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, the face of a human being has a plane of symmetry down its centre, or a pine cone displays a clear symmetrical spiral pattern. Internal features can also show symmetry, for example the tubes in the human body which are cylindrical and have several planes of symmetry.
A body plan, Bauplan, or ground plan is a set of morphological features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many.
The (pan)arthropod head problem is a long-standing zoological dispute concerning the segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other. While the dispute has historically centered on the exact make-up of the insect head, it has been widened to include other living arthropods, such as chelicerates, myriapods, and crustaceans, as well as fossil forms, such as the many arthropods known from exceptionally preserved Cambrian faunas. While the topic has classically been based on insect embryology, in recent years a great deal of developmental molecular data has become available. Dozens of more or less distinct solutions to the problem, dating back to at least 1897, have been published, including several in the 2000s.
Protostomia is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's members, although the reverse is typically true of its sister clade, Deuterostomia. Well-known examples of protostomes are arthropods, molluscs, annelids, flatworms and nematodes. They are also called schizocoelomates since schizocoely typically occurs in them.
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 pattern of early development found in most members of the Lophotrochozoa.
Nematoida is a clade of pseudocoelomate free living or parasitic animals. It consists of phyla Nematoda and Nematomorpha. The two groups share a number of features in common; the presence of a cloaca in both sexes, aflagellate sperm, and a cuticle made of collagen.
James A. Lake is an American evolutionary biologist and a Distinguished Professor of Molecular, Cell, and Developmental Biology and of Human Genetics at UCLA. Lake is best known for the New Animal Phylogeny and for the first three-dimensional structure of the ribosome. He has also made significant contributions to understanding genome evolution across all kingdoms of life, including discovering informational and operational genes, elucidating the complexity hypothesis for gene transfer, rooting the tree of life, and understanding the early transition from prokaryotic to eukaryotic life.
Tactopoda or Arthropodoidea is a proposed clade of ecdysozoan animals that includes the phyla Tardigrada and Euarthropoda, supported by various morphological observations. The cladogram below shows the relationships implied by this hypothesis.
The annelids, also known as the segmented worms, comprise a large phylum called Annelida. It contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecologies – some in marine environments as distinct as tidal zones and hydrothermal vents, others in fresh water, and yet others in moist terrestrial environments.
Maldanidae is a family of more than 200 species of marine polychaetes commonly known as bamboo worms or maldanid worms. They belong to the order Capitellida, in the phylum Annelida. They are most closely related to family Arenicolidae, and together form the clade Maldanomorpha.
Pleistoannelida is a group of annelid worms that comprises the vast majority of the diversity in phylum Annelida. Discovered through phylogenetic analyses, it is the largest clade of annelids, comprised by the last common ancestor of the highly diverse sister groups Errantia and Sedentaria and all the descendants of that ancestor. Most groups in the Clade find their ancestors within the Cambrian explosion when Annelid diversity expanded dramatically. The Pleistoannelida clade covers a variety of traits. However, the evolution of simple to complex eyes, developed papillae for burrowing, and for some specialized radioles for feeding can be seen universally across every species. New findings have discovered the range of Annelid diversity have led to uncertainty if groups with developed ancestral traits should remain within the clade. Furthermore There's been a lack of recently discovered Annelid traits being used in the categorization of groups within the clade, leading to many hypothesis on how to do so and which should remain within the clade. Currently three smaller clades that were originally a part of the groups Errantia and Sedentaria have been proven to fall outside while still being connected to the basal groups.