Spiralia Possible Ediacaran fossil | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Subkingdom: | Eumetazoa |
Clade: | ParaHoxozoa |
Clade: | Bilateria |
Clade: | Nephrozoa |
(unranked): | Protostomia |
(unranked): | Spiralia sensu Edgecombe et al. 2011 |
Phyla | |
The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other taxa. [4] The term Spiralia is applied to those phyla that exhibit canonical spiral cleavage, a pattern of early development found in most (but not all) members of the Lophotrochozoa. [5]
Members of the molluscs, annelids, platyhelminths and nemerteans have all been shown to exhibit spiral cleavage in its classical form. Other spiralian phyla (rotifers, brachiopods, phoronids, gastrotrichs, and bryozoans) are also said to display a derived form of spiral cleavage in at least a portion of their constituent species, although evidence for this is sparse. [6]
Previously, spiral cleavage was thought to be unique to the Spiralia in the strictest sense—animals such as molluscs and annelids which exhibit classical spiral cleavage. The presence of spiral cleavage in animals such as platyhelminths could be difficult to correlate with some phylogenies. [7]
Evidence of a close relationship between molluscs, annelids and lophophorates was found in 1995 and Lophotrochozoa was defined as the group containing these taxa and all the descendants of their last common ancestor. [8] More recent research has established the Lophotrochozoa as a superphylum within the Metazoa. [9] With this understanding, the presence of spiral cleavage in polyclad platyhelminths, as well as the more traditional Spiralia, has led to the hypothesis that spiral cleavage was present ancestrally across the Lophotrochozoa as a whole. [6] With the introduction of Platytrochozoa and Rouphozoa, the cladogram is as follows, with an indication approximately how many million years ago (Mya) the clades radiated into newer clades. [10] [11] [12] [13] [14] [15] [16]
Protostomia | |
610 mya |
An alternative phylogeny was given in 2019, with a basal grouping Mollusca with Entoprocta grouping named Tetraneuralia, and a second grouping of Nemertea with Platyhelminthes named Parenchymia as sister of Annelida. In their proposal and according to the original definition, Lophotrochozoa may become a senior synonym for Platytrochozoa. [17] [18] [19] [20]
In 2019 the Rouphozoa was recovered again as a basal Platytrochozoa clade. [21]
A 2022 study supported the Trochozoa and Platyzoa hypotheses, as shown below. [22] The same year another study placed bryozoans, entoproctans and cycliophorans in the group Polyzoa as one of the earliest branches among Lophotrochozoa. [23]
The flatworms, flat worms, Platyhelminthes, or platyhelminths are a phylum of relatively simple bilaterian, unsegmented, soft-bodied invertebrates. Being acoelomates, and having no specialised circulatory and respiratory organs, they are restricted to having flattened shapes that allow oxygen and nutrients to pass through their bodies by diffusion. The digestive cavity has only one opening for both ingestion and egestion ; as a result, the food can not be processed continuously.
The Chaetognatha or chaetognaths are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, they are mostly nektonic; however about 20% of the known species are benthic, and can attach to algae and rocks. They are found in all marine waters, from surface tropical waters and shallow tide pools to the deep sea and polar regions. Most chaetognaths are transparent and are torpedo shaped, but some deep-sea species are orange. They range in size from 2 to 120 millimetres.
The Lophophorata or Tentaculata are a Lophotrochozoan clade consisting of the Brachiozoa and the Bryozoa. They have a lophophore. Molecular phylogenetic analyses suggest that lophophorates are protostomes, but on morphological grounds they have been assessed as deuterostomes. Fossil finds of the "tommotiid" Wufengella suggest that they evolved from worm-like animals that resembled annelids.
Nemertea is a phylum of animals also known as ribbon worms or proboscis worms, consisting of about 1300 known species. Most ribbon worms are very slim, usually only a few millimeters wide, although a few have relatively short but wide bodies. Many have patterns of yellow, orange, red and green coloration. The foregut, stomach and intestine run a little below the midline of the body, the anus is at the tip of the tail, and the mouth is under the front. A little above the gut is the rhynchocoel, a cavity which mostly runs above the midline and ends a little short of the rear of the body. All species have a proboscis which lies in the rhynchocoel when inactive but everts to emerge just above the mouth to capture the animal's prey with venom. A highly extensible muscle in the back of the rhynchocoel pulls the proboscis in when an attack ends. A few species with stubby bodies filter feed and have suckers at the front and back ends, with which they attach to a host.
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.
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.
The Trochozoa are a proposed Lophotrochozoa clade that is a sister clade of Bryozoa and Platyzoa. The clade would include animals in five phyla: the Nemertea, the Annelida, the Mollusca, and the two Brachiozoan phyla, Brachiopoda and Phoronida. Both annelids and molluscs have been suggested as the sister group of Brachiozoa. It has also been proposed that nemerteans are actually a clade of annelids.
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.
The lophophore is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Hyolitha, and Phoronida, which collectively constitute the protostome group Lophophorata. All lophophores are found in aquatic organisms.
The evolution of the molluscs is the way in which the Mollusca, one of the largest groups of invertebrate animals, evolved. This phylum includes gastropods, bivalves, scaphopods, cephalopods, and several other groups. The fossil record of mollusks is relatively complete, and they are well represented in most fossil-bearing marine strata. Very early organisms which have dubiously been compared to molluscs include Kimberella and Odontogriphus.
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. 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.
Deuterostomes are bilaterian animals of the superphylum Deuterostomia, typically characterized by their anus forming before the mouth during embryonic development. Deuterostomia is further divided into 4 phyla: Chordata, Echinodermata, Hemichordata, and the extinct Vetulicolia known from Cambrian fossils. The extinct clade Cambroernida is also thought to be a member of Deuterostomia.
Phoronids are a small phylum of marine animals that filter-feed with a lophophore, and build upright tubes of chitin to support and protect their soft bodies. They live in most of the oceans and seas, including the Arctic Ocean but excluding the Antarctic Ocean, and between the intertidal zone and about 400 meters down. Most adult phoronids are 2 cm long and about 1.5 mm wide, although the largest are 50 cm long.
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.
Gnathifera is a clade of generally small spiralians characterized by complex jaws made of chitin. It comprises the phyla Gnathostomulida, Rotifera and Micrognathozoa. Chaetognatha has recently been recognised as closely related to the group, with it either being included within Gnathifera or the broader group Chaetognathifera.
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.
The annelids, 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 ecologies – some in marine environments as distinct as tidal zones and hydrothermal vents, others in fresh water, and yet others in moist terrestrial environments.
The Platytrochozoa are a proposed basal clade of spiralian animals as the sister group of the Gnathifera. The Platytrochozoa were divided into the Rouphozoa and the Lophotrochozoa. A more recent study suggests that the mesozoans also belong to this group of animals, as sister of the Rouphozoa.
The Kryptotrochozoa are a proposed Lophotrochozoa clade. It consists of the Nemertea and Lophophorata. It is controversial.
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.