Acoela | |
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Many flatworm-like, orange individuals of the Waminoa acoel on a Plerogyra coral (whitish bubbles) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Xenacoelomorpha |
Subphylum: | Acoelomorpha |
Order: | Acoela Uljanin, 1870 |
Acoela, or the acoels, is an order of small and simple invertebrates in the subphylum Acoelomorpha of phylum Xenacoelomorpha, a deep branching bilaterian group of animals, which resemble flatworms. Historically they were treated as an order of turbellarian flatworms. [1] [2]
The etymology of "acoel" is from the Ancient Greek words ἀ (a), the alpha privative , expressing negation or absence, and κοιλία (koilía), meaning "cavity". [3] [4] This refers to the fact that acoels have a structure lacking a fluid-filled body cavity.
Acoels are very small flattened worms, usually under 2 millimetres (0.079 in) in length, but some larger species, such as Symsagittifera roscoffensis , may reach up to 15 millimetres (0.59 in). [5] They are bilaterally symmetric and microscopic. [6]
They are found worldwide in marine and brackish waters, usually having a benthic lifestyle, although some species are epibionts. [7] Two species, Limonoposthia polonica and Oligochoerus limnophilus, lives in freshwater. [8]
Members of the class Acoela lack a conventional gut, so that the mouth opens directly into the mesenchyme, i.e., the layer of tissue that fills the body. [9] [10] Digestion is accomplished by means of a syncytium that forms a vacuole around ingested food. There are no epithelial cells lining the digestive vacuole, but in the families Diopisthoporidae, Hallangiidae, Hofsteniidae, and Solenofilomorphidae, and the genera Oligochoerus (Convolutidae) and Proporus (Proporidae), there is a short pharynx leading from the mouth to the vacuole. [11] All other bilateral animals (apart from tapeworms) have a gut lined with epithelial cells. As a result, the acoels appear to be solid-bodied.
As the basal lineage of bilateral animals, the Acoela provide interesting insights into early animal evolution and development. [12] [13] The most thoroughly studied animal in this group is the species Isodiametra pulchra . Acoela used to be classified in the phylum Platyhelminthes. However, Acoela was separated from this phylum after molecular analyses showed that it had diverged before the three main bilaterian clades had formed.
The following sub-taxa are recognised in the order Acoela: [14]
The flatworms, flat worms, Platyhelminthes, or platyhelminths are a phylum of relatively simple bilaterian, unsegmented, soft-bodied invertebrates. Unlike other bilaterians, they are acoelomates, and have no specialised circulatory and respiratory organs, which restricts them 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 cannot be processed continuously.
Bilateria is a large clade/superphylum 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 achieve secondary pentaradial symmetry as adults, but are 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.
The Turbellaria are one of the traditional sub-divisions of the phylum Platyhelminthes (flatworms), and include all the sub-groups that are not exclusively parasitic. There are about 4,500 species, which range from 1 mm (0.039 in) to large freshwater forms more than 500 mm (20 in) long or terrestrial species like Bipalium kewense which can reach 600 mm (24 in) in length. All the larger forms are flat with ribbon-like or leaf-like shapes, since their lack of respiratory and circulatory systems means that they have to rely on diffusion for internal transport of metabolites. However, many of the smaller forms are round in cross section. Most are predators, and all live in water or in moist terrestrial environments. Most forms reproduce sexually and with few exceptions all are simultaneous hermaphrodites.
Acoelomorpha is a subphylum of very simple and small soft-bodied animals with planula-like features which live in marine or brackish waters. They usually live between grains of sediment, swimming as plankton, or crawling on other organisms, such as algae and corals. With the exception of two acoel freshwater species, all known Acoelomorphs are marine.
Cephalization is an evolutionary trend in which, over many generations, the mouth, sense organs, and nerve ganglia become concentrated at the front end of an animal, producing a head region. This is associated with movement and bilateral symmetry, such that the animal has a definite head end. This led to the formation of a highly sophisticated brain in three groups of animals, namely the arthropods, cephalopod molluscs, and vertebrates.
Xenoturbella is a genus of very simple bilaterians up to a few centimeters long. It contains a small number of marine benthic worm-like species.
Convolutidae is a family of acoels. It contains more than a third of all known acoel species.
Deuterostomia or deuterostomes are bilaterian animals typically characterized by their anus forming before their mouth during embryonic development. The three major clades of extant deuterostomes include chordates, echinoderms and hemichordates.
The embryological origin of the mouth and anus is an important characteristic, and forms the morphological basis for separating bilaterian animals into two natural groupings: the protostomes and deuterostomes.
Actinoposthiidae is a family of acoels.
Dakuidae is a family of acoels.
Nemertodermatidae is a family of wormlike animals in the phylum Acoelomorpha. They are similar to the flatworms of the phylum Platyhelminthes, and were traditionally classified as such.
Isodiametridae is a family of acoels.
Nephrozoa is a major clade of bilaterians, divided into the protostomes and the deuterostomes, containing almost all animal phyla and over a million extant species. Its sister clade is the Xenacoelomorpha. The Ambulacraria are occasionally thought to be sister to the Xenacoelomorpha, forming the Xenambulacraria as basal Deuterostomia, or basal Bilateria invalidating Nephrozoa and Deuterostomia in multiple studies. The coelom, the digestive tract and excretory organs (nephridia), and nerve cords developed in the Nephrozoa. It has been argued that, because protonephridia are only found in protostomes, they cannot be considered a synapomorphy of this group. This would make Nephrozoa an improper name, leaving Eubilateria as this clade's name.
Xenacoelomorpha is a small phylum of bilaterian invertebrate animals, consisting of two sister groups: xenoturbellids and acoelomorphs. This new phylum was named in February 2011 and suggested based on morphological synapomorphies, which was then confirmed by phylogenomic analyses of molecular data.
ParaHoxozoa is a clade of animals that consists of Bilateria, Placozoa, and Cnidaria. The relationship of this clade relative to the two other animal lineages Ctenophora and Porifera is debated. Some phylogenomic studies have presented evidence supporting Ctenophora as the sister to Parahoxozoa and Porifera as the sister group to the rest of animals. Some studies have presented evidence supporting Porifera as the sister to Parahoxozoa and Ctenophora as the sister group to the rest of animals.
Xenoturbella bocki is a marine benthic worm-like species from the genus Xenoturbella. It is found in saltwater sea floor habitats off the coast of Europe, predominantly Sweden. It was the first species in the genus discovered. Initially it was collected by Swedish zoologist Sixten Bock in 1915, and described in 1949 by Swedish zoologist Einar Westblad. The unusual digestive structure of this species, in which a single opening is used to eat food and excrete waste, has led to considerable study and controversy as to its classification. It is a bottom-dwelling, burrowing carnivore that eats mollusks.
Xenambulacraria is a proposed clade of animals with bilateral symmetry as an embryo, consisting of the Xenacoelomorpha and the Ambulacraria.
Meara is a genus of bilaterally symmetric, small aquatic worms in the phylum Acoelomorpha. This genus contains only one species, Meara stichopi, a parasite of the sea cucumber Parastichopus tremulus. It occurs in Norway and Sweden.