Acoela Temporal range: [1] | |
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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. [2] [3] About 400 species are known, but probably many more not yet described. [4]
The etymology of "acoel" is from the Ancient Greek words ἀ (a), the alpha privative , expressing negation or absence, and κοιλία (koilía), meaning "cavity". [5] [6] 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). [7] They are bilaterally symmetric and microscopic. [8]
They are found worldwide in marine and brackish waters, usually having a benthic lifestyle, although some species are epibionts. [9] Two species, Limonoposthia polonica and Oligochoerus limnophilus, live in freshwater. [10]
Species in the family Convolutidae often form endosymbiotic relationship with microalgae. In one of the genera, Waminoa, the algae are transmitted vertically from parents to offspring. In addition to Convolutidae, there appears to be a potential new and yet unnamed family of acoels that also live in relationships with microalgal endosymbionts. [11]
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. [12] [13] 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. [14] 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. [15] [16] 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, making flatworms as traditionally understood an evolutionary grade from which higher animals had evolved.
The following sub-taxa are recognised in the order Acoela: [17]
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.
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.
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 animals that, over many generations, the special sense organs and nerve ganglia become concentrated towards the rostral end of the body where the mouth is located, often producing an enlarged head. This is associated with the animal's movement direction and bilateral symmetry, and cephalization of the nervous system led to the formation of a functional centralized brain in three groups of bilaterian animals, namely the arthropods, cephalopod molluscs, and vertebrates (craniates).
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, belonging to the phylum Xenacoelomorpha. This family contains more than a third of all known acoel species.
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.
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 proposed major clade of bilaterian animals. It includes all bilaterians other than Xenacoelomorpha. It contrasts with the Xenambulacraria hypothesis, which instead posits that Xenacoelomorpha is most closely related to Ambulacraria. Which hypothesis is correct is controversial. Authors supporting the Xenambulacraria hypothesis have suggested that the genetic evidence used to support Nephrozoa may be due to systematic error.
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.
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.
Hofstenia, or panther worms, is a genus of worms belonging to the family Hofsteniidae.