Acoelomorpha is a

subphylum In zoological nomenclature, a subphylum is a taxonomic rank below the rank of phylum. The taxonomic rank of "subdivision (rank), subdivision" in fungi and plant taxonomy is equivalent to "subphylum" in zoological taxonomy. Some plant taxonomists ...

of very simple and small soft-bodied animals with

planula A planula is the free-swimming, flattened, ciliated, bilaterally symmetric larval form of various cnidarian species and also in some species of Ctenophores. Some groups of Nemerteans also produce larvae that are very similar to the planula, which ...

-like features which live in marine or

brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estua ...

s. They usually live between grains of

sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand an ...

, swimming as

plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...

, or crawling on other organisms, such as

algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...

and

coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and sec ...

s.Cannon, L. R. G. (1986) ''Turbellaria of the World. A guide to families and genera''. Brisbane, Queensland Museum, 136 p. With the exception of two acoel freshwater species, all known Acoelomorphs are marine.

Systematics

Etymology

The term "acoelomorph" derives from the

Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...

words (), the ''

alpha privative An alpha privative or, rarely, privative a (from Latin ', from Ancient Greek ) is the prefix ''a-'' or ''an-'' (before vowels) that is used in Indo-European languages such as Sanskrit and Greek and in words borrowed therefrom to express negat ...

'', expressing negation or absence, (), meaning "cavity", and (), meaning "form". This refers to the fact that acoelomorphs have a structure lacking a fluid-filled body cavity.

Classification

The subphylum Acoelomorpha is divided into two classes. There are at least 408 described species, with a majority of these falling within the Crucimusculata infraorder in Acoela. *

Acoela 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 o ...

comprise small flattened worms, classified into a dozen of families. *

Nemertodermatida Nemertodermatida is a class of Acoelomorpha, comprising 18 species of millimetre-sized turbellariform, mostly interstitial worms. Taxonomy The order ''Nemertodermatida'' contains two families with 6 genera. The high level of cryptic diversity ...

comprise millimetre-sized, mostly interstitial worms, distributed into two families.

Phylogeny

The soft bodies of acoelomorphs and the lack of some of the key bilaterian traits make them difficult to classify. Traditionally, based on phenotypic features, acoelomorphs were considered to belong to the phylum

Platyhelminthes The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek language, Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a Phylum (biology), phylum of relati ...

, which was long seen as the sister group to all other bilaterian phyla.Conway-Morris, S.; George, J. D.; Gibson R.; Platt, H. M. (1985) ''The Origins and relationships of lower invertebrates''. Oxford, Clarendon Press, 397 p. However, a series of molecular phylogenetics studies at the hinge between the 20th and 21st centuries demonstrated that they are fast-evolving organisms not closely related to platyhelminthes, therefore involving the

polyphyly A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...

of flatworms. Actually, Acoelomorpha appeared to constitute a separate, deep-branching phylum, kingpin of bilaterian evolution. Yet their evolutionary affinities remain enigmatic as they might be the sister-group either to all other bilateral animals or to all deuterostomes. Resolving this debate would indicate whether acoelomorphs are simple or simplified. If they are the sister group to Bilateria, it would point to a simple

body plan A body plan, ( ), 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. This term, usually applied to animals, envisages a "blueprin ...

for the first bilaterian. Alternatively, if acoelomorphs are related to

deuterostomes Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exampl ...

, this would imply that their organisation is the result of secondary simplification. Xenoturbella japonica

In addition, comparative analyses of morphological, developmental, and molecular characters raised two points. *

Xenoturbellida ''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. The first known species ('' Xenoturbella bocki'') was discovered in 1915 by Sixten Bock, but it w ...

is the sister group to acoelomorphs, constituting the so-called

Xenacoelomorpha 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 (physical ...

clade. The close evolutionary relationship between Acoelomorpha and ''Xenoturbella'' is supported by the morphology (structure of epidermal cilia), the embryology (direct development without a feeding larval stage), and the concatenation of hundreds of proteins. * The phylogenetic placement of Xenacoelomorpha among bilaterian animals is not yet well defined, despite increased taxon and gene sampling, (re)-analyses of published data sets, and use of more sophisticated models of sequence evolution in

phylogenomic Phylogenomics is the intersection of the fields of evolution and genomics. The term has been used in multiple ways to refer to analysis that involves genome data and evolutionary reconstructions. It is a group of techniques within the larger fields ...

studies. There is a conflict between two evolutionary hypotheses, with Xenacoelomorpha being the sister group to Ambulacraria within

(i.e., the

Xenambulacraria Xenambulacraria is a proposed clade of animals with bilateral symmetry as an embryo, consisting of the Xenacoelomorpha (i.e., ''Xenoturbella'' and acoelomorphs) and the Ambulacraria (i.e., echinoderms and hemichordates). If confirmed, the cla ...

hypothesis) on the one hand, and Xenacoelomorpha as sister group to all other bilaterians (i.e., the

Nephrozoa 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 (conventionally deut ...

hypothesis) on the other. However, the Nephrozoa hypothesis might reflect methodological errors resulting from model violations in the phylogenomic inference.

Anatomy

Acoelomorphs resemble flatworms in many respects, but have a simpler anatomy, not even having a gut. Like flatworms, they have no circulatory or respiratory systems, but they also lack an excretory system. They lack body cavities (acoelomate structure), a hindgut or an anus. The

epidermal cells The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...

of acoelomorphs are unable to proliferate, a feature that is only shared with

rhabditophora Rhabditophora (from ''rhabdito''-, rhabdite + Greek -φορος ''phoros'' bearer, i.e., "rhabdite bearers") is a class of flatworms. It includes all parasitic flatworms (clade Neodermata) and most free-living species that were previously group ...

n flatworms and was for some time considered a strong evidence for the position of Acoelomorpha within Platyhelminthes. In both groups, the epidermis is renewed from mesodermal stem cells. The

nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...

of acoelomorphs is formed by a set of longitudinal nerve bundles beneath the ciliated epidermis. Close to the anterior end, these bundles are united by a ring commissure, but do not form a true brain, although it is hypothesized that such organization was the precursor of the

cephalization 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, ...

of the nerve system in more derived bilaterians. After decapitation, such a "brain" (rather, a ''cerebroid ganglion'') regenerates in a few weeks. The sensory organs include a

statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, and crustaceans. A similar structure is also found in '' Xenoturbella''. The statocyst c ...

– which presumably helps them orient to gravity –, and, in some cases,

ancestral An ancestor, also known as a forefather, fore-elder or a forebear, is a parent or ( recursively) the parent of an antecedent (i.e., a grandparent, great-grandparent, great-great-grandparent and so forth). ''Ancestor'' is "any person from wh ...

pigment-spot

ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...

capable of detecting light. Acoelomorphs are simultaneous

hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...

s, but have no

gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...

s and no ducts associated with the female reproductive system. Instead,

gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...