Xenoturbella Bocki
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''Xenoturbella bocki'' is a marine
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
worm Worms are many different distantly related bilateria, bilateral animals that typically have a long cylindrical tube-like body, no limb (anatomy), limbs, and usually no eyes. Worms vary in size from microscopic to over in length for marine ...
-like species from the genus ''
Xenoturbella ''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 collected in 1878 and 1879 in the Gullmar fi ...
''. It is found in saltwater sea floor habitats off the coast of Europe, predominantly
Sweden Sweden, formally the Kingdom of Sweden, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. It borders Norway to the west and north, and Finland to the east. At , Sweden is the largest Nordic count ...
. 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 (likely larval forms, as opposed to hard-shelled adults).


Systematics


Etymology

For the genus name ''Xenoturbella'',
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
''xénos'', means foreign or strange, and
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
, means a bustle or turbulence in water. Genus ''Xenoturbella'' is a member of sub-phylum
Xenoturbellida ''Xenoturbella'' is a genus of very simple bilaterians up to a few centimeters long. It contains a small number of marine benthos, benthic worm-like species. The first known species (''Xenoturbella bocki'') was collected in 1878 and 1879 in the G ...
, which are known as – Swedish for "paradox worms" (a term that some popular media have applied to the species), because if it is classified as a
deuterostome Deuterostomes (from Greek: ) are bilaterian animals of the superphylum Deuterostomia (), typically characterized by their anus forming before the mouth during embryonic development. Deuterostomia comprises three phyla: Chordata, Echinodermata, ...
, it would be more closely related to humans than other, more complex,
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s such as
lobster Lobsters are Malacostraca, malacostracans Decapoda, decapod crustaceans of the family (biology), family Nephropidae or its Synonym (taxonomy), synonym Homaridae. They have long bodies with muscular tails and live in crevices or burrows on th ...
s. Deuterostomes are a
superphylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclat ...
of animals whose anus forms before their mouth does during embryonic development. It includes humans, other
chordate A chordate ( ) is a bilaterian animal belonging to the phylum Chordata ( ). All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics ( synapomorphies) that distinguish them from ot ...
s,
echinoderm An echinoderm () is any animal of the phylum Echinodermata (), which includes starfish, brittle stars, sea urchins, sand dollars and sea cucumbers, as well as the sessile sea lilies or "stone lilies". While bilaterally symmetrical as ...
s and
hemichordate Hemichordata ( ) is a phylum which consists of triploblastic, eucoelomate, and bilaterally symmetrical marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and incl ...
s. The species signifier ''bocki'' refers to Sixten Bock, who first collected the organism in 1915. It was assigned by Swedish zoologist Einar Westblad, who described the species in 1949.


Taxonomy

In 1999, examination of ''X. bocki'' specimens held at the Swedish Museum of Natural History showed that a small subset of them must belong to another species. This population differed from specimens identified as ''X. bocki'' in internal fertilization, its small size of at most, and its pink coloration'' ''– in contrast to yellow-white coloration identified for ''X. bocki''. The new taxon was named after Westblad, who collected the specimens from coarser and shallower habitats in the same range as ''X. bocki''. However,
mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
sequencing from the specimens identified with both species suggested that the two populations belonged to the same species, involving that ''X. westbladi'' is a junior synonym to ''X. bocki''.


Phylogeny


Species level

Comparison of
mitochondrial DNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondrion, mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the D ...
and protein sequences showed that the species ''Xenoturbella bocki ''– often found off the coast of Sweden'' ''– is the sister group to ''X. hollandorum'', a species discovered in 2016 in eastern
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five Borders of the oceans, oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is ...
. In turn, these two species share evolutionary affinities with ''X. japonica'' into a clade of 'shallow-water' taxa.


Above the genus level

When it was discovered, ''X. bocki'' was placed in a new genus ''
Xenoturbella ''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 collected in 1878 and 1879 in the Gullmar fi ...
''. Above the genus level, the classification of this animal is controversial. Westbald placed it in the
phylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...
Platyhelminthes Platyhelminthes (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") is a phylum of relatively simple bilaterian, unsegmented, soft-bodied invertebrates commonly called f ...
in the
class Class, Classes, or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used d ...
Turbellaria 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 to large freshwater forms mo ...
(free-living flatworms). In 1999, based on genetic analysis, it was placed in
Protostomia 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 memb ...
by Israelsson, grouped with the
bivalve Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
s. Protostomia is a large clade including
worms The World Register of Marine Species (WoRMS) is a taxonomic database that aims to provide an authoritative and comprehensive catalogue and list of names of marine organisms. Content The content of the registry is edited and maintained by scien ...
,
mollusks Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The num ...
and
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s. In embryonic development, their mouth develops prior to the development of the anus for most protostomes, though some have evolved other developmental pathways. If placed in this clade, ''
Xenoturbella ''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 collected in 1878 and 1879 in the Gullmar fi ...
'' would also be among these exceptions. However, today, this is understood as a misclassification due to contaminating DNA from its shellfish food. Swedish scientist Sarah J. Bourlat and her coauthors in 2006 placed it in its own phylum,
Xenoturbellida ''Xenoturbella'' is a genus of very simple bilaterians up to a few centimeters long. It contains a small number of marine benthos, benthic worm-like species. The first known species (''Xenoturbella bocki'') was collected in 1878 and 1879 in the G ...
. More recent studies suggest on the basis of genetic and developmental evidence (e.g.
Hox genes Hox genes, a subset of homeobox genes, are a group of related genes that specify regions of the body plan of an embryo along the head-tail axis of animals. Hox proteins encode and specify the characteristics of 'position', ensuring that the c ...
) that it should be grouped with
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 ...
and
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 in ...
into
Acoelomorpha Acoelomorpha is a subphylum of very simple and small soft-bodied organism, soft-bodied animals with planula-like features which live in marine (ocean), marine or brackish waters. They usually live between grains of sediment, swimming as plankton, ...
. These three taxa are sometimes placed within the
deuterostome Deuterostomes (from Greek: ) are bilaterian animals of the superphylum Deuterostomia (), typically characterized by their anus forming before the mouth during embryonic development. Deuterostomia comprises three phyla: Chordata, Echinodermata, ...
s (a large
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
that includes humans and other
chordates A chordate ( ) is a bilaterian animal belonging to the phylum Chordata ( ). All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics ( synapomorphies) that distinguish them from ot ...
,
sea stars A sea is a large body of salt water. There are particular seas and the sea. The sea commonly refers to the ocean, the interconnected body of seawaters that spans most of Earth. Particular seas are either marginal seas, second-order section ...
and others), while others classify these organisms as a basal offshoot that resembles a common ancestor of deuterostomes and protostomes. A 2016 analysis of many genetic data sets supports the latter, and suggests that, like ''Xenoturbella bocki'', the common ancestor of protostomes and deuterostomes likely had one opening, ciliated locomotion and a wormlike body. However, if the deuterostome hypothesis is correct, then ''Xenoturbella'' must have lost many ancestral traits, such as an anus.


Description

This animal usually grows to in length, though individuals as long as have been reported. Its nervous system consists of a
nerve net ''Nerve Net'' is the eleventh solo studio album by Brian Eno, released on 1 September 1992 on Opal and Warner Bros. Records. It marked a return to more rock-oriented material, mixed with heavily syncopated rhythms, experimental electronic com ...
with no defined
brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
or
ganglia A ganglion (: ganglia) is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system, this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system, there a ...
. The nerve net is found on the basal (away from the animal's surface) side of the skin. This animal lacks a
coelom The coelom (or celom) is the main body cavity in many animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, i ...
. It also lacks an anus, excreting waste through the same opening as it intakes food. Thus, the digestive organ is sac-like. The opening is on the belly of the animal, near the front. The animal is simultaneously
hermaphroditic A hermaphrodite () is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic. The individuals of many ...
. A furrow runs along the circumference of the body in the middle of the animal. There are also side furrows. On its sides there are numerous tiny cilia that aid in locomotion. Small cells contain
vesicle Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features ...
s which may act as glands. An organ of unknown function, preliminarily called a
statocyst The statocyst is a balance sensory receptor present in some aquatic invertebrates, including bivalves, cnidarians, ctenophorans, echinoderms, cephalopods, crustaceans, and gastropods, A similar structure is also found in '' Xenoturbella''. T ...
, has been observed on the front end of the animal. Two leading hypotheses are that it aids in balance, as statocysts do in other invertebrates, or that it has
endocrine The endocrine system is a messenger system in an organism comprising feedback loops of hormones that are released by internal glands directly into the circulatory system and that target and regulate distant organs. In vertebrates, the hypotha ...
functions. Experiments in which the animal was observed to cleave into two after a wound show that the statocyst is essential for normal behavior and long-term survival.


Ecology

The animal moves through the water via rhythmic muscle contraction, aided by its side cilia, and a tuft of longer cilia on its back. The organism can also use its musculature to roll up into a ball, and maintain that form for several months. Adults are known to have a symbiotic relationship with Chlamydiae and Gammaproteobacteria, two bacterial
endosymbiont An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
s found in their
gastrodermis Gastrodermis (from Ancient Greek: , , "stomach"; , , "skin") is the inner layer of Cell (biology), cells that serves as a lining membrane of the gastrovascular cavity in cnidarians. It is distinct from the outer epidermis and the inner dermis and ...
. Genetic data confirms that its diet includes bivalve mollusks. However, it has never been observed feeding, so it is unknown if it eats bivalve carcasses, eggs, sperm, mucus, feces, or live larval or adult bivalves. It lacks any visible means to get through the shells of adult bivalves. Captive specimens survived for several months without food, and showed no interest in any of the proposed food items afterwards. This has led some to suggest that it feeds by absorbing dissolved organic matter through its skin. At least one specimen that has been proposed to show a consumed bivalve larvae is preserved in the Swedish Natural History Museum. This species burrows, and has been observed to make tunnels as deep as into substrate in a laboratory aquarium.


Range

This species has been found in ocean habitats off the coast of Europe, most often off the coast of Sweden. It is often collected using a Warén’s dredge from mud on the sea floor, at depths of .


Reproduction

''X. bocki'' has only been observed to reproduce sexually. In the wild, this species spawns in the winter. It lays small, mucus-coated eggs, which sink in the water column. The eggs have a pale-orange color, and are opaque. Young, upon hatching, are yellowish, nearly spherical, and move to the surface of the water. Larvae lack a
blastopore Gastrulation is the stage in the early embryonic development of most animals, during which the blastula (a single-layered hollow sphere of cells), or in mammals, the blastocyst, is reorganized into a two-layered or three-layered embryo known as ...
and do not feed until they are fully developed. They may derive nourishment from the yolk which would make them lecithotrophic. Within five days muscular contractions are observed in a laboratory setting, which may aid locomotion. ''X. bocki'' is a direct developer. As of 2013, this animal is extremely challenging to grow in captivity.


References


External links

* {{Use dmy dates, date=March 2018 Animals described in 1949 bocki