Lophotrochozoa (, "crest/wheel animals") is a
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 ...
of
protostome
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 ...
animals within the
Spiralia
The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other Taxon, taxa. The term ''Spiralia'' is applied to those phyla that exhibit canonical spiral clea ...
. The taxon was established as a monophyletic group based on molecular evidence.
The clade includes animals like
annelid
The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to vario ...
s,
molluscs
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
,
bryozoa
Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
ns, and
brachiopod
Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear e ...
s.
Groups
Lophotrochozoa was defined in 1995 as the "last common ancestor of the three traditional
lophophorate taxa (
brachiopods,
bryozoa
Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
ns, and
phoronid
Phoronids (Taxonomy (biology), taxonomic name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that Filter feeder, filter-feed with a lophophore (a "crown" of tentacles), and build upright tubes of chitin to suppo ...
worms), the
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 the
annelid
The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to vario ...
s, and all of the descendants of that common ancestor".
It is a
cladistic definition (a node-based name), so the affiliation to Lophotrochozoa of spiralian groups not mentioned directly in the definition depends on the topology of the
spiralia
The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other Taxon, taxa. The term ''Spiralia'' is applied to those phyla that exhibit canonical spiral clea ...
n tree of life, and in some phylogenetic hypotheses, Lophotrochozoa may even be synonymous to Spiralia.
Nemertea
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. ...
and
Orthonectida
Orthonectida () is a small phylum (biology), phylum of poorly known parasites of marine invertebrates that are among the simplest of multi-cellular organisms. Members of this phylum are known as orthonectids.
Biology
The adults, which are the se ...
(if not directly considered as part of
Annelida
The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to variou ...
) are probably lophotrochozoan phyla;
Dicyemida,
Gastrotricha
The gastrotrichs (phylum Gastrotricha), common name, commonly referred to as hairybellies or hairybacks, are a group of Microscopic scale, microscopic (0.06–3.0 mm), cylindrical, acoelomate animals, and are widely distributed and abundant ...
, and
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 ...
may be lophotrochozoans or placed in the
Rouphozoa clade outside Lophotrochozoa;
Chaetognatha
The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, they are mostly pelagic; however about 20% of the known species ...
,
Gnathostomulida
Gnathostomulids, or jaw worms, are a small phylum of nearly microscopic marine animals. They inhabit sand and mud beneath shallow coastal waters and can survive in relatively anoxic environments. They were first recognised and described in 1956.
...
,
Micrognathozoa, and
Syndermata are probably
gnathiferans and so placed as a basal spiralian clade outside Lophotrochozoa;
Cycliophora could be a gnathiferan or a lophotrochozoan phylum. One of the candidate hypotheses is presented below.
The Lophotrochozoa has basal
Cycliophora and
Mollusca
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
groups, and more derived
Lophophorate,
Nemertea
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. ...
and
Annelid
The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to vario ...
a groups.
With the introduction of
Platytrochozoa and
Rouphozoa, one candidate phylogeny is pictured below – though other studies recover a range of alternative possibilities:
In the most recent research, the three phyla Cycliophora, Entoprocta and Bryozoa makes up a single clade and are the first to branch off from the other lophotrochozoans. The second split is the molluscs, and the third consists of two sister phyla, annelids and nemerteans. Lastly remains the clade that consist of the phoronids and the brachiopods.
Another study recovers Lophotrochozoa as equivalent to Platytrochozoa, forming a sister group with
Gnathifera at the base of
Spiralia
The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other Taxon, taxa. The term ''Spiralia'' is applied to those phyla that exhibit canonical spiral clea ...
.
A number of fossil taxa can be identified as early Lophotrochozoans, even if their precise affinity remains contested. However, relevant Cambrian fossils are debated.
Characteristics
The clade Lophotrochozoa is named after the two distinct characteristics of its members; the
lophophore, a feeding structure consisting of a ciliated crown of tentacles surrounding a mouth, and the developmental stage of the
trochophore
A trochophore () is a type of free-swimming planktonic marine larva with several bands of cilia.
By moving their cilia rapidly, they make a water eddy to control their movement, and to bring their food closer in order to capture it more easily.
...
larva.
Lophophorata such as
Brachiozoa and
Bryozoa
Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
have lophophores, while members of
Trochozoa such as
molluscs
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
and
annelids
The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to vario ...
have trochophore larvae, although some may have none.
References
Further reading
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{{Taxonbar, from=Q321481
Extant Cambrian first appearances
Superphyla
Spiralia