Lancelet - Wikipedia

Lancelet - Wikipedia
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Cephalochordata
Subphylum of chordates
Not to be confused with
Lancet
or
Lancelot
Cephalochordata (Lancelets)
Temporal range:
Recent
PreꞒ
Pg
Possible
Cambrian
and
Permian
records
Branchiostoma lanceolatum
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Subphylum:
Cephalochordata
Haeckel
, 1866
Class:
Leptocardii
Müller
, 1845
Genera
Cathaymyrus
Palaeobranchiostoma
Branchiostomatidae
Bonaparte
, 1846
Asymmetron
Branchiostoma
Epigonichthys
Synonyms
Subphylum or class:
Acrania Haeckel, 1866
Order:
Amphioxi Bonaparte, 1846
Amphioxiformes Berg, 1937
Branchiostomiformes Fowler, 1947
Family:
Amphioxidae Gray, 1842
Asymmetrontidae Whitley, 1932
Epigonichthyidae Hubbs, 1922
The
lancelets
ɑː
-/
LA(H)N
-slit
), also known as
amphioxi
sg.
amphioxus
AM
-fee-
OK
-səs
), consist of 32 described species of somewhat fish-like
benthic
filter-feeding
chordates
in the subphylum
Cephalochordata
, class
Leptocardii
, and family
Branchiostomatidae
10
Lancelets diverged from other chordates during or prior to the
Cambrian
period. A number of fossil chordates have been suggested to be closely related to lancelets, including
Pikaia
and
Cathaymyrus
from the Cambrian and
Palaeobranchiostoma
from the
Permian
, but their close relationship to lancelets has been doubted by other authors.
11
12
Molecular clock
analysis suggests that modern lancelets probably diversified much more recently, during the
Cretaceous
or
Cenozoic
13
14
They are of interest to
zoologists
as lancelets contain many organs and organ systems that are
homologous
to those of modern fish. Therefore, they provide a number of examples of possible evolutionary
exaptation
. For example, the gill-slits of lancelets are used for feeding only, and not for respiration. The circulatory system carries food throughout their body, but does not have
red blood cells
or
hemoglobin
for transporting oxygen.
Comparing the
genomes
of lancelets and vertebrates and their differences in gene expression, function and number can shed light on the origins of vertebrates and their
evolution
15
The genome of a few species in the genus
Branchiostoma
have been sequenced:
B. floridae,
16
B. belcheri
17
and
B. lanceolatum
18
In Asia, lancelets are harvested commercially as food for humans. In Japan, amphioxus (
B. belcheri
) has been listed in the registry of "Endangered Animals of Japanese Marine and Fresh Water Organisms".
19
Ecology
edit
Habitat
edit
Adult amphioxus typically inhabit the seafloor, burrowing into well-ventilated substrates characterized by a soft texture and minimal organic content. While various species have been observed in different types of substrate, such as fine sand, coarse sand, and shell deposits, most exhibit a distinct preference for coarse sand with low levels of fine particles. For instance,
Branchiostoma nigeriense
along the west coast of Africa,
Branchiostoma caribaeum
in
Mississippi Sound
and along the coast from South Carolina to Georgia,
B. senegalense
in the Atlantic Ocean on the
shelf
region off
North West Africa
, and
B. lanceolatum
along the Mediterranean coast of southern France all demonstrate this preference.
20
21
22
23
24
25
However,
Branchiostoma floridae
from
Tampa Bay
, Florida, appears to be an exception to this trend, favoring fine sand bottoms instead.
26
Feeding
edit
Their habitat preference reflects their feeding method: they only expose the front end to the water and filter-feed on
plankton
by means of a branchial ciliary current that passes water through a mucous sheet.
Branchiostoma floridae
is capable of trapping particles from microbial to small phytoplankton size,
27
while
B. lanceolatum
preferentially traps bigger particles (>4 μm).
28
Reproduction and spawning
edit
Lancelets are
gonochoric
animals, i.e. having two sexes, and they reproduce via external
fertilisation
. They only reproduce during their
spawning
season, which varies slightly between species — usually corresponding to spring and summer months.
29
All lancelets species spawn shortly after sunset, either synchronously (e.g.
Branchiostoma floridae
, about once every two weeks during spawning season
30
) or asynchronously (
Branchiostoma lanceolatum
, gradual spawning through the season
31
). Rare instances of
hermaphroditism
have been reported in
Branchiostoma lanceolatum
and
B. belcheri
, where a small number of female gonads were observed within male individuals, typically ranging from 2 to 5 gonads out of a total of 45–50.
citation needed
Nicholas and
Linda Holland
were the first researchers to describe a method of obtaining amphioxus embryos by induction of spawning in captivity and in vitro fertilization.
32
Spawning can be artificially induced in the lab by electric or thermal shock.
33
History
edit
The first representative organism of the group to be described was
Branchiostoma lanceolatum
. It was described by
Peter Simon Pallas
in 1774 as
molluscan
slugs in the genus
Limax
34
It was not until 1834 that
Oronzio Gabriele Costa
brought the phylogenetic position of the group closer to the
agnathan vertebrates
hagfish
and
lampreys
), including it in the new genus
Branchiostoma
(from the Greek,
branchio-
= "gills",
-stoma
= "mouth").
35
36
In 1836,
William Yarrell
renamed the genus as
Amphioxus
(from the Greek: "pointed on both sides"),
37
now considered an obsolete
synonym
of the genus
Branchiostoma
. The term "amphioxus" is still used as a
common name
along with "lancelet", especially in the English language. All extant lancelets are all placed in the family Branchiostomatidae, class Leptocardii, and subphylum Cephalochordata.
38
Observations of amphioxus anatomy began in the middle of the 19th century.
Alexander Kovalevsky
first described the key anatomical features of the adult amphioxus (hollow
dorsal nerve tube
endostyle
, segmented body, postanal tail).
39
Armand De Quatrefages
first completely described the nervous system of amphioxus.
40
Kovalevsky also released the first complete description of amphioxus embryos,
39
while
Max Schultze
was the first
specify
to describe the larvae.
41
Anatomy
edit
Anatomy of the lancelet
The larvae are extremely asymmetrical, with the mouth and anus on the left side, and the gill slits on the right side.
42
43
Organs associated with the pharynx are positioned either exclusively on the left or on the right side of the body. In addition, segmented muscle blocks and parts of the nervous system are asymmetrical.
44
After metamorphosis the anatomy becomes more symmetrical, but some asymmetrical traits are still present also as adults, such as the nervous system and the location of the gonads which are found on the right side in Asymmetron and Epigonichthys (in Branchiostoma gonads develop on both sides of body).
45
46
Depending on the exact species involved, the maximum length of lancelets is typically 2.5 to 8 cm (1.0–3.1 in).
47
48
Branchiostoma belcheri
and
B. lanceolatum
are among the largest.
47
Except for the size, the species are very similar in general appearance, differing mainly in the number of
myotomes
and the pigmentation of their larvae.
47
They have a translucent, somewhat fish-like body, but without any paired fins or other limbs. A relatively poorly developed tail fin is present, so they are not especially good swimmers. While they do possess some
cartilage
material stiffening the
gill slits
, mouth, and tail, they have no true complex skeleton.
49
Nervous system and notochord
edit
In common with vertebrates, lancelets have a hollow nerve cord running along the back,
pharyngeal slits
and a tail that runs past the anus. Also like vertebrates, the muscles are arranged in blocks called
myomeres
50
Unlike vertebrates, the dorsal nerve cord is not protected by bone but by a simpler
notochord
made up of a
cylinder
of
cells
that are closely packed in collagen fibers to form a toughened rod. The lancelet notochord, unlike the vertebrate
spine
, extends into the head. This gives the subphylum, Cephalochordata, its name (
κεφαλή
kephalē
means 'head'). The fine structure of the notochord and the cellular basis of its adult growth are best known for the Bahamas lancelet,
Asymmetron lucayanum
51
The nerve cord is only slightly larger in the head region than in the rest of the body, so that lancelets do not appear to possess a true brain. However, developmental gene expression and
transmission electron microscopy
indicate the presence of a
diencephalic
forebrain
, a possible
midbrain
, and a
hindbrain
52
53
Recent studies involving a comparison with vertebrates indicate that the vertebrate
thalamus
pretectum
, and
midbrain
areas jointly correspond to a single, combined region in the amphioxus, which has been termed
di-mesencephalic primordium
(DiMes).
54
Visual system
edit
Lancelets have four known kinds of light-sensing structures: Three are respectively called
Joseph cells
Hesse organs
and
lamellar body
further explanation needed
The fourth is an unpaired anterior eye. All of them utilize
opsins
as light receptors. All of these organs and structures are located in the neural tube, with the frontal eye at the front, followed by the
lamellar body
, the
Joseph cells
, and the
Hesse organs
55
47
56
Joseph cells
and
Hesse organs
edit
Joseph cells are bare photoreceptors surrounded by a band of
microvilli
. These cells bear the opsin
melanopsin
. The
Hesse organs
(also known as dorsal ocelli) consist of a photoreceptor cell surrounded by a band of microvilli and bearing melanopsin, but half enveloped by a cup-shaped pigment cell. The peak sensitivity of both cells is ~470 nm
57
(blue).
Both the
Joseph cells
and
Hesse organs
are in the neural tube, the
Joseph cells
forming a dorsal column, the
Hesse organs
in the ventral part along the length of the tube. The
Joseph cells
extend from the caudal end of the anterior vesicle (or cerebral vesicle) to the boundary between myomeres three and four, where the
Hesse organs
begin and continue nearly to the tail.
58
59
Frontal eye
edit
The frontal eye consists of a pigment cup, a group of photoreceptor cells (termed
Row 1
), three rows of neurons (
Rows 2–4
), and
glial cells
. The frontal eye, which expresses the
PAX6
gene, has been proposed as the homolog of either the paired eyes or the
pineal eye
on vertebrates, the pigment cup as the homolog of the RPE (
retinal pigment epithelium
), the putative photoreceptors as homologs of vertebrate
rods
and
cones
, and Row 2 neurons as homologs of the
retinal ganglion cells
60
The pigment cup is oriented concave dorsally. Its cells contain the pigment
melanin
60
61
The putative photoreceptor cells, Row 1, are arranged in two diagonal rows, one on either side of the pigment cup, symmetrically positioned with respect to the ventral midline. The cells are flask-shaped, with long, slender ciliary processes (one cilium per cell). The main bodies of the cells lie outside of the pigment cup, while the cilia extend into the pigment cup before turning and exiting. The cells bear the opsin
c-opsin 1
, except for a few which carry
c-opsin 3
60
62
The Row 2 cells are
serotonergic
neurons in direct contact with Row 1 cells. Row 3 and 4 cells are also neurons. Cells of all four rows have
axons
that project into the left and right ventrolateral nerves. For Row 2 neurons, axon projections have been traced to the
tegmental
neuropil
. The tegmental neuropil has been compared with
locomotor
control regions of the vertebrate
hypothalamus
, where
paracrine
release modulates locomotor patterns such as feeding and swimming.
60
Fluorescent proteins
edit
Green fluorescence in Lancelets. (a.
Branchiostoma floridae
GFP near the eye spot and in the oral tentacles.) (b.
Asymmetron lucayanum
green fluorescence in the gonads.)
Lancelets naturally express
green fluorescent proteins
(GFP) inside their oral tentacles and near the eye spot.
63
Depending on the species, it can also be expressed in the tail and
gonads
, though this is only reported in the
Asymmetron
genus.
64
Multiple fluorescent protein
genes
have been recorded in lancelet species throughout the world.
Branchiostoma floridae
alone has 16 GFP-encoding genes. However, the GFP produced by lancelets is more similar to GFP produced by
copepods
than jellyfish (
Aequorea victoria
).
citation needed
It is suspected GFP plays multiple roles with lancelets such as attracting plankton towards their mouth. Considering that lancelets are filter feeders, the natural current would draw nearby plankton into the digestive tract. GFP is also expressed in
larvae
, signifying it may be used for
photoprotection
by converting higher energy blue light to less harmful green light.
citation needed
Live lancelet (
B. floridae
) under a fluorescent microscope.
The fluorescent proteins from lancelets have been adapted for use in molecular biology and microscopy. The
yellow fluorescent protein
from
Branchiostoma lanceolatum
exhibits unusually high
quantum yield
(~0.95).
65
It has been
engineered
into a
monomeric
green fluorescent protein known as
mNeonGreen
, which is the brightest known monomeric green or yellow fluorescent protein.
Feeding and digestive system
edit
Lancelets are passive
filter feeders
14
spending most of the time half-buried in sand with only their frontal part protruding.
66
They eat a wide variety of small
planktonic
organisms, such as bacteria,
fungi
diatoms
, and
zooplankton
, and they will also take
detritus
67
Little is known about the diet of the lancelet
larvae
in the wild, but captive larvae of several species can be maintained on a diet of
phytoplankton
, although this apparently is not optimal for
Asymmetron lucayanum
67
Lancelets have oral cirri, thin
tentacle
-like strands that hang in front of the mouth and act as sensory devices and as a filter for the water passing into the body. Water passes from the mouth into the large
pharynx
, which is lined by numerous gill-slits. The ventral surface of the pharynx contains a groove called the
endostyle
, which, connected to a structure known as
Hatschek's pit
, produces a film of
mucus
Ciliary
action pushes the mucus in a film over the surface of the gill slits, trapping suspended food particles as it does so. The mucus is collected in a second, dorsal groove, known as the
epipharyngeal groove
, and passed back to the rest of the digestive tract. Having passed through the gill slits, the water enters an atrium surrounding the pharynx, then exits the body via the atriopore.
49
Both adults and larvae exhibit a "cough" reflex to clear the mouth or throat of debris or items too large to swallow. In larvae the action is mediated by the pharyngeal muscles while in the adult animal it is accomplished by atrial contraction.
68
69
The remainder of the digestive system consists of a simple tube running from the pharynx to the anus. The
hepatic caecum
, a single blind-ending
caecum
, branches off from the underside of the gut, with a lining able to
phagocytize
the food particles, a feature not found in vertebrates. Although it performs many functions of a liver, it is not considered a true liver but a
homolog
of the vertebrate liver.
70
71
72
Other systems
edit
Lancelets have no respiratory system, breathing solely through their skin, which consists of a simple
epithelium
. Despite the name, little if any respiration occurs in the "gill" slits, which are solely devoted to feeding. The circulatory system does resemble that of primitive fish in its general layout, but is much simpler, and does not include a
heart
. There are no blood cells, and no
hemoglobin
49
The excretory system consists of segmented "kidneys" containing
protonephridia
instead of
nephrons
, and quite unlike those of vertebrates. Also unlike vertebrates, there are numerous, segmented
gonads
49
Model organism
edit
Lancelets became famous in the 1860s when
Ernst Haeckel
began promoting them as a model for the ancestor of all vertebrates. By 1900, lancelets had become a
model organism
. By the mid-20th century they had fallen out of favor for a variety of reasons, including a decline of comparative anatomy and embryology, and due to the belief that lancelets were more derived than they appeared, e.g., the profound asymmetry in the larval stage.
73
74
More recently, the fundamental symmetric and twisted development of vertebrates is the topic of the
axial twist theory
. According to this theory, there is a deep agreement between the vertebrates and cephalochordates, and even all chordates.
75
76
With the advent of molecular genetics lancelets are once again regarded as a model of vertebrate ancestors, and are used again as a model organism.
77
36
As a result of their use in science, methods of keeping and breeding lancelets in captivity have been developed for several of the species, initially the European
Branchiostoma lanceolatum
, but later also the West Pacific
Branchiostoma belcheri
and
Branchiostoma japonicum
, the
Gulf of Mexico
and West Atlantic
Branchiostoma floridae
and the circumtropical (however, genetic evidence suggest the Atlantic and
Indo-Pacific
populations should be recognized as separate
14
Asymmetron lucayanum
67
78
They can reach an age of up to 7–8 years.
78
As human food
edit
The animals are edible and harvested in some parts of the world. They are eaten both fresh, tasting like herring, and as a food additive in dry form after being roasted in oil.
citation needed
When their gonads start to ripen in the spring it affects their flavor, making them taste bad during their breeding season.
79
Phylogeny and taxonomy
edit
The lancelet is a small, translucent, fish-like animal that is one of the closest living invertebrate relatives of the vertebrates.
80
81
The lancelets were traditionally seen as the sister lineage to the
vertebrates
; in turn, these two groups together (sometimes called Notochordata) were considered the sister group to the
Tunicata
(also called Urochordata and including
sea squirts
). Consistent with this view, at least ten morphological features are shared by lancelets and vertebrates, but not tunicates.
82
Newer research suggests this pattern of evolutionary relationship is incorrect. Extensive
molecular phylogenetic
analysis has shown convincingly that the Cephalochordata is the most
basal
subphylum of the chordates, with tunicates being the sister group of the vertebrates.
83
84
This revised phylogeny of chordates suggests that tunicates have secondarily lost some of the morphological characters that were formerly considered to be
synapomorphies
(shared, derived characters) of vertebrates and lancelets. Lancelets have turned out to be among the most genetically diverse animals sequenced to date, due to high rates of genetic changes like
exon shuffling
and domain combination.
17
Among the three extant (living)
genera
Asymmetron
is basal.
Molecular clock
studies have come to different conclusions on their divergence, with some suggesting that
Asymmetron
diverged from other lancelets more than 100 million years ago
13
while others have suggested that it occurred about
46
million years ago.
14
According to the younger estimation,
Branchiostoma
and
Epigonichthys
have been estimated to have diverged from each other about
38.3
million years ago.
14
Despite this deep separation,
hybrids
between
Asymmetron lucayanum
and
Branchiostoma floridae
are viable (among the deepest split species known to be able to produce such hybrids).
67
The following are the species recognised by
WoRMS
. Other sources recognize about thirty species.
74
14
85
It is likely that currently unrecognized
cryptic species
remain.
67
Class Leptocardii
Family
Branchiostomatidae
Bonaparte 1846
Genus
Asymmetron
Andrews 1893
Amphioxides
Gill 1895
Asymmetron inferum
Nishikawa 2004
Asymmetron lucayanum
Andrews 1893
(Sharptail lancelet)
Genus
Branchiostoma
Costa 1834 non Newport 1845 non Banks 1905
Amphioxus
Yarrell 1836
Limax
Pallas 1774 non Linnaeus 1758 non Férussac 1819 non Martyn 1784
Dolichorhynchus
Willey 1901 non Mulk & Jairajpuri 1974
Branchiostoma africae
Hubbs 1927
Branchiostoma arabiae
Webb 1957
Branchiostoma bazarutense
Gilchrist 1923
Branchiostoma belcheri
(Gray 1847)
(Belcher's lancelet)
Branchiostoma bennetti
Boschung & Gunter 1966
(Mud lancelet)
Branchiostoma bermudae
Hubbs 1922
Branchiostoma californiense
Andrews 1893
(Californian lancelet)
Branchiostoma capense
Gilchrist 1902
Branchiostoma caribaeum
Sundevall 1853
(Caribbean lancelet)
Branchiostoma elongatum
(Sundevall 1852)
Branchiostoma floridae
Hubbs 1922
(Florida lancelet)
Branchiostoma gambiense
Webb 1958
Branchiostoma indicum
(Willey 1901)
Branchiostoma japonicum
(Willey 1897)
(Pacific lancelet)
Branchiostoma lanceolatum
(Pallas 1774)
(European lancelet)
Branchiostoma leonense
Webb 1956
Branchiostoma longirostrum
Boschung 1983
(Shellhash lancelet)
Branchiostoma malayanum
Webb 1956
Branchiostoma moretonense
Kelly 1966
nomen dubium
86
87
Branchiostoma nigeriense
Webb 1955
Branchiostoma platae
Hubbs 1922
Branchiostoma senegalense
Webb 1955
Branchiostoma tattersalli
Hubbs 1922
Branchiostoma virginiae
Hubbs 1922
(Virginian lancelet)
Genus
Epigonichthys
Peters 1876
Amphipleurichthys
Whitley 1932
Bathyamphioxus
Whitley 1932
Heteropleuron
Kirkaldy 1895
Merscalpellus
Whitley 1932
Notasymmetron
Whitley 1932
Paramphioxus
Haekel 1893
Zeamphioxus
Whitley 1932
Epigonichthys australis
(Raff 1912)
Epigonichthys bassanus
(Günther 1884)
Epigonichthys cingalensis
(Kirkaldy 1894)
nomen dubium
88
Epigonichthys cultellus
Peters 1877
Epigonichthys hectori
(Benham 1901)
(Hector's lancelet)
Epigonichthys maldivensis
(Foster Cooper 1903)
The
cladogram
presented here illustrates the
phylogeny
(family tree) of lancelets, and follows a simplified version of the relationships found by Igawa and colleagues (2017):
74
14
85
Chordata
Olfactores
Lancelet
Cathaymyrus
? († 518 mya)
Palaeobranchiostoma
? († 273.01 mya)
Lancelet
(crown group)
Asymmetron
Asymmetron inferum
Asymmetron lucayanum
Asymmetron maldivense
16.4 mya
Epigonichthys
Epigonichthys cultellus
Epigonichthys maldivensis
27.0 mya
Branchiostoma
Branchiostoma lanceolatum
Branchiostoma floridae
Branchiostoma japonicum
Branchiostoma belcheri
28.7 mya
46.0 mya
518 mya ?
See also
edit
Phylliroe
References
edit
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Further reading
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Stach, T.G. (2004). "Cephalochordata (Lancelets)". In M. Hutchins; Garrison, R.W.; Geist, V.; Loiselle, P.V.; Schlager, N.; McDade, M.C.; Duellman, W.E. (eds.).
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External links
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Wikimedia Commons has media related to
Amphioxiformes
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Cephalochordata
"Cephalochordata"
Museum of Paleontology
. Berkeley, CA:
U.C. Berkeley
"Branchiostoma japonicum and B. belcheri are Distinct Lancelets (Cephalochordata) in Xiamen Waters in China"
– via
ResearchGate
"Error in the genealogy of humans"
Sars International Centre for Marine Molecular Biology
sars.no
(Press release). Bergen, Norway:
University of Bergen
"A special issue of Amphioxus research"
biolsci.org
. Archived from
the original
on 2012-03-05
. Retrieved
2006-06-08
"A special issue of Amphioxus research"
biolsci.org
II
. Archived from
the original
on 2012-03-05
. Retrieved
2006-06-08
"Amphioxus and the T-box gene"
news-info.wustl.edu
(Press release). St. Louis, MO:
Washington University in St. Louis
A movie of the amphioxus embryonic development
on
"Scripps scientists discover fluorescence in key marine creature"
Scripps Institute
scrippsnews.ucsd.edu
(Press release). San Diego, CA:
U.C. San Diego
. Archived from
the original
on 2013-05-15
. Retrieved
2007-11-01
"Amphioxus"
Encyclopædia Britannica
. Vol. I (9th ed.). 1878. p. 774.
"Amphioxus: Taxonomy, brief facts, life cycle and embryology"
GeoChemBio
. Archived from
the original
on 2018-10-10
. Retrieved
2011-03-05
View the
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Evolution of fish
Fish
Timeline of fish evolution
Forerunners
Basal member
Pikaia
Cephalochordate
Cathaymyrus
Lancelet
Olfactores
Haikouella
Tunicate
Myllokunmingiidae
? (
Haikouichthys
Myllokunmingia
Zhongxiniscus
Jawless fish
Cyclostomata
Hagfish
Hyperoartia
Lamprey
Conodonts
Protoconodonta
Paraconodontida
Conodonta
Ostracoderms
Pteraspidomorphi
Thelodonti
Anaspida
Cephalaspidomorphi
Galeaspida
Pituriaspida
Osteostraci
Jawed fish
Placoderms
Antiarchi
Arthrodira
Brindabellaspida
Petalichthyida
Phyllolepida
Ptyctodontida
Rhenanida
Acanthothoraci
Pseudopetalichthyida
Stensioellida
Acanthodii
Acanthodiformes
†"
Climatiiformes
Diplacanthiformes
Ischnacanthiformes
Cartilaginous
Elasmobranchii
Ctenacanthiformes
Hybodontiformes
Phoebodontiformes
Xenacanthida
Ray
Shark
Holocephali
Eugeneodontida
Iniopterygiformes
Petalodontiformes
Symmoriiformes
Chimaera
Bony
Lobe-finned
Onychodontiformes
Coelacanth
Rhipidistia
Porolepiformes
Lungfish
Tetrapodomorpha
Ray-finned
Palaeonisciformes
Cladistia
Polypteriformes
Chondrostei
Acipenseriformes
Neopterygii
Pycnodontiformes
Halecomorphi
Ginglymodi
Teleost
Lists
Lists of prehistoric fish
spiny sharks
placoderms
cartilaginous
bony
lobe-finned
List of transitional fossils
Related
Prehistoric life
Transitional fossils
Vertebrate paleontology
† extinct
Extant
animal
phyla
Domain
Archaea
Bacteria
Eukaryota
(major groups
Metamonada
Discoba
Diaphoretickes
Hacrobia
Cryptista
Rhizaria
Alveolata
Stramenopiles
Plants
Amorphea
Amoebozoa
Opisthokonta
Animalia
Fungi
Mesomycetozoea
Animalia
Porifera (sponges)
Ctenophora (comb jellies)
ParaHoxozoa
Planulozoa
Placozoa (
Trichoplax
and relatives)
Cnidaria (jellyfish and relatives)
Bilateria
(Triploblasts)
(see below↓)
The
phylogeny
of the animal root
is disputed
; see also
Eumetazoa
Benthozoa
Bilateria
Bilateria
Xenacoelomorpha (acoels and relatives)
Chordata (vertebrates and relatives)
Ambulacraria
Echinodermata (starfish and relatives)
Hemichordata (acorn worms and relatives)
Protostomia
Ecdysozoa
Scalidophora
Kinorhyncha (mud dragons)
Priapulida (penis worms)
N+L+P
Nematoida
Nematoda (roundworms)
Nematomorpha (horsehair worms)
L+P
Loricifera (corset animals)
Panarthropoda
Onychophora (velvet worms)
Arthropoda (insects and relatives)
Tardigrada (waterbears)
Spiralia
Gnathifera
Chaetognatha (arrow worms)
Gnathostomulida (jaw worms)
M+R
Micrognathozoa
(Limnognathia)
Rotifera (wheel animals inc. acanthocephalans)
Platytrochozoa
R+M
Rouphozoa
Platyhelminthes (flatworms)
Gastrotricha (hairybacks)
Mesozoa
Orthonectida
Dicyemida or Rhombozoa
Monoblastozoa (
Salinella
Lophotrochozoa
Cycliophora (
Symbion
Annelida (earth worms and relatives)
M+K
Mollusca (snails and relatives)
Kryptotrochozoa
Nemertea (ribbon worms)
Lophophorata
Bryozoa s.l.
Entoprocta or Kamptozoa
Ectoprocta (moss animals)
Brachiozoa
Brachiopoda (lamp shells)
Phoronida (horseshoe worms)
The
phylogeny
of Bilateria
is disputed
; see also
Nephrozoa
Deuterostomia
Xenambulacraria
Centroneuralia
Major groups
within phyla
Sponges
Demosponges
Glass sponges
Calcareous sponges
Cnidarians
Anthozoans inc. corals
Medusozoans inc. jellyfish
Myxozoans
Chordates
Lancelets
Tunicates
Vertebrates
Echinoderms
Sea lilies
Asterozoans inc. starfish
Echinozoans inc. sea urchins
Hemichordates
Acorn worms
Pterobranchs
Nematodes
Chromadorea
Enoplea
Secernentea
Arthropods
Chelicerates inc. arachnids
Myriapods
Pancrustaceans inc. hexapods
Rotifera
Bdelloidea
Monogononta
Seisonidae
Acanthocephala
Platyhelminths
Turbellaria
Trematoda
Monogenea
Cestoda
Ectoproctans
Phylactolaemata
Stenolaemata
Gymnolaemata
Annelids
Polychaetes
Clitellata
Sipuncula
Molluscs
Gastropods
Cephalopods
Bivalves
Chitons
Tusk shells
Phyla with ≥1000 extant species
bolded
Potentially
dubious phyla
Extant
chordate
classes
Kingdom
Animalia
Clade
Bilateria
Superphylum
Deuterostomia
Cephalochordata
Leptocardii (lancelets)
Olfactores
Tunicata
(Urochordata)
Appendicularia (larvaceans)
Acopa
Stolidobranchia
Thaliacea (pyrosomes, salps, doliolids)
Enterogona
Phlebobranchia
Aplousobranchia
Vertebrata
Cyclostomata
Myxini (hagfish)
Hyperoartia (lampreys)
Gnathostomata
(jawed vertebrates)
Chondrichthyes (cartilaginous fish: sharks, rays, chimaeras)
Euteleostomi
(bony vertebrates)
Actinopterygii (ray-finned fish)
Sarcopterygii
(lobe-finned fish)
Actinistia (coelacanths)
Rhipidistia
Dipnoi (lungfish)
Tetrapoda
Lissamphibia (modern amphibians: frogs, salamanders, caecilians)
Amniota
Mammalia (mammals)
Sauria
Lepidosauria
Rhynchocephalia (tuatara)
Squamata (scaled reptiles: snakes, lizards, worm lizards)
Archelosauria
Testudines (turtles)
Archosauria
Crocodilia (crocodilians)
Aves (birds)
¹orders of class
Ascidiacea (sea squirts)
²classes of clade
Sarcopterygii (lobe-finned fish and descendants)
³orders of traditional class
Reptilia (reptiles)
italics
denote paraphyletic groups
Taxon identifiers
Amphioxiformes
Wikidata
Q1156226
ADW
Amphioxiformes
BOLD
343
EoL
2773790
EPPO
1AMPXO
GBIF
730
iNaturalist
152875
IRMNG
12169
ITIS
685568
NBN
NHMSYS0020787064
NCBI
2682553
NZOR:
4c2b8f61-5b7a-4caf-aa4b-1037907a0bb6
Leptocardii
Wikidata
Q20722244
BOLD
359990
CoL
DR
EPPO
1LEPTC
GBIF
7375758
IRMNG
1045
NBN
NHMSYS0020787058
NCBI
2682552
Open Tree of Life
5665514
WoRMS
104897
Leptocardia
Wikidata
Q21286775
Wikispecies
Leptocardia
ITIS
201890
Branchiostomidae
Wikidata
Q2747865
Wikispecies
Branchiostomidae
ADW
Branchiostomatidae
AFD
Branchiostomidae
EoL
2773791
EPPO
1BRNSF
GBIF
2016
iNaturalist
318011
IRMNG
109182
ITIS
159680
NBN
NHMSYS0000065830
Open Tree of Life
176551
Paleobiology Database
352170
WoRMS
104899
Branchiostomatidae
Wikidata
Q61882392
BOLD
178131
CoL
7NG6D
GBIF
5296
IRMNG
106210
ITIS
685569
NBN
NHMSYS0021053186
NCBI
7736
Open Tree of Life
176551
WoRMS
196078
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