Subphylum of arthropods
Crustaceans
are a group of
arthropods
that are a part of the
subphylum
Crustacea
(
), a large, diverse group of mainly
aquatic
arthropods
including
decapods
(
shrimps
,
prawns
,
crabs
,
lobsters
and
crayfish
),
seed shrimp
,
branchiopods
,
fish lice
,
krill
,
remipedes
,
isopods
,
barnacles
,
copepods
,
opossum shrimps
,
amphipods
and
mantis shrimp
.
[1]
The crustacean group can be treated as a subphylum under the
clade
Mandibulata
. It is now well accepted that the
hexapods
(
insects
and
entognathans
) emerged deep in the Crustacean group, with the completed group referred to as
Pancrustacea
.
[2]
The three classes
Cephalocarida
,
Branchiopoda
and
Remipedia
are more closely related to the hexapods than they are to any of the other crustaceans (
oligostracans
and
multicrustaceans
).
[3]
The 67,000 described species range in size from
Stygotantulus stocki
at 0.1 mm (0.004 in), to the
Japanese spider crab
with a leg span of up to 3.8 m (12.5 ft) and a mass of 20 kg (44 lb). Like other
arthropods
, crustaceans have an
exoskeleton
, which they
moult
to grow. They are distinguished from other groups of arthropods, such as
insects
,
myriapods
and
chelicerates
, by the possession of
biramous
(two-parted) limbs, and by their
larval forms
, such as the
nauplius
stage of
branchiopods
and
copepods
.
Most crustaceans are free-living
aquatic animals
, but some are
terrestrial
(e.g.
woodlice
,
sandhoppers
), some are
parasitic
(e.g.
Rhizocephala
,
fish lice
,
tongue worms
) and some are
sessile
(e.g.
barnacles
). The group has an extensive
fossil record
, reaching back to the
Cambrian
. More than 7.9 million tons of crustaceans per year are harvested by fishery or farming for human consumption,
[4]
consisting mostly of
shrimp and prawns
.
Krill
and
copepods
are not as widely fished, but may be the animals with the greatest
biomass
on the planet, and form a vital part of the food chain. The scientific study of crustaceans is known as
carcinology
(alternatively,
malacostracology
,
crustaceology
or
crustalogy
), and a scientist who works in carcinology is a
carcinologist
.
Anatomy
[
edit
]
A shed
carapace
of a
lady crab
, part of the hard
exoskeleton
Body structure of a typical crustacean ? krill
The body of a crustacean is composed of segments, which are grouped into three regions: the
cephalon
or
head
,
[5]
the
pereon
or
thorax
,
[6]
and the
pleon
or
abdomen
.
[7]
The head and thorax may be fused together to form a
cephalothorax
,
[8]
which may be covered by a single large
carapace
.
[9]
The crustacean body is protected by the hard
exoskeleton
, which must be
moulted
for the animal to grow. The shell around each somite can be divided into a dorsal
tergum
, ventral
sternum
and a lateral pleuron. Various parts of the exoskeleton may be fused together.
[10]
: 289
Each
somite
, or body segment can bear a pair of
appendages
: on the segments of the head, these include two pairs of
antennae
, the
mandibles
and
maxillae
;
[5]
the thoracic segments bear
legs
, which may be specialised as
pereiopods
(walking legs) and
maxillipeds
(feeding legs).
[6]
Malacostraca and Remipedia (and the hexapods) have abdominal appendages. All other classes of crustaceans have a limbless abdomen, except from a
telson
and
caudal rami
which is present in many groups.
[11]
[12]
The abdomen in malacostracans bears
pleopods
,
[7]
and ends in a telson, which bears the
anus
, and is often flanked by uropods to form a
tail fan
.
[13]
The number and variety of
appendages
in different crustaceans may be partly responsible for the group's success.
[14]
Crustacean
appendages
are typically
biramous
, meaning they are divided into two parts; this includes the second pair of antennae, but not the first, which is usually
uniramous
, the exception being in the Class Malacostraca where the antennules may be generally biramous or even triramous.
[15]
[16]
It is unclear whether the biramous condition is a derived state which evolved in crustaceans, or whether the second branch of the limb has been lost in all other groups.
Trilobites
, for instance, also possessed biramous appendages.
[17]
The main body cavity is an
open circulatory system
, where blood is pumped into the
haemocoel
by a
heart
located near the dorsum.
[18]
Malacostraca have
haemocyanin
as the oxygen-carrying pigment, while copepods, ostracods, barnacles and branchiopods have
haemoglobins
.
[19]
The alimentary canal consists of a straight tube that often has a gizzard-like "gastric mill" for grinding food and a pair of digestive glands that absorb food; this structure goes in a spiral format.
[20]
Structures that function as kidneys are located near the antennae. A brain exists in the form of ganglia close to the antennae, and a collection of major ganglia is found below the gut.
[21]
In many
decapods
, the first (and sometimes the second) pair of pleopods are specialised in the male for sperm transfer. Many terrestrial crustaceans (such as the
Christmas Island red crab
) mate seasonally and return to the sea to release the eggs. Others, such as
woodlice
, lay their eggs on land, albeit in damp conditions. In most decapods, the females retain the eggs until they hatch into free-swimming larvae.
[22]
Ecology
[
edit
]
Abludomelita obtusata
, an
amphipod
Most crustaceans are aquatic, living in either marine or
freshwater
environments, but a few groups have
adapted
to life on land, such as
terrestrial crabs
,
terrestrial hermit crabs
, and
woodlice
. Marine crustaceans are as ubiquitous in the oceans as insects are on land.
[23]
[24]
Most crustaceans are also
motile
, moving about independently, although a few taxonomic units are
parasitic
and live attached to their hosts (including
sea lice
,
fish lice
,
whale lice
,
tongue worms
, and
Cymothoa exigua
, all of which may be referred to as "crustacean lice"), and adult barnacles live a
sessile
life ? they are attached headfirst to the substrate and cannot move independently. Some branchiurans are able to withstand rapid changes of
salinity
and will also switch hosts from marine to non-marine species.
[25]
: 672
Krill
are the bottom layer and the most important part of the food chain in
Antarctic
animal communities.
[26]
: 64
Some crustaceans are significant
invasive species
, such as the Chinese mitten crab,
Eriocheir sinensis
,
[27]
and the Asian shore crab,
Hemigrapsus sanguineus
.
[28]
Since the piercing of the
Suez Canal
, close to 100 species of crustaceans from the Red Sea and the Indo-Pacific realm have established themselves in the eastern Mediterranean sub-basin, with often significant impact on local ecosystems.
[29]
Life cycle
[
edit
]
Eggs of
Potamon fluviatile
, a freshwater crab
Zoea
larva of the
European lobster
,
Homarus gammarus
Mating system
[
edit
]
Most crustaceans have
separate sexes
, and
reproduce sexually
. In fact, a recent study explains how the male
T. californicus
decide which females to mate with by dietary differences, preferring when the females are algae-fed instead of yeast-fed.
[30]
A small number are
hermaphrodites
, including
barnacles
,
remipedes
,
[31]
and
Cephalocarida
.
[32]
Some may even change sex during the course of their life.
[32]
Parthenogenesis
is also widespread among crustaceans, where viable eggs are produced by a female without needing fertilisation by a male.
[30]
This occurs in many
branchiopods
, some
ostracods
, some
isopods
, and certain "higher" crustaceans, such as the
Marmorkrebs
crayfish.
Eggs
[
edit
]
In many crustaceans, the fertilised eggs are released into the
water column
, while others have developed a number of mechanisms for holding on to the eggs until they are ready to hatch. Most
decapods
carry the eggs attached to the
pleopods
, while
peracarids
,
notostracans
,
anostracans
, and many
isopods
form a
brood pouch
from the
carapace
and thoracic limbs.
[30]
Female Branchiura do not carry eggs in external ovisacs but attach them in rows to rocks and other objects.
[33]
: 788
Most
leptostracans
and
krill
carry the eggs between their thoracic limbs; some
copepods
carry their eggs in special thin-walled sacs, while others have them attached together in long, tangled strings.
[30]
Larvae
[
edit
]
Crustaceans exhibit a number of larval forms, of which the earliest and most characteristic is the
nauplius
. This has three pairs of
appendages
, all emerging from the young animal's head, and a single naupliar eye. In most groups, there are further larval stages, including the
zoea
(pl. zoeæ or zoeas
[34]
). This name was given to it when naturalists believed it to be a separate species.
[35]
It follows the
nauplius
stage and precedes the
post-larva
. Zoea larvae swim with their thoracic
appendages
, as opposed to nauplii, which use cephalic appendages, and megalopa, which use abdominal appendages for swimming. It often has spikes on its
carapace
, which may assist these small organisms in maintaining directional swimming.
[36]
In many
decapods
, due to their accelerated development, the zoea is the first larval stage. In some cases, the zoea stage is followed by the mysis stage, and in others, by the megalopa stage, depending on the crustacean group involved.
Providing camouflage against predators, the otherwise black eyes in several forms of swimming larvae are covered by a thin layer of crystalline
isoxanthopterin
that gives their eyes the same color as the surrounding water, while tiny holes in the layer allow light to reach the retina.
[37]
As the larvae mature into adults, the layer migrates to a new position behind the retina where it works as a backscattering mirror that increases the intensity of light passing through the eyes, as seen in many nocturnal animals.
[38]
DNA repair
[
edit
]
In an effort to understand whether
DNA repair
processes can protect crustaceans against
DNA damage
, basic research was conducted to elucidate the repair mechanisms used by
Penaeus monodon
(black tiger shrimp).
[39]
Repair of DNA double-strand breaks was found to be predominantly carried out by accurate
homologous recombinational
repair. Another, less accurate process,
microhomology-mediated end joining
, is also used to repair such breaks. The expression pattern of DNA repair related and DNA damage response genes in the intertidal copepod
Tigriopus japonicus
was analyzed after ultraviolet irradiation.
[40]
This study revealed increased expression of proteins associated with the DNA repair processes of
non-homologous end joining
,
homologous recombination
,
base excision repair
and
DNA mismatch repair
.
Classification and phylogeny
[
edit
]
Copepods, from
Ernst Haeckel
's 1904 work
Kunstformen der Natur
Decapods, from Ernst Haeckel's 1904 work
Kunstformen der Natur
The name "crustacean" dates from the earliest works to describe the animals, including those of
Pierre Belon
and
Guillaume Rondelet
, but the name was not used by some later authors, including
Carl Linnaeus
, who included crustaceans among the "
Aptera
" in his
Systema Naturae
.
[41]
The earliest
nomenclatural
valid work to use the name "Crustacea" was
Morten Thrane Brunnich
's
Zoologiæ Fundamenta
in 1772,
[42]
although he also included
chelicerates
in the group.
[41]
The subphylum Crustacea comprises almost 67,000 described
species
,
[43]
which is thought to be just
1
⁄
10
to
1
⁄
100
of the total number as most species remain as yet
undiscovered
.
[44]
Although most crustaceans are small, their morphology varies greatly and includes both the largest arthropod in the world ? the
Japanese spider crab
with a leg span of 3.7 metres (12 ft)
[45]
? and the smallest, the 100-
micrometre
-long (0.004 in)
Stygotantulus stocki
.
[46]
Despite their diversity of form, crustaceans are united by the special
larval
form known as the
nauplius
.
The exact relationships of the Crustacea to other taxa are not completely settled as of April 2012
[update]
. Studies based on morphology led to the
Pancrustacea
hypothesis,
[47]
in which Crustacea and
Hexapoda
(
insects
and allies) are
sister groups
. More recent studies using
DNA sequences
suggest that Crustacea is
paraphyletic
, with the hexapods nested within a larger Pancrustacea
clade
.
[48]
[49]
The traditional classification of Crustacea based on morphology recognised four to six classes.
[50]
Bowman and Abele (1982) recognised 652 extant families and 38 orders, organised into six classes:
Branchiopoda
,
Remipedia
,
Cephalocarida
,
Maxillopoda
,
Ostracoda
, and
Malacostraca
.
[50]
Martin and Davis (2001) updated this classification, retaining the six classes but including 849 extant families in 42 orders. Despite outlining the evidence that Maxillopoda was non-monophyletic, they retained it as one of the six classes, although did suggest that Maxillipoda could be replaced by elevating its subclasses to classes.
[51]
Since then phylogenetic studies have confirmed the polyphyly of Maxillipoda and the paraphyletic nature of Crustacea with respect to Hexapoda.
[52]
[53]
[54]
[55]
Recent classifications recognise ten to twelve classes in Crustacea or Pancrustacea, with several former maxillopod subclasses now recognised as classes (e.g.
Thecostraca
,
Tantulocarida
,
Mystacocarida
,
Copepoda
,
Branchiura
and
Pentastomida
).
[56]
[57]
The following cladogram shows the updated relationships between the different extant groups of the paraphyletic
Crustacea
in relation to the class
Hexapoda
.
[53]
According to this diagram, the Hexapoda are deep in the Crustacea tree, and any of the Hexapoda is distinctly closer to e.g. a Multicrustacean than an Oligostracan is.
Fossil record
[
edit
]
Eryma mandelslohi
, a fossil
decapod
from the
Jurassic
of
Bissingen an der Teck
,
Germany
Crustaceans have a rich and extensive
fossil record
, which begins with animals such as
Canadaspis
and
Perspicaris
from the
Middle Cambrian
age
Burgess Shale
.
[58]
[59]
Most of the major groups of crustaceans appear in the fossil record before the end of the Cambrian, namely the
Branchiopoda
,
Maxillopoda
(including
barnacles
and
tongue worms
) and
Malacostraca
; there is some debate as to whether or not Cambrian animals assigned to Ostracoda are truly
ostracods
, which would otherwise start in the
Ordovician
.
[60]
The only classes to appear later are the
Cephalocarida
,
[61]
which have no fossil record, and the
Remipedia
, which were first described from the fossil
Tesnusocaris goldichi
, but do not appear until the
Carboniferous
.
[62]
Most of the early crustaceans are rare, but fossil crustaceans become abundant from the
Carboniferous period
onwards.
[58]
Norway lobsters
on sale at a Spanish market
Within the Malacostraca, no fossils are known for
krill
,
[63]
while both
Hoplocarida
and
Phyllopoda
contain important groups that are now extinct as well as extant members (Hoplocarida:
mantis shrimp
are extant, while
Aeschronectida
are extinct;
[64]
Phyllopoda:
Canadaspidida
are extinct, while
Leptostraca
are extant
[59]
).
Cumacea
and
Isopoda
are both known from the
Carboniferous
,
[65]
[66]
as are the first true mantis shrimp.
[67]
In the
Decapoda
,
prawns
and
polychelids
appear in the Triassic,
[68]
[69]
and
shrimp
and
crabs
appear in the
Jurassic
.
[70]
[71]
The fossil burrow
Ophiomorpha
is attributed to ghost shrimps, whereas the fossil burrow
Camborygma
is attributed to crayfishes. The Permian?Triassic deposits of Nurra preserve the oldest (Permian: Roadian) fluvial burrows ascribed to ghost shrimps (Decapoda: Axiidea, Gebiidea) and crayfishes (Decapoda: Astacidea, Parastacidea), respectively.
[72]
However, the great radiation of crustaceans occurred in the
Cretaceous
, particularly in crabs, and may have been driven by the
adaptive radiation
of their main predators,
bony fish
.
[71]
The first true
lobsters
also appear in the Cretaceous.
[73]
Consumption by humans
[
edit
]
Many crustaceans are consumed by humans, and nearly 10,700,000
tons
were harvested in 2007; the vast majority of this output is of
decapod crustaceans
:
crabs
,
lobsters
,
shrimp
,
crawfish
, and
prawns
.
[74]
Over 60% by weight of all crustaceans caught for consumption are shrimp and prawns, and nearly 80% is produced in Asia, with China alone producing nearly half the world's total.
[74]
Non-decapod crustaceans are not widely consumed, with only 118,000 tons of
krill
being caught,
[74]
despite krill having one of the greatest
biomasses
on the planet.
[75]
See also
[
edit
]
References
[
edit
]
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(1911).
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- Powers, M., Hill, G., Weaver, R., & Goymann, W. (2020). An experimental test of mate choice for red carotenoid coloration in the marine copepod Tigriopus californicus. Ethology., 126(3), 344?352.
An experimental test of mate choice for red carotenoid coloration in the marine copepod Tigriopus californicus
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