Isopoda

Isopoda is an order of crustaceans that includes woodlice and

their relatives. Isopods live in the sea, in fresh water, or on land. Isopoda
All have rigid, segmented exoskeletons, two pairs of antennae, Temporal range: Latest
seven pairs of jointed limbs on the thorax, and five pairs of Carboniferous to present
branching appendages on the abdomen that are used in
respiration. Females brood their young in a pouch under their
thorax.

Isopods have various feeding methods: some eat dead or

decaying plant and animal matter, others are grazers, or filter
feeders, a few are predators, and some are internal or external
parasites, mostly of fish. Aquatic species mostly live on the
seabed or bottom of freshwater bodies of water, but some taxa
can swim for a short distance. Terrestrial forms move around by
crawling and tend to be found in cool, moist places. Some Eurydice pulchra, a carnivorous
species are able to roll themselves into a ball as a defense isopod found on sandy shores
mechanism or to conserve moisture.
Scientific classification
There are over 10,000 species of isopod worldwide, with around Kingdom: Animalia
4,500 species found in marine environments, mostly on the
seabed, 500 species in fresh water, and another 5,000 species on Phylum: Arthropoda
land. The order is divided into eleven suborders. The fossil Subphylum: Crustacea
record of isopods dates back to the Carboniferous period (in the
Pennsylvanian epoch), at least 300 million years ago, when Class: Malacostraca
isopods lived in shallow seas. The name Isopoda is derived from Superorder: Peracarida
the Greek roots iso- (from ἴσος ísos, meaning "equal") and -pod
(from ποδ-, the stem of πούς poús, meaning "foot").[2][3] Order: Isopoda
Latreille, 1817 [1]

Suborders
Contents
Asellota
Description
Calabozoida
Diversity and classification
Cymothoida
Evolutionary history
Limnoriidea
Locomotion
Feeding and nutrition Microcerberidea

Reproduction and development Oniscidea

Terrestrial isopods Phoratopidea
References Phreatoicidea
External links Sphaeromatidea
Tainisopidea
Description Valvifera

Classified within the arthropods, isopods have a chitinous

exoskeleton and jointed limbs.[4] Isopods are typically flattened
dorsoventrally (broader than they are deep),[5] although many
species deviate from this rule, particularly parasitic forms, and those
living in the deep sea or in ground water habitats. Their colour may
vary, from grey to white,[6] or in some cases red, green, or
The woodlouse Oniscus asellus brown.[7] Isopods vary in size, ranging from some Microcerberidae
showing the head with eyes and species of just 0.3mm to the deep sea Bathynomus spp. of nearly
antennae, carapace and relatively 50 cm (20 in).[3] Isopods lack an obvious carapace (shell), which is
uniform limbs reduced to a "cephalic shield" covering only the head. This means
that the gill-like structures, which in other related groups are
protected by the carapace, are instead found on specialised limbs on
the abdomen. [3][8] The dorsal (upper) surface of the animal is covered by a series of overlapping,
articulated plates which give protection while also providing flexibility. The isopod body plan consists of a
head (cephalon), a thorax (pereon) with seven segments (pereonites), and an abdomen (pleon) with six
segments (pleonites), some of which may be fused.[5] The head is fused with the first segment of the thorax
to form the cephalon. There are two pairs of unbranched antennae, the first pair being vestigial in land-
dwelling species. The eyes are compound and unstalked and the mouthparts include a pair of maxillipeds
and a pair of mandibles (jaws) with palps (segmented appendages with sensory functions) and lacinia
mobilis (spine-like movable appendages).[9]

The seven free segments of the thorax each bear a pair of unbranched pereopods (limbs). In most species
these are used for locomotion and are of much the same size, morphology and orientation, giving the order
its name "Isopoda", from the Greek equal foot. In a few species, the front pair are modified into gnathopods
with clawed, gripping terminal segments. The pereopods are not used in respiration, as are the equivalent
limbs in amphipods, but the coxae (first segments) are fused to the tergites (dorsal plates) to form epimera
(side plates). In mature females, some or all of the limbs have appendages known as oostegites which fold
underneath the thorax and form a brood chamber for the eggs. In males, the gonopores (genital openings)
are on the ventral surface of segment eight and in the females, they are in a similar position on segment
six.[9]

One or more of the abdominal segments, starting with the sixth segment, is fused to the telson (terminal
section) to form a rigid pleotelson.[9][10][11] The first five abdominal segments each bear a pair of biramous
(branching in two) pleopods (lamellar structures which serve the function of gas exchange, and in aquatic
species serve as gills and propulsion),[3][12] and the last segment bears a pair of biramous uropods
(posterior limbs). In males, the second pair of pleopods, and sometimes also the first, are modified for use in
transferring sperm. The endopods (inner branches of the pleopods) are modified into structures with thin,
permeable cuticles (flexible outer coverings) which act as gills for gas exchange.[9] In some terrestrial
isopods, these resemble lungs.[3]

Diversity and classification

Isopods belong to the larger group Peracarida, which are united by the presence of a special chamber under
the thorax for brooding eggs. They have a cosmopolitan distribution and over 10,000 species of isopod,
classified into 11 suborders, have been described worldwide.[3][13] Around 4,500 species are found in
marine environments, mostly on the sea floor. About 500 species are found in fresh water and another
5,000 species are the terrestrial woodlice, which form the suborder
Oniscidea.[14] In the deep sea, members of the suborder Asellota
predominate, to the near exclusion of all other isopods, having undergone a
large adaptive radiation in that environment.[14] The largest isopod is in the
genus Bathynomus and some large species are fished commercially for
human food in Mexico, Japan and Hawaii.[15]

Some isopod groups have evolved a parasitic lifestyle, particularly as

external parasites of fish.[9] They can damage or kill their hosts and can
cause significant economic loss to commercial fisheries.[16] In reef
aquariums, parasitic isopods can become a pest, endangering the fish and
possibly injuring the aquarium keeper. Some members of the family
Numbers of marine Isopoda
Cirolanidae suck the blood of fish, and others, in the family Aegidae,
(except Asellota and
consume the blood, fins, tail and flesh and can kill the fish in the
crustacean symbionts) in
process.[17] biogeographic regions

The World Marine, Freshwater and Terrestrial Isopod Crustaceans database

subdivides the order into eleven suborders:[1]

Asellota – This suborder contains the superfamily

Aselloidea, a group that contains most of the freshwater
isopods in the northern hemisphere, and the
superfamilies Stenetrioidea, Gnathostenetroidoidea and
Janiroidea, which are mostly marine. The latter
superfamily, Janiroidea, has a massive radiation of
deepsea families, many which have taken bizarre forms.
Calabozoida – A small suborder consisting of two
marine species in the family Calabozoidae and one Representative marine isopod forms
freshwater species in the family Brasileirinidae which is
found in subterranean locations.[18]
Cymothoida – Chiefly marine isopods[9] with over 2,700 species. Members are mostly
carnivorous or parasitic. Includes the family Gnathiidae, the juveniles of which are parasitic
on fishes.[19] The previously recognised suborder Epicaridea is included as two
superfamilies within this suborder and Cymothoida now includes part of the formerly
recognised suborder Flabellifera.[20] Also includes the former suborder Anthuridea, a group
of worm-like isopods with very long bodies.
Limnoriidea – Mainly tropical isopods, some of which are herbivorous.[20]
Microcerberidea – Tiny, worm-like isopods that live between particles on the bed of
freshwater and shallow marine habitats.[9]
Oniscidea – Semi-terrestrial and terrestrial isopods fully adapted for life on land.[9] There are
over 4,000 species of woodlice inhabiting forests, mountains, deserts and the littoral
zone.[21]
Phoratopidea – A single marine species, Phoratopus remex, which warrants its own
suborder because of its unique characteristics.[20]
Phreatoicidea – Small suborder of freshwater isopods resembling amphipods, limited to
South Africa, India, Australia and New Zealand.[9]
Sphaeromatidea – Benthic isopods mostly from the southern hemisphere with respiratory
pleopods inside a branchial chamber. This suborder now includes part of the formerly
recognised suborder Flabellifera.[20][22]
Tainisopidea – Freshwater isopods in a "relictual environment".[20]
 Valvifera – A large group of benthic, marine isopods with respiratory pleopods inside a
branchial chamber under the abdomen.[9]

Evolutionary history
Isopods first appeared in the fossil record during the Carboniferous period of the Paleozoic some 300
million years ago.[23] They were primitive, short-tailed members of the suborder Phreatoicidea. At that
time, Phreatoicideans were marine organisms with a cosmopolitan distribution. Nowadays, the members of
this formerly widespread suborder form relic populations in freshwater environments in South Africa, India
and Oceania, the greatest number of species being in Tasmania. Other primitive, short-tailed suborders
include Asellota, Microcerberidea, Calabozoidea and the terrestrial Oniscidea.[14]

The short-tailed isopods have a short pleotelson and terminal, stylus-like uropods and have a sedentary
lifestyle on or under the sediment on the seabed. The long-tailed isopods have a long pleotelson and broad
lateral uropods which can be used in swimming. They are much more active and can launch themselves off
the seabed and swim for short distances. The more advanced long-tailed isopods are mostly endemic to the
southern hemisphere and may have radiated on the ancient supercontinent of Gondwana soon after it broke
away from Laurasia 200 million years ago. The short-tailed forms may have been driven from the shallow
seas in which they lived by increased predatory pressure from marine fish, their main predators. The
development of the long-tailed forms may also have provided competition that helped force the short-tailed
forms into refugia. The latter are now restricted to environments such as the deep sea, freshwater,
groundwater and dry land. Isopods in the suborder Asellota are by far the most species-rich group of deep
sea isopods.[14]

Locomotion
Unlike the amphipods, marine and freshwater isopods are entirely benthic. This gives them little chance to
disperse to new regions and may explain why so many species are endemic to restricted ranges. Crawling is
the primary means of locomotion, and some species bore into the seabed, the ground or timber structures.
Some members of the Flabellifera can swim to a limited extent and have their front three pairs of pleopods
modified for this purpose, with their respiratory structures limited to the hind pleopods. Most terrestrial
species are slow-moving and conceal themselves under objects or hide in crevices or under bark. The semi-
terrestrial sea slaters (Ligia spp.) can run rapidly on land and many terrestrial species can roll themselves
into a ball when threatened, a feature that has evolved independently in different groups and also in the
marine sphaeromatids.[9]

Feeding and nutrition

Isopods have a simple gut which lacks a midgut section; instead
there are caeca connected to the back of the stomach in which
absorption takes place. Food is sucked into the esophagus, a
process enhanced in the blood-sucking parasitic species, and passed
by peristalsis into the stomach, where the material is processed and
filtered. The structure of the stomach varies, but in many species
there is a dorsal groove into which indigestible material is
channelled and a ventral part connected to the caeca where
Anilocra (Cymothoidae) parasitising intracellular digestion and absorption take place. Indigestible
the fish Spicara maena, Italy material passes on through the hindgut and is eliminated through
the anus, which is on the pleotelson.[9]
Isopods are detritivores, browsers, carnivores (including predators and scavengers), parasites, and filter
feeders, and may occupy one or more of these feeding niches. Only aquatic and marine species are known
to be parasites or filter feeders.[24][25] Some exhibit coprophagia and will also consume their own fecal
pellets.[25] Terrestrial species are in general herbivorous, with woodlice feeding on moss, bark, algae, fungi
and decaying material. In marine isopods that feed on wood, cellulose is digested by enzymes secreted in
the caeca. Limnoria lignorum, for example, bores into wood and additionally feeds on the mycelia of fungi
attacking the timber, thus increasing the nitrogen in its diet. Land-based wood-borers mostly house
symbiotic bacteria in the hindgut which aid in digesting cellulose. There are numerous adaptations to this
simple gut, but these are mostly correlated with diet rather than by taxonomic group.[9]

Parasitic species are mostly external parasites of fish or crustaceans and feed on blood. The larvae of the
Gnathiidae family and adult cymothoidids have piercing and sucking mouthparts and clawed limbs adapted
for clinging onto their hosts. In general, isopod parasites have diverse lifestyles and include Cancricepon
elegans, found in the gill chambers of crabs; Athelges tenuicaudis, attached to the abdomen of hermit crabs;
Crinoniscus equitans living inside the barnacle Balanus perforatus; cyproniscids, living inside ostracods
and free-living isopods; bopyrids, living in the gill chambers or on the carapace of shrimps and crabs and
causing a characteristic bulge which is even recognisable in some fossil crustaceans; and entoniscidae living
inside some species of crab and shrimp.[9][26] Cymothoa exigua is a parasite of the spotted rose snapper
Lutjanus guttatus in the Gulf of California; it causes the tongue of the fish to atrophy and takes its place in
what is believed to be the first instance discovered of a parasite functionally replacing a host structure in
animals.[27]

Reproduction and development

In most species, the sexes are separate and there is little sexual dimorphism, but a few species are
hermaphroditic and some parasitic forms show large differences between the sexes.[9] Some Cymothoidans
are protandrous hermaphrodites, starting life as males and later changing sex, and some Anthuroideans are
the reverse, being protogynous hermaphrodites that are born female. Some Gnathiidans males are sessile
and live with a group of females.[24] Males have a pair of penises, which may be fused in some species.
The sperm is transferred to the female by the modified second pleopod which receives it from the penis and
which is then inserted into a female gonopore. The sperm is stored in a special receptacle, a swelling on the
oviduct close to the gonopore. Fertilisation only takes place when the eggs are shed soon after a moult, at
which time a connection is established between the semen receptacle and the oviduct.[9]

The eggs, which may number up to several hundred, are brooded by the female in the marsupium, a
chamber formed by flat plates known as oostegites under the thorax. This is filled with water even in
terrestrial species.[9] The eggs hatch as mancae, a post-larval stage which resembles the adult except for the
absence of the last pair of pereopods. The lack of a swimming phase in the life cycle is a limiting factor in
isopod dispersal, and may be responsible for the high levels of endemism in the order.[14] As adults,
isopods differ from other crustaceans in that moulting occurs in two stages known as "biphasic
moulting".[3] First they shed the exoskeleton from the posterior part of their body and later shed the anterior
part. The giant Antarctic isopod Glyptonotus antarcticus is an exception, and moults in a single process.[28]

Terrestrial isopods
The majority of crustaceans are aquatic and the isopods are one of the few groups of which some members
now live on land.[29][30] The only other crustaceans which include a small number of terrestrial species are
amphipods (like sandhoppers) and decapods (crabs, shrimp, etc.).[29] Terrestrial isopods play an important
role in many tropical and temperate ecosystems by aiding in the decomposition of plant material through
mechanical and chemical means, and by enhancing the activity of microbes.[31] Macro-detritivores,
including terrestrial isopods, are absent from arctic and sub-arctic
regions, but have the potential to expand their range with increased
temperatures in high latitudes.[32]

The woodlice, suborder Oniscidea, are the most successful group of

terrestrial crustaceans[9] and show various adaptations for life on land.
They are subject to evaporation, especially from their ventral area, and
as they do not have a waxy cuticle, they need to conserve water, often
living in a humid environment and sheltering under stones, bark,
Armadillidium vulgare on the
debris or leaf litter. Desert species are usually nocturnal, spending the
move ...
day in a burrow and emerging at night. Moisture is achieved through
food sources or by drinking, and some species can form their paired
uropodal appendages into a tube and funnel water from dewdrops
onto their pleopods. In many taxa, the respiratory structures on the
endopods are internal, with a spiracle and pseudotrachaea, which
resemble lungs. In others, the endopod is folded inside the adjoining
exopod (outer branch of the pleopod). Both these arrangements help
to prevent evaporation from the respiratory surfaces.[9]

Many species can roll themselves into a ball, a behaviour used in

defence that also conserves moisture. Members of the families ... and rolled into a ball
Ligiidae and Tylidae, commonly known as rock lice or sea slaters, are
the least specialised of the woodlice for life on land. They inhabit the
splash zone on rocky shores, jetties and pilings, may hide under debris
washed up on the shore and can swim if immersed in water.[9]

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du/abs/2011Oecol.167.1163V). doi:10.1007/s00442-011-2051-8 (https://doi.org/10.1007%2
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External links
Media related to Isopoda at Wikimedia Commons
 Data related to Isopoda at Wikispecies
World List of Marine Freshwater and Terrestrial Isopod Crustaceans (http://www.marinespeci
es.org/isopoda/)

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