Chapter 7

BENTHOS

BENTHOS
(Benthic Animals)

Benthic animals (or zoobenthos) are

separated into two ecological categories based
on where they live relative to the substrate.

➢ The infauna are those species that live

wholly or partly within the substrate;
this category includes many clams and
worms (polychaetes) as well as other
invertebrates.

➢ Infaunal species usually dominate

communities in soft substrates, and
they are most diverse and abundant in
subtidal regions. Representative infauna, showing their
burrows and living positions. (a) Hydrobia,
a snail; (b) burrow of Pygospio, a polychaete;
➢ There are a few infaunal species in hard (c) burrow of Corophium, an amphipod; (d)
Arenico/a, a polychaete; and the clams (e)
substrate communities as well, rock- Cardium, (f) Macoma, (g) Scrobicularia, and
boring clams being one example. (h) Mya,

➢ The epifauna are those animals living on or attached to the seafloor;

about 80% of the larger zoobenthos belong to this category.

➢ A few common examples of

epifauna include corals,
barnacles, mussels, many
starfish, and sponges.

➢ Epifauna are present on all

substrate types, but they are
particularly richly developed
on hard substrates, and
they are most abundant and
diverse in rocky intertidal
areas and coral reefs. Representative epifauna and epiflora.

➢ A third category can be added to include those animals that live in

association with the seafloor but also swim temporarily above it; animals
such as prawns and crabs, or flatfish such as sole, form the epibenthos.

CIMAbraham
MBIO 102
 Chapter 7
BENTHOS

The following categories encompass all sizes of benthos:

➢ Macrofauna (or macrobenthos): those animals retained by a 1.0-mm-
mesh sieve. These are the largest benthic animals, including starfish,
mussels, most clams, corals, etc.
➢ Meiofauna (or meiobenthos): those animals retained by a 0.1-1.0-mm-
mesh sieve. These are small animals commonly found in sand or mud.
The group includes very small molluscs, tiny worms, several small
crustacean groups (including benthic copepods), as well as less familiar
invertebrates.
➢ Microfauna (or microbenthos): those animals that are smaller than 0.1
mm in dimension. This smallest size category is largely made up of
protozoans, especially ciliates.

SYSTEMATICS AND BIOLOGY

Benthic communities contain
an extremely diverse assemblage of
zoobenthos. Many of these marine
species have no terrestrial or
freshwater counterparts and are
unfamiliar animals. Some of the
dominant types of animals in benthic
communities are listed in Table 7.1
and illustration below.

Examples of microfauna: ciliate

protozoans showing diversity of form.

CIMAbraham
MBIO 102
 Chapter 7
BENTHOS

The most primitive

multicellular animals are the
sponges, which may constitute a
large fraction of the
macrobenthos in some marine
regions.
➢ Known to exist from late
Precambrian times (>600
million years ago), this
ancient group now has
roughly 10000 species,
almost all of them marine.
➢ They are named for their
porous nature (Phylum
Porifera), and the many
cavities of sponges provide
protective refuges for myriads
of small animals such as
worms and crustaceans. All
sponges are sessile, that is,
they are attached and
immobile. Most filter-feed by
producing currents that draw
suspended particles through
the sponge.

Within the Phylum Cnidaria, the Class Representative benthic animals:

Hydrozoa includes the colonial hydroids, (a) unicellular xenophyophore; (b) sponge;
(c) sea anemone; (d) nemertean; (e)
formed of unions of structurally and polychaete; (f) flatworm; (g) enteropneust
functionally different types of individuals. hemichordate; (h) echiurid; (i) sipunculid;
and (j) pogonophoran. (All scales in mm.)
Although they are usually small and
inconspicuous, a large part of the marine growth attached to rocks, shells, and
wharf pilings and usually called 'seaweed' is actually composed of hydrozoan
colonies.

The much larger Class Anthozoa, with over 6000 species, includes sea
anemones and a variety of corals, as well as less familiar forms such as sea
whips and sea fans. Sea anemones are common residents of intertidal and
subtidal communities, but are also found at over 10000 m depth; they are
solitary animals, ranging in diameter from about 1 cm to more than 1 m.

Benthic worms belong to a number of different phyla.

➢ The threadlike nematodes (Phylum Nematoda) constitute one of
the most numerous and widespread groups of marine (and

CIMAbraham
MBIO 102
 Chapter 7
BENTHOS

terrestrial) animals, although most of the species are inconspicuous

inhabitants of soft sediments.
➢ A single square meter of bottom mud off the Dutch coast was
reported to contain about 4500000 individual meiobenthic
nematodes.

➢ The Phylum Nemertea, encompasses about 600 species of

elongated worms, all characterized by having a long eversible
proboscis that is used to capture food. Nemerteans are more
abundant in temperate seas than in tropical areas, and they are
more common in shallow zones.

➢ Free-living flatworms (Phylum Platyhelminthes) reside in sand or

mud, under stones and shells, or on seaweeds, but they are seldom
present in large numbers.

➢ Sipunculids (Phylum Sipuncula), also called peanut worms, are

unsegmented worms ranging in length from about 2 mm to more
than 0.5 m. Many of the 250 or so species burrow into sand or mud,
using movements of their large proboscis to force their way through
the sediments; others inhabit rock or coral crevices, or even empty
snail shells. They are mostly deposit feeders.

➢ Echiurids (Phylum Echiura) are somewhat similar to sipunculids

in size and general habit. Most species use their large nonretractible
proboscis to forage for food contained in sediments. The majority of
deep-sea echiurid species have dwarf parasitic males attached to
the female, a mode of reproduction that is reminiscent of that of
the deep-sea angler-fish.

➢ More than 10000 species belonging to the Phylum Annelida, Class

Polychaeta, make up the largest and most diverse group of marine
worms. Polychaetes are the segmented worms with multiple
appendages called parapodia. Size ranges from a few millimetres
to 3 m in length. Ecologically, polychaetes can be separated into
those that move actively over the seafloor or burrow into sand and
mud, and those that inhabit permanent tubes or burrows.

➢ Members of the Phylum Mollusca include over 50 000 marine

species, among them the familiar snails and related nudibranchs or
sea slugs (Class Gastropoda), and the bivalved clams and mussels
(Class Bivalvia). This phylum also includes the flattened chitons
(Class Polyplacophora), with a shell divided into eight plates. Less
well-known members are the burrowing scaphopods (Class
Scaphopoda) with tusk-shaped shells, and the wormlike, shell-less
aplacophorans (Class Aplacophora) found within sediments. Most

CIMAbraham
MBIO 102
 Chapter 7
BENTHOS

of the octopus species (Class Cephalopoda) are also essentially

benthic species, although they are capable of swimming. The great
diversity in this phylum is expressed in the facts that molluscs
inhabit all depths of the sea, are found both on and within
sediments, have representative species in all trophic levels, and are
present in all benthic communities.

Echinoderms (Phylum Echinodermata) are exclusively marine animals.

Although differing in external appearance, all echinoderms are characterized by
having radial symmetry, with the body divided into five parts around a central
axis; a skeleton composed of calcareous plates; and tube feet. The approximately
5600 species are divided into
five classes.

➢ The Class Asteroidea

includes roughly 2000
species of starfish (or
seastars) whose habitats
range from intertidal zones
to about 7000 m. Many
starfish are carnivorous,
and they may have
considerable ecological
impact on cultivated
shellfish beds as well as in
natural habitats.

➢ Class Ophiuroidea
comprises almost 2000
species of long-armed
brittle stars and basket
stars. Deep-sea
photographs often show
ophiuroids carpeting the
seafloor, where they feed
on deposited sediments, on Representative benthic echinoderms: (a) feather star
(crinoid); (b) sea cucumber (holothurian); (c) brittle star
small dead or living (ophiuroid); and (d) sea urchin (echinoid). (All scales in
animals, or on suspended mm.)

organic material.

➢ Some 800 species of spiny sea urchins and flattened sand dollars are
placed in the Class Echinoidea. Urchins are conspicuous members of the
macrobenthos of rocky shores, kelp beds, and coral reefs; they use a
special chewing apparatus to feed on all types of organic material, but
most shallow-water species are regarded as basically herbivorous and
deep-sea species (to about 7000 m) are considered to be deposit feeders.

CIMAbraham
MBIO 102
 Chapter 7
BENTHOS

➢ The Class Holothuroidea (with 500 species) includes the elongated sea
cucumbers, named because of their resemblance to the vegetable. The
epibenthic species of holothurians can be either deposit or suspension
feeders; infaunal species swallow the sand or mud in which they live.
Although also found in shallow waters, the greatest number of abyssal
echinoderms are sea cucumbers, and they feature prominently in deep-
sea photographs.

➢ The Class Crinoidea is the most ancient echinoderm group, and presently
includes about 650 species of animals known commonly as feather stars
and sea lilies. Feather stars live mostly in depths above 1500 m, and
although they often cling to the seafloor, they are mobile animals that are
capable of crawling as well as of swimming temporarily.

Bryozoa or moss animals, belong to the

Phylum Ectoprocta. Like the hydroids,
bryozoans are colonial and sessile animals
that form inconspicuous encrustations or
seaweed-like growths on intertidal rocks,
shells, or artificial surfaces. A few species
have also been recorded from depths of
more than 8000 m.
Representative benthic echinoderms: (a)
Brachiopods constitute a phylum feather star (crinoid); (b) sea cucumber
(Brachiopoda) of somewhat less than 300 (holothurian); (c) brittle star (ophiuroid); and
(d) sea urciiin (echinoid). (All scales in mm.)
marine species that superficially resemble
molluscs in having a bivalved calcareous
shell (5-80 mm in diameter), though their
fundamental body-plan is quite different.
Most live in the upper 200 m, and most are
cemented to a hard substrate.

Tunicates (or ascidians) are benthic Representative benthic animals:

relatives of the pelagic larvaceans and (a) scaphopod; (b) bryozoan colony; (c)
stalked brachiopod; (d) two tunicates; (e)
salps. These sessile barrel-shaped animals tanaid; (f) isopod; and (g) a horseshoe crab
belong to the Phylum Chordata, Class (not related to true crabs). (All scales in mm.)
Ascidiacea. Most of the common tunicates
are solitary organisms, but there are also many colonial species that develop by
asexual budding. Ascidians are commonly found in intertidal waters, attached
to rocks, shells, wharves, or other firm substrates, but they also inhabit depths
to at least 8000 m. The free end of a tunicate has two siphons that provide
passage for a current of water drawn through the animal by cilia.

CIMAbraham
MBIO 102
 Chapter 7
BENTHOS

The segmented Crustacea are well represented on the seafloor.

Meiobenthic species include ostracods and the cyclopoid and harpacticoid
copepods. Harpacticoids are an especially abundant group whose members
crawl over or burrow through soft sediments. Also included in this size category
are the tanaids.

Barnacles (Cirripedia) are familiar marine animals and the only sessile
crustaceans. There are about 800 species, including a large number that are
parasitic in other marine invertebrates. These shrimplike animals live within
an external covering of calcareous plates. Some attach directly to substrates,
others are stalked. The more familiar barnacles form crowded aggregations in
rocky intertidal regions, but some species have become specially adapted to
attach to mobile surfaces and live on the bodies of whales, sharks, sea snakes,
manatees, fish, or crabs.

Benthic decapod crustaceans include the familiar crabs, lobsters, and shrimp,
and the group has both epifaunal and infaunal representatives. Decapods show
their greatest diversity in shallower waters, but a few species live at depths of
5000-6000 m. The group includes predators, omnivores, and scavengers. Some
are filter feeders (e.g. burrowing mud shrimp and mole crabs), but detritus rather
than plankton is often the dominant food.

CIMAbraham
MBIO 102