Brachiopoda Lecture Notes

A S Maurya, IITR

The brachiopods are a phylum of benthonic marine organisms in which the soft parts
are enclosed in a pair of hinged calcareous valves. Although similar to bivalves in this
and in their filter-feeding habit, they are in fact not closely related to them. In
brachiopods the two valves are different in size but symmetrical about a median
plane, whereas in bivalves they are equal in size but inequilateral. Brachiopods are
very abundant in the fossil record, particularly in Palaeozoic shallow-water marine
deposits, and range from the Cambrian to the Present. They are not, however, as
abundant today as they were in the past, and of the 3000 described genera only 100
are modern. Their organization reflects their fixed mode of life; they have no
locomotory devices and no highly developed sense-organs. Their only protection
from predators or other dangers, apart from being inconspicuous, is to close their
valves. Like most other sessile or attached organisms they feed by filtering nutritive
particles from sea water. Most fossil brachiopods appear to have been limited to the
same way of life and their basic anatomical and physiological organization has
remained unchanged throughout their geological history. Within these limits,
however, the brachiopods display a remarkable diversity of form and habit.

Morphology
The brachiopod shell is usually anchored to the sea floor by a muscular stalk, the
pedicle, which projects through the pedicle opening in the posterior end of one valve
(Fig. 1). This pedicle valve is ventral in relation to the organs of the animal and is
thus sometimes called the ventral valve. The other dorsal valve is frequently termed
the brachial valve because of the presence of the brachidium, a calcareous internal
structure that supports the lophophore or food-gathering apparatus. The two valves
fit closely along the commissure when closed, giving protection to the tissues of the
organism. The hinge mechanism at the posterior end of the valves consists of
interlocking teeth and sockets. Internal muscles control valve movement; adductor
muscles connect the valves and operate to close them; didductor muscles which
operate around the hinge, can open them. Hinges may be strophic, in which case the
hinge axis is coincident with the edge of the valve, or non-strophic in which case the
edges of the shell are curved. Between the hinge and the initial growing area of the
valve, the beak, lies an area termed the interarea. This is the site of the pedicle
opening in the pedicle valve; in some brachiopods this opening may be represented
by a broad triangular notch termed the delthyrium, through which the pedicle
projects.

a
 b

Fig. 1 a, b. Hard-part morphology and symmetry element of a typical brachiopod.

Taxonomy
Brachiopods have long been divided into two main classes: the Inarticulata, in which
there are no teeth and sockets and the pedicle projects between the valves, and the
Articulata, which possess a functional hinge and a pedicle opening. More recently,
however, the inarticulates have been further subdivided and a new class, the
Lingulata, has been recognized. These are brachiopods in which the valves are
chitinophosphatic rather than being calcareous, as in all other brachiopods, probably
because of their early adaptation to coastal environments with extreme salinity
changes that most marine organisms cannot tolerate. They include the well-known
genus Lingula, which has remained essentially unchanged since the Cambrian.
Lingulates are the only known burrowing brachiopods. They use the two valves in
rotary and scissoring motions to dig down anteriorly and then follow a U-shaped
trail, coming to the surface in feeding position. Although they were important
initially, lingulates and inarticulates form only 5 per cent of the known brachiopod
genera, and it is the articulates that are most important and abundant. They are
divided into six orders based on features of the gross morphology (Fig. 2). The
Orthida were the earliest; they are biconvex forms with non-strophic hinges, large
interareas, and delthyrium. These were followed by the Strophomenida, with convex
pedicle and concave brachial valves, and strophic hinge-lines. The Pentamerida were
biconvex with incurved beaks and were characterized by an internal muscle platform,
the spondylium. The Rhynchonellida are small non-strophic forms, typically with
strongly ribbed valves. The Spiriferida also had strongly ribbed valves but were
strophic and possessed spiral brachidia. Finally, the Terebratulida are biconvex and
non-strophic. The differences in morphology are in all cases related closely to the
mode of life of the organisms.
.

Fig. 2. Typical members of the main orders of brachiopods, illustrating the general
character of the shells. In each case the anterior or posterior and brachial views are
shown.

Shell Ornamentation
Internal Morphology
Shell Structure
Evolutionary history
The earliest brachiopods appeared in the Early Cambrian and include abundant
lingulates and the first articulates, the orthids. This fauna became more abundant
and diverse through the Cambrian and pentamerids are first seen in Upper Cambrian
rocks. All the classes became more diverse in the Ordovician, although the articulates
were greatly at an advantage and assumed dominance from the inarticulates and
lingulates at this time. The inarticulates and lingulates declined from this point and
by the Devonian were reduced to a few stable stocks, which have survived almost
unchanged to the present day. The articulates achieved their most significant
evolutionary radiation during the Ordovician, adding the strophomenids,
rhynchonellids, and spiriferids. The earlier orders reduced somewhat during the
Silurian and, towards the end of the period, the last new order, the terebratulids, was
added. The Devonian was chiefly a period of expansion, but major extinctions of
invertebrate faunas at the end of the Devonian took their toll and the pentamerids
were lost. The later Palaeozoic was a time of expansion, particularly for the
strophomenids, since they were able to colonize quasi-infaunal habitats. However,
the severe extinctions of the Late Permian affected the brachiopods as they did all
other invertebrate groups, and by the end of the Jurassic the only articulate orders
left were the terebratulids and the rhynchonellids, both of which are still extant. The
decline of brachiopods in the Mesozoic is probably related to competition with
bivalves. Once the strophomenids had become extinct the bivalves were able to
colonize habitats within the sediment (infaunal), which were thenceforth denied to
brachiopods. These habitats provided the bivalves with protection from predators,
particularly starfish, while the brachiopods remained vulnerable.

Ecology
Brachiopods are normally part of the sessile epifauna and as such they show many
and varied adaptations to different types of benthonic existence. Many were attached
through life by the pedicle, either directly to hard substrates or into soft sediments by
the division of the end of the pedicle into numerous threads. It has been
demonstrated that some brachiopods with long thin pedicles would act like kites,
‘flying’ in water currents. This would result in a region of reversed flow round the
anterior edge that would provide them with feeding currents. Some forms attached
themselves directly to the substrate by the pedicle valve. In some species this was a
juvenile development; once the brachiopod became large enough it would break off
and become free-lying. Attached forms are prominent members of reef associations
in the Permian of Texas. Here conical forms known as richthofeniids attached
themselves to the hard substrate by long tubular spines, forming an interlacing
network that contributed to the reef fabric. They may have generated currents by
flapping the small brachial valve. Many brachiopods had a closed pedicle opening
and must have been free-lying on the sea floor. This resulted in adaptations such as
dish-like valves to spread weight on unstable substrates and radiating spines that
supported the animal. In the spiriferids the interareas were developed as broad
stable platforms, and in some species the shell was thickened in the basal area to
provide additional stability. Burrowing forms are rare; only the lingulids exploited
this way of life; many of the strophomenids, however, particularly the productids,
lived a partially buried or quasi-infaunal existence, held in place by strong spines.
Short spines were present on the brachial valves to hold sediment and prevent it
being winnowed away; the animals were thus effectively hidden from predators.
Communities and distribution
Brachiopods are found in recurring associations commonly composed of several
species. These appear to be characteristic of particular environments and are
presumably related to physical parameters such as temperature, water depth,
salinity, and substrate type. Work on the Ordovician of Wales in 1983 by Martin
Lockley of the University of Colorado has demonstrated the presence of eight
palaeocommunities that persisted until the Silurian. He found that coarse sediments
were inhabited by coarse-ribbed orthids, while the fine sediments supported an
assemblage of inarticulates and small strophomenids. A similar picture is seen in the
Silurian, with five communities inhabiting the same area and forming concentric
belts parallel to what was then the shoreline. This seems to indicate a relationship to
water depth, but factors that vary with depth, such as temperature, pressure,
turbulence, or food supply, may be more important.

On a global basis the distribution of brachiopods is best known from the Palaeozoic,
where they are most abundant, and is similar to the distributions of trilobites and
graptolites. In the Ordovician an early fauna of lingulates and inarticulates occupied
the polar regions while a separate fauna of orthids inhabited the more equatorial
areas. The closer approach of the continental areas during movements of the
lithospheric plates resulted in a lessening of provincialism during the Ordovician and
this was further reduced by the expansion of polar ice-caps at the end of the
Ordovician, which severely affected shallow-water marine faunas. Silurian faunas
were generally more cosmopolitan, but by the Devonian continental movements had
once more resulted in the development of five provinces.

Brachiopods are generally too long-ranging in the Mesozoic to be of more than local
stratigraphic use. They are, however, useful in the Palaeozoic, despite their facies
dependence and provinciality. They have proved to be useful, together with trilobites,
for zoning the shallow-water shelly sequences of the Ordovician and Silurian, where
they provide, in some instances, a greater degree of accuracy than can be provided by
the standard graptolite zones.