Brachiopoda and Bivalvia
Brachiopoda and Bivalvia
Brachiopoda and Bivalvia
Joachim de Fourestier
Student Number: 101022736
27/03/2017
Brachiopods and Bivalves are superficially similar, but are rather distantly related. Both
are twin-valved and are filter-feeders. That said, what caused these morphological
convergences or possibly divergences? What similarities are there? What distinguishes
them from one to another?
Joachim de Fourestier Winter 2017
SN#101022736 Brachiopoda and Bivalvia ERTH2312A
Abstract
The Brachiopoda phylum is compared to the Bivalvia class. An attempt is made at
explaining what could have influenced their morphology as well as to explore what
similarities and differences these two groups exhibit. They are commonly confused due
to their similar external shells. Both of these are organisms with “valves” (somewhat
symmetrical), but many differences that exist between the two: these might not seem
quite as obvious at first sight. The most common way of differentiating them is using the
symmetry content these two groups expose and express differently. Brachiopoda have
their plane of symmetry perpendicular to the hinge of the valves. In contrast, bivalvia
(with exceptions from scallops and oysters) have a plane of symmetry parallel to this
hinge. The morphological changes are assumed to be the direct result of natural selection.
However, organisms are known to able to change morphologically during their lifespan.
It has been concluded that the superficial resemblance is from the same ancestors rather
than a result of convergent evolution. Both of them are found to occupy similar niches
but exhibit a few differences. Brachiopods mainly focus on simply keeping a single
location that is stable and have developed many different apparatus in order to achieve
this goal. Bivalves are more flexible in that are mobile. Both are able to burrow down
into the sediment, but bivalves are able to go much deeper or change their lateral location
altogether if necessary. Knowing the significant advantages that bivalves have over
brachiopods and that they can share similar lifestyles, it is somewhat surprising that both
still coexist to this day. This is most likely due to the slightly different environments that
these two can occupy.
Introduction
Brachiopoda and Bivalvia are often confused by their similar exterior, but have some
differences in live styles and internal anatomy. Although the two are not closely related,
their morphology of a twin-valved shell is possibly due to similarities in their ecological
niches. That being said, it is most likely that they originated in a similar niche and
became divergent more internally then externally as they evolved with time. They have
can have similar lifestyles but have come to different adaptations to arising external or
environmental pressures such as salinity, competition, temperature changes, substrate
type, etc.
Brachiopod Morphology
The term ‘Brachiopoda’ comes from Ancient Greek, the prefix brachio– meaning
something that is related to an arm and the suffix –pod meaning foot. Brachiopods,
commonly known as lampshells, are amongst the most successful invertebrate phyla.
Their first appearance datum is within the Early Cambrian which collides with the well-
known and so-called “Cambrian Life Explosion”. They have ever since evolved,
diversified and lived on throughout the Paleozoic dominating low-level benthic
suspension feeding. They are mainly composed of a pedicle (resembling an “arm” or
“lamp stand”) and a shell with two valves (resembling a “foot” or “lamp”). From the
outside, they might seem almost the same as bivalves. However, their feeding methods,
life styles and internal structures (such as soft tissues) are quite definitely different from
one another (see Figure 1).
The two valves of the shell are different: one ventral and one dorsal. One of the
valves is termed the pedicle valve where the stalk-like appendage is attached: this can be
the ventral or dorsal valve depending on the species. Dorsal and ventral are terms used in
relation to the internal morphological features (soft-body parts) of the organism. Bearing
this in mind, brachiopods generally have their pedicle exiting from the ventral valve. The
currently living Magellania is an example of this. The other valve is referred to as the
brachial valve where it contains supports for the lophophores. Following with our last
example, this would be the upper or dorsal valve. Evidently, the pedicle valve is larger
than the brachial valve due to the pedicle opening. Some extinct brachiopods have
evolved and lost their pedicles: a free-living mode. However, they were still benthic,
lying idle on or partially buried in the seafloor. Even though brachiopods tend to have
shells with different size valves, all of them exhibit bilateral symmetry. The plane of
symmetry is perpendicular to the hinge line. In other words, this plane cuts
perpendicularly through both valves of the shell.
The umbo or beak is the initial point where the valve starts growing (or secreting).
Valves with radial ornamentation generally point towards this origin. Opposite to the
posterior or apex (where the beak and hinge is located) is the opening of the valves: this
is the anterior. The margin or trace between this “opening” is termed the commissure. In
one species this can be a relatively straight feature whereas in another, this can be
sinuous. For example, Waconella wacoensis has a relatively linear commissure whereas
the Zygospira modesta has a zig-zag trace. Most brachiopod shells today have a mineral
composition of calcium phosphate and chitin (a complex, long-chain polymer). This is
usually easily recognisable by its enamel-like lustre. Others have calcitic or calcium
carbonate shells like many marine organisms.
Brachiopods use a system of muscles to open and close their valves. As with most
fossils, soft tissue parts such as these muscles are not commonly preserved. However, the
impressions or scars are usually still visible. The adductor scars are where the closing
muscles were attached. Its opposite, the diductor scars, is where the opening muscles
were attached.
Some brachiopods exhibit dentition and were archaically grouped under two
classes: Inacticulata (no teeth or sockets) and Articulata (those with both teeth and
sockets). These teeth are knob-like protrusion located on the hinge of the pedicle valve.
Sockets are small depressions on the hinge of the brachial valve. The teeth fit into the
sockets to assure the appropriate opening and closing of the valves. That being said,
brachiopod dentition is nowhere near as complex as bivalvian dentition (discussed later).
Brachiopod life styles can be classified based on its relation with the substrate.
When the animal lives completely buried within the seafloor, it is known as Infaunal.
Those that do live this way commonly have their posterior oriented downward and can
stabilize themselves by projecting their pedicle further downwards. For those that are
only partially buried are known as Semi Infaunal and are not necessarily attached to the
substrate by their pedicle. Those that are Epifaunal essentially live on or above the
substrate rather than in it. They are generally attached to the seafloor or other objects
(such as marine plants) by their pedicle (discussed later). Reclining or free-living
brachiopods (essentially an unattached lifestyle) are those that are horizontally floating
on (or partial in) the substrate (not to be confused with epifaunal). These will generally
have a plano-convex or concavo-convex shell shape. Some of them are modified to have
spines or larger surface area to help float atop the sediment. Additionally, some of these
have no pedicle opening, but will expose attachment point for those with external spines.
Although brachiopods are sessile, they have developed many different adaptations
(see Figure 2 for following morphologies). Some cement themselves to rocks or other
hard-shell animals such as Craniops or Schuchertella. Others such as Craniids or
Disciniids live together by encrusting hard surfaces (similarly to barnacles). Some species
have clasping spines can attach and cling on to other marine lifeforms and feed from
there instead of on the substrate acting as an epibiont such as Linoproductus and
Brachiopods seem to be able to change their shell shape and life mode during
ontogeny (from birth to maturity). A study from the San Juan Islands, show the shells to
be more lopsided and asymmetric with increasing hydroenergy. (Schumann, 1991)1 This
strongly suggests that morphologic changes not only occur as a result of evolution but
also as a result of ecology.
Bivalve Morphology
The term ‘Bivalvia’ quite evidently comes from prefix bi- meaning two and valve,
and that is quite frankly what they look like. In contrast to Brachiopoda, Bivalvia is a
class, not a phylum. From a taxonomic and ecological standpoint, they are far more
diverse: a taxonomically lower rank expressing more heterogeneity. Like brachiopods,
bivalves (with exceptions from scallops and oysters) are also bilaterally symmetrical with
a twin-valve shelly exoskeleton (see Figure 3). Bivalves first appeared in the Early
Cambrian and started out as burrowers. Their diversity was originally fairly limited. It
was not until the Mesozoic that radiated becoming very successful burrowers and the
now second largest class within the Mollusca phylum. The section is slightly briefer as
many parts found in bivalves are analogous to those of brachiopods.
1
From Introduction to Paleobiology and the Fossil Record
However, some bivalves produce nacre (discussed later). In addition, the valves are
attached together by a toothed hinge (somewhat analogous to interlocking door hinges).
This structure ensures the valves close correctly without misalignment. There are eight
different types of teeth that can be observed with bivalves: desmodont (reduced to
absent), dysodont (small, simple teeth near the edge of the valve), taxodont (multiple
teeth, subequal, subparallel), actinodont (multiple, radially fanning out towards the
umbo), isodont (large teeth located on both side of the ligament’s depression), schizodont
(large, diverging, sometimes grooved), pachyodont (rather large, blunt) and heterodont
(cardinal, very large, lateral).
There are also Epifaunal bivalves. Instead of a pedicle to hold their ground, they
have the ability (that all bivalves have) to secrete byssal threads. These are sticky threads
that are typical used during infancy for when additional stabilization is needed. That
being said, there are some that use this ability during adulthood. The Mytilus is example
of this. They are known as epibyssate bivalves because they use these threads (that reach
down in the substrate) for stability and maintain this life style. This is somewhat
analogous to spider webs. Semi Infaunal or Endobyssates are similar but are partially in
the sediment: they combine the substrate and these thread for even more stability. These
typically have a pointed anterior. Given these points, bivalves that employ byssal threads
during their adulthood are typically found in high current environments. Next, cementing
and reclining bivalves are commonly inequivalved. Just like reclining brachiopods,
reclining bivalves can also have spines to help with stabilization. Unique to bivalve is
swimming. This offers a huge advantage over brachiopods. Swimming bivalves are
usually equilateral but they are inequivalved. The lower valve is usually larger. They
typically exhibit a wider umbonal angle (greater to 105°) and have a single, large,
centrally located adductor muscle.
Although most bivalves have shells that are primarily composed of calcium
carbonate, some can produce a different material known as nacre (such as Pinctada also
known as the “pearl oyster”). Calcareous shells are hard, robust and provide generally
good protection. However, it is very susceptible to chemical weathering and is relative
brittle. Chitin (complex, long-chain polymer) is quite resilient and pliable (so not very
brittle). With this in mind, nacre has the advantage of both calcium carbonate and chitin.
This biomaterial (organic-inorganic) composite consist of multilayered structures of
calcium carbonate and an elastic biopolymer such as chitin. More precisely, it is
comprised of micro aragonite tablets that usually have a rectangular, hexagonal or
rounded shape. This combination gives it a tough, hard structure that will rather deform
that shatter. Nacre has even been used in the material science industry to produce tougher
materials. It is considered to be almost as resilient as silicon. An interesting fact is that
pearls are the result of an autoimmune reaction known as “Nacrezation”: when foreign
material such as an irritant (such as sand) or a parasite, the host will secrete nacre around
it. Sometimes it is incorporated and creates bump within the animal. Otherwise, the
irritant essentially become the nucleus of a pearl. The goal of this behaviour to isolate the
irritant or to the very least creates a smoother surface. This seems to be the next big step
in their evolution to have a greater tolerance towards both mechanical and chemical
weathering.
Conclusion
To summarize, lophopores are too complex and distinctive of a structure for it to
be implicated in convergent evolution. A twin-valved exoskeleton currently seems to
primarily involve the specific lifestyle of filter feeding. Species that have this
morphology are more likely to be from one common point of origin: Lophotrochozoa.
Both Bivalvia and Brachiopoda are members of this superphylum. However, Bivalvia use
siphons for feeding whereas Brachiopoda use lophophores. This is one of the most
important distinctions between the two. Both of them have a shell that can be composed
of calcium carbonate. However, brachiopods tend to have a more phosphoritic
composition (such as chitin which is less susceptible to chemical deterioration). And
bivalve shells are primarily calcareous. That being said, the shell can provide protection
against predators and desiccation. Both have adopted similar forms of burrowing
lifestyles and tend to remain near the substrate. For filter feeding, brachiopods use
lophophores whereas bivalves use siphons. Both started to appear in the early Cambrian:
bivalves already had toothed hinges, but articulate brachiopods only started to appear in
the late Cambrian. Bivalves seem to have taken this evolution step much sooner. The
Jurassic involved losses of both brachiopods and bivalves, but free swimming animals
were not affected. This is most likely due to other lifeforms starting to also take
advantage of the substrate disturbing pre-existing life. It is possible that many of the
brachiopods were tipped over and bivalves were completely submerged by “sediment-
bulldozing” organisms, ultimately leading to the extinction of some species. Bivalves can
live in anywhere from freshwater to brackish waters. With this in mind, they are
commonly found in the shallower subtidal environments whereas brachiopods will prefer
deeper, calmer ocean waters. Both exhibit similar external morphology but their internal
anatomies are quite different. Brachiopods have developed many different complex
structures to achieve stability such as clasping spines and various shell shapes. In
contrast, bivalves have adopted different life modes such as swimming or moving to
different areas using their muscular foot. The concluded distinction is brachiopods
changed morphologically to maintain a lifestyle whereas bivalves changed
morphologically to adopt new lifestyles. That said, both are well adapted to their
environments and have been generally very successful.
References
- “Brachipoda”, Michael J. Benton and David A.T. Harper, “Introduction to
Paleobiology and the Fossil Record”, 2009, p. 298-309
- “Inarticulata: Characters and Anatomy”, University of Bristol, Earth Sciences,
http://palaeo.gly.bris.ac.uk/Palaeofiles/Fossilgroups/Inarticulate/Characters%20an
d%20Anatomy.html , retrieved March 2017
- “The Lophophore”, University of California - Berkeley, Earth Sciences,
http://www.ucmp.berkeley.edu/glossary/gloss7/lophophore.html ,
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- “Brachiopoda: Morphology and Ecology”, State University of New York,
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http://paleo.cortland.edu/tutorial/Bivalves/bivalvemorph.htm ,
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- “Bivalvia”, Michael J. Benton and David A.T. Harper, “Introduction to
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“Introduction to the Lophotrochozoa”, University of California - Berkeley, Earth
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http://palaeo.gly.bris.ac.uk/Palaeofiles/Fossilgroups/Bivalvia/Characters.html,
retrieved March 2017
- “Mollusk shell formation: Mapping the distribution of organic matrix components
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- “Evolution of Immune Reactions”, Petr Sima, Vaclav Vetvicka, CRC Press, 1990,
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Appendix
Figure 1
Figure 2
Retrieved (March 2017) from:“Brachipoda”, Michael J. Benton and David A.T. Harper,
“Introduction to Paleobiology and the Fossil Record”, 2009,
http://www.blackwellpublishing.com/paleobiology/figure.asp?chap=12&fig=Fig12-
9&img=c12f009
Figure 3
Retrieved (March 2017) from: “Bivalvia”, Michael J. Benton and David A.T. Harper,
“Introduction to Paleobiology and the Fossil Record”, 2009,
http://www.blackwellpublishing.com/paleobiology/figure.asp?chap=13&fig=Fig13-
5&img=c13f005
Figure 4
Retrieved (March 2017) from: “Bivalvia”, Michael J. Benton and David A.T. Harper,
“Introduction to Paleobiology and the Fossil Record”, 2009,
http://www.blackwellpublishing.com/paleobiology/figure.asp?chap=13&fig=Fig13-
9&img=c13f009