ZOOLOGY (H) : SEMESTER III

ZOOACOR05T
Unit 3: Origin of Chordata
Dipleurula concept and the Echinoderm theory of origin of chordates

Evolution of chordate was one of the most important event in the history of chordate as it was the
beginning of evolution of more advanced chordates like bird and mammals. It is the story of origin
from a primitive invertebrate like creatures to early chordates. Though first fossil of the first vertebrate
the ostrachoderm was discovered from the Ordovician period but it might have originated in late
Cambrian period. As early chordates were soft bodied, their fossil records are not preserved. Hence, to
trace their ancestry, we have to find out the similarity among different deuterostomes to trace the
origin of chordates. Some structural features shared by them such as bilateral symmetry, antero-
posterior body axis, triploblastic coelomate condition, etc., may he because of their common ancestry.

Rituparna Maity
Dept.of Zoology
HMMCW, Dakshineswar
 The Phylum Chordata encompasses a vast group of diverse animals ranging from ascidians to man. A few characteristics like Notochord, Dorsal hollow tubular nerve cord and
Pharyngeal gill-slits unite these diverse animals under a common phylum. They differ from the non-chordates significantly by the position of nerve cord. The non-chordates possess a
ventral solid nerve cord below the alimentary canal, while a dorsal hollow nerve cord above the gut is the diagnostic character of the chordates.

• Time of Origin: Chordates evolved sometime during late Cambrian period, 500 million years ago during Cambrian explosion, almost at the same time when invertebrates were beginning to
evolve.

• Place of Origin: Fresh water - Chamberlain (1900) pointed out that all modern chordates possess glomerular kidneys that are designed to remove excess water from body so they have evolved
from freshwater forms. Modern sherks, myxinoidea as well as primitive protochordates posses glomerular kidney making it plausible for chordate origin.
• Chordates evolved from some deuterostome ancestor (echinoderms, hemichordates, pogonophorans etc.) as they have similarities in embryonic development, type of coelom and larval stages.
Fossils of the earliest vertebrates are known from the Silurian-Devonian period, about 400 million years ago.
• DIVISION OF BILATERIA: The Bilateria is divided into two major divisions (1) Protostomia and (2) Deuterostornia. This division is based on the differences in embryonic and larval developments.
Protostomia includes from Annelida to Arthropoda while deuterostomia includes Echinodermata, Pogonophora and Chordate.
• DEUTEROSTOME LINE OF CHORDATE EVOLUTION Following common features of all Deuterostomes suggests strong evidence of a closer evolutionary relationship between the three principal
deuterostome phyla – Echinodermata, Hemichordata and Chordata. i) Early cleavage of zygote is indeterminate ii) Blastopore of gastrula develops into anus. iii) Coelom (enterocoelous except
vertebrates) is formed by the fusion of pockets developed from the endoderm of developing archenteron of the embryo iv) Pelagic larvae of echinoderms and hemichordates have a close
resemblance. Vertebrate does not have a floating larva. v) Deuterostomes use creatine as phosphagen whereas invertebrates use arginine. Some hemichordates as well as echinoids use both.
• The members of the Phylum Chordata possess many features in common.
• The most important features are:
(a) The notochord,
(b) The dorsal tubular nerve cord,
(c) The pharyngeal gill-slits and
(d) A post-anal tail.
These four characteristics are unique to the phylum. Existence of such common structures is interpreted as a result of inheritance from a common ancestry. Besides these four basic structures, there are
a few other characteristics which have less diagnostic value. 2
Chordate Features Shared by the Non-Chordates:
Besides the four unique features of the chordates, there are many characteristics which are also present in many higher invertebrate chordates. The significance of the structural similarities
is very difficult to interpret from the phylogenetic point of view. However, it may be suggested that the chordates as a group evolved from some higher groups of non-chordates and, hence,
the structural resemblances are due to remote common ancestry.
Bilateral Symmetry: Both the chordates and most of the non-chordates like annelids, arthropods, etc. exhibit distinct bilateral symmetry.
Axiate Organization: All the chordates have a distinct polar axis. The anterior end is marked by the presence of head and the posterior end is characterised in most cases by the tail. The
axis extending from the head to the tail end is regarded as the anteroposterior axis.
The anteroposterior axis of the chordates corresponds to that of most of the higher non-chordates. The axiate organization is not strictly homologous, because many fundamental
differences exist between the two groups.
Triploblastic Condition: All animals above the rank of cindarian coelenterates have a third germ layer besides ectoderm and endoderm. This third layer is known as mesoderm.
Although the embryonic formation of the mesoderm is different in non-chordates, its formation is similar in chordates, echinoderms, brachiopods, chaetognaths and in some other
enterocoelous forms. The triploblastic condition has added more weight to the phylogenetic relationship of the chordates with the non-chordates.
Metamerism: Segmental organisation is characteristic of most of the non-chordates and the chordates. In annelids and arthropods, segmentation is well-marked both internally as well as
externally but in chordates the external segmentation is not seen. The segmental arrangement of the body wall musculature is prominent in chordates.
Coelom:The eucoelom or true coelom is the secondary body cavity of triploblastic animals, situated between the gut and body wall. The space of body cavity is lined by coelomic epithelium
and contains coelomic body fluid.The mode of origin of coelom is different among the different groups of invertebrates and chordates. In annelids, arthropods and molluscs the coelom
formation is of schizocoelic type, because the coelom develops by the splitting of the embryonic mesoderm layer.In echinoderms, hemichordates and in other chordates the coelom
formation is of enterocoelic type, or the coelom is called enterocoel, because the coelom develops from the embryonic archenteron or enteron. Here mesoderm arises in the embryo as
paired lateral pouches growing out from the endoderm.These pouches gradually lose continuity with the endoderm and grow downwards and inwards until they meet and fuse. The inner
splanchnic part remains against the wall of developing gut and outer somatic part of the mesoderm becomes applied against the developing body wall.
Embryonic Development: Protostome and deuterostome are the two groups of animals which differ in the embryonic origin of the mouth. Among protostomes, the mouth is formed from
the blastopore, hence protostome means ‘first mouth’. Among the deuterostomes, the mouth does not form from blastopore. Instead it may give rise to anus.
In this group the mouth is the second opening, hence called deuterostome. The differences on the basis of embryological development have strongly supported by analysis of phosphate-
containing storage molecules that are found in muscles and are used in the synthesis of ATP.
Protostomes (e.g., Annelids, arthropods and molluscs) contain arginine phosphate and deuterostomes (e.g., echinoderms and chordates) contain creatine phosphate.

Rituparna Maity
Dept.of Zoology 3
HMMCW, Dakshineswar
Concept of Dipleurula larva : The term dipleura or dipleurula was coined by Semon (1888) but the proper illustration of this hypothetical form was given by Bather in 1900, which
was accepted by most of the zoologists. Majority of echinoderms have indirect development with free swimming and bilaterally symmetrical larval stages. These echinoderms have small
eggs and the fertilized eggs develop in seawater. The cleavage is holoblastic, nearly equal, radial and intermediate to form a hollow one layered ciliated blastula. The blastula transforms
into a gastrula by invagination. The cilia of the gastrula are restricted to (i)a large pre oral band present around the mouth on the ventral side and (ii)a small adoral band lining the mouth
or stomodeum. This larval stage is called Dipleurula larva. This dipleura larval form is regarded as the hypothetical ancestral form of echinoderms as this larva is universally present in
all echinoderms and from it all the larvae of echinoderms have been derived. All well known forms of larvae of echinodermata are derived from the hypothetical dipleurula. Among them
fall the Bipinnaria and the Brachiolaria of the sea stars, the Auricularia of the sea rollers and the plutei of the sea hedgehog etc.

• Echinoderm theory : It was given by Johannes Muller(1860) and is based on the

comparative studies of larval stages of echinoderms and hemichordates. Garstang
and DeBeers proposed the echinoderm larvae gave rise to chordates by
neoteny.

• The theory was given by Muller is based on the comparative studies of larval stages
of echinoderms and hemichordates. Tornaria larva of hemichordates resembles
echinoderm larvae such as Bipinnaria, Auricularia, Dipleurula and Doliolaria, which all
possess ciliary bands and apical tuft of cilia.

• Fossil evidence : The discovery of fossil echinoderms called Calcichordata from

Ordovician period (450 mya) further confirms echinoderm ancestry of chordates.
Calcichordates were asymmetrical animals which demonstrate affinities with both
echinoderms and chordates but their skeleton is made of CaCO3 whereas in
vertebrates the bones are made of hydrated Ca and phosphate. They had large Figure : Hypothetical Dipleurula larva
pharynx with a series of gill slits, each covered with flaps for filter feeding, a small
segmented body and a postanal tail. A perforated pharynx for filter feeding appears to Proposition: The hemichordate larva (tornaria) is strictly similar to the larva
have evolved in diverse groups of animals during Cambrian-Orodovician periods when
planktons were abundant in water. (bipinnaria or dipleurula) of echinoderms. Both are small, transparent, free
swimming, bilaterally symmetrical. Both have similar ciliated band in loops, a
• It is believed that chordates have originated from invertebrates. It is difficult to dorsal pore, sensory cilia at the anterior end and a complete digestive system of
determine from which invertebrates group the Chordates developed. Chordate ventral mouth and posterior anus. This striking larval resemblance Johannes
ancestors were soft bodied animals. Hence they were not preserved as fossils. There Muller and Bateson to suggest a common ancestry for the echinoderms and the
are several theories have been put forwarded to explain the origin of chordates. These
hemichordates.
are directly from some invertebrate group or through the intervention of some
Protochordates. Almost every invertebrate phylum- Coelenterates, Nemertean,
Phoronida, Annelida, Arthropods and Echinoderms has been suggested. But these
theories are far from being satisfactory and convincing and have been only historical
value. Only Echinoderm Theory has received some acceptance. Rituparna Maity
Dept.of Zoology
HMMCW, Dakshineswar
Note : Fossilization of soft bodied animals is very rare occasion. If a soft bodied animal is to fossilized, it must go through the very process without being
decomposed. It means the animal either buried alive and also escape decomposition by fast fossilization in mud volcano conditions, thus a rapid death
caused by devoid of oxygen. The fast fossilization needs very low amount of oxygen and proper burying substratum, under all of this circumstances soft
animal can be fossilized.
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 Supporting Evidence: This theory infers origin of chordates, hemichordates and
echinoderms from a common ancestors. This theory is based on the following
evidence:

• A) Embryological Evidence: Both echinoderms and Chordates have enterocoelic

coelome, mesoderm and deuterostomous mouth. There is resemblances between the
bipinnaria larva of echinoderms and the tornaria larva of hemichordates.

• B) Serological Evidence: A close similarity between the proteins of the body fluid of
chordata and echinoderms. Hence the chordates are more related to echinoderms. Dissimilarity and Doubts:
The radial symmetry of adult echinoderms will disapproved the relationship with the bilaterally
symmetrical chordates. The bilateria is divided into two major divisions- Prostomia and • Presence of apical plate with eyespots in tornaria larva builds doubts the common
Deuterostomia. The division is based on the differences in embryonic and larval ancestry of echinoderms and hemichordates. Garstang and DeBeers proposed the
development. Prostomia includes from Annelida to Arthropoda while Deuterostomia includes Neotenous larva theory suggesting that probably the Auricularia larva of echinoderms
Echinodermata,Pogonophora and Chordates. became sexually mature and later this neotenic larva gave rise to chordates. Cambrian
and Ordovician fossil records of carapoid echinoderm lead Torsten and Gisten to
• C) Deuterostome line of Chordate Evolution: Following common features of assume that carapoid echinoderms might have evolved from tornaria like creatures
Deuterostome suggests strong evidence of a closer evolutionary relationship between the which have began to settle down to lead sedentary life. The water vascular system
three principal Deuterostome phyla- Echinodermata, Hemichordata and Chordata. might have developed out of ciliated grooves of these creatures. Besides this, it was
also claimed that in the lower Silurian period, one carapoid echinoderm had the calyx
(i) Early cleavage of zygote is indeterminate. perforated by a series of 16 small apertures. These apertures can be compared with the
gill slits of Branchiostoma. Some isolated biochemical studies ( Needham,1932 and
(ii) Blastopore of gastrula develops into anus. Whelmi,1942) put some weight on the concept of the diversion of Chordates from
echinodermates. Most of the non chordates use arginine phosphate for the transfer of
(iii) Coelom ( enterocoelous except vertebrates) is formed by the fusion of pockets developed energy but Ophiuronoids, Cephalochordates, ascidian and vertebrates use creatinine
from the endoderm of developing archenteron of the embryo. phosphate. On the other hand, Hemichordata and Echinoderms use both arginine and
creatinine phosphate as phosphate carrier. The descent of chordata from
(iv) Pelagic larva of Echinoderms and Hemichordates have a close resemblance vertebrate echinodermates by the direct transformation of any echinoderms or its neotenous larva
does not have a floating larva. into a chordate is no longer accepted now-a days. Instead they had a common an
immediate ancestor.
(v) Deuterostomes use creatinine as phosphogen whereas invertebrates use arginine. Some
Hemichordates as well as echinoids use both.

Rituparna Maity 5
Dept.of Zoology
HMMCW, Dakshineswar
Other theories: The geological records established beyond doubt that the chordates originated prior to Cambrian period because the relics of some lower chordate forms have
been discovered in Cambrian strata. There are various theories regarding the origin of the chordates from the non-chordate groups. Most of the theories suffer from serious defects.

Of all the theories regarding the ancestry of chordates from some non-chordates, Garstang’s suggestion that the chordates have evolved from some free-swimming echinoderm
larvae (possibly auricularian larvae) by means of paedomorphosis has been accepted by many workers.

The role of paedogenesis (reproduction in pre-adult stage) in evolutionary dynamics is emphasised by many workers on this line. But in recent years the ancestry of the chordates
from the echinoderm source is not accepted.

Recent workers regard the differences between the vertebrates and the non-chordates (invertebrates) to be artificial in nature. Inclusion of the echinoderms, pogonophores and
chordates under deuterostomia (animals where the anus develops from the blastopore and the mouth is formed anew) is accepted nowadays. The protochordates (urochordates and
cephalochordates) are the members of the Phylum Chordata.

The protochordates provide connecting link between the vertebrates with other deuterostomes. The deutorostomes are highly specialised groups and it will be improper to regard
them in the direct line of vertebrate descent.

The phylogenetic status of the hemichordates is a subject of great controversy. But the chordate nature of the urochordates and the cephalochordates is well-established though their
relationships with the vertebrates and with each other are difficult to ascertain.

Barrington (1965) suggested that the deuterostomes have evolved from sessile/semi-sessile ancestors having bilaterally symmetrical and tripartite body and coelom. The
echinoderms have departed a long way from the ancestors, while the hemichordates remained closer.

The hemichordates have developed pharyngotremy (i.e., existence of openings in the pharyngeal wall) which is associated with its ciliary mode of feeding. In course of time a group
with internal food collection mechanism by elaborate and complicated pharynx gave rise to the urochordates, cephalochordates and vertebrates.

Rituparna Maity
Dept.of Zoology
HMMCW, Dakshineswar
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