Phylum Chordata: Meaning,

Features and Ancestry |

Biology
In this article we will discuss about Phylum Chordata:- 1.
Meaning of Phylum Chordata 2. Important Features of
Phylum Chordata 3. Non-Chordates 4. Ancestry 5. Outline
Classification.

Contents:

1. Meaning of Phylum Chordata

2. Important Features of Phylum Chordata
3. Chordate Features Shared by the Non-Chordates
4. Ancestry of Phylum Chordata
5. Outline Classification of Phylum Chordata

1. Meaning of Phylum Chordata:

The Phylum Chordata encompasses a vast group of diverse

animals ranging from ascidians to man (Fig. 1.1). 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 pos​sess a ventral solid nerve
cord below the ali​mentary canal, while a dorsal hollow nerve cord
above the gut is the diagnostic character of the chordates. The
study of Phylum Chordata is of special interest to us, because
we, the human beings, are also included with​in this group.

2. Important Features of Phylum Chordata:

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 pha​ryngeal gill-slits (Fig. 1.2) 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 inheri​tance
from a common ancestry. Besides these four basic structures,
there are a few other cha​racteristics which have less diagnostic
value.

Notochord or Chorda Dorsalis:

The notochord is the prime identifying structure of the chordates

and the group derives its name from it. It is a rod-like elastic
structure situated just above the alimentary canal and
immediately beneath the dorsal tubular nerve cord. The
notochord is composed of special type of vacuolated
notochordal cells (Fig. 1.3) and remains covered over by one or
two connective tissue cove​rings, called notochordal sheath.
The noto​chord (Stomochord) in the hemichordates originates
from the embryonic endoderm but ​in other chordates, it is a
mesodermal deriva​tive. The stomochord is not supportive and
possesses a cavity that opens into the pharynx. The notochord
may persist even in the adult primitive chordates, but in
vertebrates it is either partially or wholly replaced by the
vertebral column.

Dorsal Tubular Nerve Cord:

In chordates, a single dorsal tubular nerve cord extends along the

anteroposterior axis of the body. It is located just above the
notochord (Fig. 1.4). This nerve cord originates from the
embryonic dorsal ectoderm. It develops as an oval dorsal plate
called neural or medullary plate. The neural plate is formed as a
result of assemblage of (neural) cells.
The neural plate subsequently transforms into a neural or
medullary groove possibly due to the lateral pressure exerted by
the surrounding non-neu​ral tissue. The dorsal tips of the neural
groove become fused with each other and a neural tube is
formed.

The cavity within the neural tube is called neurocoel. In the

invertebrate chordates the dorsal tubular nerve cord remains
almost unchanged, but in the verte​brates the anterior region of
the nerve cord becomes specialised into the brain and the
remaining part is modified into spinal cord. These two parts
remain connected by a short bridge called isthmus.

The development of dorsal nerve cord is as follows:

Pharyngeal Gill-slits:

The gill-slits have many alternative names, such as pharyngeal or

branchial clefts, visce​ral clefts, visceral or branchial pouches. In
the majority of the primary aquatic chordates, the pharyngeal
wall is perforated by gill-slits which are meant for the exit of
water current from the pharyngeal cavity to the outside.
In the majority of such forms the respiratory organs (gills) are
lodged in the gill-chamber. Exchange of gases takes place during
the tran​sit of the water current that passes through the mouth
and goes out through the gill-slits.

The gill-slits, as such, are persistent in invertebrate chordates,

fishes, amphibian larvae and in the neotenic adult amphibians.
But the transition from the aquatic environment to the land-life
has changed the nature of the gill-slits.

The gill-slits lose their importance due to the acquisition of

pulmonary respiration. The res​piratory function of the gill-slits is
taken up by the lungs, the respiratory organs of the land
chordates. The initial stages of formation of gill-slits are clearly
seen in the embryos of the land chordates (Fig. 1.5). In the
adults, these structures become transformed into other organs,
specially the endocrine systems.

The gill-slits are developed as a result of subsequent fusion of

the in-pouchings of the body ectoderm with the corresponding
out-pouchings of the pharyngeal endoderm.

Initially both the in-pouchings and the out- pouchings meet with
each other and then the regions of union breakthrough to form
conti​nuous passages between the pharyngeal cavi​ty and the
exterior. The number of the gill-slits varies considerably in
different chordates. Cephalodiscus bears only one pair of gill-
slits whereas Branchiostoma possesses as many as two hundred.

Remarks:

According to Kardong (2002) — “The term gill-slits is often used

in place of ‘pharyn​geal slits’. A ‘gill’ proper is a structure that is
composed of tiny plates and harbour capilla​ries, used for aquatic
respiration. In vertebrates, the gills are situated adjacent to the
pharyn​geal slits. The slits do not play any significant role in
respiration, only remain as openings. In many primitive
chordates, the openings only serve primarily in feeding but in
embryos they play no respiratory role, hence gill-slits is a
misleading term.”

Post anal Tail:

Chordates possess a post anal muscular tail containing

extensions of the notochord and nerve cord. Though it is a
characteristic feature of the vertebrates but it may or may not
present in adult stage.

In protochordates, the tail is absent in adult ascidians and

hemichordates, whereas in amphioxus, it is present in adult
stage. In primitive chordates the tail plays as a locomotors organ
but in higher chor​dates or vertebrates the tail is employed in
various functions such as swimming, balan​cing, prehensile and
food capturing.

3. 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
inverte​brate chordates. The significance of the struc​tural
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 resem​blances 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 organiza​tion 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 chor​dates 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 exter​nally
but in chordates the external segmenta​tion 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 endo​derm.

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
deve​loping body wall.

Embryonic Development:

Protostome and deuterostome are the two groups of animals

which differ in the embry​onic 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.

Protos​tomes (e.g., Annelids, arthropods and molluscs) contain

arginine phosphate and deuterostomes (e.g., echinoderms and
chordates) con​tain creatine phosphate.
Although the chordates share many com​mon features with the
non-chordates, the fun​damental organisation is different. The
diffe​rences are shown in Table 4 Chordata.

Nature of Sexes:

Majority of chordates are dioecious (the sexes are separate) but

in a few hemichordates and in majority of urochordates the
hermaphroditism predominates.
4. Ancestry of Phylum Chordata:

The chordates include organisms having a notochord, a dorsal

hollow nerve cord, pharyn​geal slits or pouches and a few
features like bilateral symmetry, axial organisation, triplo​blastic
condition, segmentation, etc., that are common with the non-
chordates. The question of the origin of the chordates still
remains unan​swered and considerable controversy exists on this
issue.

The geological records established beyond doubt that the

chordates originated prior to Cambrian period because the relics
of some lower chordate forms have been discov​ered in Cambrian
strata. There are various the​ories 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 ver​tebrates

and the non-chordates (invertebrates) to be artificial in nature.
Inclusion of the echino​derms, pogonophores and chordates
under deuterostomia (animals where the anus deve​lops 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 bet​ween 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
sym​metrical and tripartite body and coelom. The echinoderms
have departed a long way from the ancestors, while the
hemichordates remai​ned 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 com​plicated
pharynx gave rise to the urochordates, cephalochordates and
vertebrates.

5. Outline Classification of Phylum Chordata:

Like many other phyla, the taxonomical subdivisions of the

Phylum Chordata are a prob​lematic issue. Diverse opinions exist
on this particular aspect. The systematic status of
hemichordates has long been a debated issue. In recent years
the hemichordates have been separated from the Phylum
Chordata and given the rank of a separate phylum.

In this present text the Phylum Chordata is divided into:

Subphylum I. Cephalochordata

Subphylum II. Urochordata

Subphylum III Conodontophorida

Subphylum IV. Vertebrata (Craniata)

The classification of vertebrates is a very difficult task. Despite

remarkable advancement of knowledge on this topic, the
classification of vertebrates is open to refinement. The term
Vertebrata was introduced by Lamarck into the Science of
Zoology. But the classification of the Vertebrata into different
classes remained still a problem.

Aristotle and Linnaeus established four classes [Pisces,

Amphibia (including the reptiles also), Aves and Mammalia]
under the Vertebrata. The class Amphibia included the naked
Amphibia and the scaly animals (Reptilia).

Pisces and naked amphibia have many features (e.g. branchial

respiration, lack of amnion and allantois, persistent notochord,
etc.) in common. Similarly the reptiles and birds share many
common characters. Huxley adopted three principal groups of
Verte​brata.

They are:

I. Ichthyopsida (Pisces and Amphibia);

II. Sauropsida (Reptilia and Aves) and

III. Mammalia.

The Class Pisces has been divided into five subclasses by many
authors like Sedgwick (1905). The subclasses are:
Marsipobranchii, Elasmobranchii, Ganoidei, Dipnoi and Teleostei.

Chordata is divided into two major groups — protochordates and

vertebrates or craniates. Hemichordates, urochordates and
cephalo​chordates are collectively recognised as proto​chordates.
The members of this group do not possess vertebral column. The
protochordates often referred to as invertebrate chordates or
primitive chor​dates which are considered as transitional group in
between invertebrates and verte​brates.

At present some authors are of opinion to remove Hemichordata

from Protochordata or Chordata and treat Hemichordata as a
sepa​rate Phylum. Vertebrata is recognised for the possession of
vertebral column. The diffe​rences between protochordates and
vertebrates are shown in Table 5.

Vertebrata is divided into two super cla​sses Agnatha and

Gnathostomata. Agnatha means mouth without jaws and
Gnathostomata means jawed mouth vertebrates. Agnathans are
petromyzontids (lampreys) and myxinoids (hagfishes), and
gnathostomes are cartilaginous and bony fishes, amphibians,
reptiles, birds and mammals.

The terms Tetrapoda (tetrapods) or quadrupeds are often used

within gnathos​tomes. Tetrapoda means four-footed verte​brates
which include amphibians, reptiles, birds and mammals but
proper meaning for four-footed should be quadrupeds.
From embryological point of view, gnathostomes are divided into
Anamniota and Amniota. Amniota is a group of vertebrates
whose embryos possess a delicate, transparent mem​brane, the
amnion that remains around the developing embryo.

The sac-like amnion con​tains a fluid, the amniotic fluid, which

protects the developing embryos from extreme cold, desiccation
and also from external injury. Vertebrates are without any
amnion, called anamniota (fishes and amphibians), and amniotes
are reptiles, birds and mammals.

The term “Pisces” is often used for cartilaginous and bony

fishes. Pisces are aquatic gnathos​tomes having gills in adult
state, and median fins are supported by a special skeleton. These
terms are used by various taxonomists to classify the
vertebrates.

Inclusion of the subphyla Urochordata, Cephalochordata and

Vertebrata under the Phylum Chordata is of special interest to us
from the point of view of evolution. The associ​ation of the first
two subphyla with the verte​brates suggests the possibility of
intergradation between these groups and the sequences for
evolutionary transition from one to the other.

This biological association dates back to Lamarck (1809, 1815)

and Cuvier (1816) who sponsored the scheme for the first time
and recognised the uniqueness of the vertebrates possessing the
vertebral column and the inver​tebrates who lack this feature.

Costa (1938) and Yarrel (1835) recognised Branchiostoma

(Amphioxus) lanceolatum as a ‘low vertebrate’ and Kowalevski
(1869, 1871) demonstrated the existence of basic similarities
between verte​brates and tunicate larvae.
Balfour (1880) esta​blished the phylum Chordata which included
the vertebrates, Amphioxus and tunicates. Bateson (1885) added
the hemichordates to the Phylum Chordata. Fowler (1892)
suggested the inclusion of Rhabdopleura in the Subphylum
Hemichordata and Spengel (1932) added Planctosphaera to it.

The current opinion is in favour of removal of the Hemichordata

from the Phylum Chordata leaving the Urochordata,
Cephalochordata, Conodontophorida and Vertebrata as the
Subphyla of the Phylum Chordata. The Hemichordata is now
given the status of a sep​arate phylum.

However, it is still a common practice in many text-books to

designate the hemichordates, urochordates and the
cephalochordates as the protochordates/lower
chordates/invertebrate chordates. The naming of Hemichordata
as the Invertebrate chordates appears to be most convincing.

In the present text, the Hemichordata is described as a separate

phylum, recognised as a transitional group between the non-
chordates and protochordates (urochordates and
cephalochordates). The Phylum Chordata unites the diverse
forms having a notochord, hollow dorsal nerve cord and
pharyngeal gill-slits.

Patterns of Classification:

Nowadays, more or less two types of classification are used in

the classification of animals. The older one is evolutionary classi​-
fication which depends maximally on fossil record. This
classification reconstructs phylogenetic sequences using
judgement and makes evolutionary trees based on available data
including fossil record.
In this type of classi​fication, the different animals belonging to a
single group have a common ancestor. The phylogenetic
classification or cladistic pays little attention on fossil record.

It is one method, related to the branching sequences of the

phylogenesis, and the phylogeny is recon​structed on the basis of
shared derived charac​ters that analyse the primitive and
derivative characters. This system of classification performs a
good degree of objectivity in the selection of morphological
characteristic features that are employed in reconstructing
phylogenetic relationship.

A clade is the phylogenetic lineage evol​ving from a common

ancestor. A clade may be monophyletic, polyphyletic or
paraphyletic. A monophyletic of all the organisms of a lineage is
evolved from a common evolutio​nary ancestor or ancestral
group.

Polyphyletic is one which is characterized by features but is not

homologous. Groups of different lineages which have a common
ancestor and some members, but not all of its descendants, are
paraphyletic. The evolutionary history of a lineage is called
phylogeny and when it rep​resents in a graphic scheme is called
dendro​gram. The cladogram is a simply branching diagram that
depicts the relationships of diffe​rent clades.

Table 6 presents a conventional type of classification of the

Phylum Chordata in which Hemichordata is treated as
subphylum. Table 7 presents a “Cladogram of Chordata” where
species or groups of species have shown their relationships.
Table 8 presents the “characteris​tics of the living chordate
classes” and Table 9 presents the “Geological time scale”.