Some advanced features of vertebrates over protochordates

 The notochord is either replaced entirely by the vertebral column or is present along with
the vertebral column.
 The heart is divided into chambers and has RBC in the blood.
 Each verterbra has a distinct sex. The only exception is in the case of some fish.
 When fertilization takes place, the sperm and the ovum unite in the animal pole pathway.
 Endostyle is totally absent in the adult stages of vertebrates, (only found in the ammocoetes
larval stage of Petromyzon).
 Presence of neural crest cells in the development of nervous system.

Primitive features of protochordates

 No cranium or vertebral column.

Origin of Chordates

Chordates evolved sometime during Cambrian period, 500 million years ago during Cambrian

explosion, almost at the same time when invertebrates were beginning to evolve. They may have

evolved from some freshwater forms as Chamberlain (1900) pointed out that all modern chordates

possess glomerular kidneys that are designed to remove excess water from body. However, early

fossils of chordates have all been recovered from marine sediments and even modern

protochordates are all marine forms. Also glomerular kidneys are also found in some marine forms

such as myxinoids and sharks. This hints to the Marine origin of chordates.
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. The following theories have been given to explain the origin of chordates:

Theories

Dipleurula concept

The term 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

dipleurula 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.

Dipleurula represents an ancestral form for the the primitive deuterostomes. We can see that 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. The Dipleurula concept was propounded by Bather in 1900. The

common ancestors didn’t possess all the common characters of the free swimming bilateral larvae
of different groups of echinoderms and it might add one or two characters which none of them

possess. Hence it appears that dipleurula concept cannot illustrate the common ancestor of the
echinoderms or can explain the characteristic features of the ancestor of echinoderms. Dipleurula

is merely a name for features common to present echinoderm larvae.

Echinoderm Theory of Origin of Chordates:

It is believed that chordates have originated from invertebrates. It is difficult to determine from

which invertebrates group the Chordates developed. Chordate ancestors were soft bodied animals.

Hence they were not preserved as fossils. There are several theories have been put forwarded to
explain the origin of chordates. These 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.

Echinoderm theory 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.

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.

symmetrical chordates. The bilateria is divided into two major divisions- Prostomia and

Deuterostomia. The division is based on the differences in embryonic and larval development.

Prostomia includes from Annelida to Arthropoda while Deuterostomia includes

Echinodermata,Pogonophora and Chordates.

Deuterostome line of Chordate Evolution:

Following common features of Deuterostome 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 larva of Echinoderms and Hemichordates have a close resemblance vertebrate does not

have a floating larva.

(v) Deuterostomes use creatinine as phosphogen whereas invertebrates use arginine. Some

Hemichordates as well as echinoids use both.