Chordate Origin

ORIGIN OF CHORDATES

(Dr. Girish Chandra)

 SUMMARY

            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. That makes the marine origin of chordates more plausible.

            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:

1. Echinoderm Origin. The theory was given by Johannes Muller (1860) and 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. Johannes Muller, W. Garstang and DeBeers proposed that echinoderm larvae gave rise to chordates by neoteny. Also like chordates, echinoderms are also deuterostomes and possess mesodermal skeletal elements.

            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 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 have evolved in diverse groups of animals during Cambrian-Orodovician periods when planktons were abundant in water.

2. Hemichordate Origin.  Romer (1959) suggested that ancestral deuterostomes were sedentary tentacle feeders whose mucous-laden ciliated tentacles served to trap planktons as they were waved in water as do the modern lophophorates and pterobranch hemichordates, Cephalodiscus and Rhabdopleura. By some mutation pharyngeal gill slits evolved in these ancestors, which made the pharynx sieve-like to trap planktons as the water current passed through it.  Extant pterobranchs possess both ciliated arms and pharyngeal gill slits. Tornaria larva of hemichordates shows phylogenetic relationship with echinoderm larvae and hemichordates also show affinities with chordates.

3. Urochordate Origin. W. Garstang (1928) and N.J. Berrill (1955) gave importance to the tadpole-like larva of urochordates which carries typical chordate characters, namely, a notochord in tail along with segmented myotomes, dorsal hollow nerve cord, sense organs and pharyngeal gill slits. Garstang (1928) suggested that chordates evolved from some sessile filter feeding urochordate by the larval stage evolving into adult by neoteny and by losing the sedentary adult stage. 

4. Cephalochordate Origin. Chamberlain (1900) studied the primitive and advanced characters of cephalochordates and proposed that while extant cephalochordates possess all chordate characters in typical state, they also show some primitive features of non-chordates, such as, absence of heart, head, sense organs, respiratory pigment, filter-feeding mode of food capture and excretion by solenocytes.  Fossils of 60 specimens from mid-Cambrian of the earliest chordate, Pikaia gracilens have been discovered from Burgess Shale in British Columbia, Canada. The Amphioxus-like fossils show streamlined, ribbon-shaped, 5 cm long body having notochord in the posterior two-third of body and myomeres. It has a small head with two tentacles and gill slits in the neck region. Other chordate-like fossils are: Cathaymyrus from early Cambrian sediments in China and Palaeobranchiostomata from early Permian from South Africa that appears to be more similar to Amphioxus.

5. Combined theory.  E.J.W. Barrington (1965) combined all the above theories and proposed that the common ancestor of echinoderms and chordates was a sessile ciliary arm feeder that lived in the plankton-rich environment of the Cambrian. Modern Crinoidea (Echinodermata), Pogonophora and Pterobranch hemichordates evolved from a similar ancestor by retaining the original mode of feeding, perhaps because they continued to inhabit the same environment as occurred in ancestral days. However, pharyngotremy (perforation of pharynx with gill slits) must have evolved in a large number of groups at that time, which must have been much more superior method of food gathering by filtering water through pharynx as compared to ciliated arm feeding.  Hence, the sedentary Protoascidians of that time lost ciliated arm feeding and adopted pharyngeal filter feeding as the only method of food gathering. Sometime later, when the plankton population in water declined, free-swimming tailed larva of these urochordates did not metamorphose and became a neotenic adult, since free-swimming mode was superior in food searching at a time of food scarcity. Cephalochordate-like ancestors evolved by perfection and expansion of chordate characters that were already present in the ascidian tadpole larva. We already have fossils of such primitive chordates, e.g. Pikaia gracilens from mid-Cambrian.