Chordates and the Evolution of Vertebrates

Two clades of chordates are invertebrates: Cephalochordata and Urochordata. Members of these groups also possess the five distinctive features of chordates at some point during their development.


Members of Cephalochordata possess a notochord, dorsal hollow tubular nerve cord, pharyngeal slits, endostyle/thyroid gland, and a post-anal tail in the adult stage (Figure). The notochord extends into the head, which gives the subphylum its name. Although the neural tube also extends into the head region, there is no well-defined brain, and the nervous system is centered around a hollow nerve cord lying above the notochord. Extinct members of this subphylum include Pikaia, which is the oldest known cephalochordate. Excellently preserved Pikaia fossils were recovered from the Burgess shales of Canada and date to the middle of the Cambrian age, making them more than 500 million years old. Its anatomy of Pikaia closely resembles that of the extant lancelet in the genus Branchiostoma.

The lancelets are named for their bladelike shape. Lancelets are only a few centimeters long and are usually found buried in sand at the bottom of warm temperate and tropical seas. Cephalochordates are suspension feeders. A water current is created by cilia in the mouth, and is filtered through oral tentacles. Water from the mouth then enters the pharyngeal slits, which filter out food particles. The filtered water collects in a gill chamber called the atrium and exits through the atriopore. Trapped food particles are caught in a stream of mucus produced by the endostyle in a ventral ciliated fold (or groove) of the pharynx and carried to the gut. Most gas exchange occurs across the body surface. Sexes are separate and gametes are released into the water through the atriopore for external fertilization.

The illustration shows a lancelet with a head protruding form the sand, and the rest of the body buried. On the head, tentacles surround the mouth. The mouth leads to a digestive tract. The anus is just before the post anal tail. The pharyngeal slits are next to the atrium, which empties into the atriopore. The body has segmented muscles running along it from top to bottom.
Cephalochordate anatomy. In the lancelet and other cephalochordates, the notochord extends into the head region. Adult lancelets retain all five key characteristics of chordates: a notochord, a dorsal hollow nerve cord, pharyngeal slits, an endostyle, and a post-anal tail.


The 1,600 species of Urochordata are also known as tunicates (Figure). The name tunicate derives from the cellulose-like carbohydrate material, called the tunic, which covers the outer body of tunicates. Although tunicates are classified as chordates, the adults do not have a notochord, a dorsal hollow nerve cord, or a post-anal tail, although they do have pharyngeal slits and an endostyle. The “tadpole” larval form, however, possesses all five structures. Most tunicates are hermaphrodites; their larvae hatch from eggs inside the adult tunicate’s body. After hatching, a tunicate larva (possessing all five chordate features) swims for a few days until it finds a suitable surface on which it can attach, usually in a dark or shaded location. It then attaches via the head to the surface and undergoes metamorphosis into the adult form, at which point the notochord, nerve cord, and tail disappear, leaving the pharyngeal gill slits and the endostyle as the two remaining features of its chordate morphology.

Photo A shows tunicates, which are sponge-like in appearance and have holes along the surface. Illustration B shows the tunicate larval stage, which resembles a tadpole, with a post anal tail at the narrow end. A dorsal hollow nerve cord run along the upper back, and a notochord runs beneath the nerve cord. The digestive tract starts with a mouth at the front of the animal connected to a stomach. Above the stomach is the anus. The pharyngeal slits, which are located in between the stomach and mouth, are connected to an atrial opening at the top of the body. Illustration C shows an adult tunicate, which resembles a tree stump anchored to the bottom. Water enters through a mouth at the top of the body and passes through the pharyngeal slits, where it is filtered. Water then exits through another opening at the side of the body. A heart, stomach and gonad are tucked beneath the pharyngeal slit.
Urochordate anatomy. (a) This photograph shows a colony of the tunicate Botrylloides violaceus. (b) The larval stage of the tunicate possesses all of the features characteristic of chordates: a notochord, a dorsal hollow nerve cord, pharyngeal slits, an endostyle, and a post-anal tail. (c) In the adult stage, the notochord, nerve cord, and tail disappear, leaving just the pharyngeal slits and endostyle. (credit: modification of work by Dann Blackwood, USGS)

Adult tunicates may be either solitary or colonial forms, and some species may reproduce by budding. Most tunicates live a sessile existence on the ocean floor and are suspension feeders. However, chains of thaliacean tunicates called salps (Figure) can swim actively while feeding, propelling themselves as they move water through the pharyngeal slits. The primary foods of tunicates are plankton and detritus. Seawater enters the tunicate’s body through its incurrent siphon. Suspended material is filtered out of this water by a mucous net produced by the endostyle and is passed into the intestine via the action of cilia. The anus empties into the excurrent siphon, which expels wastes and water. Tunicates are found in shallow ocean waters around the world.

The image displays a group of salps near a coral reef.
Salps. These colonial tunicates feed on phytoplankton. Salps are sequential hermaphrodites, with younger female colonies fertilized by older male colonies. (credit: Oregon Department of Fish & Wildlife via Wikimedia Commons)