9.2 Chordates

Characteristics of Chordata

Animals in the phylum Chordata share five key characteristics that appear at some stage during their development: a notochord, a dorsal hollow (tubular) nerve cord, pharyngeal gill arches or slits, a post-anal tail, and an endostyle/thyroid gland (see figure below). In some groups, some of these key characteristics are present only during embryonic development.

The chordates are named for the notochord, which is a flexible, rod-shaped mesodermal structure that is found in the embryonic stage of all chordates and in the adult stage of some chordate species. It is strengthened with glycoproteins similar to cartilage and covered with a collagenous sheath. The notocord is located between the digestive tube and the nerve cord, and provides rigid skeletal support as well as a flexible location for attachment of axial muscles. In some chordates, the notochord acts as the primary axial support of the body throughout the animal’s lifetime. However, in vertebrates (craniates), the notochord is present only during embryonic development, at which time it induces the development of the neural tube and serves as a support for the developing embryonic body. The notochord, however, is not found in the postembryonic stages of vertebrates; at this point, it has been replaced by the vertebral column (that is, the spine).

Visual Connection: Chordate features.

In the chordates figure below, four common features appear at some point during development: a notochord, a dorsal hollow nerve cord, pharyngeal slits, and a post-anal tail. The endostyle is embedded in the floor of the pharynx.

 

The dorsal hollow nerve cord is derived from ectoderm that rolls into a hollow tube during development. In chordates, it is located dorsally to the notochord. In contrast, the nervous system in protostome animal phyla is characterized by solid nerve cords that are located ventrally and/or laterally to the gut. In vertebrates, the neural tube develops into the brain and spinal cord, which together comprise the central nervous system (CNS). The peripheral nervous system (PNS) refers to the peripheral nerves (including the cranial nerves) lying outside of the brain and spinal cord.

Pharyngeal slits are openings in the pharynx (the region just posterior to the mouth) that extend to the outside environment. In organisms that live in aquatic environments, pharyngeal slits allow for the exit of water that enters the mouth during feeding. Some invertebrate chordates use the pharyngeal slits to filter food out of the water that enters the mouth. The endostyle is a strip of ciliated mucus-producing tissue in the floor of the pharynx. Food particles trapped in the mucus are moved along the endostyle toward the gut. The endostyle also produces substances similar to thyroid hormones and is homologous to the thyroid gland in vertebrates. In vertebrate fishes, the pharyngeal slits are modified into gill supports, and in jawed fishes, into jaw supports. In tetrapods (land vertebrates), the slits are highly modified into components of the ear, tonsils, and thymus glands. In other vertebrates, pharyngeal arches, derived from all three germ layers, give rise to the oral jaw from the first pharyngeal arch, with the second arch becoming the hyoid and jaw support.

The post-anal tail is a posterior elongation of the body, extending beyond the anus. The tail contains skeletal elements and muscles, which provide a source of locomotion in aquatic species, such as fishes. In some terrestrial vertebrates, the tail also helps with balance, courting, and signaling when danger is near. In humans and other great apes, the post-anal tail is reduced to a vestigial coccyx (“tail bone”) that aids in balance during sitting.

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.

Urochordata

The 1,600 species of Urochordata are also known as tunicates. 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.

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 (see figure below) 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.

Subphylum Vertebrata (Craniata)

A cranium is a bony, cartilaginous, or fibrous structure surrounding the brain, jaw, and facial bones (see figure below). Most bilaterally symmetrical animals have a head; of these, those that have a cranium comprise the clade Craniata/Vertebrata, which includes the primitively jawless Myxini (hagfishes), Petromyzontida (lampreys), and all of the organisms called “vertebrates.” (We should note that the Myxini have a cranium but lack a backbone.)

The subphylum Craniata (or Vertebrata), including this placoderm fish (Dunkleosteus sp.), are characterized by the presence of a cranium, mandible, and other facial bones.

 

Members of the phylum Craniata/Vertebrata display the five characteristic features of the chordates; however, members of this group also share derived characteristics that distinguish them from invertebrate chordates. Vertebrates are named for the vertebral column, composed of vertebrae—a series of separate, irregularly shaped bones joined together to form a backbone (see figure below). Initially, the vertebrae form in segments around the embryonic notochord, but eventually replace it in adults. In most derived vertebrates, the notochord becomes the nucleus pulposus of the intervertebral discs that cushion and support adjacent vertebrae.

Vertebrata are characterized by the presence of a backbone, such as the one that runs through the middle of this fish.

The relationship of the vertebrates to the invertebrate chordates has been a matter of contention, but although these cladistic relationships are still being examined, it appears that the Craniata/Vertebrata are a monophyletic group that shares the five basic chordate characteristics with the other two subphyla, Urochordata and Cephalochordata. Traditional phylogenies place the cephalochordates as a sister clade to the chordates, a view that has been supported by most current molecular analyses. This hypothesis is further supported by the discovery of a fossil in China from the genus Haikouella. This organism seems to be an intermediate form between cephalochordates and vertebrates. The Haikouella fossils are about 530 million years old and appear similar to modern lancelets. These organisms had a brain and eyes, as do vertebrates, but lack the skull found in craniates.^[1] This evidence suggests that vertebrates arose during the Cambrian explosion.

Vertebrates are the largest group of chordates, with more than 62,000 living species, which are grouped based on anatomical and physiological traits. More than one classification and naming scheme is used for these animals. Here we will consider the traditional groups Agnatha, Chondrichthyes, Osteichthyes, Amphibia, Reptilia, Aves, and Mammalia, which constitute classes in the subphylum Vertebrata/Craniata. Virtually all modern cladists classify birds within Reptilia, which correctly reflects their evolutionary heritage. Thus, we now have the nonavian reptiles and the avian reptiles in our reptilian classification. We consider them separately only for convenience. Further, we will consider hagfishes and lampreys together as jawless fishes, the Agnatha, although emerging classification schemes separate them into chordate jawless fishes (the hagfishes) and vertebrate jawless fishes (the lampreys).

Animals that possess jaws are known as gnathostomes, which means “jawed mouth.” Gnathostomes include fishes and tetrapods. Tetrapod literally means “four-footed,” which refers to the phylogenetic history of various land vertebrates, even though in some of the tetrapods, the limbs may have been modified for purposes other than walking. Tetrapods include amphibians, reptiles, birds, and mammals, and technically could also refer to the extinct fishlike groups that gave rise to the tetrapods. Tetrapods can be further divided into two groups: amphibians and amniotes. Amniotes are animals whose eggs contain four extraembryonic membranes (yolk sac, amnion, chorion, and allantois) that provide nutrition and a water-retaining environment for their embryos. Amniotes are adapted for terrestrial living, and include mammals, reptiles, and birds.