Lecture 2 Origins of Vertebrates

Chapter 2, KK or MH

Chordate Characteristics
Vertebrates are phylum Chordata, subphylum Vertebrata. Phylum Chordata can be defined as those animals with a notochord. Subphylum Vertebrata are those chordates with a brain (cephalization) surrounded by a cranium.
Fig. 2.4 KK, Other chordate characteristics: - hollow dorsal nerve cord - gill slits in pharynx - complete gut - post-anal tail

Vertebrate Characteristics
bilaterally symmetry, cephalization cranium surrounding brain hollow dorsal nerve cord above notochord, with anterior brain adjacent to 3 special sense organs anterior mouth, complete gut, post-anal tailgill slits in the anterior part of gut or pharynx coelom (around gut, heart, and lungs if present) segmented (metamerism) large size

Vertebrate Origins
The origin of vertebrates, and their relationship to other animal taxa, is poorly known.

This is because vertebrate ancestors were small, soft organisms that do not fossilize well and lived long ago (>500 mybp). Good fossils are rare. On the other hand, we know vertebrate evolution comparatively well.
Calendar analogy. If the earth is 4.6 billion years old, and we express the timing of events on a calendar year:
the first vertebrates arose in late November (500 million years ago) land animals in early December mammals in mid-December our genus (Homo) about 8 pm on New Years Eve.

Homo sapiens appears during the last hour before midnight

The Geologic Time Scale

KK Fig. 1.31

Life on Earth 4.6 billion years

Vertebrates appear in the fossil record about 500 MYBP early Paleozoic Era (Cambrian Period)
Era Period Neogene Paleogene Cretaceous Jurassic Triassic Permian Carboniferous Devonian Silurian Ordovician Cambrian MYBP 23 66 146 200 251 291 359 416 444 488 542 first appearance Homo mammals mammals,birds reptiles reptiles amphibians fish with jaws agnathans chordates amphibians fish with jaws prominent Group

Cenozoic Mesozoic

Paleozoic

Pre-Cambrian

The 3 subphyla of the phylum Chordata were all present in the Cambrian period
Urochordata

Cephalochordata
Vertebrata

Subphylum Cephalochordata
Amphioxus

Nature 453: 957 (June 19, 2008) The Amphioxus genome.

KK Figure 2.16, MH Fig. 2.2

A fossil chordate from the Cambrian period, western Canada.

Vertebrate Characteristics shared by Amphioxus

1.notochord under hollow dorsal nerve chord 2.complete gut and post-anal tail 3.midgut cecum (homologue of liver?) 4.perforated pharynx 5.segmentation 6.coelomate

Features of Amphioxus that are not vertebrate-like

no cranium (or other skeleton) no special sense organs and associated brain no kidneys no heart

Subphylum - Urochordata
(tunicates or sea squirts)
KK Fig. 2.24

Why are tunicates classified as Chordata?

Because their planktonic larval stage has a notochord. (KK Fig. 2.23, H&G Fig. 2.1)

More distant relatives: Deuterostomes

Animals can be categorized as deuterostomes or protostomes based on their early embryonic development. Chordates are among the deuterostomes.

KK. Fig. 2.3

Three characteristics of Deuterostomes:

- radial (as opposed to spiral) cleavage - indeterminate cleavage (as opposed to determinate) - the blastopore is at the posterior, and becomes the anus rather than the mouth

Phylum Hemichordata (acorn worms)

Hemichordates are detuterostomes and used to be considered Chordata, but the stomochord is no longer considered to be homologous with the notochord of chordates. They do have a perforated pharynx.

KK Fig. 2.8

Phylum Echinodermata
The largest phylum of deuterostomes other than chordates.

Sea Urchin Crinoid (sea lily)

Starfish

KK Fig. 2.1

Chordata is a small phylum compared to Arthropoda, and the other deuterostome phyla are even smaller

Possible Origins of Vertebrates

KK 2.32

One school of thought has held that vertebrate ancestors were motile filterfeeders from which several more sedentary filter feeding groups evolved, like echinoderms, hemichordates, cephalochordates and tunicates evolved.

KK 2.31

Another view emphasizes the evolution of larval forms, and suggests that a cephalochordate-like ancestor of vertebrates arose from tunicate-like ancestor by paedomorphosis.

The Origin of Vertebrates (According to Kardong)

KK 2.33

New molecular studies point to Urochordates as being the closest living relatives of vertebrates, not cephalochordates.