Vertebrates (Vertebrata) S. Blair Hedges Vertebrates are treated here as a separate phylum Department of Biology, 208 Mueller Laboratory, Pennsylvania State rather than a subphylum of Chordata. 7 e morpho- University, University Park, PA 16802-5301, USA (
[email protected]) logical disparity among the chordates (urochordates, cepahalochordates, and vertebrates), and their deep time of separation based on molecular clocks (5) is as great Abstract as that among other groups of related animal phyla (e.g., The vertebrates (~58,000 sp.) comprise a phylum of mostly arthropods, tardigrades, and onycophorans). 7 e phyl- mobile, predatory animals. The evolution of jaws and ogeny of the lineages covered here is uncontroversial, for limbs were key traits that led to subsequent diversifi cation. the most part. Evidence from nuclear genes and morph- Atmospheric oxygen change appears to have played a major ology (1, 2, 6, 7) agree in the backbone phylogeny of ver- role, with an initial rise in the late Precambrian (~580–542 tebrates represented by these nested groups: Tetrapoda million years ago, Ma) permitting larger body size, followed (Lissamphibia, Amniota), Sarcopterygii (Actinistia, by two Paleozoic pulses affecting prey. The First Pulse Dipnoi, Tetrapoda), Osteichthyes (Actinopterygii, (~430–390 Ma) brought fi shes to brackish and freshwater Sarcopterygii), and Gnathostomata (Chondrichthyes, environments where they diversifi ed, with one lineage giv- Osteichthyes). ing rise to tetrapods. The Second Pulse (~340–250 Ma) led to Cyclostomata wa s or ig i na l ly considered a ba sa l, mono- a Permo-Carboniferous explosion of tetrapods, adapting to phyletic group based on morphology (8), but later mor- diverse terrestrial niches.