[1] Univ Otago, Dept Geol, POB 56, Dunedin 9054, New Zealand
[2] Columbia Univ, Lamont Doherty Earth Observ, POB 1000, Palisades, NY 10964 USA
[3] Univ Adelaide, Australian Ctr Evolutionary Biol & Biodivers, Sch Biol Sci, Benham Bldg DX 650 312, Adelaide, SA 5005, Australia
[4] Univ Adelaide, Sprigg Geobiol Ctr, Sch Biol Sci, Benham Bldg DX 650 312, Adelaide, SA 5005, Australia