[1] Univ Cape Town, Dept Oceanog, Rondebosch, South Africa
[2] South African Environm Observat Network, Gqeberha, South Africa
[3] Nelson Mandela Univ, Inst Coastal & Marine Res, Gqeberha, South Africa
[4] Florida State Univ, Earth Ocean & Atmospher Sci Dept, Tallahassee, FL USA
[5] CSIR, Southern Ocean Carbon Climate Observ, Cape Town, South Africa
[6] Rhodes Univ, Dept Biochem & Microbiol, Makhanda, South Africa
[7] Univ Cape Town, Dept Biol Sci, Rondebosch, South Africa
[8] Univ Cape Town, Marine & Antarctic Res Ctr Innovat & Sustainabil M, Rondebosch, South Africa