1 Institute of Biology, Environmental and Rural Sciences (IBERS), Aberystwyth University, Wales, UK.
2 Interdisciplinary Centre for Environmental Microbiology (iCEM), Aberystwyth University, Wales, UK.
3 Department of Geography and Earth Sciences (DGES), Aberystwyth University, Wales, UK.
4 Present address: Group for Aquatic Microbial Ecology (GAME), University of Duisburg-Essen, Campus Essen - Environmental Microbiology and Biotechnology, Universitätsstr. 5, 45141 Essen, Germany.