Institute of Biological Environmental and Rural Sciences, Aberystwyth University, Aberystwyth SY23 3DA, UK. jhd@aber.ac.uk|Institute of Biological Environmental and Rural Sciences, Aberystwyth University, Aberystwyth SY23 3DA, UK. jhd@aber.ac.uk|Institute of Biological Environmental and Rural Sciences, Aberystwyth University, Aberystwyth SY23 3DA, UK. jhd@aber.ac.uk|Institute of Biological Environmental and Rural Sciences, Aberystwyth University, Aberystwyth SY23 3DA, UK. jhd@aber.ac.uk|Institute of Biological Environmental and Rural Sciences, Aberystwyth University, Aberystwyth SY23 3DA, UK. jhd@aber.ac.uk|Institute of Biological Environmental and Rural Sciences, Aberystwyth University, Aberystwyth SY23 3DA, UK. jhd@aber.ac.uk|Institute of Biological Environmental and Rural Sciences, Aberystwyth University, Aberystwyth SY23 3DA, UK. jhd@aber.ac.uk