Fog and dew deposition on Ruschia bolusiae in the Knersvlakte.

Fog and dew in the Succulent Karoo

An indispensible source of water for arid Succulent Karoo shrubs
by Ignatious Matimati, Charles Musil, Lincoln Raitt and Edmund February

f all the earths resources, water is the most fundamental to life, especially in the arid Succulent Karoo along South Africas West Coast where the low average annual rainfall is augmented by substantial amounts of fog generated by the cold Benguela Current of the Atlantic Ocean and nocturnal dew resulting from atmospheric water condensation. Fog and dew may provide a vital source of moisture for many of the rare succulent shrubs that are limited to the fog belt along our arid West Coast. Indeed, experimental fog and dew collection devices placed along South Africas West Coast have shown that fog and dew constitute a considerable portion of the hydrological input which concur with the substantial water yields generated in established fog water collection projects in Chile and other developing countries. Plants are ideal fog and dew interceptors. Water condenses on leaves and stems, and drips or runs off onto the soil surface where it is absorbed by plant root systems. It is thought that the leaves of some succulents such as Crassula species may absorb water vapour directly from a saturated atmosphere or from wet leaf surfaces. However, for both processes, the pathways of foliar absorption of water are poorly understood and as yet the relevance of fog and dew as sources of moisture for succulent plants in the arid South African ecosystems has not been substantiated. Fog and dew frequency and precipitation from them was higher than rainfall when we measured both at our study sites in the Knersvlakte and at Port Nolloth from July 2007 to June 2008, using weighing microlysimeters. The total recorded fog and dew net amounts of 252.9 mm year -1 exceeded those of 149.5 mm year VELD&FLOrA | SEPTEMBER 2010

recorded for rainfall by almost 70%. More fascinating results were that where Agyroderma pearsonii plants grew in quartz soil substrates, they channelled a very high net amount of 460mmyear1, which was more than twice the rainfall amount. This implied that these miniature succulents play an active role in trapping fog and dew water sources into the plant-soil system. With the expected increase in global temperatures, Agyroderma pearsonii is expected to die in large numbers since it has already reached the maximum range of temperatures that it can tolerate. We were also keen to know the contributions of fog and dew to succulent shrubs growing on shale-based soils. Since these succulent shrubs have deeper roots and bigger canopies to fit in weighing microlysimeters, the most suitable alternative technique was of stable hydrogen (2H/H) and oxygen (18O/16O) isotopes. Rain, fog and dew have different hydrogen and oxygen isotopic signatures or fingerprints which can be used to identify which of these sources of water are used by plants. Numerous studies have confirmed that fog, a condensate of saturated air close to the earths surface, has a higher proportion of heavy to light hydrogen (2H/H) and oxygen (18O/16O) isotopes than rain, a condensate from saturated air at higher levels in the atmosphere. Analysis of these proportions of isotopes in plant tissues may therefore provide an indirect measure of the fraction of total precipitation contributed by rain, fog and dew. Leaf tissues are unsuitable for such isotopic assays as there is a greater loss of lighter water isotopes than heavy isotopes during transpiration and photosynthesis, a feature not apparent in non-photosynthetic water-conducting stem xylem tissues which provide more stable sources of water isotopes.

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TOP: A fog-collecting device. TOP RIGhT: Fog and dew deposition on Agyroderma pearsonii in the Knersvlakte. RIGhT: Fog in Port Nolloth condenses on leaves and drips onto the soil below. The wet zone is demarcated with a line. ABOVE: Annual rainfall amounts at a study site in the Knersvlakte (149 mm per year) compared with fog and dew deposition amounts on A - bare quartz gravel soil (253 mm year) and B - on quartz gravel containing a dwarf succulent Agyroderma pearsoni (460 mm year). To identify what fraction of the total precipitation utilized by plants in the arid Succulent Karoo is from rainfall as opposed to fog and dew, we selected four evergreen and two partly drought-deciduous succulent shrubs. These are prominent in the Knersvlakte Bioregion, an acknowledged centre of diversity and endemism in the Succulent Karoo. Stem samples were collected from each of the eight shrubs at monthly intervals and sealed in airtight tubes. At the same time daytime samples of rainwater and night time samples of fog and dew water were collected and the water samples sealed individually in airtight tubes. In the laboratory the values of 18O/16O and 2H/Hisotopes were assayed in the samples of stem, rain, fog and dew water on a mass spectrometer. Over an entire year our results signified a predominance of water derived from fog and dew precipitation especially in the wet winter. This implies that fog and dew are even more important sources of water than rain in winter. Indeed, if plants depend on fog and dew more than they do on rainfall, then fog and dew must be vital for succulent plant survival. Restoration efforts along the succulent Karoo could capitalize on the importance of fog and dew as moisture sources in these arid environments. Such moisture sources can be harnessed for supplementing moisture to plants in disturbed ecosystems. (In Nepal, Chile and other developing countries, fog water has been harnessed using mesh grids for household use whilst passive condensers such as sea stones and roof surfaces have also been used for harvesting dew.) Moreover, fog harvesting along the West Coast of South Africa has already been considered feasible for household consumption purposes and further efforts should develop harvesting techniques for enhancing plant available moisture in these arid ecosystems. Whilst these results are the first report on the reliance of succulent shrubs on fog and dew for their survival in South Africa, their implications are phenomenal. Climatic warming has been predicted to increase by 2 to 6 oC by the year 2100, a phenomenon that can raise average night-time temperatures, thereby reducing the availability of these moisture sources. In such a scenario, will succulent shrubs that rely on fog and dew survive?

ACKNOWLEDGEMENTS This work is part of the BIOTA Southern Africa Project which was funded by the German Federal Ministry of Education and Research (Promotion number 01LC0624A2). READING hulme, M., R.M. Doherty, T. Ngara, M.G. New, & D. Lister. 2001. African climate change: 1900-2100. Climate Research 17, 145-168. Olivier, J. 2004. Fog harvesting: An alternative source of water supply on the West Coast of South Africa. GeoJournal 61, 203-214. Musil, C.F., P.D.R. Van heerden, C.D. Cilliers, & U. Schmiedel. 2009. Mild experimental climate warming induces metabolic impairment and massive mortalities in Southern African quartz field succulents. Environmental and Experimental Botany. 66, 79-87.
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