Avian nectarivory and pollination in Aloe marlothii Berger : interactions between bird communities and a winter-flowering succulent

Aloe marlothii is a winter-flowering succulent that is widespread in the savanna biome of northern and north-eastern South Africa. Plants grow up to 8 m in height and are commonly found on rocky north-facing slopes. Nectar production occurs through a 24 h period with flowers producing copious amounts (c. 250 µl) of dilute nectar (c. 12%). This abundant nectar supply, that is available for a 5-10 week period during June-August, is utilised by numerous opportunistic avian nectarivores. At a study site in Suikerbosrand Nature Reserve, 60 km south-east of Johannesburg, at least 59% (38 species) of birds recorded during census transects fed on nectar; throughout the range of A. marlothii at least 85 species feed on nectar. This diversity surely far exceeds the number of species ever recorded feeding on nectar of a single plant. During the flowering period an influx of birds at the aloe forest occurred, with an overall increase in abundance and diversity. Pollinator exclusion experiments supported the hypothesis that A. marlothii is pollinated by generalist birds; specialist nectarivores are possibly excluded as inefficient pollinators by the nectar of low concentration and high volume. Fruit set was higher in plants that had avian visitors and very low when pollinators were absent. Stable carbon isotope analysis of whole blood was used to quantify the importance of nectar sugars for opportunistic nectarivores. During flowering there was an enrichment in the δ13C isotopic signature of whole blood of nectar-feeding birds towards that of nectar (δ13C = -12.6‰). This shift was most prominent in frugivores, insectivores and omnivores that typically fed on a diet depleted in 13C when nectar was not available. The C4 grass seed diet of granivores was similar to the isotopic signature ofA. marlothii nectar, so we were unable to determine to what degree granivores benefitted from nectar. Stable nitrogen isotopes in whole blood may suggest that many nectar-feeding birds shift their trophic position during flowering. However, we interpret these results with caution because of insufficient knowledge on diet-tissue fractionation factors of wild birds and/or temporal changes in vegetation isotopic values. Stable carbon isotope analysis of breath samples was used to show that A. marlothii nectar is a readily available income energy source for nectar-feeding birds. Because A. marlothii nectar is so dilute we expected it to be an important water source for many opportunistic nectar-feeding bird species. There was no correlation between the enrichment of δ13C of breath CO2 (representing metabolised nectar sugars) and the δ18O in breath CO2 (representing a highly evaporated water source in nectar); for most birds the δ18O in breath CO2 was more similar to that of free-standing water sources. However, because our knowledge on the relationship between δ18O of ingested water and body water, and fractionation processes when CO2 is exhaled is limited, we were unable to quantify water obtained from nectar. The sugars of A. marlothii nectar are probably more important, as a food source for opportunistic nectarivores during dry winter months when insect abundance is low, than the water in nectar, because birds are able to source water from other drinking sites.