Granivorous Birds in the Sahel: Is Seed Supply Limiting Bird Numbers?

Abstract

During the dry season four billion African and European granivorous birds in the Sahel consume, by grand average, 15 g seeds/ha/day, equivalent to an average annual consumption of 4.5 kg/ha. This represents only 4–15% of the estimated average total soil seed bank of some 30–100 kg/ha in the early dry season. Despite this apparent abundance of food, there are many reasons to presume that the number of seed-eating birds is limited by their food supply. First, the birds have to share the seed supply with rodents and insects that eat more seeds than all the birds combined. Second, granivorous birds are constrained by foraging time available to them. They avoid foraging during the midday heat and feeding time is mostly restricted to the early morning and late afternoon, totalling about 4 h per day. This forces them to achieve high intake rates and thus to select feeding sites where the available seeds can be handled quickly and/or are so abundant that the encounter rate is high. Third, only a proportion of the seeds lies on the surface where they are easy to find. Most grass seeds are tiny and even small birds need to eat thousands per day. Because they have so little time to look for food, they cannot afford to search for seeds hidden in the sand. Doves rapidly swallow seeds whole, but all smaller seedeaters have to separate the husk from the seed, a process that takes time too. Fourth, seed-eating birds in the Sahel discriminate between seeds. They ignore ‘empty seeds’ (husks) and also avoid feeding on common graminoids whose seeds have long awns (Aristida) or spines (Cenchrus) and which are time-consuming to process. Occasionally, granivorous birds may select seeds from forbs, but these, being low in digestibility, are not the preferred choice. Granivorous birds prefer the seeds of Panicum grass and other grass species with highly soluble carbohydrate fractions. Birds switch to marginal seed types at the end of the dry season, when the seed bank of the preferred species is depleted. Fifth, soil seed bank of preferred grass species is much reduced in dry years. Panicum and other preferred annual grasses are found mostly on riverine floodplains and in depressions that are prone to ephemeral flooding during the rainy season. Such sites attract many seed-eating birds, but the total surface area of floodplains is relatively small compared to the extensive drylands, on top of being very much smaller in dry years, circumstances that account for high mortality among seed-eating birds in drought years. The final argument for food-limitation is that the mounting grazing pressure of livestock over the last decades has severely reduced the annual soil seed bank and changed the plant community (preferred grass species replaced by non-preferred grasses and forbs). The combination of these factors caused a very large decline of seed-eating bird populations in the Sahel between the 1970s and 2010, including a handful of Eurasian species. The Sahel is still home to some four billion granivorous birds during the dry season, but just half a century ago the numbers must have been much higher.