IntroductionThe reasons for variations in the clutch size of birds are poorly understood (Ricklefs 1970) and this is more surely the case in the Charadrii where no species normally has a clutch of more than foul' (MacLean 1972). One of the trends which is often discussed is the variation with respect to latitude where, generally, tropical species have smaller clutches than temperate species (Lack 1968). Part of the problem in these comparisons is that clutch sizes are compared in isolation whereas other aspects of the biology should also be considered. For example, Graul (1974) showed that the Mountain Plover Cha'~udrius montanv~ lays two successive clutches of two eggs, one of which is incubated by the male and the other by the female. In this case clutch size is not related to egg production. However, information on the biology of tropical species is often lacking. This paper describes the breeding biology of the White-fronted Plover Charadrius marginatus, a species which breeds mainlv on coastal beaches of Africa and normally has a clutch of two (MeLaehlan and Liversidge 1970). Those aspects which may relate to clutch size, such as the mating system, length of breeding season, frequency of clutch replacement and breeding success are examined as well as other aspects of the biology. Comparisons are then made with the Ringed Plover Charadrius hiaticula, a closely related species Which breeds on beaches or stony areas in temperate and arctic regions and normally has a clutch of four.
Study area and methodsThe study was carried out at Langebaan Lagoon (33°S 18°E), a large inlet of the sea on the west coast of South Africa ( fig. 1). The lagoon has extensive sand flats rich in invertebrates and is surrounded by either salt marshes or sandy beaches and low dunes. White-fronted Plovers nest on the dunes, and the study area extended along 900 m of beach and its adjacent sand flats, dunes, salt marsh, and salt pans (figs. 2 and 3 (a)). The salt marshes and salt pans were inundated only by spring tides. The Present addresses: t Culterty Field Station, Aberdeen University, Scotland. ++ t~yeeroft, Frampton Cotterell, Bristol, England.