THE BREEDING ECOLOGY OF ROYAL TERNS STERNA (THALASSEUS) MAXIMA MAXIMA

Summary The breeding ecology of the New World race of the Royal Tern Sterna maxima maxima was studied at colonies in Virginia and North Carolina, U.S.A., from 1967 to 1970. Colony sites are quite varied, but isolation, good distance visibility and (especially) freedom from quadruped predators seem important if not essential prerequisites. In Virginia, most adults arrive at the colony site in the last few days of March. Courtship, displaying and copulation take place at, near, and some distance from the colony site. In this the Royal Tern differs from its near relative S. sandvicensis sandvicensis which carries out these activities away from the colony site, presumably as an anti-predator device. Courtship displays are not described, but in their essentials are similar to those of other terns. Copulations continue throughout incubation and gradually disappear when the eggs hatch. No post-copulatory displays are known. Some synchrony of egg-laying is evident, but no “mass laying” occurs, the colony increasing in size steadily over a period of weeks and months. Defaecation on the nest is normal and probably serves to strengthen the nest rim against flooding by high tides. Contrary to published reports, the normal clutch is one; the largest is two, probably often from two different females. All incubating adults examined had two brood patches. Average egg measurements are 63 × 44-5 mm, and average egg weight is 64.3 g. Egg colour varies greatly, and evaluation is difficult. Eggs are probably not cryptically coloured and individual variation, as well as nest-site, are used by returning adults to identify their own eggs. Average maximum nest density is c. 7/m2. Sterna sandvicensis acuflavida nests regularly, if not obligatorily, with m. maxima; interactions between the two, and possible benefits accruing to each, are discussed. Unhatched eggs were significantly nearer other nests than were successfully hatched eggs, and possible explanations are given. Incubation lasts about 30–31 days, a week longer than in most terns; both sexes incubate. Broken eggshells are never removed by the adults. Instead, after 2–3 days, the chicks leave the nest permanently to join a creche that roams freely about the immediate vicinity of the colony. Chicks remain in the creche for about 25–30 days, leaving it at fledging, approximately 30 days after hatching. While in the creche, chicks are normally fed only by their parents, who probably recognise them both vocally and visually, using the extensive variation in voice and colour of chicks characteristic of the species. Sandvicensis acuflavida chicks also are highly variable, and join Royal creches, adults of both species attending. Variation also occurs in Royals' juvenal plumage, and seems associated with extended parental care. Feeding adults normally wander up to 40 km from their colony, and this probably facilitates the intercolonial exchange of breeding birds we recorded. They feed inshore, in shallow waters, taking small fish by dives which do not go below the surface. They regularly take small soft-shell crabs in this way, and frequently water-skim like skimmers Rynchops spp., sometimes capturing food while doing so. A relationship between crab capture and water-skimming is established for the first time, and water-skimming in non-feeding contexts is mentioned. Besides the quadruped predators which they normally avoid by fleeing the colony site, Royals have no known predators beyond the egg stage. Eggs, although not chicks, are readily taken by Laughing Gulls. Relationships between the two species are discussed, emphasising their constant association the year round.