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A PATERNAL ROLE FOR BULLS OF THE GALAPAGOS ISLANDS SEA LION

Bartholomew (970) proposed a model for the evolution of polygyny in pinnipeds. Males were said to have a minimal parental role. Observations on the Galapagos sea lion, however, suggest that this may not hold for all polygynous pinnipeds. According to Peterson and Bartholomew (1967) the bull of the California sea lion, Zalopkus californianus californianus, establishes a territory when breeding and excludes other males. The chief function of the territorial bull is inseminating post-partum females. A minor consequence is that his activities often interrupt hostile interactions among his females. The bull is tolerant of pups but provides no protection, even treading on them while reacting to other males. This account applies to anyone of a number of species. Once the male has transmitted his genes, he becomes inconsequential to their survival. If the male protected pups, however, the situation would be more complicated. The pups in his territory have issued from inseminations of the previous year. Further, pups and their mothers move between territories in the rookery. Thus he could be promoting survival of another bull's offspring. Eibl-Eibesfeldt (1955) suggested that the bull of the Galapagos sea lion, Z. c. wollebaeki, protects the young. Once a bull threatened but he did not attack the investigators until they approached a female with a very young pup. However, this is also a bull's response to a rival approaching an estrus female. A bull patrolling in the water just off the beach was said to prevent the young from moving into deep water. Some young attempted to approach a diver. The bull interposed himself between diver and young, and drove the young out of the water. This behavior was interpreted as protecting the pups from sharks (I saw pups that bore severe shark wounds). These observations, however, were in response to intrusion by humans. Also, details are lacking. The behavior in a rookery on Champion Island was observed from 15:00 to 16:30 hr. on 13 February, 1967. I sat perhaps 100 m. above the sea, looking down directly into a cove (Fig. 1). Later, I approached to within 25 m. There were about 100 sea lions on the sand beach, including young pups, some of them newborn. About an equal number of sea lions were in the water over the dark reef. Five or six large bulls were evenly spaced in the water over the 100 m. long dark reef, thus about one per 20 m. of shoreline (Eibl-Eibesfeldt, 1955, reported one per 50-100 m. at a different island). Territorial boundaries could not be recognized, probably because the bulls stayed in the water; also, the sand beach lacked distinctive features. There seemed a continuous assemblage of sea lions in which all moved about except the bulls. The rookery therefore resembled more that described by Peterson and Bartholomew (1967) than that by Eibl-Eibesfeldt (1955). Every 5-10 min. one of two 3-m. long sharks (Carcharinus galapagensis) appeared from the blue water and swam slowly across the green reef, then parallel to the edge of the dark reef, doubling back to sea (Fig. 1). Occasionally, it passed across part of the dark reef in water no more than 1 m. deep (Fig. 1, path a). Adult male and female sea lions returning from the sea seemed to detect sharks at 20-30 m. They slowed, then swam underwater, gliding past the shark. Then they continued on to the rookery. The sharks made no detectable response. When a shark approached to 5-10 m. of the edge of the dark reef, one or two, sometimes three bulls swam toward it. Each sea lion's approach was rapid and initially on the surface. When 5-7 m. away the bull slowed and coasted underwater on an intercept course. At the leading bull's approach, the shark started to turn away. The bull then adjusted course to parallel that of the veering shark. The bull thus deflected the shark away from the rookery. The closest bull, and often one or two others, followed the shark 10-30 m. Once one bull bit at a shark's tail but apparently missed. The sea lions behind the bulls responded to the sharks only when they entered the dark reef. Then the smaller animals dashed shoreward, pausing in water about 30 cm. deep with land 1-5 m. distant. Much later, a person suddenly appeared on the beach and fired a shot pistol at a lizard. Panic followed, and all but the extremely young pups dashed into the water; the rookery surged out over the green reef, but quickly returned. A large shark soon appeared swimming at the edge of the dark reef. Then it turned and moved in over the dark reef to within 5 m. of shore (b in Fig.